Monographs on Australian Lepidoptera
Monographs on Australian Lepidoptera Volume 10
10 Olethreutine Moths of Australia
Olethreutine Moths of Australia (Lepidoptera : Tortricidae)
Marianne Horak With contributions by Furumi Komai PUBLISHING
ISBN 0 643 09093 2
PUBLISHING
Olethreutine Moths of Australia (Lepidoptera: Tortricidae)
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Monographs on Australian Lepidoptera Editorial Board Editor-in-Chief M. Horak, Canberra, Australia Editorial Panel I. F. B. Common, Toowoomba, Australia E. D. Edwards, Canberra, Australia R. L. Kitching, Brisbane, Australia S. E. Miller, Washington DC, USA G. S. Robinson, London, UK M. J. Scoble, London, UK M. S. Upton, Canberra, Australia
Spilonota constrictana (Meyrick, 1881), male Drawing by R. Nielsen, Copenhagen
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Monographs on Australian Lepidoptera Volume 10
Olethreutine Moths of Australia (Lepidoptera: Tortricidae)
Marianne Horak With contributions by Furumi Komai
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© CSIRO 2006 All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO PUBLISHING for all permission requests. National Library of Australia Cataloguing-in-Publication entry Horak, M. Olethreutine moths of Australia. Bibliography. Includes index. ISBN 0 643 09093 2. 1. Lepidoptera – Australia – Classification. I. Komai, Furumi. II. Title. Series: Monographs on Australian Lepidoptera; 10. 595.78 Available from CSIRO Publishing 150 Oxford Street (PO Box 1139) Collingwood Vic. 3066 Australia Telephone: Local call: Fax: Email: Web site:
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Monographs on Australian Lepidoptera Previous volumes in this series: Volume 1, 1989 Primitive Ghost Moths Morphology and Taxonomy of the Australian Genus Fraus Walker (Lepidoptera: Hepialidae s. lat.) E.S. Nielsen and N.P. Kristensen Volume 2, 1993 Tineid Genera of Australia (Lepidoptera) G.S. Robinson and E.S. Nielsen Volume 3, 1994 Oecophorine Genera of Australia. I. The Wingia Group (Lepidoptera: Oecophoridae) I.F.B. Common Volume 4, 1996 Checklist of the Lepidoptera of Australia E.S. Nielsen, E.D. Edwards and T.V. Rangsi (Editors) Volume 5, 1997 Oecophorine Genera of Australia II The Chezala, Philobota and Eulechria Groups (Lepidoptera: Oecophoridae) I.F.B. Common
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Volume 6, 1999 Biology of Australian Butterflies R.L. Kitching, E. Scheermeyer, R.E. Jones and N.E. Pierce (Editors) Volume 7, 1999 Heliothine Moths of Australia A Guide to Pest Bollworms and Related Noctuid Groups M. Matthew Volume 8, 2000 Oecophorine Genera of Australia III The Barea Group and Unplaced Genera (Lepidoptera: Oecophoridae I.F.B. Common Volume 9, 2004 Zygaenid Moths of Australia A revision of the Australian Zygaenidae (Procridinae: Artonini) Gerhard M. Tarmann
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Dedication This volume is dedicated to Ian Common and to the memory of Alex Diakonoff who both so generously shared their knowledge on Australian and Oriental Tortricidae and provided unstinting encouragement and inspiration early on when I was learning my trade as a young scientist.
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Contents
Abstract
1
Introduction
3
Material and methods
7
Material studied
7
Taxonomic treatment
7
Preparatory techniques
8
Abbreviations
8
Acknowledgments 1. Phylogeny of the Olethreutinae
11
2. Morphology
21
3. Biology
49
4. Diversity and distribution
53
5. Australian olethreutine genera
57
Key to olethreutine genera based on Australian species
57
Microcorsini
67
Cryptaspasma Walsingham
68
Collogenes Meyrick
73
Olethreutini Oxysemaphora-group
79 81
Oxysemaphora Diakonoff
82
Cnecidophora, gen. nov.
89
Eudemis-group Sorolopha Lower Sycacantha-group
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9
93 93 99
Sycacantha Diakonoff
100
Atriscripta, gen. nov.
104
Demeijerella Diakonoff
108
Lobesia Guenée
111
Gnathmocerodes-group
120
Gnathmocerodes Diakonoff
121
Rhectogonia Diakonoff
125
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viii
CONTENTS
Olethreutes-group
128
Ophiorrhabda Diakonoff
128
Dudua Walker
134
Trachyschistis Meyrick
139
Archilobesia Diakonoff
142
Euobraztsovia Diakonoff
146
Eremas Turner
148
Zomaria-group
151
Zomariana Diakonoff
152
Podognatha Diakonoff
156
Neopotamia-group
161
Temnolopha Lower
162
Diakonoffiana Koçak
165
Megalota Diakonoff
170
Costosa Diakonoff
173
Gatesclarkeana-group
177
Gatesclarkeana Diakonoff Statherotis-group
178 183
Metrioglypha Diakonoff
183
Dactylioglypha Diakonoff
187
Statherotis Meyrick
190
Aterpia Guenée
196
Proschistis Meyrick
199
Rhodacra Diakonoff
203
Bactra Stephens
206
Syntozyga Lower
212
Endothenia Stephens
216
Bactrini
205
Enarmoniini
219
Aglaogonia, gen. nov.
221
Anathamna Meyrick
225
Ancylis Hübner
229
Ancylophyes Diakonoff
235
Anthozela Meyrick
239
Balbidomaga Diakonoff
242
Cyphophanes Meyrick
248
Eucosmogastra Diakonoff
252
Helictophanes Meyrick
257
Irianassa Diakonoff
262
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CONTENTS
Loboschiza Diakonoff
266
Metaselena Diakonoff
271
Oriodryas Turner
275
Periphoeba Bradley
278
Pseudancylis, gen. nov.
282
Pternidora Meyrick
285
Tetramoera Diakonoff
288
Thysanocrepis Diakonoff
292
Toonavora, gen. nov.
295
Eucosmini Epinotia-group Crocidosema Zeller Epiblema-group
300 301 306 307
No available genus: absconditana-group
310
Coenobiodes Kuznetzov
313 317
Melanodaedala, gen. nov.
317
Rhopobota Lederer
321
Fibuloides Kuznetzov
326
Acroclita Lederer
334
Tritopterna Meyrick
338
Noduliferola Kuznetzov
341
Heleanna Clarke
345
Spilonota-group
349
Spilonota Stephens
350
Strepsicrates Meyrick
355
Holocola Meyrick
362
Hermenias Meyrick
371
Eccoptocera Walsingham
375
Unassigned to genus-group
380
Herpystis Meyrick
380
Icelita Bradley
383
Eucosmophyes Diakonoff
388
Whittenella, gen. nov.
392
Unassigned to genus (‘Acroclita’ tothastis Meyrick)
395
Grapholitini Loranthacydia-group Loranthacydia Horak, Common & Komai
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298
Epiblema Hübner
Acroclita-group
ix
396 398 399
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x
CONTENTS
Dichrorampha-group Pammenemima Diakonoff Cydia-group Leguminivora Obraztsov Notocydia, gen. nov. Fulcrifera Danilevsky & Kuznetzov Apocydia, gen. nov. Cydia Hübner Grapholita-group Thaumatotibia Zacher Gymnandrosoma Dyar Archiphlebia, gen. nov. Cryptophlebia Walsingham Acanthoclita Diakonoff Pammenopsis Kuznetzov Grapholita Treitschke Grapholita Treitschke Aspila Stephens Microsarotis Diakonoff Commoneria, gen. nov. Parapammene Obraztsov Ixonympha, gen. nov. Unplaced taxa
403 403 406 407 411 415 419 422 426 427 430 433 435 439 443 448 448 452 454 459 461 464 467
6. References
468
7. Appendices
477
8. Index
511
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Abstract
The family Tortricidae, with just over 9000 described species (Brown 2005), is the second largest lineage in the microlepidoptera, second only to the Gelechioidea, and by far the most important economically. A division into three subfamilies, Tortricinae, Chlidanotinae and Olethreutinae, has become generally accepted, though only the latter two are natural groupings defined by reliable characters (Horak 1998). The majority of tortricid caterpillars are leafrollers, but internal feeding is found in all subfamilies. Caterpillars boring in fruit, shoots, stems or roots are particularly prevalent in the Olethreutinae and the group includes many horticultural pests. This volume reviews the 90 olethreutine genera occurring in Australia, presenting revised concepts of the six recognised olethreutine tribes (Microcorsini, Olethreutini, Bactrini, Enarmoniini, Eucosmini and Grapholitini) and a refined classification based on informal genus-groups for the Olethreutini, Eucosmini and Grapholitini. The revision of the Grapholitini and the new grapholitine taxa are jointly authored by F. Komai and M. Horak. The Australian Olethreutinae are largely derived from the Oriental fauna, and results from this study reflect on the classification of the Oriental Olethreutinae, with one revised and 80 new generic combinations established for nonAustralian taxa. The treatment of each genus includes synonymy, a detailed morphological description, diagnosis, reviews of distribution and biology, discussion of taxonomic history, monophyly and relationships of the genus, and a checklist for all named Australian species. Illustrations of wing venation, adult moths, head, genitalia, and diagnostic secondary sexual structures are provided, wherever possible, for more than one species. A general section contains brief reviews of olethreutine phylogeny, adult morphology and biology, diversity and distribution of the Australian fauna, and a key to genera. Twelve new genera are proposed: Olethreutini: Cnecidophora, gen. nov., Atriscripta, gen. nov.; Enarmoniini: Aglaogonia, gen. nov., Pseudancylis, gen. nov., Toonavora, gen. nov.; Eucosmini: Melanodaedala, gen. nov., Whittenella, gen. nov.; Grapholitini (Komai & Horak): Notocydia, gen. nov., Apocydia, gen. nov., Archiphlebia, gen. nov., Commoneria, gen. nov. and Ixonympha, gen. nov.
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Ancylophyes is raised to genus level. Sixteen new species are named: Microcorsini: Collogenes albocingulata, sp. nov.; Olethreutini: Oxysemaphora notialis, sp. nov., O. hacobiani, sp. nov., Sorolopha johngreeni, sp. nov., Costosa australis, sp. nov.; Enarmoniini: Balbidomaga uptoni, sp. nov., Cyphophanes gracilivalva, sp. nov.; Eucosmini: Fibuloides phycitipalpia, sp. nov., F. minuta, sp. nov.; Eccoptocera australis, sp. nov., Eucosmophyes commoni, sp. nov.; Grapholitini (Komai & Horak): Leguminivora longigula, sp. nov., Notocydia niveimacula, sp. nov., Fulcrifera persinuata, sp. nov., Pammenopsis barbata, sp. nov., Microsarotis sanderyi, sp. nov. One revised and 41 new generic combinations are proposed for Australian species. Lectotypes are designated for Argyroploce thystas Meyrick, Helictophanes uberana Meyrick and Hermenias epidola Meyrick. Notable outcomes of this study are the inclusion of the Gatesclarkeana-group in the Olethreutini, the reassessment of the Bactrini as a highly derived rather than an ancestral group, and the discovery of a very diverse enarmoniine fauna in Australia, allowing a much more comprehensive definition of this tribe. Recognition of two seemingly monophyletic genus-groups, the predominantly Oriental Acroclita-group and the Australia-centered Spilonota-group, is a major step towards understanding the phylogeny of the Eucosmini. The Australian Grapholitini are surprisingly diverse with several endemic genera, two with a unique biology, feeding on Loranthaceae. Whereas the bulk of the Australian Olethreutinae have strong Oriental relationships, the eucosmine Spilonotagroup on Myrtaceae and several endemic genera in the Grapholitini represent Australian radiations. M. Horak CSIRO Entomology GPO Box 1700 Canberra, ACT 2601 Australia F. Komai Department of General Education Osaka University of Arts Kanan-cho, Minamikawachi-gun, Osaka 585–8555 Japan
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Introduction
Most people’s introduction to olethreutine moths would have been an unpleasant experience, such as finding an unwelcome codling moth caterpillar in their apple. The large majority of tortricid caterpillars are so-called leaf-rollers, feeding between rolled or folded leaves spun together to form a shelter, but internal feeding has been developed several times within the family, with the larvae boring into shoots, roots or fruit. Several groups in the subfamily Olethreutinae have fruitboring larvae and some are of great economic importance. The Tortricidae, the sole family in the superfamily Tortricoidea, have just over 9000 described species (Brown 2005) and are the second largest lineage in the microlepidoptera, second only to the Gelechioidea. They belong to the Ditrysia, the higher Lepidoptera comprising over 95% of all species and defined by a separate copulatory orifice and ovipore in the female, the two internally connected for sperm transfer. Together with the Galacticoidea, Zygaenoidea, Cossoidea, Sesioidea, Choreutoidea, Urodoidea, Schreckensteinioidea, Epermenioidea, Alucitoidea and Pterophoroidea, they form the Apoditrysia (Kristensen and Skalski 1998), based on the structure of the second abdominal sternum with ‘tortricoid’ apodemes and produced anterolateral corners. However, it is not confirmed that the Apoditrysia are a cohesive, monophyletic group, and with the Tortricoidea by far their largest superfamily the assemblage is often referred to as the ‘tortricoid’ grade. The tortricids are a well-defined family easily recognisable by a combination of characters. Adult tortricids are mostly small to medium-sized moths folding their wings roof-like above the body at rest, sometimes nearly flat in some Tortricinae. In many Eucosmini the folded wings are longitudinally bent at a right angle so that the cross section of a sitting adult is rectangular. The tortricid frons scaling is characteristic with a large, downcurved tuft just beneath the antennae but with the rest of the frons scaling very short, appressed and upwardly oriented. The labial palpi are relatively stout, either porrect and sinuate or gently upcurved, but never with a long, slender and tapering terminal segment. The proboscis never has scales at its base, and the
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maxillary palpi are small and usually not visible on the undissected head. Ocellus and chaetosema are nearly always present, and the antenna has a basal notch in the males of some groups. The thorax often has a posterior, dorsal tuft of raised scales, and in the male especially the hind tibia may be modified, in particular with a scale pencil from its base in many Olethreutinae. The flat ovipositor lobes of the female are unmistakable and characteristic for the family. A division into three subfamilies has become generally accepted for the Tortricidae (Horak 1998), with reliable derived characters defining both the Chlidanotinae and the Olethreutinae. The Tortricinae are a collection of ancestral as well as derived groups, and the two other subfamilies are presumably derived from within this assemblage. The Olethreutinae are characterised by two unique derived character states: 1, fusion of juxta and caulis in the male genitalia into a single functional unit, and 2, reduction of the antennal scaling to one ring of scales per segment (from two rings in other Tortricidae). Many other features, often present in various combinations, produce the usually characteristic ‘olethreutine look’. Among these are typical wing pattern elements such as the costal strigulae and ocelloid patch, costal fold and antennal notch in some groups, cubital pecten on the hindwing, scale pencil on the male hind tibia, and several character states of the genitalia in both sexes. The book provides an overview and key to the 90 olethreutine genera occurring in Australia, represented by 249 named and about 200 unnamed Australian species. Each genus is described in detail, diagnosed and defined by unique, derived characters, and its relationships are discussed. A special effort has been made to provide copious illustrations of high quality of adults, heads, genitalia and other diagnostic structures. Whenever possible more than one species was illustrated to convey generic concepts and allow identification of common or pest species. Literature, biology and distribution are reviewed for each genus, and a checklist of all Australian species is given. This study is based on the named Australian Olethreutinae wherever possible; new species are
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4
OLETHREUTINE MOTHS OF AUSTRALIA
only described if no named species was available or if a species added significantly to the generic concept. Several more olethreutine genera are known to be present in Australia, but most are represented by insufficient material to allow description. This book follows the pattern of several earlier volumes in the series by providing a ‘top-down’ approach (Robinson and Nielsen 1989), primarily addressing the generic level and refraining from resolving difficult species-groups. A sound generic framework based on demonstrably monophyletic and thus unambiguously definable genera will enable efficient species-level studies as demand arises. The current study also shows that congeneric species very often have similar biology and host preferences. This means that information about the biology of just one species provides a hypothesis for the biology of its congeners, as long as the genus is a monophyletic grouping. It would not have been possible to complete this volume without Furumi Komai’s willingness to be first author of the Grapholitini chapter. Only his knowledge, expertise and long involvement with this extremely difficult group made it possible to deal with the surprising diversity of the Australian fauna. The chapter in this book is based on a comprehensive species-level revision of the Australian Grapholitini that will be published at a later date, including a large number of new species. Conforming to Nielsen et al. (1996), this revision covers the fauna of continental Australia and Tasmania and includes islands close to the mainland. Distant islands under Australian control such as Ashmore Reef, Cocos-Keeling Is, Christmas I., Macquarie I., Lord Howe I. and Norfolk I. are not included. The Australian Olethreutinae are probably all derived from incursions from the Oriental and Papuan region and can only be understood in the context of this fauna. Many of the genera described from Indonesia, New Guinea and the Pacific region were regarded in isolation and earlier Australian names were ignored. Some were based on very little and often damaged material, even on only one sex, and it was impossible to form an adequate generic concept. Long series of both sexes are available in the ANIC for many Australian species allowing characterisation of some previously dubious genera and understanding of their relationships. Comparison of genera across the Oriental and Australian regions has led to several synonymies and numerous recombinations of non-Australian species.
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The earliest olethreutines to be described from Australia were included in Francis Walker’s List of the Specimens of Lepidopterous Insects in the Collection of the British Museum, published in instalments during the middle of the 19th century. The first publication entirely dedicated to the Olethreutinae was in 1881 by Edward Meyrick, who lived in Australia and New Zealand from 1877 to 1886, teaching classics (Robinson and Nielsen 1989) in Sydney, Parramatta and Christchurch. During his stay he accumulated a large collection of Australian and New Zealand microlepidoptera that he took back to Britain on his return, and that is now in The Natural History Museum, London. In a long series of papers from 1895 to 1920, O. B. Lower gradually added new species and genera to the known Australian olethreutine fauna. In 1911, long after his return to London, Meyrick revisited the Australian Olethreutinae and once again produced a comprehensive, updated revision of the subfamily. The second author to contribute greatly to the knowledge of the Australian Olethreutinae was Alfred Jefferis Turner. Turner was a highly regarded medical doctor, yet managed to publish over one hundred papers on Lepidoptera (Mackerras 1949), describing many new species from subtropical and tropical regions hardly collected during Meyrick’s times. He accepted Meyrick’s classification for the Olethreutinae but gradually added numerous new taxa, usually in papers dealing with several families or a local fauna. Late in life, Turner (1946) published an entire paper devoted to the Olethreutinae in which he described a large number of new taxa, including, uncharacteristically, many synonyms. The Olethreutinae have an estimated worldwide total of about 4300 named species (J. W. Brown personal communication). Our knowledge of the subfamily varies greatly between faunal regions. However, the recent publication of a World Catalogue of the Tortricidae (Brown 2005) has closed a large gap in our knowledge as there was no previous checklist for the Oriental region. The Holarctic region is well documented: with the still very relevant revision of the Nearctic fauna by Heinrich (1926); Miller’s (1987) ‘Guide to the Olethreutine Moths of midland North America’; Razowski’s (1989) synopsis of the Palaearctic olethreutine genera, and his Tortricidae of Europe II (Olethreutinae) (Razowski 2003a); Kuznetzov’s (1987) revision of the fauna of the European part of the former USSR, and Kawabe’s (1982) tortricid chapter in ‘Moths of Japan’. Komai’s (1999) revision of the Palaearctic genera of the Grapholita-group
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INTRODUCTION
was an invaluable base for revision of the Australian Grapholitini. There are no comprehensive treatments of the Olethreutinae of the Oriental region except for Liu and Li’s (2002) tortricid volume, and Diakonoff’s (1973) monumental monograph on the South Asiatic Olethreutini, that provided the key to the understanding of the Australian fauna. However, there is a wealth of information on Oriental Olethreutinae in innumerable publications, especially by Diakonoff, Kawabe and Bradley. Both
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5
Diakonoff’s studies of the Olethreutinae of New Guinea and Clarke’s monographs on the Polynesian and Micronesian fauna are crucial to the Australian fauna. Both the Olethreutinae of the African and Neotropical regions still await revision. Diakonoff addressed the fauna of Madagascar in a series of papers of relevance to the Australian Olethreutinae, but information about Neotropical taxa has to be largely gleaned from Nearctic literature.
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Material and methods Material studied
Taxonomic treatment
This work was deliberately focussed at the generic level and whilst, where possible, all the named Australian olethreutine species were accounted for, no attempt was made to document and describe all Australian species. Recuration of the Olethreutinae at the Australian National Insect Collection (ANIC) was the basis for the study, integrating the large V. J. Robinson collection with the ANIC collection initially curated by I. F. B. Common and including most of the types by A. J. Turner. The types were examined if they could be located, either during visits to the relevant collections or obtained on loan, unless the identity of Australian species was beyond doubt based on the collection of Lepidoptera type images in the ANIC. The Natural History Museum, London (BMNH), holding most of the types of Edward Meyrick and Francis Walker, is crucial to any study of Australian Lepidoptera, especially as the Olethreutinae there were partially curated by K. R. Tuck concurrently with this study. The olethreutine holdings of the South Australian Museum, Adelaide, containing type material of O. B. Lower and T. P. Lucas, were borrowed and curated. Olethreutine types were also examined in the Queensland Museum, Brisbane, the Australian Museum, Sydney, and the Museum Victoria, Melbourne. Comparative material was borrowed from the New Zealand Arthropod Collection, Auckland. The Australian olethreutines are largely derived from the Oriental fauna, and their genera often have a wide distribution throughout the Oriental region and the Pacific, sometimes even further. Many genera only just reach the north-eastern or north-western corner of Australia with a single species, and generic definitions had to include a study of the olethreutine fauna from well beyond Australia. The holdings of New Guinea Lepidoptera at the ANIC, both my own and the W. W. Brandt collection, and the material collected during Project Wallace in Sulawesi, were instrumental in helping to define genera. Type species of olethreutine genera from the Oriental region, New Guinea and the Pacific were examined at the BMNH, the National Museum of Natural History Naturalis, Leiden, and the Smithsonian Institution, Washington.
All genera represented by named species in the Australian fauna have been revised, their monophyly established and their relationships discussed. In many cases, the species composition of genera beyond Australia had to be adjusted, and a number of new combinations for the Oriental and Papuan fauna have been established. Where possible, more than one species has been figured for each genus to provide a visual concept of generic characters, including some non-Australian taxa if needed. New species have only been described if no named Australian species were available for commonly occurring or taxonomically interesting genera, or if the species concerned added significantly to the scope of a genus. The sequence of genera and species reflects relationships as far as this is possible in a linear arrangement. It follows the layout of the olethreutine collection at the ANIC, where arranging closely related taxa next to each other is the only way to allow identification of a fauna with at best half the species named. Misspellings were only added if they were significant, either widely used or not easily recognisable as misspellings. Three species could not be placed as their types are lost and the description is not sufficiently informative to link them to a known species. The three names are listed under ‘Unplaced taxa’ after the last tribe. Unless it was necessary to stabilise the usage and classification, no lectotypes have been designated. The world catalogue of Tortricidae (Brown 2005) provided type information and type locality for each species. It was stated in the introduction to the catalogue that in many cases the term ‘lectotype’ in the catalogue referred to a selected and thus potential rather than a designated and published lectotype. This applies to nearly all lectotypes in the ANIC and many in the BMNH. In the case of a potential lectotype, the type locality given in the catalogue refers to the specimen labelled ‘lectotype’, whereas in this work the localities for all the syntypes are given. In some cases when a species could only be tentatively identified, yet indicating its probable position provided useful information, the species name was preceded by a question mark, i.e.
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8
OLETHREUTINE MOTHS OF AUSTRALIA
?mysterica. If an undescribed species was considered to be close to, but probably not conspecific with, a named species, the convention was to use ‘near’, i.e. near metallocosma.
Preparatory techniques Head illustrations were either made by low-power scanning electron microscopy using a JEOL SM 35C, after sputter-coating with gold, or by macro photography. Wing preparations were made by the method developed by Common (1987, 1990). Dislodged wings were gently immersed for a few minutes in 100% ethanol to remove any traces of oil or fat, and then moved for a few minutes to an excavated block containing a few drops of 10% household bleach dissolved in water. Under a dissecting microscope, some of the scales were gently removed with two fine synthetic brushes during the bleaching process, and the wings then washed three times in 30% ethanol. After a final rinse in 70% ethanol, the wings were immersed overnight in the following acid fuchsin stain: 1 g of acid fuchsin dissolved in 100 ml of 50% ethanol mixed with 5 ml glacial acetic acid in which 10 g of chloral hydrate had been dissolved. Each wing was gently transferred from the stain into an excavated block with 95–100% ethanol and immediately brushed flat. After rinsing twice more in 95–100% ethanol, whilst carefully removing further scales if necessary, the wings were, with a rolling brush motion, dropped onto a dry microscope slide and covered with a small drop of Euparal as soon as most of the ethanol had evaporated. After gently lowering the coverslip onto the preparation it was weighed down for a few days to keep the wings flattened. Genitalia preparations were made according to the technique described by Robinson (1976) and Common (1990). The entire abdomen was added to a small amount of 10% KOH in a glass tube lightly stoppered with a loose cork, stood in a beaker filled with water to just beyond the KOH level, gently brought to the boil and kept at boiling point for about 10 min. The abdomen was rinsed in water several times and transferred to 30% ethanol for dissection. The genitalia were separated from the abdominal pelt, but in the female the 7th sternite was left attached to the abdominal tip as it was often fused with the sterigma. Genitalia and the cleaned abdominal pelt were stained briefly in 70%
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ethanol to which a few drops of a 1% solution of Chlorazol Black E in 70% ethanol had been added. After flattening, cleaning and dehydration in 2–3 changes of absolute ethanol, the genitalia and pelt were mounted in Euparal. The majority of images are based on conventional photography. However, some genitalic images, in particular in the Enarmoniini, were done with a Zeiss ProgRes 3012 high-resolution digital camera on a compound microscope and the software Automontage (Synoptics Ltd, Cambridge, UK), which combined a series of focus levels into a single image. Many adult and a few genitalia images have been reversed to produce comparable illustrations, this is flagged in the captions for adult images only. Wingspan measurements were made with a ruler, from wing tip to wing tip of conventionally set specimens. The wingspan is descriptively summarised as five size ranges: less than 8 mm as very small; 8–12 mm as small; 12–18 mm as medium-sized; 18–24 mm as large; over 24 mm as very large. Head and wing ratios were measured with an eyepiece graticule on a stereoscopic microscope. Labial palpus length is given as a multiple of widest eye diameter measured parallel to body axis. Forewing index is maximum wing width measured at a right angle to the costa, divided by the wing length measured from the wing base to apex. Ratio of discal cell length to wing length is also based on wing length measured from wing base to apex.
Abbreviations AM ANIC
Australian Museum, Sydney, Australia Australian National Insect Collection, CSIRO Entomology, Canberra, Australia BMNH The Natural History Museum, London, UK E. D. E. E. D. Edwards I. F. B. C. I. F. B. Common M. S. U. M. S. Upton NP National Park NTM Northern Territory Museum, Darwin, Australia QM Queensland Museum, Brisbane, Australia S sternum T tergum
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Acknowledgments
This project began with research into tortricid phylogeny funded by a CSIRO Postdoctoral Award (1986–1988). Two research grants from the Australian Biological Resources Study (ABRS) and generous support by CSIRO Entomology over many years allowed gradual revision of the 90 Australian olethreutine genera, preparation of the numerous illustrations and completion of the final manuscript. A grant from the Hermon Slade Foundation assisted with publishing costs. This volume would never have been written without the help and support of three exceptionally generous colleagues: The late Dr Ebbe Nielsen inspired and fostered the notion that a revision of all the Australian Olethreutinae genera could be done. Mr Ted Edwards, CSIRO Canberra, provided daily unstinting support, advice and guidance throughout the long haul and critically read the entire manuscript. Prof. Furumi Komai came to the rescue dealing with the difficult Grapholitini and offered his knowledge and expertise as first author for the chapter on the Australian grapholitine genera. The help of colleagues at CSIRO Entomology is gratefully acknowledged, foremost that of Ms Vanna Rangsi who most skilfully prepared many genitalia figures and all the plates for this volume, as well as setting selected specimens for illustration. The photography for most adult and genitalia images was done by Mr John Green, and by Mr David McClenaghan for heads and some smaller adults. Mr Eric Hines prepared the electron micrographs of the heads, Ms Judith Hull typed large parts of the manuscript, and Dr Adam Slipinski and Ms Ann Hastings provided invaluable technical advice, help and support. Grateful thanks go to Dr Leif Aarvik, Prof. Richard Brown, Dr John Dugdale, Dr Bruce Halliday, Prof. Furumi Komai and Dr Adam Slipinski who critically read the manuscript or parts therof, and in particular to Mr Ted Edwards,
120503•Olethreutine Moths 4pp.in9 9
Mr Murray Upton and Dr John Brown who attended to the text in detail throughout, including the Germanic English. Drs L. Aarvik, J. Baixeras, V. I. Kuznetzov, H. Li, Y. Nasu and J. Razowski kindly provided last-minute comments on the absconditana-group. Thanks are due also to CSIRO Publishing, especially to Ms Ann Crabb and Ms Briana Elwood for editorial help and to Mr Paul Reekie for his ongoing support for the Monographs on Australian Lepidoptera series. We thank the following for lending us material in their charge for study: Mr Kevin Tuck, BMNH, London, UK; Dr J. S. Dugdale, New Zealand Arthropod Collection, Auckland, New Zealand; Drs Rienk de Jong and Erik van Nieukerken, the National Museum of Natural History Naturalis, Leiden, the Netherlands; Dr Geoff Monteith, QM, Brisbane; Mr Ken Walker, MV, Melbourne. We are particularly grateful to Ms Jan Forrest, South Australian Museum, Adelaide, for extensive loans of olethreutine material in her care. A great debt of thanks is owed to Mr Kevin Tuck whose generous help, advice and support during all the years of this study have been invaluable. Dr Gaden Robinson, BMNH, kindly made an electronic file of the tortricid part of the HOSTS database available for easy interrogation. This study was based mostly on the wonderfully rich Lepidoptera collection in the Australian National Insect Collection, CSIRO Entomology, Canberra, built up over the last 50 years by Dr Ian Common and his colleagues. Ian’s work on tortricine moths also provided an important stimulus and benchmark. Mr. Ken Sandery and Dr David Rentz made special efforts to collect rare and obscure species for this study. Material collected in Sulawesi during Project Wallace, organised by the Royal Entomological Society of London and the Indonesian Institute of Sciences, has been drawn upon heavily for generic definitions.
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1. Phylogeny of the Olethreutinae
The Tortricidae are the only family in the homogeneous and undoubtedly monophyletic superfamily Tortricoidea. The group is easily recognisable by a combination of character states such as: an unscaled proboscis, presence of ocellus and chaetosema, porrect or upcurved labial palpi (but never horn-shaped with a long, slender, curved terminal segment), minute maxillary palpi generally not visible on undissected specimens, usually rough head scaling but with the scaling on the lower frons very short and upwardly oriented, ‘tortricoid’ sternum 2 with broad apodemes and at most very weak sternal rods, and flat, leaf-like ovipositor lobes. The flat ovipositor lobes of the female are a derived feature (apomorphy) characterising the family (Dugdale 1988; Horak and Brown 1991), with only few exceptions that are due to secondary modification associated with specialised oviposition. Upwardly oriented scaling on the lower part of the frons and the anal fork with straight prongs, found in most tortricid caterpillars, are possible further apomorphies for the family. The anal fork serves to eject frass from the shelter and tends to be lost in groups where caterpillars feed as borers. Three subfamilies, Tortricinae, Chlidanotinae and Olethreutinae, have become generally accepted within the Tortricidae (Horak 1998). The latter two are considered monophyletic based on several unique, derived character states whereas the Tortricinae are paraphyletic, comprising highly derived groups as well as very generalised, plesiomorphic taxa. All Olethreutinae share two unique features: 1, highly modified male genitalia with juxta and caulis fused into a single structure instead of loosely hinged as in other tortricids, and 2, each segment of the antennal shaft with only a single row of scales instead of two rows. Two rows of scales per antennal segment are considered to be the groundplan state for the Ditrysia (Robinson and Nielsen 1993). Some tortricine Sparganothini (not occurring in Australia) also have antennae with a single ring of scales per segment, but they differ in the details of their structure and are clearly independently derived. Olethreutines can usually also be recognised by the presence of a cubital pecten (Fig.
120503•Olethreutine Moths 4pp.in11 11
14), but this structure is absent in a few olethreutine genera and present in some generalised tortricines. Whilst male olethreutine genitalia are highly derived, the wing venation in this subfamily is more generalised overall than in both the Tortricinae and Chlidanotinae. M-stem and chorda are retained at least in some species of most olethreutine genera, in contrast to the large majority of the other two subfamilies. The historical development of the tribal classification of the Olethreutinae is discussed in detail in the general remarks for each of the six tribes here recognised. The aim of this revision was primarily to define the olethreutine genera present in Australia in cladistic terms, based on derived characters that would provide clear generic limits. This will allow identification at least to genus and the assignment of unnamed and yet-to-be-discovered species to the correct genus. Examination of the type species and as wide a range of species as possible for each genus, including all taxa present in Australia, was the necessary basis for such a study. On the other hand, there was no need to resolve difficult species complexes as they do not impinge on generic definitions. Generic boundaries are subjective depending on the chosen level of inclusiveness even in a cladistic framework based on monophyletic groupings that comprise all the progenies of the taxon that acquired a given derived feature. An entirely pragmatic approach was chosen, guided by practicalities and accepted usage wherever possible, resulting in the fewest possible nomenclatural changes. After the genera were individually assembled and defined they were used as building blocks for a higher classification by searching for synapomorphies, i.e. shared derived characters, linking them into groups. This process was very successful for the Olethreutini, building on the monumental monograph of the south-east Asian fauna by Diakonoff (1973), and for the Eucosmini. It was less satisfactory for the Enarmoniini. Though several pairs of sister genera were identified, it was not possible to identify cohesive higher groupings, and the genera were arranged in alphabetical order within the tribe. Komai (1999) had already provided a phylogeny for the Palaearctic
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12
OLETHREUTINE MOTHS OF AUSTRALIA
Table 1.
Characters used in cladistic analysis
(1)
Upper part of male frons, scaling: (1) with short scales; (2) with moderately long anteriorly projecting scales; (3) with long anteriorly projecting scales; (4) with modified, sculpted scaling.
(2)
Labial palpus length, as × horizontal eye diameter: (1) lower value more than 2×; (2) lower and upper values including 2×; (3) upper value less than 2×.
(3)
Labial palpus, shape: (1) porrect, sinuate; (2) upcurved.
(4)
Labial palpus, width: (1) slender throughout or postmedially at most slightly expanded; (2) postmedially significantly expanded; (3) widest in middle.
(5)
Second segment of labial palpus: (1) slender throughout; (2) slender at base, with long loose scales along dorsal and ventral margins; (3) with long loose scales along ventral margin.
(6)
Third segment of labial palpus, length: (1) short; (2) moderately long; (3) very long.
(7)
Third segment of labial palpus, width: (1) very slender; (2) moderately slender; (3) stout.
(8)
Antenna in male, notch: (1) without notch; (2) with notch.
(9)
Notch, position: (1) at base; (2) distant from base.
(10)
Antenna in male, scaling: (1) entirely scaled; (2) anterior surface without scales.
(11)
Antenna in male, cilia length: (1) with minute cilia, hardly visible in stereo microscope; (2) with short cilia, less than 56M diameter of flagellum; (3) with moderately long cilia, 56M to 56O diameter of flagellum; (4) with long cilia, clearly longer than 56O diameter of flagellum.
(12)
Thorax, crest: (1) without crest; (2) with posterior crest.
(13)
Scale pencil from anteroventral corner of tegulae: (1) normal; (2) very long and slender.
(14)
Fore or mid tibia in male: (1) unmodified; (2) enlarged by long scales; (3) with tibial pencil (fore tibia).
(15)
Hind tibia in male, scale pencil: (1) unmodified; (2) with scale pencil.
(16)
Hind tibia in male, scaling apart from pencil: (1) unmodified; (2) with scale fringes along margin; (3) with tuft of long hairscales dorsally at base.
(17)
Forewing, costal fold: (1) without costal fold; (2) with costal fold; (3) with apparent costal fold of modified scales.
(18)
Forewing, shape: (1) subtriangular; (2) subrectangular; (3) subovate.
(19)
Forewing, raised scales on dorsum: (1) without tufts of raised scales on dorsum; (2) with tufts of raised scales on dorsum.
(20)
Ocelloid patch, development: (1) well developed; (2) not apparent.
(21)
Forewing costa, shape: (1) hardly curved; (2) distinctly curved throughout; (3) curved near base and distinctly curved distally; (4) curved near base then nearly straight; (5) curved distally only; (6) lightly sinuate; (7) strongly sinuate.
(22)
Forewing apex, shape: (1) acute, even if tip rounded; (2) nearly rectangular; (3) broadly rounded; (4) with conspicuous notch below apex.
(23)
Forewing termen, whether oblique: (1) oblique; (2) not oblique; (3) outwardly oblique.
(24)
Forewing termen, whether sinuate: (1) not sinuate; (2) sinuate.
(25)
M-stem, development: (1) well developed; (2) only trace in membrane; (3) totally absent.
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PHYLOGENY OF THE OLETHREUTINAE
(26)
Chorda: (1) present; (2) absent.
(27)
Chorda, origin: (1) from between R1 and R2; (2) from R2; (3) from between R2 and R3.
(28)
Chorda, termination: (1) to R4; (2) to between R4 and R5; (3) to R5; (4) to beyond R5.
(29)
R3, position: (1) distant from R4; (2) closely approximated to R4 at base; (3) connate with R4; (4) stalked with R4; (5) modified.
(30)
R4 and R5: (1) separate; (2) connate; (3) stalked; (4) R4 and R5 entirely fused; (5) fused R4 and R5 stalked with M1.
(31)
M3, position: (1) separate and closer to M2 than to CuA1; (2) separate, equidistant between M2 and CuA1; (3) separate and closer to CuA1 than to M2; (4) connate with CuA1; (5) stalked with CuA1; (6) fused with CuA1.
(32)
Forewing, distal course of M3: (1) distally not curved towards apex; (2) distally curved towards apex.
(33)
Forewing, distal course of CuA1: (1) distally not curved towards apex; (2) distally curved towards apex.
(34)
Axillary tuft: (1) present, unmodified; (2) as scattered hairscales between 1A+2A and anal margin; (3) otherwise modified; (4) absent.
(35)
Anal region in male, whether enlarged: (1) not enlarged; (2) enlarged, but not forming projecting lobe; (3) expanded to form projecting lobe.
(36)
Conspicuous additional lobe from base of anal margin in male: (1) absent; (2) present.
(37)
Anal margin in male, development: (1) unmodified; (2) thickened.
(38)
Anal margin in male, whether folded: (1) not folded; (2) folded downwardly; (3) folded upwardly.
(39)
Upperside of anal region in male, modified scales: (1) without modified scales; (2) with scale pencil from base; (3) with patch of short, modified scales on upper side; (4) with patch of hairscales on upper side between 3A and anal margin; (5) with modified scales along anal margin; (6) with scattered hairscales above on anal region; (7) with band of modified scales along margin; (8) with hairscales along 3A, especially distally.
(40)
Underside of anal region, modified scales: (1) without modified scales; (2) with modified scales near anal margin; (3) with bundle of long scales from between CuP and 1A+2A; (4) with large round depression with modified scales.
(41)
Modified, small and more widely spaced hindwing scales in male: (1) absent; (2) present.
(42)
Persistent scale fringe on distal half of hindwing costa: (1) absent; (2) present.
(43)
Rs and M1: (1) distant and diverging from base; (2) close at base but immediately diverging; (3) adjacent and parallel at base; (4) connate; (5) stalked.
(44)
Hindwing, whether course of M2, M3 and CuA1 is sexually dimorphic: (1) same configuration in male and female, proportions may be different; (2) modified in male.
(45)
Hindwing, position of base of M2 in female: (1) parallel and distant from M3; (2) bent towards M3 but bases not close; (3) bent close to base of M3; (4) connate from angle of cell.
(46)
Hindwing, configuration of M3 and CuA1 in female: (1) separate; (2) connate; (3) stalked; (4) entirely fused; (5) CuA1 from far below angle of cell.
(47)
3A in hindwing: (1) present; (2) absent.
(48)
Anterolateral processes on S2: (1) present; (2) absent.
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13
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14
OLETHREUTINE MOTHS OF AUSTRALIA
(49)
Sternal apodemes, shape: (1) slender; (2) unremarkable; (3) stout; (4) tip slightly clubshaped.
(50)
Sternal rods on S2: (1) recognisable; (2) not present.
(51)
Paired round depressions on T2: (1) absent; (2) present.
(52)
Modified abdominal scaling dorsally in male: (1) absent; (2) present.
(53)
Modified abdominal scaling laterally in male: (1) absent; (2) present, but not at base of abdomen; (3) present, near base of abdomen.
(54)
Modified scaling ventrally on S2 in male: (1) absent; (2) present.
(55)
Hindmargin of S8 in male: (1) straight; (2) medially concave; (3) bilobed; (4) scalloped; (5) medially convex; (6) downcurved, hood-shaped.
(56)
Coremata behind S8: (1) fully developed; (2) partially developed; (3) absent.
(57)
Sclerotisation at base of T8 in male: (1) not conspicuous transverse sclerotisation; (2) transverse sclerotisation.
(58)
Shape of T8 in male: (1) unremarkable; (2) small; (3) very large; (4) hindmargin a projecting lobe.
(59)
Vinculum, whether modified: (1) unremarkable; (2) wide; (3) W-shaped at base; (4) widened at base into saccus-like process.
(60)
Tuft of long scales laterally from base of tegumen: (1) absent; (2) from lateral surface of tegumen, towards base; (3) from enlarged base of pedunculi; (4) from between longitudinal lateral folds of tegumen; (5) from small fold between tegumen and valva.
(61)
Tegumen, shape: (1) band-shaped; (2) hood-shaped.
(62)
Tegumen, outline: (1) subtriangular; (2) subrectangular; (3) semi-ovate.
(63)
Tegumen, length: (1) tegumen not longer than wide; (2) tegumen longer than wide. (Length to base of uncus.)
(64)
Tegumen shape: (1) tegumen not with pronounced shoulders; (2) tegumen with pronounced shoulders.
(65)
Tegumen, degree of sclerotisation: (1) strongly sclerotised; (2) moderately sclerotised; (3) weakly sclerotised.
(66)
Tegumen, apodemes for muscle m4: (1) with apodemes for muscle m4 present; (2) without apodemes for muscle m4.
(67)
Tegumen, concavity dorsally at base of uncus: (1) absent; (2) present.
(68)
Uncus: (1) present; (2) absent.
(69)
Uncus, whether paired: (1) paired processes from base; (2) a single process at base.
(70)
Uncus shape: (1) finger-shaped; (2) spatulate; (3). hump-shaped; (4) lobe-shaped; (5) with ‘neck’ and dilated apex; (6) distally bifid process; (7) tooth-shaped; (8) nipple-shaped.
(71)
Uncus bristles: (1) without conspicuous bristles; (2) with an apical tuft of bristles; (3) with bristles ventrally at apex; (4) with bristles on dorsal surface only; (5) with a conspicuous fringe of flattened pegs; (6) with scattered bristles throughout; (7) with persistent hairscales; (8) densely bristled throughout.
(72)
Uncus, whether ventrally curved: (1) straight; (2) ventrally curved.
(73)
Socii: (1) present; (2) absent.
(74)
Socii: (1) membranous; (2) sclerotised.
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PHYLOGENY OF THE OLETHREUTINAE
(75)
Socii: (1) small; (2) medium-sized; (3) large.
(76)
Socii shape: (1) finger-shaped; (2) band-shaped; (3) lobe-shaped; (4) knob-shaped; (5) illdefined patches only; (6) bipartite; (7) inflated.
(77)
Bristles modified to scales on socii: (1) absent (i.e. bristles unmodified); (2) scales present.
(78)
Gnathos: (1) present; (2) absent.
(79)
Gnathos: (1) membranous; (2) sclerotised.
(80)
Gnathos shape: (1) lateral bands only, at most weakly fused medially; (2) complete, transverse band; (3) both arms medially joined in free process; (4) arms medially fused with sclerotised ventral wall of anal tube (subscaphium); (5) linked with anellus; (6) two rising, parallel arms; (7) fused with socii; (8) developed medially as U-shaped band only; (9) large rising lobes with bristles or pegs.
(81)
Gnathos structure: (1) not conspicuously spinulose; (2) conspicuously spinulose; (3) with bristles.
(82)
Ventral surface of anal tube: (1) unremarkable; (2) sclerotised; (3) spinulose; (4) connected with anellus.
(83)
Valva: (1) symmetrical; (2) asymmetrical (i.e. the two valvae of different shape).
(84)
Valva: (1) without basal excavation; (2) with small basal excavation; (3) with normal basal excavation; (4) with large basal excavation.
(85)
Valva: (1) without process from ventral margin of basal excavation; (2) with naked process from ventral margin of basal excavation; (3) with bristled process from ventral margin of basal excavation. (Processes from below ventral margin are not considered homologous.)
(86)
Ventral margin of valva: (1) roughly convex throughout; (2) with conspicuous emargination.
(87)
Cucullus shape: (1) elongate ovate; (2) subrectangular; (3) narrowly band-shaped; (4) narrowing towards apex; (5) transversely ovate; (6) subtriangular; (7) semicircular; (8) bilobed.
(88)
Cucullus spines on inner surface: (1) not with isolated prominent spine(s) except for slightly larger bristles along margin; (2) with several isolated prominent spine(s); (3) with only one large ventral spike or thorn; (4) with one large apical spine.
(89)
Single spine on tip of cucullus: (1) absent; (2) simple; (3) bifid or bipronged.
(90)
Sacculus shape: (1) not conspicuously projecting; (2) a projecting lobe.
(91)
Sacculus spine groups other than those from process: (1) without discrete groups of spines; (2) with discrete group(s) of spines. (This refers to groups of strong, shortish spines on inner surface but not the tuft of bristles below the basal excavation.)
(92)
Shape of costal process: (1) without costal process; (2) at base of costa; (3) on inner surface; (4) upwardly curved; (5) reduced.
(93)
Invaginated pocket in outer valva base: (1) absent; (2) weakly developed; (3) strongly developed.
(94)
Process from sacculus or from just beyond basal excavation: (1) absent; (2) present. (Not considered homologous with processes projecting from margin of basal excavation.)
(95)
Shape of process from sacculus or from just beyond basal excavation: (1) bristled process; (2) spiny process; (3) naked process; (4) keel-shaped process.
(96)
Process other than costal process from above basal excavation: (1) absent; (2) present.
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15
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OLETHREUTINE MOTHS OF AUSTRALIA
(97)
Shape of process from above basal excavation: (1) large lobe crowned with single spike; (2) bristled knob, digit, hump or lobe; (3) knob, digit, hump or lobe with strong spines.
(98)
Other process from inner surface of valva: (1) absent; (2) knob with long bristles in excavation of ventral margin; (3) bristled blade on ventral margin; (4) raised base of cucullus.
(99)
Base of costa (apart from costal process): (1) unremarkable; (2) forming hump; (3) with projecting angle close to base; (4) with finely bristled lobes; (5) with large tuft of hairscales from depression.
(100)
Modified scales or bristles from outer surface of valva: (1) absent; (2) patch of bristles; (3) band along ventral margin; (4) conspicuous flattened bristles; (5) patch of very long scales from cucullus; (6) other modified scales.
(101)
Other modified scales or bristles from inner surface of valva: (1) absent; (2) tuft of bristles beyond basal emargination; (3) row of pegs on cucullus ‘neck’; (4) inflated scales from inner surface; (5) tuft of long bristles from centre of valva.
(102)
Caulis: (1) no caulis; (2) fused with anellus; (3) short; (4) moderately long; (5) long; (6) fused with juxta.
(103)
Anellus shape: (1) anellus not obvious; (2) ring-shaped; (3) cup-shaped; (4) wide and irregular; (5) dorsally extended; (6) ventrally expanded; (7) ventrally on each side with long spike; (8) caulis and anellus fused into elongate structure; (9) strands between anellus and base of socii.
(104)
Cornuti: (1) deciduous; (2) fixed; (3) absent.
(105)
Cornuti shape: (1) needle-shaped; (2) lanceolate; (3) long, heavy spines; (4) flattened, sinuate; (5) very small, triangular.
(106)
Cornuti numbers: (1) large bundle; (2) one to few.
(107)
Sterigma position: (1) in intersegmental membrane behind S7; (2) in excavation of hindmargin of S7; (3) behind projecting hindmargin of S7; (4) within S7.
(108)
Sterigma: (1) not connected with anterior apophyses; (2) laterally connected with anterior apophyses.
(109)
Sterigma: (1) not fused with S7; (2) fused with S7.
(110)
Sterigma shape: (1) not differentiated; (2) tube or funnel; (3) plate-shaped; (4) raised rim; (5) invaginated structure; (6) projecting ventral lip.
(111)
Sterigma: (1) membranous; (2) weakly sclerotised; (3) strongly sclerotised.
(112)
Sterigma: (1) smooth (in stereo microscope); (2) at least partially spinulose, visible in stereo microscope.
(113)
Ductus bursae: (1) short, less than 56: length of corpus bursae; (2) moderately long, 56: length to as long as corpus bursae; (3) long, longer than length of corpus bursae.
(114)
Width of ductus bursae, except at entrance to corpus bursae: (1) narrow, at most 56E width of corpus bursae; (2) moderately wide, more than 56E and less than 56: width of corpus bursae; (3) wide, at least 56: width of corpus bursae.
(115)
Colliculum: (1) normal; (2) very long; (3) of thickened membrane; (4) very weakly developed; (5) absent; (6) modified.
(116)
Ductus bursae, whether sclerotised apart from colliculum and granulation: (1) entirely membranous; (2) at least partially sclerotised.
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PHYLOGENY OF THE OLETHREUTINAE
(117)
Shape of sclerotisation of ductus bursae: (1) unspecific; (2) ring of some sort; (3) an anteriorly bifid prong.
(118)
Ductus bursae: (1) not granulose; (2) lightly granulose; (3) conspicuously granulose.
(119)
Ductus seminalis originating from: (1) neck of corpus bursae; (2) anterior third; (3) central third; (4) posterior third.
(120)
Structure of neck of corpus bursae: (1) not obviously different from adjoining ductus bursae; (2) with conspicuous, asymmetrical patch of spinules; (3) with conspicuous spinules, scattered or as ring; (4) with sclerite, smooth or spinulose; (5) with paired patches of sclerotised scobination; (6) with small, membranous appendix.
(121)
Corpus bursae: (1) at most with minute spinules; (2) with large spinules; (3) with patch of very large, thorn-shaped spinules.
(122)
Signum: (1) present; (2) absent.
(123)
Signum position: (1) close to neck of corpus bursae, originating in posterior 56M; (2) not in posterior 56M of corpus bursae.
(124)
Signum number: (1) one; (2) two.
(125)
Signa: (1) equal-sized; (2) of different size.
(126)
Signum shape: (1) horn-, thorn- or nipple-shaped; (2) blade-shaped; (3) invaginated, small scobinate plate; (4) patch of enhanced scobination or granulation; (5) scobination with individual teeth enlarged.
Grapholitini and extended this for all Australian genera of the tribe. A cladistic analysis using WinClada (Nixon 1999) and NONA (Goloboff 1999) was run on a matrix (Appendix 1) of 126 morphological characters (Table 1) for 73 olethreutine genera and two outgroup taxa. The plesiomorphic tortricine Williella sauteri Horak from New Caledonia and Templemania animosana Busck (Anacrusini) from Mexico (coll. J. A. Powell) were used as outgroups. All Australian olethreutine genera were included except for the Grapholitini as work on their revision was not based on a DELTA matrix. Also, Komai (1999) had already demonstrated their monophyly and provided a phylogeny for the tribe. Grapholita zapyrana (Meyrick) was subsequently scored as a single exemplar species to include at least one representative of the tribe. All genera were scored to represent the character variation across the genus that resulted in numerous polymorphies. All multistate characters were treated as unordered. A heuristic search using multiple TBR + TBR (multi*max*) with 10 000 maximum trees retained, 1000 replications and 10 starting trees per replication, produced two trees with a length of 1283, consistency index of 21 and retention index of 47. The strict consensus tree is given as Table 2. Ordering, weighting or deletion of selected characters significantly changed the topography of the resulting trees but was not further pursued.
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17
In the consensus tree (Table 2), the entire tribe Olethreutini, apart from Sycacantha Diakonoff and Atriscripta, gen. nov., clusters on a single branch with both the Microcorsini and Bactrini included as subordinate and paraphyletic respectively polyphyletic taxa. The majority of both Eucosmini and Enarmoniini are each clustered as a monophyletic group. A basal grade below the Olethreutini contains the single grapholitine taxon (Grapholita zapyrana), two genera of the Enarmoniini (Loboschiza Diakonoff and Irianassa Diakonoff) and the only two isolated genera of the Olethreutini, Sycacantha and Atriscripta, gen. nov. A second grade between the Olethreutini and the terminal Eucosmini and Enarmoniini consists first of four enarmoniine genera (Anthozela Meyrick, Oriodryas Turner, Eucosmogastra Diakonoff and Pternidora Meyrick), followed by four eucosmine genera (Epiblema Hübner, Coenobiodes Kuznetzov, Eucosmophyes Diakonoff, Herpystis Meyrick). The eucosmine absconditana-group is the sister taxon to the enarmoniine clade. The Microcorsini appear as a paraphyletic group subordinate within the Olethreutini, with Demeijerella Diakonoff as the sister taxon to Collogenes Meyrick. The Bactrini also are subordinate within the Olethreutini, as well as polyphyletic with Lobesia Guenée between Endothenia Stephens and the sister taxa Bactra Stephens and Syntozyga Lower.
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OLETHREUTINE MOTHS OF AUSTRALIA
Table 2 Williella sauteri Horak Templemania animosana Busck Grapholita zapyrana Meyrick Irianassa Diakonoff Loboschiza Diakonoff Sycacantha Diakonoff Atriscripta gen. nov. Oxysemaphora Diakonoff Cnecidophora gen. nov. Cryptaspasma Walsingham Collogenes Meyrick Demeijerella Diakonoff Eremas Turner Endothenia Stephens Lobesia Guenee Bactra Stephens Syntozyga Lower Aterpia Guenee Proschistis Diakonoff Gnathmocerodes Diakonoff Rhectogonia Diakonoff Megalota Diakonoff Sorolopha Lower Podognatha Diakonoff Costosa Diakonoff Gatesclarkeana Diakonoff Zomariana Diakonoff Temnolopha Lower Ophiorrhabda Diakonoff Archilobesia Diakonoff Dudua Walker Trachyschistis Meyrick Euobraztsovia Diakonoff Rhodacra Diakonoff Diakonoffiana Kocak Dactylioglypha Diakonoff Metrioglypha Diakonoff Statherotis Meyrick Anthozela Meyrick Oriodryas Turner Eucosmogastra Diakonoff Pternidora Meyrick Epiblema Huebner Coenobiodes Kuznetsov Herpystis Meyrick absconditana-group Ancylophyes Diakonoff Balbidomaga Diakonoff Periphoeba Bradley Tetramoera Diakonoff Toonavora gen. n. Anathamna Meyrick Metaselena Diakonoff Cyphophanes Meyrick Helictophanes Meyrick Ancylis Huebner Thysanocrepis Diakonoff Aglaogonia gen. n. Pseudancylis gen. n. Crocidosema Zeller Noduliferola Kuznetsov Heleanna Clarke Whittenella gen. n. Acroclita Lederer Tritopterna Meyrick Melanodaedala gen. n. Fibuloides Kuznetsov Rhopobota Lederer Eccoptocera Walsingham Hermenias Meyrick Strepsicrates Meyrick Spilonota Stephens Holocola Meyrick Icelita Bradley Eucosmophyes Diakonoff
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PHYLOGENY OF THE OLETHREUTINAE
The failure of the analysis to recover several well accepted groupings based on unique apomorphies casts some doubt on the adequacy of the matrix. It was scored for all characters encountered within each genus as a very effective basis for descriptions and keys and to facilitate future work on the Papuan and Oriental fauna. However, it possibly contained too much polymorphism for a phylogenetic analysis, and time constraints did not allow rescoring a more suitable matrix based on exemplar taxa. The resulting phylogenetic hypothesis therefore has to be regarded as very preliminary and insufficiently conclusive to make changes to the higher classification. For example,
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several probable synapomorphies linking Bactra and Endothenia with some Olethreutini and the presence of a tibial pencil in Endothenia suggest that the highly derived Bactrini are merely subordinate within the Olethreutini rather than a separate tribe. The cladistic analysis indeed places the Bactrini within the Olethreutini but is considered too flawed to provide sufficient corroboration to sink a generally accepted tribe. A more rigorous cladistic analysis, reassessing some of the characters and their states, and including molecular information, may resolve this question in the near future.
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2. Morphology
Morphological structures are here discussed largely from a taxonomic angle, with emphasis on diagnostic characters including external features and the sclerotised parts of the genitalia. An indepth treatment of lepidopteran morphology is provided by Kristensen (2003) and a summary of tortricid morphology by Horak (1991).
Head (Figs 1–9) The head capsule of Olethreutinae, as of all Tortricidae, lacks prominent sutures, but there is usually a scale-free band between the antennal sockets, with the vertex above and the frontoclypeus (termed frons in the following) below. The tentorial pits (Fig. 6) are prominent on the denuded head. The tortricid head scaling is quite distinct, with anteriorly oriented scales on the vertex and upper frons, but upwardly oriented, small, appressed scales over the lower 36: of the frons (Figs 1–3). According to Zimmerman (1978), upwardly directed scales on the lower part of the frontoclypeus occur among microlepidoptera only in the Tortricidae, many Cossidae and some Tineidae. The anteriorly oriented scales on the vertex and upper frons are usually erect and long, rarely short and closely appressed. The compound eye is large and spherical, with a band of microtrichia along its margin. Only at very high magnification are short, sparse setae visible. The ocellus and chaetosema are usually well developed, adjacent to each other behind the antenna, next to the eye margin (Fig. 4). Ocellus size varies greatly, but at least a vestigial scar is always present. The chaetosema is a pin-cushionlike domed area with narrow, modified hair scales. The antenna is filiform, reaching to about the middle of the forewing or to slightly beyond. The scape and pedicel are unmodified, densely scaled, with a small intercalary sclerite between the two segments. The flagellum always has scales across its dorsal surface and usually an unscaled band of variable width along its ventral surface bearing the majority of sensilla. More rarely, the flagellum is scaled all round. Each olethreutine flagellar segment has a single ring of scales (Fig. 8), usually ventrally interrupted, whereas all other Tortricidae,
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except for some Spargonothini not present in Australia, have two rings of scales per flagellar segment (Fig. 7). The sensory setae, termed cilia in taxonomic language, vary particularly in the male from nearly invisible on the scaled antenna to as long as the diameter of the flagellum. In some genera of the tribe Eucosmini and the tortricine tribe Archipini the flagellum in the male may be notched at or near its base, with a deep dorsal excavation (Fig. 9) involving several flagellar segments. This secondary sexual modification has not only been developed at least twice within the family, it also has been lost again many times within genera. Olethreutine mouthparts (Figs 5, 6) conform to those of other higher Lepidoptera, with the maxillae joined to form a proboscis. The narrow, unscaled labrum has a lateral pair of pilifers. The basal portion of the maxilla, the cardo and stipes, bear the grooved galea and the maxillary palpus. The small olethreutine maxillary palpus (Fig. 5) is hardly visible on the fully scaled head and varies from four-segmented and scaled, to reduced to a minute, naked remnant. The two grooved galeae form a naked proboscis, entirely unscaled. The labial palpi are well developed, three-segmented, with an invaginated vom Rath’s organ subapically on the last segment (Fig. 6). Relatively long, horizontally held, sinuate labial palpi with the second segment sinuate and distally widened, and the relatively short terminal segment angled forward or downward (Figs 1, 2), is the most common configuration for olethreutines and the plesiomorphic condition for the family. More rarely, the labial palpi are upcurved, with a rather slender and curved second segment, and a short but porrect or apically rising third segment (Fig. 3). However, upcurved olethreutine labial palpi never have a long, drawn-out tip tapering to a point as in the horn-shaped gelechioid palpi.
Thorax and appendages (Figs 10–17) The thorax scaling may be smoothly appressed above or the scales may be raised to form a single or bipartite transverse crest posteriorly. Presence or absence of this raised scale tuft, the posterior crest,
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22
OLETHREUTINE MOTHS OF AUSTRALIA
Figs 1–9. Tortricid head structure. 1, Epiphyas postvittana (Tortricinae). 2, Crocidosema plebejana (Olethreutinae) (arrow: maxillary palpus). 3, Cydia pomonella (Olethreutinae). 4, ocellus (oc) and chaetosema (ch). 5, pilifer (pi) and maxillary palpus (mp). 6, descaled head with mouthparts, tentorial pits (tp) and vom Rath’s organ (vR). 7, tortricine antennal flagellum (E. postvittana). 8, olethreutine antennal flagellum (Dudua aprobola). 9, flagellum with antennal notch (arrow) (Strepsicrates semicanella).
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MORPHOLOGY
is a taxonomically informative character, but it is also a very ephemeral structure easily lost without the damage being apparent. The tegulae (Fig. 10), curved, crescent-shaped plates pertaining to the mesothorax and covering the bases of the forewings, have a scale tuft from their downcurved anterior corner (termed anteroventral corner in descriptions) that may be lengthened to form a long scale pencil, extending posteriorly beneath the wing bases. In the Olethreutinae the legs are well developed and typical of ditrysian Lepidoptera, with the fore tibia with an epiphysis with bristles along its inner surface, the mid tibia with a pair of apical spurs, and the hind tibia with two pairs of spurs, one medial and one apical. The legs are densely scaled, with the scaling on the mid tibia modified with a spiny appearance, due to parallel, grooved, pointed scales of decreasing size packed into sheaves, each enveloped by the outermost, largest scale. Several olethreutine groups have a modified hind tibia in the male, often with a narrow, long, eversible scale pencil from the base of the inner surface (Figs 15, 16), and sometimes with large lateral scale fringes and/or various large clumps of modified scales along the inner surface (Fig. 17). The fivesegmented tarsus usually has spines near the apex of segments 1–4, reduced in some groups. Olethreutine forewing shape (Figs 10–12) ranges from triangular to subrectangular to subovate, and from rather wide to quite narrow, with the hindwings roughly as broad as the forewings. The forewing index (the maximum width of forewing divided by its maximum length) quantifies the wing ratio. The termen varies from nearly at right angles to the costa to strongly oblique and may be more or less straight or variably sinuate with an indentation beneath the apex. In some enarmoniine genera the apex is strongly projecting and followed by a deep notch, producing a so-called ‘falcate’ apex (Figs 474, 475). The hindwing shape is sometimes sexually dimorphic, usually with an expanded anal area in the male, often folded or rolled along the anal margin (Fig. 13). The forewing shape is sexually dimorphic only in a few cases (Lobesia, Archilobesia), but the base of the costa is expanded in the males of some groups and folded over the wing as a so-called costal fold (Figs 10, 12), usually enclosing modified scales. Wing coupling is effected by frenular bristles from the base of the hindwing fitting into the retinaculum beneath the forewing. Females have three (Fig. 14) (rarely two) frenular bristles, males a single composite one (Fig. 13). The retinaculum is a membranous hook arising
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from the subcosta (Sc) or a spur of Sc in males and a row of erect scales behind the base of the cubitus (CuA) in most females. The forewing scales are usually smoothly appressed, but raised scale tufts occur in several groups and can be of taxonomic significance. The olethreutine hindwing nearly always has a cubital pecten (Fig. 14), a row of hair scales on or just behind the base of CuA, sometimes modified (Crocidosema plebejana Zeller, Fig. 76), in contrast to nearly all other tortricids that lack a cubital pecten. The hindwing scaling is variously modified in the male, particularly along the anal margin, but also including very small and widely spaced scales resulting in a semi-translucent wing (Fig. 667), fields of melanic scales above and/or beneath (Fig. 48), and scale pencils in diverse positions (Figs 13, 25–27, 39). The olethreutine forewing pattern (Figs 10, 11) is derived from several basic elements, variously developed and modified. Several dark, parallel, outwardly oblique, transverse bands and derivations thereof constitute one of the basic pattern elements, present in all tortricid subfamilies but with various interpretations of their number (Brown and Powell 1991; Baixeras 2002; Razowski 2003b). Traditionally, these so-called transverse fasciae have been referred to by specific names, but as homology of a given pattern element with the relevant fascia is often unclear and there is no consensus nomenclature, no fascia names are used here. Costal strigulae (pairs of short transverse white marks along the costa) constitute the second basic pattern element, also found in all three tortricid subfamilies. Pairs of costal strigulae frame the five dark fasciae (Brown and Powell 1991; Baixeras 2002), but as some of the strigulae are usually obliterated, especially in the basal half, doubled or confluent, no attempt at numbering has been made here. Strictly speaking, the white markings are the strigulae, but on pale wings the dark spacers in between are often more conspicuous and for ease of descriptions have been referred to as dark strigulae. A complex, often metallic marking in the tornus consisting of several parallel, longitudinal, black dashes framed by an erect silvery band on each side if fully developed, is variably termed ocellus, speculum or ocelloid patch (Fig. 11), the term here used to differentiate it from the ocellus proper. Olethreutine wing venation is typically heteroneurous with a reduced number of radius branches (R-branches) in the hindwing, but otherwise only little modified in the generalised condition. The vein nomenclature adopted here
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OLETHREUTINE MOTHS OF AUSTRALIA
Figs 10–13. Olethreutine adult males and wing venation (ap, apex; cf, costal fold; ch, chorda; cs, costal strigulae; cv, closing vein; el, expanded anal lobe; fb, frenulum bristle, male; Ms, M-stem; op, ocelloid patch; sp, scale pencil; tg, tegulae; to, tornus). 10, Spilonota sp. 11, Grapholita zapyrana. 12, forewing, Spilonota constrictana, male. 13, hindwing, Holocola zopherana, male, with single frenulum bristle, M3 and CuA1 fused into single vein and 3A absent.
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MORPHOLOGY
Fig. 14. Olethreutine female hindwing with cubital pecten (cp) and three frenulum bristles (fb).
conforms to that of Common (1990), Nielsen and Common (1991) and the earlier volumes of the series ‘Monographs on Australian Lepidoptera’, with the radial sectors numbered as R2, R3, R4 and R5 rather than the phylogenetically correct Rs1, Rs2, Rs3 and Rs4. The underived forewing has a full complement of free veins, with the discal cell apically closed by the closing vein (discocellular vein) and with the media stem (M-stem) and the stem of R4 and R5 (chorda) well developed. If present, the chorda delineates a small accessory cell (Fig. 12), but M-stem and/or chorda may be reduced or absent. R4 usually runs to the costa and R5 to the termen, but rarely R5 ends at or before the apex. R4 and R5 are stalked in several groups, R3 is often approximated to R4 but only rarely are the two veins stalked or R3, R4 and R5 all borne on the one stalk. Males of Lobesia Guenée often have a pterostigma (Fig. 31) and several of the R branches may be modified in Metaselena. Presence of three free M branches and two free branches of the cubitus anterior (CuA branches) is the underived condition, with the relative position of M2 and M3 of taxonomic significance, varying from widely distant and parallel at base to having their bases closely approximated and M2 curved towards M3. In Aglaogonia, gen. nov. and Ancylophyes Diakonoff (both Enarmoniini) one forewing vein is missing. With very few exceptions the cubitus posterior (CuP) is present near the wing margin, but never as a fully developed vein. The anal
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veins 1A and 2A are fused distally, forming a basal loop. The underived olethreutine hindwing also has a full complement of free veins, but the M-stem is never developed and the cell is closed only by vestigial cross-veins. The subcosta is fused with the first radius branch (Sc+R1), with R1 often visible as a cross vein between the bases of Sc and the single radial sector (Rs). The three M branches and the two CuA branches are free from the discal cell in the generalised olethreutine hindwing, with some of these veins becoming approximated, stalked or even fused in more derived wings. With few exceptions Rs and M1 are closely approximated and parallel at their base (Fig. 13) or the two veins are stalked, and the rare cases where these two veins are somewhat distant right from their bases could possibly denote the ancestral character state (Horak 1984). Whether M3 and CuA1 are stalked or not has long been regarded as the crucial difference between Eucosmini and Olethreutini, and despite several exceptions this rule still holds to a surprising extent, especially if one considers the
Figs 15–17. Olethreutine male hind tibia. 15, tibial pencil (tp) hidden in scaling. 16, exposed tibial pencil. 17, modified hind tibia of Cryptophlebia ombrodelta.
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OLETHREUTINE MOTHS OF AUSTRALIA
least derived members of each genus. The cubitus posterior and all anal veins, the distally fused 1A+2A and 3A, are all present in the underived olethreutine hindwing, though often not as tubular veins. Presence of 3A in nearly all genera except the majority of Eucosmini seems to be taxonomically informative.
Pregenital abdomen The olethreutine pregenital abdomen comprises eight segments in the male and seven segments in the female, though in more derived groups the ostium often is fused with or located within the accordingly modified seventh sternite. The sternum of the first segment (S1) is lacking and the configuration of S2 is of the tortricoid type with well-developed ventral apodemes (Figs 224, 465) and at most weak remnants of sternal rods (venulae). The anterolateral processes are usually well developed (Figs 465, 944), but absent in some genera (Fig. 977). A pair of round dorsal depressions, often called dorsal pits (Figs 312, 314), are present on T2 in several genera of the Olethreutini, and also occurs in at least two tortricine groups. The structure of the eighth segment in the male, especially the shape of its hind margin, is taxonomically relevant in some groups. Olethreutine males are characterised by various secondary sexual modifications of the pregenital abdomen and its scale cover. These structures are often part of a functional unit involving the legs or scale pencils from the thorax or hindwing, and if they are present they may be taxonomically informative. Modified scaling on S2 is found in shallow folds laterally in Sycacantha Diakonoff and Lobesia (Fig. 222), and medially either as simple patches (Metaselena, Fig. 580) or complex tufts. Eversible deep pockets containing modified scales occur usually laterally on the abdomen between two segments (some Lobesia, Proschistis Meyrick, Pammenopsis Kuznetsov (Fig. 942)). Oxysemaphora males have longitudinal lateral folds with modified scales to hold the scale pencil from the hindwing anal margin. Males of some genera have lateral and/ or dorsal brushes of modified scales, often long and mane-like (Sorolopha Lower, Cryptophlebia Walsingham). The complex coremata of the eighth segment in the Grapholitini (Fig. 948) are discussed there.
Male genitalia (Figs 18–20) Olethreutine male genitalia are unmistakable, their fused juxta, caulis and anellus complex
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immediately identifies their subfamily position and provides a convincing derived character (apomorphy) indicating that all Olethreutinae must have had a single common ancestor with exactly this structure. In other words, this unique modification strongly supports the monophyly of the Olethreutinae and, without exception, is a diagnostic feature for the entire subfamily. In the ground plan for the family, i.e. the condition hypothesized for an ancestral tortricid, the valvae and the teguminal appendages would all have been well developed and involved in clasping the female abdomen during copulation. The functional unit of the fused juxta, caulis and anellus complex, in conjunction with strongly sclerotised, rigid valvae, obviously allowed reduction of the teguminal appendages that have been reduced and lost repeatedly in many olethreutine groups. Whilst the male genitalia provide excellent diagnostic characters at the species and generic levels, parallel reduction of the teguminal appendages can result in superficially very similar structures erroneously suggesting a closer relationship. The dorsal tegumen (tergum 9) and ventral vinculum (sternum 9) are laterally articulated to form a ring from which arise the various, mostly paired processes of the genitalia. Modification of the tegumen usually goes hand in hand with reduction of some of its appendages, in extreme cases resulting in a simple wide band. The two lateral, basalmost portions of the tegumen, the socalled pedunculi, frequently have medial apodemes, attachments for the tergal flexor muscle of the valva (m4 in Kuznetsov and Stekolnikov 1973, 1977) (Figs 219, 478, 480, 482). The vinculum always is a more or less narrow band in the Olethreutinae. The full complement of teguminal appendages in the Olethreutinae, as in all lower Ditrysia, is a paired uncus, a pair of hairy socii and paired gnathos arms (Figs 19, 20). There is little doubt that the ancestral (plesiomorphic) olethreutine uncus was a paired structure that has become fused into a single process, variously modified and/or reduced repeatedly, even within genera. The membranous, hairy, plesiomorphic olethreutine socii lobes are greatly modified in many groups, with a particular derived character state often diagnostic: like the long, slender, arm-like socii of Sorolopha, the bifid socii of Sycacantha, or the rigid, strongly sclerotised socii of Spilonota and related genera. On the other hand, the socii have been reduced repeatedly to small setose patches or entirely lost in many Grapholitini. Relatively few olethreutines have welldeveloped, strongly sclerotised gnathos arms, either joined at the tip (Aterpia Guenée) or upcurved, and
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MORPHOLOGY
fused with the ventral wall of the anal tube (Fig. 20), forming a subscaphium. Often the gnathos is reduced, with only lateral remnants contiguous with the posterior tegumen margin. The valvae, paired clasping organs, are strongly sclerotised in the Olethreutinae, with a large membranous portion at the base of their inner surface to allow muscle movement and somewhat misleadingly termed ‘basal excavation’. The valvae are articulated with the vinculum and the base of the pedunculi, and their dorsal margin (costa) proximally ends in a muscle attachment, the costal process (basal process (Komai 1999), costal hook). Numerous names have traditionally been used for various parts of the lepidopteran valva, with little consistency across the groups. These terms are here restricted to sacculus (the reinforced basal portion of the ventral margin often ending in a free process) and the cucullus (the distal part of the valva usually delineated by a ventral emargination). Additional lobes and processes of more derived valvae are simply described rather than referred to with terms such as pulvinus etc. that imply homology. Olethreutine valvae are often characterised by prominent spines, single or in groups, with an associated nomenclature. Attempts to establish the homology of the various spine groups across tribal boundaries were unsuccessful, and descriptive terms were chosen instead. The diaphragm, the membrane closing the body cavity, extends between the tegumen and the vinculum and is perforated by the aedeagus and the anus. The aedeagus is supported from below by sclerotised portions of the diaphragm, the basal juxta fused with the caulis and the anellus that surrounds the aedeagus. In olethreutines, there are only rarely sclerotisations of the diaphragm above the aedeagus other than along the ventral surface of the anal tube, and a transtilla is never present. The olethreutine aedeagus is tube-shaped, and the membranous, eversible inner vesica often bears spines (cornuti) that may be shed in the bursa during copulation. A large group of deciduous cornuti, whose sockets are visible even if they have been shed, seems to be the ancestral condition. Progressive reduction of cornuti numbers and/or modification of at least some into fixed cornuti are both derived conditions.
Female genitalia (Figs 21, 22) The female reproductive system of the Olethreutinae is of the ditrysian type with a separate copulatory opening (ostium bursae) in addition to the ovipore for oviposition. The ovipore
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27
and anus are close together between the typically tortricid, broad, flattened, and usually hairy ovipositor lobes that are a diagnostic feature for the family Tortricidae, and usually allow sexing of totricid moths in a reliable manner. Rarely, the ovipositor lobes are modified: either reduced in a telescopic ovipositor (Figs 492, 626) or with conspicuously enlarged papillae in Collogenes Meyrick (Fig. 144). The ovipositor lobes and the 8th tergite have paired long apodemes as muscle attachments (the apophyses posteriores and apophyses anteriores), further lengthened if the ovipositor is telescopic. The ostium bursae is located in the 8th sternite (S8) in generalised olethreutines (Fig. 21) but is displaced anteriorly in several groups, ending up either in a deep excavation of the hindmargin of S7 (Fig. 22), invaginated behind its hindmargin or located within S7. In its original position in the ventral membranous portion of the eighth segment, the entrance to the ostium is usually surrounded by some sort of sclerotisation, referred to as the ‘sterigma’. The posterior portion of the sterigma behind the ostium is also referred to as ‘lamella postvaginalis’ and that anterior to it as ‘lamella antevaginalis’ — practical but somewhat incorrect terms as the aperture concerned is strictly the copulatory orifice and not the vagina. In tortricids of the other two subfamilies, the sterigma is connected to the base of the apophyses anteriores by a well-sclerotised band, a connection absent in all olethreutines (Figs 21, 22) except the Microcorsini (Figs 132, 144). Absence of this connection is considered the derived condition, with the Microcorsini sharing the ancestral condition with the rest of the family. The bursa copulatrix is usually differentiated into a narrow ductus bursae and the corpus bursae proper. In generalised olethreutines the corpus bursae has two signa that are often reduced to one or entirely lost. The ductus seminalis connects the ductus bursae with the vagina and nearly always has a diverticulum (bulla seminalis). The structure of the female genitalia, i.e. the bursa copulatrix and associated structures, often provides more pertinent information on higherlevel relationships than the male genitalia that are most informative at the generic and specific levels. Frequently, portions of the ductus bursae are sclerotised and may be taxonomically informative. A small and at least ventrally split ring just below the ostium, the colliculum (Fig. 21), seems part of the tortricid groundplan and is variously modified, lengthened or reduced in the olethreutines. Several enarmoniine genera are characterised by a sclerite
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OLETHREUTINE MOTHS OF AUSTRALIA
in the bursa ‘neck’, the transition between the corpus bursae and the ductus bursae (Fig. 470). The structure of the signa, if they are present, is highly diagnostic, and Diakonoff (1973) very successfully subdivided the Olethreutini on the basis of signum shape. However, the signa are frequently reduced from two to one or completely lost, even within a
genus. In some Eucosmini the bursa wall is strongly spinulose, except near the signa, and parallel with these spinules becoming larger, the signa seem to decrease in size and eventually to disappear (Figs 754–756). Deciduous cornuti retained in the corpus bursae may provide valuable information if all available males have shed their cornuti.
Figs 18–20. Olethreutine male genitalia. 18, Eucosmophyes (ae, aedeagus; an, anellus; be, basal excavation; ca, caulis; co, cornuti; cu, cucullus; ju, juxta; so, socii; vi, vinculum). 19, Cnecidophora; cp, costal process; ca, caulis; gn, gnathos; so, socii; un, uncus). 20, Fibuloides (an, anellus; cu, cucullus; gn, gnathos; pe, pedunculus; pun, paired uncus; sa, sacculus).
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Figs 21–22. Olethreutine female genitalia. 21, Ophiorrhabda (bn, bursa ‘neck’; co, colliculum; cb, corpus bursae; db, ductus bursae; la, lamella antevaginalis; lp, lamella postvaginalis; si, signum). 22, Fulcrifera (aa, apophyses anteriores; ap, apophyses posteriores; bs, bulla seminalis; di, diverticulum; ds, ductus seminalis; lp, lamella postvaginalis; os, ostium).
Figs 23–24. Microcorsini, wing venation, males. 23, Cryptaspasma sordida, slide ANIC 2747. 24, Collogenes loricata, slide ANIC 13768.
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OLETHREUTINE MOTHS OF AUSTRALIA
Figs 25–27. Olethreutini, wing venation, males. 25, Oxysemaphora hacobiani, slide ANIC 1224. 26, Cnecidophora ochroptila, slide ANIC 1228. 27, Sorolopha cyclotoma, slide ANIC 1280.
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31
Figs 28–33. Olethreutini, wing venation, males. 28, Sycacantha exedra, slide ANIC 1265. 29, Atriscripta arithmetica, slide ANIC 1267. 30, Demeijerella chrysoplea, slide ANIC 1266. 31, Lobesia physophora (pt, pterostigma), slide ANIC 1260. 32, Gnathmocerodes euplectra, slide ANIC 2717. 33, Rhectogonia electrosema, slide ANIC 1285.
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OLETHREUTINE MOTHS OF AUSTRALIA
Figs 34–39. Olethreutini, wing venation, males. 34, Ophiorrhabda phaeosigma, slide ANIC 2699. 35, Dudua phyllanthana, slide ANIC 1256. 36, Trachyschistis hians, slide ANIC 1244. 37, Archilobesia drymoptila, slide ANIC 1257. 38, Euobraztsovia chionodelta, slide ANIC 1280. 39, Eremas leucotrigona, slide ANIC 1269.
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33
Figs 40–45. Olethreutini, wing venation, males. 40, Zomariana doxasticana, slide ANIC 17119. 41, Podognatha vinculata, slide ANIC 1281. 42, Temnolopha mosaica, slide ANIC 1259. 43, Diakonoffiana tricolorana, slide ANIC 1262. 44, Megalota uncimacula, slide ANIC 1264. 45, Costosa australis, slide ANIC 1270.
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OLETHREUTINE MOTHS OF AUSTRALIA
Figs 46–51. Olethreutini, wing venation, males. 46, Metrioglypha phyllodes, slide ANIC 1272. 47, Dactylioglypha tonica, slide ANIC 2467. 48, Statherotis sp., slide ANIC 1276. 49, Aterpia protosema, slide ANIC 17115. 50, Proschistis polyochtha, slide ANIC 2700. 5l, Rhodacra pyrrhocrossa, slide ANIC 17090.
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Figs 52–55. Olethreutini and Bactrini, wing venation, males. 52, Gatesclarkeana sp. (Papua New Guinea), slide ANIC 2698. 53, Bactra venosana, slide ANIC 1252. 54, Syntozyga anconia, slide ANIC 1249. 55, Endothenia polymetalla, slide ANIC 3126.
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OLETHREUTINE MOTHS OF AUSTRALIA
Figs 56–61. Enarmoniini, wing venation, males. 56, Aglaogonia sp., slide ANIC 12960. 57, Anathamna sp., slide ANIC 3125. 58, Ancylis anguillana, slide ANIC 12961. 59, Ancylophyes monochroa, slide ANIC 1268. 60, Anthozela hemidoxa (Papua New Guinea), slide BM 24920. 61, Balbidomaga uptoni, slide ANIC 1274.
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Figs 62–67. Enarmoniini, wing venation, males except 66. 62, Cyphophanes sp., slide ANIC 1240. 63, Eucosmogastra pyrrhopa, slide ANIC 2748. 64, Helictophanes uberana, slide ANIC 1273. 65, Helictophanes prospera, slide ANIC 1219. 66, Irianassa sp., female, slide ANIC 13810. 67, Loboschiza koenigiana, slide ANIC 1271.
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OLETHREUTINE MOTHS OF AUSTRALIA
Figs 68–73. Enarmoniini, wing venation, males except 71. 68, Metaselena lepta, slide GP 550. 69, Oriodryas olbophora, slide ANIC 1235. 70, Periphoeba trepida, slide ANIC 1295. 71, Pseudancylis acrogypsa, female, slide ANIC 17120. 72, Pternidora sp., slide ANIC 12931. 73, Tetramoera gracilistria, slide ANIC 1229.
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39
Figs 74–75. Enarmoniini, wing venation, males. 74, Thysanocrepis crossota, slide ANIC 17116. 75, Toonavora aellaea, slide ANIC 1251.
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40
OLETHREUTINE MOTHS OF AUSTRALIA
Figs 76–81. Eucosmini, wing venation, males. 76, Crocidosema plebejana, slide ANIC 1209. 77, Epiblema strenuana, slide ANIC 13767. 78, Epinotia absconditana, slide ANIC 1263. 79, Coenobiodes melanocosma, slide ANIC 1247. 80, Melanodaedala scopulosana, slide ANIC 1286. 81, Rhopobota honesta, slide ANIC 1254.
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MORPHOLOGY
41
Figs 82–86. Eucosmini, wing venation, males except additional female hindwing 83 and 86. 82, Fibuloides minuta, slide ANIC 17103. 83, Acroclita bryopa, slide ANIC 1257, slide ANIC 18806 (female hindwing). 84, Noduliferola neothela, slide ANIC 8664. 85, Heleanna chloreis, slide ANIC 17114. 86, Tritopterna capyra, slide ANIC 1279, slide ANIC 18807 (female hindwing).
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42
OLETHREUTINE MOTHS OF AUSTRALIA
Figs 87–93. Eucosmini, wing venation, males. 87, Spilonota constrictana, slide ANIC 1237. 88, Strepsicrates semicanella, slide ANIC 1218. 89, Holocola nr thalassinana, slide ANIC 1245. 90, Holocola deloschema, slide ANIC 1246. 91, Holocola zopherana, slide ANIC 1248. 92, Hermenias sp., slide ANIC 1231. 93, Eccoptocera australis, slide ANIC 1253.
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43
Figs 94–97. Eucosmini, wing venation. 94, Herpystis sp., female, slide ANIC 13799. 95, Icelita indentata, male, slide ANIC 13785. 96, Eucosmophyes commoni, male, slide ANIC 13786. 97, Whittenella peltosema, female, slide ANIC 13798.
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44
OLETHREUTINE MOTHS OF AUSTRALIA
Figs 98–102. Grapholitini, wing venation, males except additional female hindwing 99 and 102. 98, Loranthacydia nr sinapichroa, slide ANIC 4942. 99, Notocydia nr niveimacula, slide ANIC 4920, slide ANIC 4921 (female hindwing). 100, Pammenemima tetramita, slide ANIC 8622. 101, Leguminivora sp., slide ANIC 19285. 102, Notocydia nr atripunctis, slide ANIC 4922, slide ANIC 4923 (female hindwing).
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MORPHOLOGY
45
Figs 103–105. Grapholitini, wing venation, male fore- and hindwing, female hindwing. 103, Fulcrifera sp., slide ANIC 4929, slide ANIC 4933 (female hindwing). 104, Cydia pomonella, slide ANIC 4934, slide ANIC 4935 (female hindwing). 105, Apocydia pervicax, slide ANIC 4926, slide ANIC 4924 (female hindwing).
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46
OLETHREUTINE MOTHS OF AUSTRALIA
Figs 106–112. Grapholitini, wing venation, males. 106, Thaumatotibia zophophanes, slide ANIC 8605. 107, Thaumatotibia sp., hindwing only, slide ANIC 8607. 108, Cryptophlebia sp., slide ANIC 17026. 109, Gymnandrosoma gonomela, syntype [SAMA]. 110, Cryptophlebia ombrodelta, slide ANIC 8603. 111, Archiphlebia endophaga, slide ANIC 17010. 112, Archiphlebia rutilescens, slide ANIC 8608.
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47
Figs 113–118. Grapholitini, wing venation, males except additional female hindwing of 117. 113, Acanthoclita sp. C, slide ANIC 8610. 114, Grapholita zapyrana, slide ANIC 8625. 115, Acanthoclita sp. B, slide ANIC 8612. 116, Microsarotis sp. (Thailand), slide F Komai 1120. 117, Pammenopsis barbata, slide ANIC 4925, slide ANIC 4936 (female hindwing). 118, Commoneria cyanosticha, slide ANIC 8635.
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OLETHREUTINE MOTHS OF AUSTRALIA
Figs 119–121. Grapholitini, wing venation. 119, Parapammene sp. A, male, slide ANIC 4941. 120, Parapammene sp. A, female, slide ANIC 4943. 121, Ixonympha sp., male fore- and hindwing, slide ANIC 4938; female hindwing, slide ANIC 4940.
Fig. 122. Posterior crest (pc), dorsally on thorax, Aterpia protosema.
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3. Biology Life histories The Olethreutinae have by far their greatest diversity in the warmer and wetter parts of Australia, where rain or monsoon forests are their dominant habitats. Interpretation of label data in collections suggests that the great majority of species have several generations per year, especially in the moist tropics and subtropics. Species with a wide distribution and clearly several generations in the north are nevertheless rarely collected during winter in the temperate region, suggesting a slowing of their development during the cooler months. However, there may well be univoltine species in cooler and drier regions, especially among the Eucosmini in temperate southern latitudes or higher altitudes and possibly among the Grapholitini in seasonally dry habitats. No work has been done to establish whether any of the Australian Olethreutinae have a strict diapause to accomodate a seasonal food supply, such as availability of suitable fruit or seeds for some of the borers. The flat tortricid ovipositor lobes are characteristic for the family and possibly an adaptation for a specific egg-laying mode. Nearly all tortricid eggs are round to oval, strongly flattened, dome-shaped to scale-like. In contrast to the Australian Tortricinae (Powell and Common 1985), there is very little information about olethreutine oviposition. Olethreutine eggs are usually deposited singly on the host plant, rarely in small, irregular clusters (Peterson 1965; van der Geest and Evenhuis 1991). In some species and genera with modified, telescopic ovipositors, the eggs are inserted into crevices in the host plant. None of the Australian species have ovipositors that would suggest a piercing mode of oviposition. Feeding habits within the Olethreutinae are very diverse, ranging from leaf rolling and spinning foliage together to boring in shoots, fruit, seeds, stems, roots and woody branches. Some produce blister mines in leaves. The larvae of a few Australian species including Spilonota constrictana (Meyrick) live in a portable case made of hollowed flower buds or seed capsules spun together to form a tube (Fig. 730). Unlike some Tortricinae, none of the Australian Olethreutinae have been reared from dead plant tissue, i.e. leaf litter.
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The ancestral tortricid larva is presumed to have been an external feeder, given the presence of an anal fork in all three tortricid subfamilies (Horak 1989). This structure serves to eject faeces from the feeding shelter and has frequently been lost in groups with internally feeding (i.e. boring) larvae. Had an internally feeding larva without the anal fork been ancestral, it would be very unlikely that later derived external feeders in all three subfamilies would have developed exactly the same type of anal fork. A considerable proportion of Australian olethreutine larvae are borers, often in seeds and fruit: one of the main reasons for their economic importance. However, both feeding modes, leaf rolling and boring, are found in several olethreutine tribes, and parallel shifts from one mode to the other have obviously occurred. It had been hoped that mapping internal and external feeding onto the cladogram resulting from this study would provide an indication of the shifts in feeding modes, but the cladistic analysis turned out to be too flawed to expect critical insights in this respect. However, the trends within tribes strongly suggest that: 1, external feeding is the more ancestral state within those tribes using both feeding modes, except possibly for the Grapholitini, and 2, switches between the two feeding modes have happened repeatedly, even within a tribe. In both the Microcorsini and Bactrini all larvae are borers, the former endosperm feeders in fruit and nuts and the latter predominantly in stems, often of monocotyledons. The known larvae of the Australian Olethreutini are nearly all leaf rollers or leaf tiers except for some species within Lobesia Guenée, Ophiorrhabda Diakonoff, Gnathmocerodes Diakonoff, and Gatesclarkeana Diakonoff, genera from four different genus-groups. The feeding mode is known for only a few enarmoniine genera, but it ranges from leaf rolling to fruit boring, and in Loboschiza koenigiana (Fabricius) both feeding modes occur in the same species. The larvae of Ancylis Hübner, probably the sister group to the remaining Enarmoniini, are usually leaf rollers, but one Australian species produces blotch mines in Banksia leaves. The Eucosmini also include external as well as internal feeders, with the more ancestral groups (Epinotia Hübner) predominantly leaf
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OLETHREUTINE MOTHS OF AUSTRALIA
rolling, but with some borers even within Epinotia. The majority of Grapholitini larvae are internal feeders and, according to Komai’s (1999) hypothesis, this was the ancestral feeding mode of the Grapholita-group, with several subsequent reversals to external feeding. Almost all species of the remaining grapholitine genus-groups are borers as far as their biology is known. The Loranthacydiagroup, the Dichrorampha-group (except for Pammenemima Diakonoff with leaf rolling larvae) and most species of the Cydia-group, in fact, all species with known biology in Australia, are borers. This poses the question as to whether internal feeding was the ancestral feeding mode for the entire tribe Grapholitini, though some species such as Grapholita molesta (Busck) have retained a welldeveloped anal fork. Exceedingly little information is available on the biology of the pupal stage of Australian Olethreutinae. Most leaf rolling larvae pupate in their shelter, whereas boring larvae may pupate in the ground. Olethreutine pupae have rows of dorsal spines on the mobile abdominal segments and they partially extrude themselves from their cocoon before the adult emerges. Adult olethreutines are mostly active early in the evening and at night, and they are successfully attracted to light. However, some Grapholitini fly during the day and may not readily come to light.
Larval host plants For a worldwide study of larval food preferences of microlepidoptera, Powell (1980) analysed host plant usage of 540 tortricine and 841 olethreutine species. The data were mainly drawn from literature but included a list of food plants for the Australian fauna by I. F. B. Common, representing c. 950 Lepidoptera species. Even though the Holarctic region was heavily overrepresented in this sample, it is a useful departure point for looking at host plants of Australian Olethreutinae. Powell (1980) found that 24% of tortricine and only 6% of olethreutine species were polyphagous, i.e. defined as feeding on three or more plant families in two or more orders. The total number of host plant families recorded was high for both subfamilies, 81 for the Tortricinae and 86 for the Olethreutinae. Looking for host plant preferences at the family level within the Lepidoptera, Powell (1980) observed the failure of the tortricid subfamilies to specialise markedly on any plant family.
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Known host plant families for the Australian olethreutine genera have been tabulated by genus (Appendix 2), based on literature references (see Biology for the relevant genera) and rearing records at the ANIC. Olethreutine species have been reared from 48 plant families in Australia, confirming the wide range of olethreutine host plants reported by Powell (1980). Host plant information for at least one species is available for 75% of the genera, either from Australia or the Oriental region, but often it is only a single record and thus not informative with regard to mono- or polyphagy. However, enough species have been reared repeatedly from the same plant genus or family to assume that a sizable proportion of Australian olethreutine genera use a single host plant family exclusively or at least predominantly. And in contrast to the observed absence of host plant specialisation of the Tortricidae at the family and subfamily levels, the picture changes as one looks at tribes, genus-groups and genera. Several groups of Australian Olethreutinae, at various levels, feed on plants of the same family or order. The Microcorsini are characterised by their feeding mode rather than their host plant choice. The larvae of both genera are endosperm borers, the polyphagous Cryptaspasma Walsingham reported from seven families in six orders, including Lauraceae, but Collogenes Meyrick only from Lauraceae, based on a single record. Several species and genera in the Olethreutini are known to be polyphagous, among them Sorolopha Lower, Lobesia Guenée, Zomariana Diakonoff, Gatesclarkeana Diakonoff, Statherotis Meyrick and Aterpia Guenée, the last based on host data from the Holarctic fauna. The two Australian genera of the Oxysemaphora-group have both been reared from Myrsinaceae, Oxysemaphora Diakonoff also from Rutaceae. Dudua Walker is almost exclusively confined to Glochidion (Euphorbiaceae), except for the polyphagous Dudua aprobola (Meyrick). Glochidion is also the suspected host plant for the closely related Archilobesia Diakonoff. The three Bactrini genera with known biology are all borers, Bactra Stephens in Cyperaceae, Juncaceae and Poaceae, Syntozyga Lower in Commelinaceae, and Endothenia Stephens polyphagous in various non-monocots. Host plants are known for only half the enarmoniine genera in Australia, partly because most species have a restricted distribution and are often rare. A prevalence of borers is probably another reason for the scarcity of host records as out of eight genera with biological information
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BIOLOGY
available, three are known to be borers and in three more the larvae of some species are borers in certain situations, or during part of their development. The widely distributed Ancylis Hübner is highly polyphagous, elsewhere as well as in Australia, and for the other genera we have mostly just a single record. However, both species of Toonavora, gen. nov. have been reared from seeds of Toona (Meliaceae) only, and several species of Loboschiza Diakonoff have been reported from several meliaceous genera with one report each from Oleaceae and Malvaceae. Host plant records are available for all except three eucosmine genera present in Australia. Nearly all the species in the five genera of the Spilonotagroup feed on Myrtaceae, probably representing a rather recent radiation on this plant family in Australia. The largest genus, Holocola Meyrick, numbers 42 nominal and many unnamed species. Several of the remaining Australian eucosmine genera are polyphagous, especially Rhopobota Lederer, but also Heleanna Clarke, Herpystis Meyrick and Crocidosema Zeller, though a speciesgroup including Crocidosema plebejana Zeller seems to be restricted to the Malvaceae. The sister genera Acroclita Lederer and Tritopterna Meyrick have both been reared from host plants in the Euphorbiaceae, with Acroclita also feeding on Elaeagnaceae in Japan. With the exception of Apocydia, gen. nov., at least one host record is available for all grapholitine genera present in Australia. In Australia, even more so than elsewhere (Komai 1999), the Fabaceae, Caesalpiniaceae and Mimosaceae are the
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51
predominant host plant families of the Grapholitini. Nine out of the seventeen Australian genera with at least one host plant record have been reared predominantly or exclusively from these three plant families, and they also feature prominently for the very polyphagous Cryptophlebia Walsingham. Two endemic genera, Loranthacydia Horak, Common & Komai and Ixonympha, gen. nov., have been reared from Loranthaceae only, with very specialised biologies. According to Meyrick (1911b), F. P. Dodd reared two species of Loranthacydia from larvae boring in stems of mistletoes, and one species of Ixonympha, gen. nov. was reared from a deployed seed of the mistletoe Amyema quandang (Reid 1984). The very specialised feeding behaviour of Ixonympha, gen. nov., and the fact that 7–36% of deployed mistletoe seeds and seedlings were destroyed by its larvae (Reid 1987), point to a well-honed adaptation to this unusual niche, reflected in the very small size of the (at least) three Ixonympha, gen. nov. species. Both Loranthacydia and Ixonympha, gen. nov. have been collected from climatically very diverse localities scattered widely across Australia, and the assumption that both genera are linked with Loranthaceae could explain their broad climatic range as well as their often single occurrence, especially in the case of the very rare Ixonympha, gen. nov. The two genera belong to very different grapholitine clades and the move onto Loranthaceae must have happened twice independently, with the relatively high species diversity in both genera suggesting subsequent radiations on this unusual host family.
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4. Diversity and distribution
The diversity and distribution of the Australian Olethreutinae present a diametrically opposite picture to those of the Oecophorinae discussed by Common (1994). Whilst Australia has a relatively high count of olethreutine genera, they are heavily concentrated in the north-eastern part of the continent and their relationships are with the Oriental fauna. Furthermore, despite the high generic diversity, only a small number of olethreutine genera appear to be endemic to Australia. A comparison of Australian olethreutine diversity with other geographic areas (Table 3) shows that at the generic level Australia has a diverse olethreutine fauna compared with those of Europe, the Nearctic or the Neotropical regions. However, the picture changes if any fauna containing a sizeable Oriental element is considered, even if the number of species known indicates that the figures reported can only be a preliminary tally. For a total of only 135 species, 69 olethreutine genera are reported from Taiwan (Kawabe, Komai and Razowski 1992), and 77 genera for the 259 known species of the Chinese fauna (Liu and Li 2002). Kawabe (1982) reported 390 olethreutine species in 97 genera for the wellstudied Japanese fauna, and many further genera
have undoubtedly been recognised since. And yet all these countries are around the fringe of the Oriental Region. As an indication of the very high olethreutine diversity of the Oriental region proper the number of genera of the tribe Olethreutini from Diakonoff’s (1973) revision of the South Asiatic fauna is included in Table 3 and contrasted with the tally for the genera of the Olethreutini in the other areas. If the ratio of Olethreutini genera in Taiwan and China can be used as an approximation, the fauna of the Oriental Olethreutinae will number well in excess of 200 genera. The more detailed analysis given below of the olethreutine fauna within Australia shows that the strong north/south diminution in the number of Oriental genera continues within the Australian continent. Indeed, it carries on to New Zealand with only eleven olethreutine genera (Dugdale 1988). The number of genera of the Olethreutinae within Australia was scored by States, dividing Queensland into north of Townsville and south of Townsville; New South Wales into north of Sydney and south of Sydney, and Western Australia into the Kimberley and the remaining southern part of the state. Of a total of 90 genera, 86 are present in the northern half of Queensland, dropping to 77 for the southern half, 45 for the northern half of New
Table 3. Olethreutine diversity in various geographic areas *in addition about 200 unnamed species known; **many species not placed to genera Country or region
Number of genera
Olethreutini genera
Number of species
Australia
90
28
249*
New Zealand
11
1
21
Dugdale 1988
Taiwan
69
33
135
Kawabe, Komai and Razowski 1992
China
77
35
259
Liu and Li 2002
Japan
97
43
390
Kawabe 1982
Europe
64
24
546
Razowski 2003
USA and Canada
59
16
800
Powell 1983
Neotropics
41
9
341**
south Asia
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88
Reference Horak in preparation
Powell, Razowski and Brown 1995 Diakonoff 1973
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54
OLETHREUTINE MOTHS OF AUSTRALIA
South Wales and only 32 still present in its southern half. Only eight olethreutine genera are reported from Tasmania. Of the 39 genera present in the Northern Territory, 38 are also recorded from northern Queensland, and there is little doubt that the only exception, the rarely collected Eucosmophyes Diakonoff with one small species taken by Ian Common in his garden in Toowoomba and present in New Guinea, will eventually also be found in north-eastern Australia. Only six genera occur in all States and Territories, the three eucosmine genera Spilonota Stephens, Strepsicrates Meyrick and Holocola Meyrick, and one genus each in the Bactrini (Bactra Stephens), Enarmoniini (Ancylis Hübner) and Grapholitini (Grapholita Treitschke). Zomariana Diakonoff (Olethreutini) is present throughout the mainland but absent from Tasmania. Three genera seem to be genuinely absent from the northern parts of Australia: Trachyschistis Meyrick in the Olethreutini, Hermenias Meyrick in the Eucosmini, and Gymnandrosoma Dyar in the Grapholitini. Given the rudimentary knowledge about the olethreutine fauna of New Guinea, especially of its lowland areas with Australian floral elements, it is not possible to establish how many olethreutine genera may be endemic to Australia. However, the very restricted distribution of Trachyschistis from Clyde Mtn in southern New South Wales to the Bunya Mtns in southern Queensland suggests that this Olethreutini genus is indeed endemic. This also applies to the eucosmine Hermenias, which is restricted to the southern half of the continent. The remaining ten genera, so far only known from Australia, consist of six grapholitine genera and the olethreutine Eremas Turner, the eucosmine Whittenella, gen. nov., and the two enarmoniine genera Aglaiogona, gen. nov. and Oriodryas Turner. The latter four genera all are rarely collected and occur in, or are restricted to, northern Queensland, hence they could well be present in New Guinea. It is more likely that some genuinely endemic genera are found among the six grapholitine genera that are mostly distributed in the drier regions of the continent. Of these, Apocydia, gen. nov., Notocydia, gen. nov. and Loranthacydia Horak, Common & Komai all have a very wide distribution, that includes most of the continent except Victoria. Archiphlebia, gen. nov. and Ixonympha, gen. nov. are both known from widely disjunct localities inland. Commoneria, gen. nov. is the only grapholitine genus restricted to northern Queensland and perhaps the least likely to be endemic to Australia. The diversity in the tribe Olethreutini shows a very steep gradient in the numbers of genera from
120503•Olethreutine Moths 4pp.in54 54
north to south, with the 27 genera present in northern Queensland gradually declining to nine genera in New South Wales south of Sydney. Zomariana is the only genus reaching Victoria and extending west to southern Western Australia, though six other genera reach into the Kimberley from the Northern Territory. The distribution pattern of the Enarmoniini, except for the widely distributed Ancylis, is very similar to that of the Olethreutini. All 19 Australian genera are found in northern Queensland but only four of these are present south of Sydney, and only Ancylis reaches Victoria. The Microcorsini and Bactrini, except for the widely distributed Bactra, show the same distribution pattern. Eucosmine diversity and distribution suggest two very different elements in the Australian fauna. The majority of genera, including the Acroclitagroup, are clustered in the northern half of Australia, with only six genera reaching south to Sydney out of a total of 15 genera in northern Queensland. Their relationships are clearly Oriental, in parallel with those of the Olethreutini and Enarmoniini. However, four genera of the Spilonota-group, i.e. Spilonota, Strepsicrates, Hermenias and Holocola, are widely distributed throughout Australia and Tasmania with their highest species diversity in the southern part of the continent. Hermenias occurs in the southern half of the continent only, mainly at higher altitudes, and the very large genus Holocola has its highest species diversity in the south, also at higher altitudes. Both these genera have endemic species in Tasmania. Eccoptocera Walsingham, the only genus in the Spilonota-group restricted to northern Queensland, is widely distributed beyond Australia from Sri Lanka to Hawaii. Apart from Hermenias, which is endemic to Australia, the other genera of the Spilonota-group are strongly represented in New Guinea. However, they extend into the Pacific rather than deep into the Oriental region, with the exception of a few species of Strepsicrates. Nearly all food plants used by the Spilonota-group belong to the Myrtaceae, and this is true also beyond Australia. There is no doubt that the high diversity of species resulted from a radiation on this plant family in Australia. Species diversity in the Oriental region decreases with distance from Australia, in stark contrast to the Olethreutini, Enarmoniini and other Eucosmini. The Australian grapholitine fauna also comprises at least two elements, though they are less clear-cut than in the Eucosmini. Seventeen of the 18 Australian grapholitine genera are present in northern Queensland but some are restricted to dry
5/5/06 2:39:34 PM
DIVERSITY AND DISTRIBUTION
habitats. The distribution of eight of these 17 genera reaches to south of Sydney and five of them (Loranthacydia, Notocydia, gen. nov., Apocydia, gen. nov., Cryptophlebia Walsingham and Grapholita) are also found in southern Western Australia, representing a dry-adapted Australian element. Notocydia, gen. nov. and Loranthacydia are both endemic speciose genera of the dry inland, indicating an Australian radiation. Pammenemima
120503•Olethreutine Moths 4pp.in55 55
55
Diakonoff, Leguminivora Obraztsov, Fulcrifera Danilevsky & Kuznetzov, Thaumatotibia Zacher, Acanthoclita Diakonoff, Pammenopsis Kuznetzov, Microsarotis Diakonoff and Parapammene Obraztsov are centered on northern Queensland and have their highest species diversity outside Australia. Commoneria, gen. nov., though so far known only from Australia, could well belong to this group.
5/5/06 2:39:35 PM
120503•Olethreutine Moths 4pp.in56 56
5/5/06 2:39:35 PM
5. Australian olethreutine genera Key to olethreutine genera based on Australian species (some genera for males only) Olethreutinae are externally very similar except for their secondary sexual characters, having a conservative venation in contrast to their derived genitalia. As far as possible, external characters, mainly wing venation, have been used for the key and for this reason several genera key out more than once. However, identification based on external characters often requires males and often does not allow resolution to genera unless wing pattern is included. Conversely, the genitalia alone, especially the male, can be diagnostic for the genus. Where two or three genera key out together, the illustrations should be consulted, keeping in mind that there are undescribed species and genera. It is suggested that, prior to using the key, specimens for identification should first be checked against the frequently encountered pest species such as Cydia pomonella and Cryptophlebia ombrodelta, and common species such as Zomariana doxasticana, Crocidosema plebejana and Strepsicrates and Holocola species. Diagnostic characters for the subfamily Olethreutinae: one ring of scales per flagellar segment; olethreutine juxta/caulis complex; hindwing nearly always with cubital pecten; with tortricid ovipositor lobes but without sclerotised connection between sterigma and anterior apophyses (except in Microcorsini). 1. Hindwing with M2 and M3 widely separate and nearly parallel to base (Fig. 101) (or if M3 and CuA1 are entirely fused then this fused vein widely separate from M2), M2 at most faintly bent towards M3; male never with costal fold, antennal notch or tibial pencil (if in doubt proceed to 2) ............................................................. 96 – Hindwing with M2 clearly bent towards M3 at base or the two converging (Fig. 27), but not necessarily closely approximated ....................................... 2 2(1). Forewing R3 stalked with R4 or with stalk of R4 and R5 ......................................... 3 – Forewing R3 and R4 not stalked ............. 5
120503•Olethreutine Moths 4pp.in57 57
3(2). – 4(3).
–
5(2).
–
6(5). –
7(6).
–
8(7).
–
9(7).
10(6).
Forewing R4 and R5 stalked ...................... ....................... Heleanna (part) (p. 345) Forewing R4 and R5 separate .................. 4 Chorda absent; wing pattern of welldefined dark grey and yellow marks ..... ............................ Demeijerella (p. 108) Chorda present; wing never with yellow marks, often with a pink semicircle on costa bordered by black .......................... ..................... Statherotis (part) (p. 190) Forewing R4 and R5 distinctly stalked (rarely fused and stalked with nearest vein) ...................................................... 6 Forewing R4 and R5 separate or connate, if very briefly stalked never beyond 56E ............................................................. 31 Hindwing M3 and CuA1 distinctly stalked, rarely entirely fused ............. 10 Hindwing M3 and CuA1 separate or connate, if very briefly stalked never beyond 56E .............................................. 7 R3 closely adjacent to and parallel with stalk of R4 and R5 in basal 56M; forewing apex strongly projecting; males only ............................................ 8 R3 distant from stalk of R4 and R5 at base; forewing apex hardly projecting, nearly square ........................................ 9 Hindwing costa with deep indentation beyond base, with long scale pencil; green species (Fig. 83 ) .......................... ................................Acroclita (p. 334) Hindwing costa without deep indentation in basal half; brownish species (Fig. 86 ) .................. Tritopterna (p. 338) Forewing with R3 distally swollen and at base usually approximated to R2 or the two veins stalked; wingspan 7.0–13.5 mm (Fig. 68) ........................................... .................... Metaselena (part) (p. 271) Forewing with R3 not modified; wingspan 15–22 mm (Fig. 57) ............... ...............................Anathamna (p. 225) R3 very close to stalk of R4 and R5 and parallel at least in basal 56M; often with raised scales on dorsum (Acroclitagroup) ................................................. 11
5/5/06 2:39:35 PM
OLETHREUTINE MOTHS OF AUSTRALIA
58
–
11(10). – 12(11).
– 13(12).
– 14(13).
–
15(11). –
16(15). – 17(16).
18(15).
19(10).
–
R3 distant from stalk of R4 and R5, and if close at base then divergent and not parallel ................................................ 19 Males ....................................................... 12 Females ................................................... 15 Forewing upperside with deep depression in basal half of cell, filled with whitish modified scales and covered by a dome of large scales .......................................... ................... Heleanna (part) (p. 345) Forewing cell without depression with modified scales .................................. 13 Male genitalia with large scale tuft from fold between tegumen and valva ........... ........................ Noduliferola (p. 341) Male genitalia without scale tuft between tegumen and valva ............................ 14 Socii upwardly directed; lower margin of basal excavation with a finger-like process; never with antennal notch ...... ............................ Rhopobota (p. 321) Socii drooping; lower margin of basal excavation without finger-like process, but with large flattened bristles from outer surface of valva; often with notch at base of antenna (Fig. 9) ...................... .................. Fibuloides (part) (p. 326) Entire ductus bursae sclerotised, without anterior prongs .................................. 16 Only anterior part of ductus bursa sclerotised, as a bifid prong (Fig. 691).... ............................................................. 18 Signa absent ......... Tritopterna (p. 338) Signa present .......................................... 17 Genitalia asymmetrical, ductus bursae long, slender, sinuate and originating laterally; sterigma not spinulose ........... ................... Heleanna (part) (p. 345) Genitalia symmetrical, ductus bursae rather short and wide; sterigma strongly spinulose Acroclita (p. 334) Tips of prong reaching halfway around corpus bursae .... Rhopobota (p. 321) Tips of prong only just reaching into bursa .................. Fibuloides (p. 326), ........................ Noduliferola (p. 341) R3 modified, distally swollen, at base closer to R2 than to stalk of R4 and R5; forewing broad, with nearly rectangular apex and straight termen; cream except for dark markings along costa (Fig. 576)......................................... .................... Metaselena (part) (p. 271) R3 not modified; wing not broad with rectangular apex ................................ 20
120503•Olethreutine Moths 4pp.in58 58
20(19).
– 21(20). –
22(21).
–
23(20).
– 24(23).
–
25(24).
– 26(25). – 27(26).
One vein missing in forewing (but if forewing blackish and intricately marked (Fig. 621) see Toonavora, gen. nov.); narrow-winged, wingspan 6–13 mm ............................................ 21 No forewing vein missing (except rarely in Toonavora, gen. nov.) .................... 23 Forewing strongly falcate with black line along termen below apex (Fig. 455) . 22 Forewing subrectangular, with large rounded costal fold and antennal notch in male (Fig. 801) Eccoptocera (p. 375) Forewing pale ochreous with irregular dark brown band along costa; antenna ringed with black scales (Fig. 487) ............................. Ancylophyes (p. 235) Forewing with dark markings not only along costa, fringe below apex conspicuously yellow Fig. 455) .............. .............. Aglaogonia, gen. nov. (p. 221) Forewing lightly notched below apex, grey or brown with black spot in apex; hindwing usually partially translucent near base and in male always with patches of modified scales above and/ or below in anal region (Fig. 927); wingspan 11–14 m .................................. .................. Acanthoclita (part) (p. 439) Not with this character combination.....24 Discal cell at least 36: length of forewing (Fig. 12); wings usually slender, parallel-sided; males often with costal fold (Fig. 10) and/or antennal notch (Fig. 9) (Eucosmini) .......................... 25 Discal cell less than 36: length of forewing (Fig. 58); usually with projecting to falcate apex (Fig. 58) (Enarmoniini) ............................................................... 30 Forewing pale grey with greenish tinge, with dark grey-brown streak (male) or large triangle (female) along distal 36: costa, with nearly falcate apex followed by white and then black scales on termen (Figs 833, 834); tegumen with two long, very slender, downcurved arms, widened and bristled in distal third; no costal fold ................................ .............................. Whittenella (p. 392) Wing pattern different .......................... 26 Males ....................................................... 27 Females ................................................... 29 Frons scaling modified, upwardly oriented; forewing widest in middle, without costal fold; complex valva with
5/5/06 2:39:36 PM
AUSTRALIAN OLETHREUTINE GENERA
28(27).
–
29(26).
30(24).
31(5).
– 32(31). – 33(32).
–
34(33).
–
band- to mushroom-shaped process from sacculus . Icelita (part) (p. 383) Frons scaling not modified, often with costal fold and notched antenna ...... 28 Costal fold always present and to well beyond middle of wing, base of R3 very close to stalk of R4 and R5 but divergent ............ Hermenias (p. 371) Costal fold, if present, at most to middle of wing, base of R3 not adjacent to base of stalk of R4 and R5 ........ Spilonota ................ (p. 350), Holocola (p. 362) Ovipositor extremely telescopic, ovipositor lobes very small .................... ......................... Icelita (part) (p. 383) Ovipositor not telescopic, ovipositor lobes normal............... Hermenias (p. 371), Spilonota (p. 350), Holocola (p. 362) Forewing apex strongly falcate (Fig. 598), wingspan 6–13 mm; forewing widest below middle; male with modified, small black scales in centre of hindwing .................................................. ........... Pseudancylis, gen. nov. (p. 282) Apex weakly falcate (Fig. 472); forewing not narrowing distally; wingspan 7–17 mm .................... Ancylis (part) (p. 229) Hindwing cell short (ratio of hindwing cell length to hindwing length 0.38– 0.46); often with prominent pretornal spot (Fig. 918) ..................................... 32 Hindwing cell long, at least 56O hindwing length .................................................. 35 Accessory cell long, from below R2 (Fig. 111) ...................................................... 33 Accessory cell small, from beyond R2 (Fig. 108) or absent ............................ 34 Forewing dark grey to blackish, hindwing pale grey (Fig. 905) ................................. .................... Gymnandrosoma (p. 430) Either forewing grey with blackish subapical spot and hindwing light orange with grey margin, or forewing orange and grey and hindwing grey ........... Archiphlebia, gen. nov. (p. 433) T8 in male a subtriangular or Y-shaped plate, with a pair of scale tufts in shallow membranous pockets on each side; female genitalia with corpus bursae spinulose in anterior 56O–169 ........ .......................... Cryptophlebia (p. 435) T8 in male a broadly sclerotised plate with a medially convex posterior margin laterally produced into curved points, with paired patches of long
120503•Olethreutine Moths 4pp.in59 59
35(31). –
36(35).
– 37(36).
–
38(37).
– 39(38).
– 40(39).
41(37).
–
59
mane-like scales but without paired lateral scale tufts in shallow pockets; female genitalia with a ring of spinules at the juncture of the ductus and corpus bursae Thaumatotibia (p. 427) Hindwing M3 and CuA1 distinctly stalked, rarely entirely fused ............. 36 Hindwing M3 and CuA1 separate or connate, if very briefly stalked never beyond 56E ............................................ 55 Forewing with large yellow-brown or redbrown triangle based along entire costa, partially edged with white, remainder of wing grey (Figs 244, 245); wingspan 16–19 mm; anellus with two antler-shaped spikes ............................... ............................. Rhectogonia (p. 125) Not with this character combination ... 37 Male hind tibia with scale pencil (Figs 15, 16); never with costal fold or antennal notch ................................................... 38 Male hind tibia without scale pencil; costal fold (Fig. 10) and/or antennal notch (Fig. 9) may be present ........... 41 Forewing venation modified in male, with R1 deflected parallel to costa by variably thickened pterostigma on costa (Fig. 31), shape and venation of male hindwing often modified (Fig. 31) ........................... Lobesia (part) (p. 111) Not with this character combination ... 39 Hind tibia in male hugely enlarged by scale pads with inner surface smooth and containing the tibial pencil (Fig. 340); scale pencil from anal margin .............. Diakonoffiana (part) (p. 165) Hind tibia not with these characters .... 40 Wingspan 9.0–13.5 mm; forewing dark grey with large pinkish ochreous semicircle from 56: costa to before apex, reaching to middle of wing and bordered with black, male with large hairpencil on upperside of hindwing within cell (Fig. 302) .............................. ........................... Eremas (part) (p. 148) Wingspan 11–22 mm; not with the above character combination............................ ............................ Dudua (part) (p. 134) Forewing with R3 closely approximated and parallel to R4 at least in basal 56M ............................................................. 42 Forewing with R3 not close and parallel to R4, if close at base then divergent ............................................................. 43
5/5/06 2:39:36 PM
OLETHREUTINE MOTHS OF AUSTRALIA
60
42(41).
–
43(41).
– 44(43).
–
45(44).
– 46(45).
– 47(46).
Wingspan 7.0–13.5 mm; forewing light greyish ochreous with red-brown projecting apex and silvery lines (Fig. 590), without raised scales on dorsum; male with anal roll enclosing scale pencil .................... Periphoeba (p. 278) Wingspan 9–21 mm; forewing dark greenish or brownish, strongly marked, with paired white strigulae before blackish apex, with raised scales on dorsum; male usually with notch at base of antenna ....................................... ...................... Fibuloides (part) (p. 326) Dark greyish ochreous forewing with a greenish tinge with two dark marks on dorsum, one along basal third and a semicircle before tornus; hindwing translucent dark grey, thinly scaled but with dense black scales along veins on underside, especially in male (Fig. 667); short, porrect labial palpi with black sclerotisation at tip in male .................... ....... Melanodaedala, gen. nov. (p. 317) Not with this character combination ... 44 Subtriangular forewing with broadly rounded or angled apex, if not dark blackish brown with metallic blue scales then with curved parallel silvery and yellow lines from costa before apex to middle of termen; male hindwing often with ribbon from base of anal margin ............. Helictophanes (p. 257) Not with above character combination; with apex projecting and male often with costal fold and/or antennal notch ............................................................. 45 Forewing lightly notched below apex, grey or brown with black spot in apex; hindwing usually partially translucent near base and in male always with patches of modified scales above and/ or below (Fig. 927); wingspan 11–14 mm Acanthoclita (part) (p. 439) Not with this character combination ... 46 Hindwing in male with M3 and CuA1 stalked nearly to wing margin or entirely fused and CuA2 close and parallel to stalk of M3 and CuA1 (Fig. 75); blackish brown moth with nearly grapholitine wing pattern (Fig. 621); female with telescopic ovipositor ............... Toonavora, gen. nov. (p. 295) Not with this character combination ... 47 Male either with cubital pecten hugely enlarged into a prominent tuft or with
120503•Olethreutine Moths 4pp.in60 60
– 48(47).
–
49(48).
– 50(49).
51(48).
–
52(51).
– 53(52).
a combination of a concave frons enhanced by modified scaling and a scale pencil from hindwing margin inserted in lateral abdomen scaling ...... ............................ Crocidosema (p. 301) Not with this character combination ... 48 Male genitalia with process from lower margin of basal excavation (Fig. 652); never with strongly falcate apex or scale pencil in anal roll; never with antennal notch ................................... 49 Male genitalia not with process from lower margin of basal excavation; some taxa with strongly falcate apex, scale pencil in anal roll or antennal notch in male .................................................... 51 Bifid uncus present, socii broad membraneous lobes; no costal fold ...... ................ absconditana-group (p. 310) Uncus at most a rounded lobe or hump in Australian species .............................. 50 Male with narrow costal fold; nearly uniformly dark grey with an indistinct paler triangular patch near middle of dorsum, ocelloid patch pale ochreous (Fig. 641) .................. Epiblema (p. 307) Male without costal fold; strongly contrasting pale and dark wing markings ............ Coenobiodes (p. 313) Discal cell at least 36: length of forewing; usually slender, parallel-sided wings; males often with costal fold and/or antennal notch (Eucosmini) ............. 52 Discal cell less than 36: length of forewing; usually with projecting to falcate apex and sinuate termen partly edged with black scales (Fig. 472); never with costal fold or antennal notch ... Ancylis (part) (p. 229) Frons scaling modified, upwardly oriented; forewing widest in middle, transversely sharply banded (Fig. 816), without costal fold, wingspan 9.0–13.5 mm; complex valva with band- to mushroom-shaped process from sacculus .............. Icelita (part) (p. 383) Not with this character combination ... 53 Forewing without costal fold, with outwardly curved transverse pale and dark grey bands, wingspan 6.5–10.0 mm; white-tipped dark labial palpi; simple genitalia with small round socii and a valva with a deep V-shaped emargination in ventral margin............. .......................... Eucosmophyes (p. 388)
5/5/06 2:39:37 PM
AUSTRALIAN OLETHREUTINE GENERA
–
54(53).
55(35). – 56(55). –
57(55). –
58(57).
– 59(58).
–
60(59).
Not with this character combination; costal fold and antennal notch often present ................................................ 54 Valva with a triangular to finger-like process from ventral margin of valva next to a single spine ........ Strepsicrates (p. 355) Valva not with a process and an associated spine from ventral margin.... ... Spilonota (p. 350), Holocola (p. 362) CuA1 distant from M3 and from well below angle of cell (Fig. 94) .............. 56 CuA1 not distant from M3, never from below angle of cell ............................. 57 Forewing narrow, brownish, wingspan 6– 10 mm ...................... Herpystis (p. 380) Forewing broad, subtriangular, R3 and R4 strongly approximated and parallel at base, wing pattern white and bluish blackish (Figs 411, 412), wingspan 11– 15 mm ......................... Aterpia (p. 196) Forewing costa not doubly sinuate and concave in middle .............................. 58 Forewing costa doubly sinuate; wingspan 16.5–19.0 mm; ochreous to grey forewing with dark triangle in concave medial portion of costa (Fig. 616); male with anal roll with pencil from wing base ................. Thysanocrepis (p. 292) Forewing broadly triangular, glossy black with deep notch below round apex and with long silvery blue and white strigulae in apical half of costa (Fig. 552); hindwing often partly unscaled and transparent, with Rs and M1 distant from base; labial palpi slender, upcurved ...... Irianassa (part) (p. 262) Not with the above character combination ....................................... 59 Hindwing blackish with conspicuous areas of whitish or orange scales in basal half, forewing dark bluish or reddish brown or orange, subrectangular, wingspan 12.5–18.5; male genitalia with ventrally bristled uncus and socii with hand-shaped scales ............... Eucosmogastra (p. 252) Not with the above character combination (female Diakonoffiana with grey hindwing with orange base, but forewing with short and triangular accessory cell and CuA1 strongly recurved) ............................................ 60 Forewing with R2 and R3 both sinuate, widely distant at base and strongly
120503•Olethreutine Moths 4pp.in61 61
–
61(60).
–
62(61). – 63(62).
–
64(63).
–
61
converging towards costa (Fig. 69); broadly subtriangular whitish forewing with indistinct silvery, grey and black markings (Figs 584, 585), hindwing dark grey with enlarged anal area in male, wingspan 14–16 mm .................... ................................. Oriodryas (p. 275) Not with the above character combination, R2 and R3 not distally converging .......................................... 61 R2, R3 and R4 all very close and parallel nearly to costa; forewing subtriangular, greyish-ochreous with red-brown mark on middle of costa and red-brown projecting apex (Fig. 501), with fake costal fold of large scales at 569 costa in male, wingspan 8.5–13.0 mm ................ ........................... Balbidomaga (p. 242) Not with the above character combination, at most R3 and R4 close and parallel ........................................ 62 Forewing markings parallel, with welldefined, longitudinal lines ................ 63 Forewing markings not with mainly parallel, longitudinal lines ................ 65 Forewing with bases of R3, R4 and R5 close together and with a conspicuous leaden curved line from before apex to termen; forewing subrectangular with a blunt apex and termen sinuate; wingspan 9–14 mm; wing pattern of well-defined, glossy, cream, grey and brown lines (Fig. 607) ............................ .............................. Tetramoera (p. 288) Forewing with bases of R3, R4 and R5 not close together and without a leaden curved line from before apex to termen, termen at most apparently sinuate due to pale fringe, wing pattern with less clearly defined lines ................ ............................................................. 64 Forewing pointed, with acute apex and oblique termen; wingspan 9–23 mm; wing pattern of variable ochreous to dark grey-brown striation, with a white dot on closing vein .................................. ............................ Bactra (part) (p. 206) Forewing elongate subrectangular, blotchy and longitudinally striate with beige and brown, base of abdomen with shallow paired round depressions dorsally on T2; wingspan 10–14 mm; male hindwing modified, narrowly triangular as in some Lobesia (Fig. 287) . Archilobesia (part) (p. 142)
5/5/06 2:39:37 PM
OLETHREUTINE MOTHS OF AUSTRALIA
62
65(62).
– 66(65).
– 67(66).
– 68(67).
– 69(68).
– 70(69).
Base of abdomen with a pair of deep round depressions dorsally on T2, visible only if scaling is brushed off (dorsal pits, Fig. 312) (Olethreutini) ..... ............................................................. 66 T2 dorsally without paired round depressions ......................................... 71 Forewing broad, intricately patterned dark brownish with pink tinge and metallic leaden streaks or spots (Figs 362–367); male with greatly expanded hind tibia and usually modified hindwing anal region; wingspan 11–15 mm ...... Gatesclarkeana (p. 178) Not with the above character combination ....................................... 67 Forewing subrectangular, brown, with a strongly sinuous pale grey band from base of costa to termen below black apex, edged with white, a large pale ochreous mark in tornus and an outwardly oblique brown mark, edged with white, from near base of dorsum (Fig. 329); wingspan 12–18 mm; with long, pointed labial palpi ....................... ............................. Temnolopha (p. 162) Not with the above character combination ....................................... 68 Hindwing shape and venation in male modified, narrowly subtriangular as in some Lobesia; forewing elongate subrectangular, blotchy and longitudinally striate with beige and brown (Fig. 287); dorsal pits shallow, weakly developed .................................... .................. Archilobesia (part) (p. 142) Not with the above character combination ....................................... 69 Forewing subtriangular, pale grey with distally red-brown costal strigulae, variably suffused with dark grey or red-brown (Figs 307–310); male with dark hairpencil from base of hindwing, at rest enclosed in groove of anal region (Figs 311, 312); wingspan 12–16 mm ..... ............................... Zomariana (p. 152) Not with the above character combination ....................................... 70 Tibia and first tarsal segment of male hind leg with long loose scale fringes, tibia with large scale pencil; welldefined, complex wing pattern always with erect mark from tornus and often a large roundish blotch below middle of costa; gnathos paired spinulose pads
120503•Olethreutine Moths 4pp.in62 62
–
71(65).
72(71).
–
73(72).
– 74(73).
75(74).
or cusps; single large horn-shaped signum ............ Ophiorrhabda (p. 128) Tibia and first tarsal segment of male hind leg greatly enlarged by dense scale pads, inner surface smooth with a small tibial pencil (Fig. 340); forewing often with raised scales, usually dark with indistinct pattern except for variable white suffusion; hindwing in female often with orange base; gnathos two smooth arms fused with anal tube; two plate-shaped signa ........................... .............. Diakonoffiana (part) (p. 165) Male with costal fold, of robust build; forewing dark grey to blackish brown (Figs 123–126); in male labial palpi densely scaled, antennal base sometimes thickened or hindwing anal margin folded over cluster of swollen scales, terminal abdominal segments with dorsolateral scale tufts; valva swollen with large process from near base; sterigma connected with anterior apophyses ......... Cryptaspasma (p. 68) Male not with costal fold, if forewing blackish brown then broadly triangular ............................................................. 72 Forewing venation modified in male, with R1 deflected parallel to costa by variably thickened pterostigma on costa (Fig. 31), shape and venation of male hindwing often modified (Fig. 31) ........................... Lobesia (part) (p. 111) Not with this character combination (if hindwing shape modified check next couplet) ............................................... 73 Hindwing shape and venation in male modified, narrowly subtriangular as in some Lobesia; forewing elongate subrectangular, blotchy and longitudinally striate with beige and brown (Fig. 287); dorsal pits shallow, weakly developed .................................... .................. Archilobesia (part) (p. 142) Not with the above character combination ....................................... 74 Male hind tibia with tibial pencil (Figs 15, 16) (often partially concealed in tibial scaling) ............................................... 75 Male hind tibia without tibial pencil ... 87 Forewing broad with costa strongly curved throughout, intricately patterned from grey to red-brown usually with no large marks; male hind tibia with large white fringe and a
5/5/06 2:39:37 PM
AUSTRALIAN OLETHREUTINE GENERA
– 76(75).
– 77(76).
– 78(77).
– 79(78).
– 80(79).
– 81(80).
double tibial pencil, one black and one pale; labial palpi upcurved, small, slender .................. Sycacantha (p. 100) Not with the above character combination ....................................... 76 Forewing with R3 closely approximated and parallel to R4 in basal ¼, or the two veins connate; brownish with pinkish triangle or semicircle over middle of costa often edged with black, or otherwise intricately marked with ochreous, grey and reddish brown, costal half often suffused with blackish scales ........... Statherotis (part) (p. 190) Not with the above character combination ....................................... 77 Forewing broad, costa angled or strongly curved at about 36: (Fig. 318–321), male hindwing modified with margin convex between CuP and 1A+2A, then deeply concave before the distally projecting anal roll (Fig. 41); forewing ochreous or grey with few dark spots along costa and scattered silvery dots or dark blackish brown with variable ochreous patches and scattered leaden blue dots .............. Podognatha (p. 156) Not with the above character combination ....................................... 78 Forewing dark grey with large pinkish ochreous semicircle from 56: costa to before apex, reaching to middle of wing and bordered by black; male with large hairpencil on upperside of hindwing within cell (Fig. 302), wingspan 9.0– 13.5 mm ............ Eremas (part) (p. 148) Not with the above character combination ....................................... 79 Forewing subrectangular, dark brown with olive tinge and conspicuous reddish fringe (Fig. 424); male hindwing with large but not distally projecting anal roll . Rhodacra (p. 203) Not with above character combination.........................................80 Forewing conspicuously patterned, either with green areas or grey-brown with apical area pink and containing large round subapical mark; male hind tibia usually with large scale fringes .............. ........................ Sorolopha (part) (p. 93) Not with green colouring and not with above wing pattern ............................ 81 Labial palpi upcurved, slender (Fig. 275) .. ............................................................. 82
120503•Olethreutine Moths 4pp.in63 63
–
82(81).
– 83(82).
–
84(81).
– 85(84).
–
86(85).
63
Labial palpi porrect, sinuate and second segment distally strongly widened (Fig. 352) ............................................. 84 Forewing subrectangular, grey-brown with one or two large snow-white marks, one from tornus nearly to costa and usually a semicircle over basal half of dorsum (Figs 293, 294) ....................... .......................... Euobraztsovia (p. 146) Wing pattern not as above .................... 83 Labial palpi slender, upcurved and sinuate, with second segment widest at tip (Fig. 284); forewing broadly triangular with strongly curved costa, ochreous and dark grey with conspicuous small white dash in middle of wing across closing vein (Fig. 281); hindwing with CuA2 from just beyond middle of cell and CuP absent .......................... Trachyschistis (p. 139) Labial palpi slender and upcurved with second segment curved throughout and widest in middle (Fig. 275); forewing elongate subrectangular, with costa weakly curved; wing pattern diverse but ocelloid patch usually present as a series of parallel black lines; male hindwing with anal roll and often modified, triangular, with sinuate or scalloped termen ................... ............................ Dudua (part) (p. 134) Forewing subrectangular, vivid redbrown with yellowish ochreous wing base and apex and with an outwardly oblique band of scattered iridescent blue scales from costa to dorsum, hindwing blackish (Fig. 357); wingspan 15.5–18.0 mm ............. Costosa (p. 173) Wing pattern not as above .................... 85 Forewing subrectangular, dark brown, intricately transversely marked and with conspicuous metallic leaden scales, wingspan 8–10 mm; hindwing with Rs and M1 stalked and M2 bent towards but still distant from M3 ......... ..................... Endothenia part. (p. 216) Forewing without metallic scales, ochreous or grey with intricate brown markings, wingspan 10.5–18.0 mm .................... 86 Forewing greyish ochreous with a large dark brown triangle on costa and often an oblique comma-shaped brown mark in terminal area; male genitalia with paired large hairy uncus lobes and very small socii; female genitalia with a
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OLETHREUTINE MOTHS OF AUSTRALIA
64
87(74).
– 88(87).
–
89(88).
– 90(89).
single signum composed of several shark-tooth-shaped sclerites .................. ........................ Megalota (part) (p. 170) Forewing with the same pattern elements as above and additional markings, with most of the wing speckled with brown; male genitalia with a single broad dorsally hairy uncus and long drooping socii; female genitalia with a single horn-shaped signum .............................. ........... Gnathmocerodes (part) (p. 121) Forewing subtriangular, costa distinctly angled at 169, grey with greenish tinge and several pale bands converging at 169 costa, apex black and fringe conspicuously reddish (Fig. 388), wingspan 10–12 mm .............................. ........................ Dactylioglypha (p. 187) Forewing costa not angled at 169, wing pattern not as above .......................... 88 Forewing subtriangular, conspicuously patterned, either with green or bluish green areas, or grey-brown with apical area pink and containing large round subapical mark; male hind tibia sometimes with large scale fringes ........ ........................ Sorolopha (part) (p. 93) Not with above wing pattern, if with green colouring then forewing subrectangular, with conspicuous white costal strigulae and a small white-edged black spot in apex (Metrioglypha) ................................... 89 Forewing dark brown with white streak along dorsum in male (Figs 148, 150), in female mottled brown with darker apex and two blackish marks along dorsum separated by some scattered white scales (Figs 149, 151); male in one species with a pencil of a few ochreous hairs from anal margin recurved and inserted in abdominal scaling (Fig. 153), wingspan 17–26 mm ............ .......................... Oxysemaphora (p. 82) Not with the above character combination ....................................... 90 Forewing elongate-subovate with strongly curved costa, pinkish grey with a curved band of parallel, white-edged minute black bars from tornus to beyond middle of costa and often a paler ochreous blotch along dorsum (Fig. 194); palpi sickle-shaped ............... .............. Atriscripta, gen. nov. (p. 104)
120503•Olethreutine Moths 4pp.in64 64
– 91(90).
– 92(91).
– 93(92).
– 94(93).
–
95(94).
–
Not with the above character combination ....................................... 91 Forewing broadly triangular, dark greybrown speckled with pale leaden bluish grey in basal and distal thirds (Fig. 417), wingspan 11–14 mm; male with antennal cilia ½ diameter of antenna ... ................................ Proschistis (p. 199) Not with the above character combination ....................................... 92 Forewing slender subrectangular, either green with blackish marks, or ochreous, reddish ochreous and blackish with white scalloped lines in rounded brown apex; male with antennal cilia ¼–½ diameter of antenna ............. Metrioglypha (p. 183) Not with above character combination.........................................93 Forewing narrowly subtriangular with strongly oblique termen and pointed apex, greyish ochreous to dark brown, nearly always with fine longitudinal striation and often with dark streak at least from minute white spot on closing vein to apex, palpi long, porrect and sinuate ............... Bactra (part) (p. 206) Not with the above character combination ....................................... 94 Hindwing with Rs and M1 stalked, forewing with M-stem absent, intricately marked with reddish or greyish brown on beige or pale yellowish ochreous, sometimes with metallic scales, wingspan 8.0–14.5 mm; male genitalia without uncus, female genitalia without signum ....................... ................................. Syntozyga (p. 212) Hindwing with Rs and M1 separate or connate, forewing with M-stem present, ochreous or grey with intricate deep brown markings, without metallic scales, wingspan 10.5–18.0 mm ....... 95 Forewing greyish ochreous with a large dark brown triangle on costa and often an oblique comma-shaped brown mark in terminal area; male genitalia with paired large hairy uncus lobes and very small socii; female genitalia with a single signum composed of several shark-tooth-shaped sclerites .................. ........................ Megalota (part) (p. 170) Forewing with the same pattern elements as above and additional markings, with most of the wing speckled with brown;
5/5/06 2:39:38 PM
AUSTRALIAN OLETHREUTINE GENERA
male genitalia with a single broad dorsally hairy uncus and long drooping socii; female genitalia with a single horn-shaped signum .............................. ........... Gnathmocerodes (part) (p. 121) 96(1). Forewing R3 and R4 not stalked ........... 97 – Forewing R3 stalked with R4 ..................... .................................. Collogenes (p. 73) 97(96). M3 and CuA1 fused into one vein; male either with hindwing with anal fold with modified fringe (Fig. 102), 3A approximated to margin and with hindtibia dorsally with long scales, or male with an elongate tubular androconial fold filled with lanceolate scales above along the posterior edge of cell (Fig. 99) ............................................ ................ Notocydia, gen. nov. (p. 411) – M3 and CuA1 not fused into one vein; male without above structures ......... 98 98(97). Hindwing pale grey, distinctly mottled and reticulate with dark grey; forewing grey variably darkened with brown in costal half, nearly always with a large pale ochreous blotch in distal third of wing (Figs 159–162); male with a large scale tuft on hindwing underside from between CuP and 1A+2A; wingspan 11–18 mm ................................................ ........... Cnecidophora, gen. nov. (p. 89) – Hindwing not distinctly mottled ......... 99 99(98). Forewing with well-defined vividly yellow and/or red pattern elements, combined with leaden metallic scaling, and hindwings dark grey, often with deep orange base or rarely entirely deep orange ............................................... 100 – Not with the above character combination, forewing at most with ill-defined yellowish pattern elements ... ............................................................... 101 100(99). Forewing with basal half yellow scattered with minute black dots and short lines, distal half black with sinuous red lines enclosing leaden spots (Figs 493, 494) .. ................................. Anthozela (p. 239) – Forewing not with above pattern .............. ............................... Loboschiza (p. 266) 101(100). Rs and M1 distant from base (Fig. 118) ........................................................... 102 – Rs and M1 approximated at base (Fig. 113) or stalked ......................... 105 102(101). Forewing with termen notched below apex (Figs 854, 964) ......................... 103
120503•Olethreutine Moths 4pp.in65 65
65
–
Forewing with termen not notched below apex; forewings glossy silvery grey with long, large white costal strigulae separated by deep black lines, meeting curved black lines of dorsal blotch in middle of wing (Fig. 956); wingspan 7– 13 mm ................. Microsarotis (p. 454) 103(102). Forewing with termen deeply notched...............................................104 – Forewing with termen shallowly notched; with two or three black dashes or dots very close to termen (Fig. 854); wingspan 7.0–9.5 mm ............................ ....................... Pammenemima (p. 403) 104(103). Ocelloid patch developed as 2–3 longitudinal, parallel black dashes between M2 and CuA2; wing tip suffused with cinnamon-tipped scales; dorsal blotch indicated as two doubled curved white lines (Fig. 964); wingspan 10.0–13.5 mm .......................................... ........... Commoneria, gen. nov. (p. 459) – No trace of ocelloid patch or dorsal blotch; forewing entirely glossy blackish with bluish sheen except for some leaden and iridescent blue lines and two white strigulae in apex ; hindwing often partially translucent (Fig. 552) ....... Irianassa (part) (p. 262) 105(101). Forewing blackish with a broad whitish band from 6M costa to tornus; hindwing conspicuously orange; wingspan 9–11 mm .................................................. ............. Ixonympha, gen. nov. (p. 464) – Not with this character combination ........ ........................................................... 106 106(105). Forewing greyish ochreous to grey-brown in males, orange ochreous to dark redbrown in females, with long costal strigulae and several oblique silver lines from costa, and with an elongate ocelloid patch (Fig. 880); antennae in male thickened with dense scales and entirely scaled ......................................... ................. Apocydia, gen. nov. (p. 419) – Not with this character combination ........ ........................................................... 107 107(106). Forewings subovate, with broadly rounded to at most rectangular apex .... ........................................................... 108 – Forewings subrectangular or subtriangular, with at least slightly projecting apex ................................ 109 108(107). Forewing dark grey-brown to bluish blackish, wingspan 7.5–15.0 mm; male
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66
OLETHREUTINE MOTHS OF AUSTRALIA
hindwing with ribbon-like process from base of anal region (Fig. 62) ......... ........................... Cyphophanes (p. 248) – Forewing ochreous grey to grey, subovate with ill-defined greyish to blackish brown longitudinal lines or suffusion and usually with a dark longitudinal mark in centre of wing, especially in female; wingspan 15–21 mm ................. ................. Centroxena (not discussed) 109(107). Forewing small (8–10 mm wingspan), subrectangular, dark brownish, intricately transversely marked, with conspicuous large patches of metallic scales throughout but with ocelloid patch not defined and without any black bars (Fig. 451); hindwing greyish; male with tibial pencil ........................... .................... Endothenia (part) (p. 216) – Not with above character combination ........................................................... 110 110(109). Forewing small (7–10 mm wingspan), subtriangular, light ochreous to greenish grey in basal half, distally blackish beyond a line from middle of dorsum to 169 costa (Fig. 602) ................. ................................ Pternidora (p. 285) – Not with above character combination....................................... 111 111(110). Forewing lightly notched below apex, grey or brown with black spot in apex; hindwing usually partially translucent near base and in male always with patches of modified scales above and/ or below (Figs 927–929); wingspan 11– 14 mm ...... Acanthoclita (part) (p. 439) – Not with this character combination...112 112(111). 3A in male with a series of granular spots distributed along either side (Fig. 890) ........................................ Cydia (p. 406) 3A in male without such granular spots – ........................................................... 113 113(112). 3A in male close to margin (Fig. 103); sacculus strongly concave ventrally near base, forming an angulate ridge; ovipositor lobes slender .................. 114 – 3A in male not close to margin (Fig. 98); sacculus not concave near base; ovipositor lobes moderate or wide ........ ........................................................... 115 114(113). Aedeagus with an often parallel, strongly sclerotised process dorsad of anellus; socii a pair of hairy patches fused with diaphragma or absent; lamella
120503•Olethreutine Moths 4pp.in66 66
postvaginalis a heart-shaped spinulose plate; ductus bursae not transversely wrinkled ................. Fulcrifera (p. 415) – Aedeagus without a dorsal process; socii a pair of long hair-pencils with converging tips and their bases from a concavity of the dorsolateral expansion of tegumen; lamella postvaginalis a weakly sclerotised area with scale sockets; ductus bursae distally transversely wrinkled ............................. .......................... Leguminivora (p. 407) 115(113). S2 with anterolateral processes absent or reduced (Fig. 977) ............................ 116 – S2 with anterolateral processes well developed (Fig. 944) ........................ 118 116(115). Valva without costal process; coremata absent .............. Loranthacydia (p. 399) – Valva with costal process; coremata present .............................................. 117 117(116). Forewing with or without median fascia and with an ocelloid patch with outer edge reduced or absent; in male hindwing Sc+R1 and Rs fused beyond outer edge of cell (Fig. 974), usually with narrow or hair-like, modified scales along Sc+R1 and Rs on upper surface .. Parapammene (part) (p. 461) – Forewing with ill-defined, rarely welldefined dorsal blotch and with typical ocelloid patch indicated by a pair of vertical leaden lines enclosing some black dots or dashes (Fig. 945); in male Sc+R1 and Rs free throughout .............. Grapholita conficitana-group (p. 448) 118(115). Bactra-like appearance without typical grapholitine pattern elements (Fig. 936); hindwing with M3 and CuA1 long-stalked and 3A a trace only in female and absent in male; male hindwing with long black hair scales along anal margin (Fig. 937) and a band of modified scales on lower surface behind M1 .................................. .......................... Pammenopsis (p. 443) – Grapholitine appearance with welldefined dorsal blotch and with ocelloid patch indicated by a single vertical leaden line; M3 and CuA1 connate or very short-stalked; anal margin of male hindwing without modified scales ........................................ ............Grapholita compositella-group (p. 448)
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AUSTRALIAN OLETHREUTINE GENERA
MICROCORSINI Kuznetsov Microcorsini Kuznetsov, 1970, Entomol. Obozr. 49: 442. Type genus: Microcorses Walsingham, 1900.
Status remarks The Microcorsini, based on Cryptaspasma Walsingham with several subgenera including Microcorses (that sometimes are treated as separate genera) have long been accepted as a basal tribe within the Olethreutinae. They are the only olethreutine group to share a sclerotised connection between sterigma and anterior apophyses with the Tortricinae, the most conspicuous of several symplesiomorphies (Kuznetsov 1970).
Evidence for monophyly There is no doubt that Cryptaspasma and its subgenera together form a monophyletic group (see Remarks). Collogenes Meyrick is here tentatively included in the Microcorsini as, apart from a similar dark-winged superficial appearance, it shares a number of unusual features with Cryptaspasma, albeit some of them symplesiomorphies. In both genera the tegumen is of similar shape and has strong medial apodemes from the pedunculi, the spinulose U- or V-shaped sterigma is laterally continuous with the ventral arms of the anterior apophyses, the signa, if present, are horn-shaped and both close to the entrance to the corpus bursae. The larvae of both are endosperm borers. There are also conspicuous differences, foremost the very different secondary sexual structures, a costal fold in Cryptaspasma versus modified mid and hind tibia with a pencil from the thorax in Collogenes, and the much larger basal excavation in Collogenes. However, the fact that the Microcorsini are the only olethreutines retaining the plesiomorphic connection between the sterigma and the ventral arm of the anterior apophyses and that the same condition is found in Collogenes, together with a shared derived position of both signa very close to the bursa ‘neck’, strongly indicates a close relationship. The little derived juxta-anellus complex of Collogenes further points to a basal position of the genus within the subfamily.
with sexually dimorphic scaling; labial palpi very variable: porrect, very long and sinuate in Collogenes, from small, short and straight to greatly enlarged, sickle-shaped in male of Cryptaspasma. Forewing with or without costal fold, with all veins separate or R3 and R4 stalked, M-stem present, chorda present or absent. Hindwing sometimes with anal fold containing modified scales; Rs and M1 usually close and parallel, rarely stalked; M2 and M3 nearly parallel and distant from each other or moderately approximated at base; M3 and CuA1 from separate to rarely short-stalked; 3A well developed. Thorax with or without posterior crest, in Collogenes with scale pencil from between mesoand metathorax. Legs usually with modified scaling, but never with a tibial pencil. Abdomen frequently with modified scaling, laterally on T5–T7 in some Cryptaspasma, on S3 in some Collogenes. Tegumen with well-developed apodemes for muscles m4. Uncus a small process with apical tuft of strong bristles (some Cryptaspasma) or absent. Socii moderate to small, membranous. Gnathos usually a band, with a median process in some species of both genera. Valva variable, with basal lobe in some Cryptaspasma. Aedeagus with coecum in Cryptaspasma, sometimes with a few large cornuti in Collogenes. Female genitalia characterised by a funnel to pouch-shaped spinulose sterigma laterally linked with anterior apophyses; with two hornshaped signa close to neck of corpus bursae, rarely absent.
Distribution Cryptaspasma is distributed throughout the southern hemisphere (South America, Africa, Madagascar, Australia, New Zealand and New Caledonia), the Oriental and Papuan regions and extends into the eastern Palaearctic region. Collogenes is known from Java across New Guinea to north-eastern Australia and Vanuatu.
Biology The larvae of all reared Microcorsini are borers in fruit and large nuts.
Diagnosis
Remarks
Microcorsini are medium-sized to large dark olethreutine moths with sparse wing pattern, brown to bluish black. Head and patagia sometimes
The fact that the Microcorsini are considered the earliest extant branch of the Olethreutinae and their occurrence on so many parts of Gondwana
120503•Olethreutine Moths 4pp.in67 67
67
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68
OLETHREUTINE MOTHS OF AUSTRALIA
makes them a possible candidate for an original southern distribution.
Cryptaspasma Walsingham
Bakia Koçak, 1981, Priamus 1: 117. Type species: Idiomorpha reticulata Turner, 1946, Trans. R. Soc. S. Aust. 70(2): 213. [Australia.] Established as an unnecessary replacement name for Idiomorpha Turner, 1946.
(Figs 23, 123–135)
Diagnosis Cryptaspasma Walsingham, 1900, Ann. Mag. Nat. Hist. (7.s.) 5: 462. Type species: Notocelia helota Meyrick, 1905: J. Bombay Nat. Hist. Soc. 16: 586, as Penthina lugubris sensu Walsingham, 1900, nec Felder & Rogenhofer, 1875, Reise öst. Fregatte Novara (Zool.) 2 (Abt. 2): pl. 138, fig. 32, by original designation. [Sri Lanka.] Brachygonia Walsingham, 1900, Ann. Mag. Nat. Hist (7.s.) 5: 464. Type species: Brachygonia angulicostana Walsingham, 1900, ibid. 5: 464, by monotypy and original designation. [Japan.] A junior homonym of Brachygonia Kirby, 1889 (Insecta, Odonata). Microcorses Walsingham, 1900, Ann. Mag. Nat. Hist. (7.s.) 5: 465. Type species: Microcorses marginifasciatus Walsingham, 1900, ibid. 5: 466, by monotypy and original designation. [Japan.] Allobrachygonia Fernald, 1908, Genera Tortricidae Types: 62. Type species: Brachygonia angulicostana Walsingham, 1900, Ann. Mag. Nat. Hist. (7.s.) 5 : 464, by original designation. [Japan.] Established as objective replacement name for Brachygonia Walsingham, 1900. Acharneodes Meyrick, 1926, Ann. S. Afr. Mus. 23: 327. Type species: Notocelia helota Meyrick, 1905, J. Bombay Nat. Hist. Soc. 16: 586, by original designation. [Sri Lanka.] A junior objective synonym of Cryptaspasma Walsingham, 1900. Idiomorpha Turner, 1946, Trans. R. Soc. S. Aust. 70(2): 213. Type species: Idiomorpha reticulata Turner, 1946, ibid. 70(2): 213, by monotypy. [Australia.] A junior homonym of Idiomorpha Förster, 1869 (Insecta, Hymenoptera). Anaphorodes Diakonoff, 1959a, Zool. Verh. (Leiden) 43: 6. Type species: Olethreutes anaphorana Walsingham, 1914. [Panama.] Established as subgenus for Cryptaspasma Walsingham, 1900. Metaspasma Diakonoff, 1959a, Zool. Verh. (Leiden) 43: 37. Type species: Acharneodes atrinodis Meyrick, 1926. [Kenya.] Established as subgenus of Cryptaspasma Walsingham, 1900.
120503•Olethreutine Moths 4pp.in68 68
Cryptaspasma species are medium-sized to very large dark grey to blackish brown moths, often mottled darker, in particular with an invertedtriangular blackish mark in centre of wing and/or suffused with ochreous, especially on dorsum, and nearly always with a minute whitish spot in centre of wing over closing vein. Some species of Collogenes Meyrick and Aterpia Meyrick with similar colouring and wing shape look deceptively alike, but Collogenes is readily recognised by stalked R3 and R4 in the forewing and slender palpi and in Aterpia CuA1 originates from well before the angle of the cell. Head and sometimes also thorax are usually sexually dimorphic with labial palpi diverse in male, dense and smoothly scaled and ranging from short, slender and straight to small, upcurved and appressed but with prominent round terminal segment (C. brachyptycha (Meyrick)) to sinuate with distally dilated second segment (C. sordida (Turner)) or greatly enlarged and horn-shaped in some Oriental species. Cryptaspasma has unusually short antennae with the basal part often expanded in male (C. brachyptycha). The wing venation of Cryptaspasma is unremarkable for an olethreutine with all veins separate except for M3 and CuA1 in the hindwing, which are usually separate or connate but very rarely stalked (not in Australian fauna). Males often have a costal fold, sometimes with a conspicuous anteriorly projecting scale brush on its outer surface (C. brachyptycha) and in most species their hindwing anal area is folded upwardly (along 3A in C. sordida), concealing a cluster of modified scales. The hind tibia never has a hairpencil in the male but the terminal abdominal segments often have dorsolateral tufts of upcurved scales meeting over the abdomen. The male genitalia are diagnostic for the genus with uncus usually ending in a tuft of long bristles and often crescentic and inflated valvae (all Australian species) with a large basal lobe, ending in a single spike in Australian species. The female genitalia with unmodified ovipositor lobes, a large funnelshaped sterigma with spinulose lamella antevaginalis connected with anterior apophyses, a short wide and partly sclerotised ductus bursae and two large hollow horn-shaped signa with
5/5/06 2:39:40 PM
AUSTRALIAN OLETHREUTINE GENERA
strongly curved and drawn-out tips in all Australian species are equally characteristic.
Description Adults (Figs 123–126) Wingspan. 12–27 mm (Australian species); medium-sized to very large, size very variable in C. sordida Turner. Head (Figs 127, 128). Ocellus well developed and globular. Vertex scaling short, anteriorly barely protruding between antennae. Frons often with sexually dimorphic and/or species-specific scaling, frequently with very dense, erect and moderately long scales over entire frons, most strongly projecting at midheight of eye; C. sordida with transverse band of anteriorly projecting scales with raised tips in uppermost part, remainder with very short, closely appressed and upwardly oriented scales. Maxillary palpus in C. sordida a minute, two-segmented and unscaled vestige. Proboscis short, no longer than labial palpus. Labial palpus very diverse in shape and size (1.2–2.0× diameter of eye), often sexually dimorphic with very dense, smooth scaling in male, ranging from short, straight, distally hardly dilated and obliquely raised (C. lugubris) to huge and sickle-shaped (Diakonoff 1959a) or upcurved, closely appressed, short and of even width but with a large, thickly scaled and roundish terminal segment (C. brachyptycha, Fig. 128); in males of some species (C. sordida, Fig. 127) and probably all females the labial palpus sinuate with second segment distally dilated and terminal segment small and downward pointing, often concealed. Antenna short, reaching to about 56: costa or shorter, in male moderately thickened and especially basal part often conspicuously expanded and flattened (C. brachyptycha, Fig. 128), anterior surface unscaled; cilia in both sexes minute. Thorax. Without posterior crest in Australian species but sexually dimorphic in C. brachyptycha with patagia in male covered with dense bundles of erect scales instead of a flat ‘collar’ and thorax lengthened by a broad band of posteriorly projecting long and lightly curled scales, less pronounced in female. Legs usually not sexually dimorphic but in male of C. brachyptycha fore and mid tibia with very long and dense scales and triangularly expanded by a dense fan of long scales from its lateral surface. Wings (Figs 23, 123–126). Forewing index 0.40–0.45. Forewing usually elongate subtriangular, widening towards termen; with or without costal fold in male, usually filled with modified scales
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69
(C. sordida), in C. brachyptycha with an anteriorly projecting large clump of dense erect scales on outer surface of costal fold (Figs 125, 128); costa lightly curved throughout (straight in basal half in males with costal fold), apex rounded-acute, termen oblique; discal cell 0.6× length of wing, with costal margin curved in distal half and accessory cell, if present, lens-shaped, with chorda and M-stem both present but non-tubular in C. sordida with chorda from between R1 and R2 (closer to R1) to between R4 and R5, M-stem to between M2 and M3, but both these veins vestigial in some species; all veins present and separate beyond discal cell; anal loop 0.19× length of 1A+2A; dark grey to blackish brown, often mottled darker, in particular with an inverted-triangular blackish mark in centre of wing and an oblique darker zig-zag line parallel to termen and/or suffused with ochreous on dorsum, in centre of wing beyond dark triangle and along termen; usually with a minute white or pale spot in centre of wing over closing vein. Hindwing with cubital pecten and axillary tuft present in females and most males, cubital pecten absent in male of C. angulicostana but particularly strongly developed in the male of an unnamed Australian species, as long scales partly arising from behind cubital vein and covered by base of forewing dorsum; males of several species with patches of modified sex scales in anal area (C. pullatana Bradley); anal margin in males folded upwardly and closely appressed, varying from a very small fold at base of dorsum to a large, deep, and semicircular pocket with foldline along 3A in C. sordida (Figs 133, 134), and often containing modified scales of diverse shape and colour (very short, broad and red-brown in C. sordida (Fig. 135), blackish in C. brachyptycha); with Rs and M1 closely approximated and parallel at base, M2 bent moderately towards M3 at base and CuA1 from just before angle of cell (distinctly separate from M3) to connate or rarely stalked with M3 (Diakonoff 1959a); CuP present only near wing margin, 1A+2A and 3A both well developed. Pregenital abdomen. Male with conspicuously modified scaling on posterior part of abdomen based on very long tufts of upwardly curved spatulate scales meeting over abdomen and arising from crescentic patches laterally on T5–T7 (or only on T7). Eighth segment in male large but weakly sclerotised except for a transverse band at base of T8, with very dense, long, spatulate scales dorsally and laterally and a small area of short scales ventrally on hind margin; genitalia usually entirely retracted into 8th segment with a longitudinally striate intersegmental membrane.
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Figs 123–128. Cryptaspasma, adults and heads. 123, 124, C. sordida, Dorrigo Nat. Park, NSW. 123, [image reversed]. 124, [image reversed]. 125, 126, C. brachyptycha. 125, , East Base Mt Bartle Frere, Q [image reversed]. 126, , West Normanby R. Crossing, Q. 127, head, C. sordida, male. 128, head, C. brachyptycha, male. Scale = 2 mm.
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Male genitalia (Figs 129, 131). Vinculum a weakly sclerotised band. Tegumen broad and rounded, weakly sclerotised, pedunculi broad with large triangular apodemes for muscle m4. Uncus a moderately sclerotised small median process, dorsoventrally flattened, downcurved and with a usually bipartite tuft of long bristles or scales projecting from its apex (absent in Sg. Metaspasma), often with two lateral ear-like projections connected to its base (C. brachyptycha). Socii from well-developed, rounded small drooping and hairy lobes to a group of bristles in membrane ventrolaterally to uncus, or entirely lacking. Gnathos a thin curved band (Sg. Microcorses, Sg. Cryptaspasma), with a small distal hook in Microcorses, or not developed. Transtilla absent. Valva ranging from flat and distally bipartite with a projecting sacculus lobe and a pad-shaped densely bristled basal lobe (pulvinus) (Sg. Anaphorodes) to crescentic and strongly inflated, without distinct sacculus but with a large flat erect basal lobe (pulvinus) crowned with a single spike and usually with blunt spines along inner margin of distal portion of valva (Sg. Allobrachygonia); with short and broad costal process and small basal excavation. Juxta and caulis fused, but caulis short; anellus a weakly sclerotised ring. Aedeagus large and tapering towards apex, with a small, but distinct, coecum; vesica without cornuti. Female genitalia (Figs 130, 132). Ovipositor lobes elongate oval to reniform, apices bluntly rounded. Sternum 7 with weakly concave hindmargin. Sterigma a large wide funnel with lamella antevaginalis forming a strongly projecting spinulose lip, sometimes medially cleft (Sg. Cryptaspasma) and/or with scattered long bristles from margin, laterally continuous with ventral arm of anterior apophyses. Ductus bursae short and rather wide, without typical colliculum but posterior half strongly tapering from its widest point at origin of ductus seminalis to below ostium and partly sclerotised, anterior half membranous; ductus seminalis with bulla well developed. Corpus bursae ovate; signa two large hollow horns originating from close to bursa ‘neck’, rather straight and distally rounded or pointed in Sg. Cryptaspasma, of unequal size and with a strongly curved and drawn out apex and a longitudinal crest on base of larger signum in Sg. Allobrachygonia.
Distribution Cryptaspasma is distributed throughout the southern hemisphere (South America, Africa,
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Madagascar, Australia, New Zealand and New Caledonia), the Oriental and Papuan regions and extends into the Palaearctic region in Japan and easternmost Russia. In Australia the genus is so far known from the Northern Territory, Queensland, New South Wales, the Australian Capital Territory and one specimen each from Western Australia (?Dampier) and Tasmania.
Biology The few species with host records have all been reared from fruits or large seeds, boring in the endosperm: C. (Microcorses) trigonana (Walsingham) from acorns of Quercus mongolica (Fagaceae) in Russia (Kuznetsov 1970); the New Zealand C. (Allobrachygonia) querula (Meyrick) from kernels of stone fruit, fruit of Beilschmiedia (Lauraceae) and Hedycarya (Monimiaceae) (Dugdale personal communication); the Australian C. sordida from fruit of Acmena ingens (Myrtaceae), nuts of Macadamia sp. (Proteaceae), and by T. D. Male in Cooloola from fruit of Endiandra sieberi (Lauraceae), and C. brachyptycha from fruit of Calamus sp. (Arecaceae).
Remarks Diakonoff (1959a) provided a comprehensive revision of Cryptaspasma, reviewing previous literature and dividing the genus into five subgenera, giving precedence to Cryptaspasma over the simultaneously described Microcorses. He figured head and wing venation for the genus and the genitalia of numerous species. Kuznetsov (1970) redescribed and figured Microcorses mirabilis Kuznetsov, raised the group to tribal level and discussed its relationships and taxonomy, treating Diakonoff’s (1959a) subgenera as genera. Photos of adults and genitalia of the types of several species were provided by Clarke (1958) and a brief diagnosis for adult, larva and pupa of Cryptaspasma was given by Horak & Brown (1991). Clarke (1976) reviewed and figured the Micronesian Cryptaspasma, describing two new species, and Razowski (1989) summarised Microcorses. Whereas the monophyly of Cryptaspasma s. lat. cannot be doubted, based on the wing pattern, the basal valva lobe, the spinulose funnel-shaped sterigma connected to the anterior apophyses and the two hollow, horn-shaped signa, it would require a study far beyond the scope of this review based primarily on the Australian fauna to establish the monophyly of each of the five subgroups currently
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Figs 129–135. Cryptaspasma, genitalia and wing structure. 129, 130, C. sordida, Lamington Nat. Park, Q. 129, male genitalia, slide T2740. 130, female genitalia, slide ANIC 2769. 131, 132, C. brachyptycha. 131, male genitalia, Kuranda, Q, slide ANIC 2779. 132, female genitalia, Papua New Guinea, slide ANIC 2770. 133–135, C. sordida, male hindwing, anal region with modified scales, progressively enlarged.
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recognised. They are here treated at subgeneric rank, as established by Diakonoff (1959a) and followed in Kawabe (1982) and Kawabe and Komai (1992). The subgenera Cryptaspasma and Anaphorodes with their less derived valva are restricted to South America and Microcorses is found only in the easternmost Palaearctic (Japan and Pacific region of Russia). The African and Malagasy Metaspasma (Diakonoff 1959a) and the Indo-Australian Allobrachygonia (from Sri Lanka to Japan, Australia, New Zealand and New Caledonia) both have an inflated valva with a series of blunt spines or pegs towards its tip but with nevertheless a different structure in each subgenus. All Australian species belong to the subgenus Allobrachygonia, characterised by a strongly swollen crescentic valva with a large single spike on apex of the basal lobe in the male and distally drawn out and strongly curved signa in the female. There is a strong suggestion that Collogenes, whose larvae are also borers, is the sister group of Cryptaspasma, as discussed for the tribe. Most species of Cryptaspasma are superficially exceedingly similar but secondary sexual characters such as labial palpus shape, modification of antennal base or patagia, length of costal fold, shape of hindwing anal fold, and structure and colour of enclosed scale cluster together with the development of modified scale patches over anal area usually allow discrimination at least in the male. Meyrick’s (1911b) report of C. helota (Meyrick) from Australia (Mt Tamborine) must be based on a misidentification of C. sordida. The two species are very similar, but different labial palpus shape and the strongly tapering valva apex of C. sordida, allow separation.
Constituent species Thirty-five named species of Cryptaspasma are known worldwide, including one from Norfolk I. (Cryptaspasma (Allobrachygonia) pullatana Bradley, 1982) and two from Australia, with an additional unidentified Australian species: sordida (Turner, 1945), Trans. R. Soc. S. Aust. 69(1): 65 (Tortrix). [Qld: Macpherson Range.] reticulata (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 213 (Idiomorpha). [Qld: Toowoomba.] brachyptycha (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 246 (Eucosma). [Qld: Cooktown.]
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Collogenes Meyrick (Figs 24, 136–147) Collogenes Meyrick, 1931, Exot. Microlepid. 4: 141. Type species: Collogenes percnophylla Meyrick, 1931, ibid. 4: 142, by monotypy. [Bismarck Arch., New Hannover.] Mesocallyntera Diakonoff, 1959a, Zool. Verh. (Leiden) 43: 47. Type species: Mesocallyntera squamosa Diakonoff, 1959a, ibid. 43: 49, pl. 10, fig. 83; pl. 11, fig. 85, by original designation. [Sulawesi.] Syn. nov.
Diagnosis Species of Collogenes are medium-sized to very large dark brownish greyish moths with broad, subtriangular forewings without costal fold in male, indistinctly mottled throughout, usually with greyish blue areas at least in tornus and often with an indistinct, wide, outwardly oblique pale band from costa near its base. Thinly scaled forewings with denser scaling along veins and with stalked R3 and R4, strongly modified male mid and hind tibia with an associated hairpencil from the thorax and very long, slender, sinuate labial palpi in the female are diagnostic for Collogenes. The male has similar but shorter labial palpi. Basal half of forewing costa thickened, R4 to apex, R5 close to stalk of R3 and R4 at its base, and M3 and CuA1 originating reasonably close together from angle of cell. The stalked R3 and R4 in Collogenes separate the genus from Cryptaspasma with a similar wing colour. The axillary tuft in the hindwing often extends along 1A+2A in both sexes and the enlarged anal area in the male has slightly modified scaling, Rs and M1 are closely approximate to connate at base and the base of M2 is roughly parallel and distant from the base of M3, which is connate with CuA1. The male genitalia of Collogenes are characterised by usually well-developed apodemes from their pedunculi, a variably developed membranous tegumen tip with scattered bristles and/or with 1–2 pairs of large sublateral bristles, medially paired socii lobes or illdefined clumps of bristles, a well sclerotised, medially constricted valva with a large basal excavation, a large, triangular costal process (sometimes partly membranous) and the cucullus either a simple, angled lobe or deeply bilobed with large spines on its inner surface, and a basally curved aedeagus often with three large, persistent cornuti. The female genitalia are characterised by
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broadly rounded ovipositor lobes often with a longitudinal row of peg-shaped papillae, the ventral arms of the anterior apophyses connected to the spinulose, U-shaped sterigma, a variably developed, tube-shaped ductus bursae with the ductus seminalis originating from close to the ostium and by two hollow, lightly curved, horn-shaped signa close to the ductus bursae, if present.
Description Adults (Figs 136–139) Wingspan. 13.5–29.0 mm; medium-sized to very large. Head (Figs 140, 141). Ocellus well developed. Vertex with shortish scales, hardly projecting between antennae; scales on upper part of frons lightly projecting, remainder of frons minutely scaled. Labial palpus strongly sinuate, diagonally rising except for base and porrect tip, moderately long in male (1.7–2.0× diameter of eye), very long in female (2.5–2.7× diameter of eye), rather slender, second segment sinuate, slender except for some longer, loose scaling either near base (C. pseusta Diakonoff) or distally along dorsal margin and apex (C. squamosa), terminal segment long, slender, porrect, even longer in female. Antenna to about middle of wing, weakly scaled in a dorsal band only in both sexes; with dense, minute cilia. Thorax. With a large posterior crest and in male with a pencil of very long, angled, yellowish hairscales laterally from between meso- and metathorax, presumably associated with modified tibial scaling. Legs strongly modified in male, fore tibia dorsally with longer, loose but unmodified scales, mid tibia hugely enlarged by modified, dark, dense, long, spatulate scales along entire dorsal surface, and hind tibia with a large tuft of long, yellowish hair-scales basally on dorsal surface; mid tibia with somewhat modified scaling even in female. Wings (Figs 24, 136–139). Forewing index 0.44–0.48. Forewing broadly subtriangular, widest before tornus, with costa straight except at base and near apex, apex rounded-acute, termen moderately to strongly oblique, tornus obliquely rounded; without costal fold, overall thinly scaled, more densely along veins, with glossy but not metallic scales scattered throughout; discal cell with curved anterior margin and distally strongly oblique, without any trace of a chorda and with Mstem to M2, in C. loricata Diakonoff distally weak and adjacent to hindmargin of cell from well below
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origin of CuA2; basal half of costa with vein-like thickening along margin, Sc curved in basal half then straight and close to costa, R1, R2 and stalk of R3 and R4 about equidistant, R3 and R4 stalked to beyond 56: with R4 to apex, R5 at base close to stalk of R3 and R4 and more distant from M1; M2 widely separate from M1 and moderately close to M3; M3 and CuA1 reasonably close from angle of cell, CuA2 from just beyond 6M of cell; CuP present at margin; anal loop very narrow, 0.29× length of 1A+2A; variably dark brownish grey, indistinctly mottled with dark greyish blue, especially an ovate-erect area in tornal region often reaching to beyond middle of wing, often with an indistinct, wide, outwardly oblique pale band from beyond base of costa to before middle of dorsum, usually not reaching dorsum, costal strigulae usually hardly developed except for a small, crescentic, reddish brown mark in apex, sometimes with similar, parallel marks on costa and termen near apex, with veins outlined by denser, darker scaling. Hindwing with cubital pecten present and anal tuft extending along 1A+2A in both sexes to a variable extent, up to 6M to margin, in male with slightly larger, more rounded scales above and below on extended anal region; R-stem only developed in distal half of cell, Rs and M1 closely approximate to connate at base, base of M2 roughly parallel and distant from base of M3, M3 and CuA1 connate from angle of cell, CuA2 from just below 36: cell; CuP well developed, 1A+2A and 3A present, with 3A halfway between 1A+2A and the anal margin; greyish. Pregenital abdomen. Some species (C. squamosa, C. loricata) with S3 crescentic and with modified, persistent scaling in male. Eighth segment in male short, dorsally strongly convex and ventrally concave; membrane linking 8th segment and genitalia flimsy. Male genitalia (Figs 142, 143, 146, 147). Tegumen weakly sclerotised, rounded to subtriangular, laterally a narrow band, apex a low and wide or narrow and high membranous triangle, variably bristled and often with 1–2 pairs of very large sublateral bristles, but no well-defined uncus; pedunculi nearly always with well-developed apodemes, often broadly rounded lobes. Socii from well-developed, medially paired lobes with outer surface densely bristled (C. squamosa, C. loricata) to ill-defined, sublateral clumps of bristles at ventral edge of scattered apical bristles on tegumen (C. pseusta, C. albocingulata, sp. nov.). Gnathos usually weak, membranous lateral arms, rarely with a lightly sclerotised medial process (C. pseusta).
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Figs 136–141. Collogenes, adults and heads. 136, 137, C. loricata. 136, , Mt Bellenden-Ker [image reversed], Q. 137, , Mt Lewis near Mt Molloy, Q. 138, 139, C. albocingulata, paratypes. 138, , The Crater, nr Atherton, Q. 139, , Davies Ck nr Mareeba, Q. 140, 141, head, C. albocingulata. 140, male. 141, female. Scale = 2 mm.
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Valva well sclerotised, variably developed, elongate and medially constricted, always with a large, basal excavation; costal process large, elongate-triangular (C. squamosa), sometimes membranous except for tip (C. pseusta); distal half of valva from a simple, angled lobe fairly evenly bristled on inner surface to deeply bilobed with large, long spines on inner surface, generally shorter and more dense on ventral lobe. Juxta a simple, triangular plate, caulis and anellus a contiguous, broad band. Aedeagus curved in basal half, from a shortish, parallel-sided tube with short teeth on outer surface near apex (C. albocingulata, sp. nov.) but without cornuti, to very long, strongly curved, distally attenuate, with bevelled apex and three large, curved, persistent cornuti (C. squamosa). Female genitalia (Figs 144, 145). Ovipositor lobes very wide with anterior lobe broadly rounded, often with a longitudinal, submedial row of 3–10 strongly enlarged, peg-shaped papillae. Sterigma in intersegmental membrane posterior to weakly sclerotised S7, laterally joined to ventral arm of anterior apophyses (not apparent in C. squamosa and C. loricata where sterigma is membranous), if well developed a large U-shaped spinulose ridge surrounding deep depression leading to ostium, with projecting ventral lip in C. albocingulata, sp. nov., but reduced and entirely membranous in C. squamosa and C. loricata. Ductus bursae a straight tube with the short portion between ostium and origin of ductus seminalis always unsclerotised but often swollen and extremely thick-walled, remainder of ductus bursae either membranous or sclerotised; corpus bursae roundish, with or without two hollow, lightly curved, horn-shaped signa close to ductus bursae.
Distribution Collogenes is known from Java, Sulawesi, New Guinea, the Bismarck Archipelago, Vanuatu and Australia where it occurs in Queensland on the east coast from the tip of Cape York to Lamington NP south of Brisbane.
Biology Collogenes loricata has been reared twice from larvae boring in the seed of Endiandra sieberi (Lauraceae) at Rainbow Beach, Queensland, by T. D. Male, confirming the assumption based on adults with greasy abdomens that Collogenes larvae are endosperm borers.
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Remarks Diakonoff (1959a) described both Collogenes and Mesocallyntera in great detail, providing figures of heads, wing venation, legs and genitalia. He remarked in the original description of Mesocallyntera (Diakonoff 1959a) that the genus could not be distinguished from Collogenes on external features. Given the species then known for the two genera, Mesocallyntera was characterised by the presence of well-developed socii, a distally bifurcate valva, ovipositor lobes with a series of peg-like papillae along their middle, a membranous ductus seminalis with a swollen and thick-walled posterior portion and absence of signa. Since then, species have become available that combine the characteristics of both groups and confirm that Mesocallyntera is merely a derived, subordinate group within Collogenes. The valva of Mesocallyntera dascia Bradley from the Vanuatu (Bradley 1962) has a structure intermediate between the two extremes: not bilobed but with well-developed spines. The Australian C. albocingulata, sp. nov. has overall Collogenes-type genitalia without well-developed socii; with a simple valva; short aedeagus without cornuti; welldeveloped sterigma and signa, but with a series of peg-like papillae along the ovipositor lobe. The monophyly of Collogenes is based on a number of unique apomorphies including the sparsely scaled forewing with stalked R3 and R4, a hindwing with a pecten extending along 1A+2A and with M2 and M3 distant and parallel at base, slender, sinuate labial palpi greatly lengthened in the female, strongly modified mid and hind tibia in the male associated with a hairpencil from the thorax, a modified tegumen tip without a distinct uncus, a valva with a very large basal excavation and a large, triangular costal process, and the ductus seminalis originating from close to the ostium. No obvious sister group is available for Collogenes, but a number of derived features shared with Cryptaspasma (including the shape of the sterigma, the position of both signa close to the bursa ‘neck’, and the fruit boring larva) together with the plesiomorphic connection between sterigma and anterior apophysis strongly suggest a position in the Microcorsini (see tribal discussion). Diakonoff’s (1959a) study of Mesocallyntera contains some inconsistencies. The type locality of Mesocallyntera siderostola Diakonoff is given as ‘Netherlands New Guinea, Snow Range, Araucaria Camp, 800 m, 17.iii.1939 (L. J. Toxopeus, Third Archbold Expedition)’, but the single female
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holotype in the RMNH bears a handwritten label ‘Ceram, 800 m, 17.iii.1939, Tox[opeus]’. The handwritten label must be in error as the date given coincides with Toxopeus’ collecting activities at the Araucaria Camp during the Third Archbold Expedition to Central Netherlands New Guinea.
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Also, the figures and description given for the male genitalia of C. squamosa squamosa Diakonoff and C. squamosa loricata Diakonoff seem to have been mixed up. Comparison with the relevant slide preparation shows that C. loricata has the shorter, thicker aedeagus, a distinction confirmed by more
Figs 142–147. Collogenes, genitalia. 142–144, C. albocingulata. 142, 143, male genitalia with aedeagus, holotype, slide T2329. 144, female genitalia, paratype, The Crater, Q, slide ANIC 16870. 145–147, C. loricata, Mt Lewis nr Mt Molloy, Q. 145, female genitalia, slide T2247. 146, 147, male genitalia and aedeagus, slide T2246.
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recent material of the two species from Sulawesi, the type locality of C. squamosa, and from Australia. This more comprehensive material, comprising both sexes, leads to the conclusion that C. squamosa loricata, based on a single male from Papua (former British New Guinea), and C. siderostola Diakonoff, based on a female from West Irian (former Dutch New Guinea), are conspecific and distinct from C. squamosa at the species level. The following two species are here referred to Collogenes: squamosa (Diakonoff, 1959a), Zool. Verh. (Leiden) 43: 49 (Mesocallyntera). [Sulawesi.] Comb. nov. dascia (Bradley, 1962), Bull. Br. Mus. (Nat. Hist.) Entomol. 12: 254 (Mesocallyntera). [Vanuatu.] Comb. nov.
from all other Collogenes species that are indistinctly mottled dark greyish brown, but it shares the sparse and glossy scales characteristic for the genus. In the male genitalia, the long and slender valva, only weakly constricted submedially and with an obliquely truncate apex, and the rather short, wide and parallel-sided aedeagus are diagnostic. The female genitalia are characterised by a spinulose sterigma with a protruding ventral lip, a strongly sclerotised, wide and parallel-sided ductus bursae and two signa slender also at base. According to the male genitalia, C. albocingulata is closest to C. percnophylla from Papua New Guinea (New Hanover), the type species of Collogenes, which, however, has completely dark forewings.
Description Constituent species Five named species of Collogenes are known from outside Australia, and one of these described from New Guinea and one described below are found in Australia: loricata (Diakonoff, 1959a), Zool. Verh. (Leiden) 43: 50 (Mesocallyntera). [Papua New Guinea.] [Established as subspecies of C. squamosa Diakonoff, 1959a.] Comb. nov., stat. nov. siderostola (Diakonoff, 1959a), Zool. Verh. (Leiden) 43: 51 (Mesocallyntera). [Irian Jaya.] Syn. nov. sp. A; Horak et al. 1996, Monogr. Aust. Lepid. 4: 135 (Mesocallyntera). albocingulata, sp. nov.
Collogenes albocingulata, sp. nov. (Figs 138–144)
Material examined Holotype. , ‘The Crater, Atherton Tableland, Qld, 3150 ft [960 m], 4 Dec. 1967, R. Dobson’, ‘Genitalia slide T2329’ (ANIC). Paratypes. Queensland: 2 , 4, same label as holotype; 17 , 6, 17°26´S 145°29´E, The Crater, 18 km S Atherton, 980 m, 24.ii.1968, 27.ii. & 17. & 21.iii.1998, D. C. F. Rentz, E. D. E., M. Horak; 1, 17°02´S 145°37´E, Davies Ck, 21 km E by S of Mareeba, 1.iii.1988, D. C. F. Rentz (ANIC, QM, BMNH).
Diagnosis With the conspicuous oblique white band across the forewing, C. albocingulata is unique and differs
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Male (Figs 138, 140) Wingspan 13.5–15.0 mm. Head brownish grey, second segment of labial palpi laterally brownish orange (Fig. 138). Thorax and tegulae of mixed white and dark grey-brown scales (unless greasy), tegulae darker in basal half. Lateral hair-tuft from base of hind coxa creamy, hind and especially mid tibia with dense, large, loose creamy scales on inner surface, forming a spreading tuft at base of hind tibia. Forewings rather thinly scaled, with conspicuous gloss; mottled greyish with a variably broad, ill-defined oblique white band from costa beyond base to middle of dorsum; wing from base to white band mottled with grey and blackish scales and beyond white band sprinkled with orange scales followed by silvery grey suffusion except near partially white termen, with orange scales in a crescent around apex and a few below middle of termen and along veins beyond cell; pairs of fine white strigulae along entire costa. Fringe whitish with some orange and few grey scales. Hindwings pale silver grey, fringe slightly paler. Abdomen grey to dark grey with pale anal tuft, often oily. Male genitalia (Figs 142, 143). Tegumen semiovate, weakly sclerotised; pedunculi narrow with a long thin muscle attachment. Uncus a nearly membranous, medially notched bristled hump. Gnathos two weakly developed, small, curved lateral bands. Valva well sclerotised, narrowly subrectangular though slightly narrowed beyond middle and widest in basal third, with ventral margin evenly sinuate, cucullus angled with distal margin oblique and lightly rounded towards costa; inner surface evenly bristled throughout with bristle size increasing towards ventral and distal edge. Juxta large, triangular. Caulis and anellus fused into large, concave, inverted teardrop-shaped structure with aedeagus high near its apex.
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Aedeagus rather short, nearly parallel-sided, curved in basal third, with short teeth on outer surface near apex, without cornuti. Female (Figs 139, 141) Wingspan 15–18 mm. As for male; mid and hind tibia with long loose scales but without spreading tuft at base of hind tibia. Female genitalia (Fig. 144). Ovipositor lobes broadly rounded, with a submedial row of three strongly enlarged, peg-shaped papillae. Sterigma sclerotised and strongly spinulose, laterally with conspicuous, sclerotised, spinulose connection to ventral arms of anterior apophyses; lamella antevaginalis a large, transversely projecting spinulose lip. Ductus bursae short and wide, strongly sclerotised throughout, nearly parallelsided but slightly widening near corpus bursae, ductus seminalis originating from just below ostium. Two signa with their base in posterior third of ovate corpus bursae, slender, tapering, curved horns with rugose surface and one with a long, slender capitulum-like structure.
Distribution Except for one specimen from near Mareeba, C. albocingulata has been collected only at ‘The Crater’, Mt Hypipamee NP, on the Atherton Tableland, where it was abundant at light on several visits between December and March.
OLETHREUTINI Walsingham Olethreutinae Walsingham, 1895, Trans. Entomol. Soc. Lond. 1895: 500 (as a subfamily of the Tortricidae). Type genus: Olethreutes Hübner, 1822. Olethreutidii Pierce & Metcalfe, 1922, The genitalia of the Tortricidae: 38. Olethreutini Obraztsov, 1946, Z. Wien. Entomol. Ges. 30: 23 (as a tribus).
Status remarks The tribe Olethreutini, established by Pierce and Metcalfe (1922) and Heinrich (1923) independently for the same grouping, gained immediate acceptance. Falkovitsh (1962) later divided the group into four tribes based on secondary sexual characters, but of these only the tribe Bactrini has been widely accepted as distinct from the remaining Olethreutini. Diakonoff (1973) regarded his Bactrae and Gatesclarkeanae merely as two of
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his twelve subtribes of the Olethreutini. Kuznetzov and Stekolnikov (1984) erected the super-tribe Gatesclarkeanidii for the Gatesclarkeanini and Endotheniini, parallel to the Olethreutidii, which include the Microcorsini, Bactrini and Olethreutini, and the Eucosmidii including the Enarmoninii, Eucosmini and Grapholitini. A slightly alternative classification (Razowski 1989; Horak and Brown 1991; Horak 1998) ranked the Gatesclarkeanini, Bactrini and Olethreutini all parallel at tribal level, but the Gatesclarkeana-group is here included as a genus-group in the Olethreutini.
Evidence for monophyly Pierce and Metcalfe (1922) (the first to isolate a grouping corresponding to the current Olethreutini) defined their Olethreutidii (among other characters) by: a long valva often with spines at the end of the sacculus; large drooping socii; aedeagus with few cornuti or without; a spiny sterigma and the signa, if present, one or two scobinate plates. Heinrich (1923) primarily characterised the Olethreutini (as the subfamily Olethreutinae) by the separate or connate (instead of stalked) M3 and CuA1 in the hindwing, whilst pointing to the few exceptions in the Holarctic fauna. In his second monograph, Heinrich (1926) provided diagnoses for his Olethreutidae, Eucosmidae and Grapholitidae, but they are not decisive, even less so for the Oriental/Australian fauna. There is no apomorphy known to be present in all olethreutine genera, and some of the characters separating the Olethreutini from other tribes, such as connate rather than stalked M3 and CuA1 in the hindwing, are merely plesiomorphies. However, some derived characters are found only in the Olethreutini: a combination of a tibial pencil with a modified, rolled anal area in the hindwing, and one or two spine clusters from or near the sacculus (Spc1 and Spc2 in Heinrich 1926). There is also a strong tendency to a lengthened colliculum but otherwise few sclerotisations of the ductus bursae, and none anteriorly to the origin of the ductus seminalis except in the Oxysemaphora-group. The tibial pencil is present in some species of all genus-groups except the Oxysemaphora-group, and it is also present in some Endothenia species, i.e. some Bactrini. A tibial pencil is also found in three genera of the Enarmoniini (Anathamna Meyrick, Metaselena Diakonoff and some species of Loboschiza Diakonoff), but never associated with a rolled anal margin of the hindwing. Both Anathamna and Metaselena have modified,
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apparently glandular scales below on the base of the abdomen, presumably forming a functional unit together with the tibial pencil. The tibial structure in Loboschiza is quite different and obviously not homologous, it is a dorsal, eversible tuft rather than a parallel pencil. As to the spine clusters on the valva, it is difficult, if not impossible, to establish homologies unless a transformation series is available, and no attempt has been made to identify Spc1 and Spc2 in this study. Among the Australian Olethreutini, only Demeijerella Diakonoff with its highly derived valva lacks a spine tuft or single large spine near the sacculus, but the structure is well developed in the other genera of the Sycacanthagroup. Prominent spines are present also in the Eucosmini and Enarmoniini, but they appear to be derived from the marginal cucullus spines (Enarmoniini, Spilonota-group) or are located on the outer surface of the valva (Acroclita-group).
Diagnosis Olethreutini are small to quite large moths with very diverse wing pattern, usually with welldeveloped costal strigulae but without ocelloid patch. Forewing without costal fold; chorda and Mstem nearly always present, R4 and R5 always separate. Hindwing in male often with modified anal region: frequently an anal roll filled with glandular scales and associated with a tibial pencil, more rarely tufts of long, modified scales (Oxysemaphora Diakonoff, Cnecidophora, gen. nov., Sorolopha Lower); sometimes modified scales in cell or across centre of wing (Eremas Turner, Statherotis Meyrick); Rs and M1 nearly always parallel and closely approximated at base, but diverging from base in Eremas, and stalked in Cnecidophora, gen. nov. and some Lobesia and Metrioglypha species; M2 at base usually curved towards and closely approximated to M3, but in Oxysemaphora, Demeijerella Diakonoff, Eremas and Proschistis Meyrick curved towards M3 but not closely approximated, and M2 and M3 distant and nearly parallel in Cnecidophora, gen. nov.; M3 and CuA1 usually close and separate, to (at most) very shortstalked, significantly stalked only in Cnecidophora, gen. nov., Rhectogonia Diakonoff, Eremas, Diakonoffiana Koçak and Costosa Diakonoff, and widely distant with CuA1 from well before angle of cell in Aterpia Guenée; 3A always present. Thorax nearly always with posterior crest (absent in Atriscripta, gen. nov., some Statherotis, Aterpia and Rhodacra Diakonoff). Hind tibia in male often with pencil. Abdomen with S2 of tortricoid type with
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anterolateral processes and a pair of apodemes, without any traces of sternal rods; often with abdominal pouches or scale brushes in male. Male genitalia usually characterised by one or two spine clusters near the sacculus and a narrow cucullus. Tegumen with apodemes for muscles m4 usually moderately developed. Uncus usually present, rarely paired (some Megalota Diakonoff and Costosa), frequently ventrally hairy or bristled. Socii nearly always present, but frequently small. Gnathos variably sclerotised, usually spinulose in the Olethreutes-group. Valva with one or two spine clusters near sacculus, usually with narrow cucullus, with costal process usually two-branched. Aedeagus rarely with small coecum (Oxysemaphora, Cnecidophora, gen. nov.); cornuti diverse, from large sheaf to few to often absent. Female genitalia with a spinulose sterigma, rarely fused with S7; with an often very long colliculum but no sclerotisation in anterior half of ductus bursa (except Oxysemaphora-group). Signa paired, less frequently single, varying from horn-shaped to scobinate and various derivations thereof.
Distribution The Olethreutini are present world-wide, with their greatest diversity in the Oriental and Holarctic regions.
Biology The larvae of the Olethreutini are nearly all leaf rollers, but boring larvae have been found in two genera of two different genus-groups (Lobesia, Ophiorrhabda Diakonoff).
Remarks There are numerous chapters on the Olethreutini in geographically localised publications. The more comprehensive treatments range from the still very relevant revision of the Nearctic fauna by Heinrich (1926), Hannemann’s (1961) diagnostically illustrated fauna of Germany, to Bentinck and Diakonoff’s (1968) fauna of the Netherlands, and the volume by Bradley et al. (1979) detailing the biology of British species with exquisite drawings. Diakonoff’s (1973) monumental monograph on the South Asiatic Olethreutini was the basis for the present study of the Australian taxa, though the absence of figures of adults makes it difficult to use for non-taxonomists. More recently, Kuznetzov (1978) provided a revision of the fauna of the
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European part of the former USSR, illustrated with excellent drawings, including wing venations. Razowski (1989) published a synopsis of the Palaearctic Olethreutinae with genitalia drawings of both sexes and a comprehensive treatment of the European olethreutine fauna (Razowski 2003a), with colour photos of all species and genitalia drawings of high quality. Kawabe (1982) provided photos of adults, in colour, and of the genitalia of some species, and Liu and Li (2002) illustrated adults and genitalia of both sexes for the Chinese fauna. Diakonoff (1973) divided his Olethreutini into twelve subtribes, including Gatesclarkeanae, Bactrae and Endotheniae, mainly on the basis of signum structure. It clearly was an inspired decision, and the majority of these groups have stood the test of time as discrete units once a few unrelated elements, mainly Enarmoniini, were removed. They have been accepted as subtribes by Kuznetzov and Stekolnikov (1984), albeit with their names emended, but not by Razowski (1989), except for the Endotheniina and Lobesiina. However, it has yet to be demonstrated to what extent these subtribes are monophyletic entities and what their relative ranks should be. For this reason, apart from the Bactrini (including Endothenia), they are here treated more informally as genusgroups, with some merged and the Oxysemaphoragroup added. Heinrich (1926) defined three distinct spine clusters on the valva of the Olethreutini, sacculus spines (ScSp) on the base of the sacculus and two spine clusters (Spc1, Spc2) from or near the sacculus. Though both Hannemann (1961) and Diakonoff (1973) followed this terminology, the homologies between the various spine groups are far from clear and the spines are here characterised in a purely descriptive manner.
Oxysemaphora-group Status remarks This genus-group is here newly defined for the genera Oxysemaphora Diakonoff, Dynatorhabda Diakonoff, Cnecidophora, gen. nov., and possibly Episimoides Diakonoff from Réunion and Episimus Walsingham in the New World. With only the male known, Diakonoff included Oxysemaphora in the subtribe Olethreutae. Dynatorhabda was included in Gnathmocerodides, presumably on the strength of the single horn-shaped signum.
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Evidence for monophyly Some of the conspicuous similarity in the genitalia of the Oxysemaphora-group is due to symplesiomorphies, such as the long slender uncus, the often very large socii, the usually large, hornshaped signa and the simple, spiny sterigma. However, several synapomorphies suggest that it is indeed a monophyletic group: 1, large spine from lower margin of valva (absent in Episimoides and a tuft of spines in Episimus); 2, ring-shaped sclerite in ductus bursae posterior of origin of ductus seminalis (often extended into sclerotisation anterior to ductus seminalis); 3, a very long caulis, and 4, large costal processes in Oxysemaphora, Cnecidophora, gen. nov., Dynatorhabda and Episimoides.
Distribution Cnecidophora, gen. nov. is known from New Guinea, Australia and several undescribed species from New Caledonia, Oxysemaphora from New Guinea and Australia, and Dynatorhabda from New Guinea only. Episimus is restricted to the New World and Episimoides to Réunion.
Biology Oxysemaphora and Cnecidophora, gen. nov. have both been reared from leaf rolling larvae on Myrsinaceae, with Oxysemaphora reported also from other families. Episimus species are leaf rollers on a range of plant families.
Remarks The position of the the Oxysemaphora-group with its overall very plesiomorphic genitalia in both sexes is unclear. Following Diakonoff (1973), the group is included here at the base of the Olethreutini, though the cladistic analysis suggests a position at the base of the Eucosmini. In his phylogenetic tree, Heinrich (1926) also placed Episimus at the base of the Olethreutini, remarking in the text on their eucosmine affinities, including a wing pattern with an ocelloid patch and a hindwing with M3 and CuA1 sometimes stalked. With the Olethreutini probably paraphyletic in terms of the Eucosmini, the Oxysemaphora-group could possibly turn out to be the sister group to the remaining Olethreutini and the Eucosmini. The distribution of the group, with Oxysemaphora and Cnecidophora, gen. nov. found in New Guinea and Australia and several species of the latter in New
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Caledonia, Episimoides on Réunion and Episimus in South America, could indicate a Gondwana origin. The Oxysemaphora-group has neither a tibial pencil nor a costal fold, and the hindwing scale tufts or pencils from the anal margin in some species of Oxysemaphora or from between CuP and 1A+2A in Cnecidophora, gen. nov. are not homologous with the pencil in the anal roll of many Eucosmini and Enarmoniini. Only a well-supported phylogeny will eventually allow to track the evolution of the various characters.
Oxysemaphora Diakonoff
spinulose uncus tip and the projecting, domeshaped bulges in the membrane on both sides of the caulis are diagnostic for Oxysemaphora. The Ushaped and faintly concave spinulose sterigma with the ostium near its centre, the dorsal sclerite along the anterior half of the ductus bursae with a ring around its middle and the two slender, pointed, curved and horn-shaped granulose signa are characteristic for the female genitalia of Oxysemaphora. The sclerite in the anterior part of the ductus is developed at least as an irregular ring in both Dynatorhabda and Cnecidophora, gen. nov., but Dynatorhabda has only one signum while the sterigma of Cnecidophora, gen. nov. has conspicuous, invaginated anterolateral corners.
(Figs 25, 148–158) Oxysemaphora Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 501. Type species: Polychrosis chionolitha Meyrick, 1938, Trans. R. Entomol. Soc. Lond. 87: 511, by original designation and monotypy. [New Guinea.]
Diagnosis Oxysemaphora species are medium-sized to very large brownish moths (Papuan O. chionolitha iridescent black) either with very fine longitudinal stripes or mottled, males with an often serrate white streak along dorsum (only along basal half in O. chionolitha) and with a white dorsal midline across head and thorax, females with blackish marks along dorsum. If present, the long narrow hair-pencil from near the base of the anal margin of the hindwing in males of some species of Oxysemaphora is diagnostic for the genus. The pencil is recurved and its tip inserted into the scaling laterally on the abdomen. The hind tibia of male Oxysemaphora is not modified. The hindwing venation with basally approximate but separate Rs and M1 and separate to at most connate M3 and CuA1 differs from the stalked condition of both these pairs of veins in the Australian species of the closely related Cnecidophora, gen. nov. The much larger anal area of Oxysemaphora with 1A+2A more distant from 3A than from CuP is the most decisive venational difference between the two genera. Male genitalia with a long, ventrally curved and apically bifid uncus, with long and slender, drooping socii and a very long and slender valva with a large basal excavation and a huge thorn or blunt spike on its ventral edge at the base of the cucullus. Dynatorhabda Diakonoff and Cnecidophora, gen. nov. have a very similar valva, but the bifid,
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Description Adults (Figs 148–151) Wingspan. 17–26 mm; medium-sized to very large. Head (Fig. 152). Ocellus well developed. Frons in uppermost part with a narrow transverse band of projecting scales, remainder with short, upwardly directed scales. Proboscis short, about 1.5× length of labial palpus. Labial palpus moderate but somewhat longer in female (1.6–2.1× diameter of eye), moderately wide, in male subascending and slightly sinuate only at apex, in female strongly sinuate and porrect; second segment lightly curved and gradually widening towards rounded apex that is usually bent forward; terminal segment porrect or drooping, very small in male and often nearly concealed, much longer in female. Antenna reaching to about middle of forewing; without scales on anterior surface and rather stout in male; cilia in male minute. Thorax. With posterior crest. Legs not modified in male. Wings (Figs 25, 148–151, 153). Forewing index 0.40–0.45; forewing subtriangular, broadest near termen, costa moderately curved throughout, apex slightly to strongly produced, termen weakly oblique and faintly concave; without conspicuously denser scaling on underside; discal cell 0.63× length of forewing, M-stem well developed, accessory cell narrow with chorda from between R1 and R2 to between R4 and R5; all veins present and separate beyond discal cell; anal loop 0.26× length of 1A+2A; brownish and longitudinally finely striped or mottled (Australian species), or iridescent black (O. chionolitha), males always with an often serrate white streak along dorsal midline across head and thorax, females with one or two
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Figs 148–153. Oxysemaphora, adults, head and hindwing pencils. 148, 149, O. hacobiani. 148, , Mt Bellenden-Ker, Q [image reversed]. 149, , nr Mt Molloy, Q [image reversed]. 150, 151, O. notialis. 150, , Clyde Mt, NSW [image reversed]. 151, , Bateman’s Bay, NSW. 152, head of O. notialis, male. 153, male hindwing pencils (arrows), O. notialis. Scale = 2 mm.
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blackish marks along dorsum. Hindwing with cubital pecten usually present (absent in O. chionolitha (Diakonoff 1973)) and with welldeveloped axillary tuft, with long loose hairscales from basal part of anal area on upper surface, males of some species with a yellowish, long and slender pencil of hairscales from upper side of anal margin close to base, with tips of pencil recurved and inserted into lateral scaling on basal third of abdomen (Fig. 153) and with lower surface of anal area often more densely scaled; Rs and M1 parallel and closely approximated at base, M2 bent towards M3 at base, M3 may be equidistant from M2 and CuA1 or connate with CuA1; CuP well developed as a non-tubular vein nearly halfway to base; anal area (between 1A+2A and anal margin) wide, with 3A well developed and distance between 3A and 1A+2A wider than that between 1A+2A and CuP; mottled grey, rarely with apex brown-tipped. Pregenital abdomen. Male with bands of dense, raised scales along posterior margin of T6 and T7; in males of those species with a hair-pencil from anal margin with (presumably associated) modified, non-deciduous scales in a band along anterior margin of T5 and in longitudinal lateral folds on each side along T4 and T5. Sternum 8 in male with faintly to distinctly bilobed hindmargin, T8 very large with a posteriorly projecting lobe densely covered with raised, long scales. Male genitalia (Figs 154, 157, 158). Vinculum a membranous band. Tegumen weakly sclerotised, with broad pedunculi without differentiated attachment points for muscle m4. Uncus a long curved median process with a dorsally bristled base and a bifid apex, densely covered with minute long spinules. Socii long, inwardly curved, drooping lobes, covered with stout, long bristles in apical part. Gnathos weakly sclerotised lateral bands, anal tube entirely membranous. Valva long and very narrow, especially beyond basal third, with large costal process and a very large basal excavation reaching nearly to ventral margin, without a welldefined sacculus but usually more or less abruptly constricted beyond basal excavation, with a very long, slender and parallel-sided curved cucullus with a huge sharp thorn or blunt spike in a socket at base of its ventral margin. Juxta small and caulis very large and greatly modified: with a deep vertical groove in midline and two posteriorly strongly projecting, dome-shaped bulges in membrane on each side, appearing as two parallel lobes below aedeagus in lateral view; anellus inconspicuous. Aedeagus usually long and slender (short and stout in O. chionolitha), base curved and without
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coecum, remainder more or less straight with bevelled apex; vesica with a large sheaf of deciduous, apically faintly curved, needle-shaped cornuti. Female genitalia (Figs 155, 156). Ovipositor lobes subtriangular with rounded lateral margin. Sclerotised part of S7 transversely crescentic with a deeply concave hindmargin with a smooth rim without scale sockets. Sterigma a large U-shaped, sclerotised, and densely spinulose shallow depression with a paler V-shaped groove or shallow funnel in posterior half leading to ostium. Ductus bursae with colliculum of two small lateral sclerites below ostium, posterior half membranous and parallel-sided, anterior half gradually widened towards corpus bursae and with a sclerotised, triangular plate along its dorsal wall with its apex ending in a ring around ductus at midlength at inception of ductus seminalis, and its base extending halfway around neck of corpus bursae. Corpus bursae roundish, membranous with concentric wrinkles around signa; with two slender, sharply pointed, hollow and curved horn-shaped signa, granulate, strongly sclerotised and laterally somewhat compressed.
Distribution New Guinea and Australia, where the genus occurs along the eastern coast from northern Queensland to southern New South Wales.
Biology The southern O. notialis, sp. nov. has been reared from several rainforest taxa (Rapanea howittiana (Myrsinaceae), Acronychia oblongifolia (Rutaceae), ?Pittosporum (Pittosporaceae)) upon which the larvae feed as leafrollers.
Remarks Up to now, Oxysemaphora has been known from the male of only one species and the long series of the two Australian species at last provides the information to bring the genus into its proper context. Diakonoff (1973) figured wing venation, head and male genitalia for his description of the genus, and photographs of the wings and genitalia of the type (as Lobesia chionolitha) can be found in Clarke (1958: pl. 232, fig. 1). Apomorphies for the genus include: the slender, band-like, distally bifid and spinulose uncus; the membranous bulges lateral to the caulis, and the
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triangular sclerite in the neck of the corpus bursae. However, the male genitalia of Oxysemaphora are overall rather plesiomorphic with well-developed uncus, large socii and a simple valva (except for the ventral spike), and the female sterigma a simple spinulose depression with a near-central ostium in
85
the membrane behind S7. This character combination is probably close to the ground plan condition for Olethreutini. The well-developed CuP in the hindwing is another obvious plesiomorphic trait for Oxysemaphora. Oxysemaphora males lack a modified hind tibia but in several species possess a
Figs 154–158. Oxysemaphora, genitalia. 154, 155, O. notialis. 154, male genitalia, upper Allyn R., NSW, slide T2259. 155, female genitalia, Clyde Mt, NSW, slide T2378. 156–158, O. hacobiani, Mt Bellenden-Ker, Q. 156, female genitalia, slide T2477. 157, 158, male genitalia and aedeagus, slide T2476.
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long slender recurved hair-pencil arising from the anal margin of the hindwing close to its base, with the tips of the scales inserted into the vestiture laterally on the abdomen. Presence of such a pencil is associated with folds and bands of modified persistent scales on T4 and T5, suggesting a functional unit for scent dissemination. Diakonoff (1973) included Oxysemaphora in his subtribe Olethreutae, but with the evidence of the female now available there is no doubt that the genus is remote from Olethreutes but closely related to Cnecidophora, gen. nov. and Dynatorhabda. All three genera share a valva with a large, strong ventral spike, a long caulis, a sclerite in the neck of the corpus bursa and horn-shaped signa The assignment of Dynatorhabda to the subtribe Gnathmocerodides (Diakonoff 1973) on the strength of the single cornute signum shared by Gnathmocerodes is not very convincing in the face of the fundamentally different valva and secondary sexual structures of the two genera. There appear to be no close relatives in the Oriental/Australian fauna for the closely-knit Oxysemaphora-group of genera. Some obvious similarities to the New World genus Episimus Walsingham and the biogeographic implications of such a relationship are alluded to in the Remarks for the Oxysemaphora-group.
Constituent species Four species of Oxysemaphora are known, two each from New Guinea and Australia, with one New Guinea species named and the two Australian species described below: notialis, sp. nov. hacobiani, sp. nov.
Oxysemaphora notialis, sp. nov. (Figs 150–155)
Material examined Holotype. , ‘Clyde Mt, 16 mls [27 km] E of Braidwood, N[ew] S[outh] W[ales], 12.12.1955, I.F.B. Common’ (ANIC). Paratypes. Queensland: 1 , Bald Mt, Emu Vale, i.1975, S. R. Monteith. New South Wales: 1, Rous, Richmond R., viii.1925, V. J. Robinson; 1 , Ebor, 7.xi.1947; 1 , Acacia Plateau, 31.x.1961, I. F. B. C. & M. S. U.; 2 , Port Macquarie, 15.i. & 11.iii.1958, I. F. B. C.; 1 , 31°32´S 152°46´E, 10 km S by E Wauchope, 24.xi.1976, I. F. B. C. & E. D. E.; 1 , Upper Allyn R., 8.ii.1961, I. F. B. C. & M. S. U.; 1 , 44 km S Singleton, 13.i.1956, I. F. B. C.; 1 , Mt
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Tomah, 1000 m, 18.xii.1967, M. S. U.; 1 , Cambewarra Mt, 7.x.1948, I. F. B. C.; 2 , Colo Vale, 21.ix.1960, D. H. Colless; 1, Wollongong, 10.xii.1961, V. J. Robinson; 11 , 3, Mt Keira, 24.i.1963, 29.i.1976, 1.ii.1963, 5 & 18. iii.1964, 13.iii.1965, 29.iv.1964, 16.ix.1970, 6.x.1968, 20. xi.1965, 3.xii.1965, 4 & 19.xii.1963, V. J. Robinson; 2 , Mt Kembla, 9.v. & 3.ix.1964, V. J. Robinson; 1 , 2, Minnamurra Falls, 22.v.1969, 2.x.1969, 22.xii.1972, V. J. Robinson; 1, Barren Grounds Fauna Reserve, 30. xii.1976, V. J. Robinson; 4 , 4, same label as holotype but also 4.i.1987, 17.x.1960, 17.x.1962, 7.xii.1955, I. F. B. C., I. F. B. C. & M. S. U.; 2 , 4, Depot Beach, 16 km NE Batemans Bay, 22.1.1963, 9–19.vii.1969, 24.xii.1967, 12. i.1987, I. F. B. C., I. F. B. C. & M. S. U., M. Horak; Durras Lake, 25.x.1956, I. F. B. C.; 1 , 29 km SSE Braidwood, 17.x.1955, I. F. B. C.; 1 , 1, 25.vi.1981, Broulee, 3.vi.1961, I. F. B. C.; 1 , Brown Mt, 18.i.1961, D. H. Colless; (ANIC, AM, BMNH, QM).
Diagnosis With their longitudinally finely striate brown wings with an irregular white band along the dorsum continued as a white band along thorax and head, the males of both O. notialis and O. hacobiani, sp. nov. are quite distinct but the two species are difficult to separate. The forewings of O. notialis are slightly more greyish and the hindwings a bit paler and have a slender yellowish pencil of a few hairscales from the base of the anal margin, but good material of both species is needed for comparison. The female of O. notialis has longitudinally striate forewings like the male but with a dark apical area and with the band along the dorsum suffused with blackish brown and only sprinkled with white scales. The male genitalia of O. notialis are characterised by an uncus with a long, band-shaped neck and with the digits of its bifid apex short, parallel-sided, distally rounded and at a right angle to each other, and by a conspicuously narrowed valva with its ventral margin strongly sinuate. The diagnostic features of the female genitalia are a sterigma with the entrance to the ostium in its posterior half and the triangular sclerotised plate anteriorly in the ductus bursae ending in a long narrow point with a shallow indentation in its left side.
Description Male (Fig. 150, 152) Wingspan 17.5–23.5 mm. Head, thorax and tegulae pale greyish brown with frons and inner surface of labial palpi white and with a broad white (very rarely pale cinnamon coloured) longitudinal stripe along vertex and thorax, including posterior
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crest, with tegulae posteriorly increasingly white. Forewing pale greyish brown, finely longitudinally striped with darker brown and sometimes with whitish and some black, sprinkled with some orange ochreous scales and indistinctly coarsely mottled with about five darker brown patches along middle of wing; with a broad whitish (very rarely cinnamon coloured) band along dorsum to tornus with a zig-zagging edge of three peaks reaching to beyond 56M wing width, sprinkled with pale greybrown scales throughout and with some scattered brown scales along dorsum and a small subtriangular brown mark at 6M dorsum; margin of brownish wing portion between first and second white peak suffused with blackish brown; costa dark with pairs of narrow pale greyish brown strigulae throughout; a transverse pair of minute white (rarely cinnamon coloured) dots above closing vein and an interrupted line of similarly pale scales along termen; fringe variably barred with pale and darker brown, increasingly whitish towards tornus. Hindwings very pale greyish brown, transversely mottled darker, increasingly so towards apex, with a long pencil of a few yellowish hairscales from base of anal margin, recurved and inserted laterally into abdominal scaling (Fig. 153). Abdomen slightly paler than hindwings. Male genitalia (Fig. 154). Tegumen elongate ovate, weakly sclerotised. Uncus with wide, dorsally bristled base tapering to a band-shaped long neck with a bifid tip of two short, parallel-sided, distally rounded digits at a right angle to each other. Socii long, inwardly curved lobes with strong bristles in apical portion. Gnathos weak, curved lateral bands. Valva conspicuously narrowed below middle with ventral margin strongly sinuate; large spike at base of parallel-sided cucullus sharply pointed. Juxta small, caulis hugely enlarged but weakly sclerotised, with projecting steeply bell-curve-shaped lobes on each side. Aedeagus long, slender, parallel-sided, curved at base, with large sheaf of needle-shaped cornuti. Female (Fig. 151) Wingspan 21–25 mm. Head and thorax as for male but white band narrower and sprinkled with brownish scales. Forewings with similar pattern but generally somewhat darker, especially apex from 6M costa to below middle of termen, proximally incorporating a well-defined dark brown mark between R4 and M1; band along dorsum dark, suffused with dark grey in basal half and ending in semicircular blackish brown mark at 36: dorsum with white scales only present as partial outlines of first and second peak, delineating triangular
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blackish suffusion between them, and scattered towards tornus as remnant of third peak. Hindwings as male, abdomen slightly darker. Female genitalia (Fig. 155). Ostium in intersegmental membrane behind transversely crescent-shaped S7, with sterigma in deep semicircular emargination of hind margin of S7. Sterigma a large, weakly concave, shield-shaped plate with entrance to ostium in posterior 56O. Ductus bursae longer than 56O length of corpus bursae, with triangular sclerotised plate ending in a narrow, drawn out point with a shallow indentation on left side. Corpus bursae roundish with two flattened, curved, horn-shaped signa each from a hollow base.
Distribution and biology Oxysemaphora notialis has been collected in rainforest habitats along the coast of New South Wales from Rous, Richmond R., to Brown Mt near Bega, and has been reared from several rainforest taxa (Rapanea howittiana (Myrsinaceae), Acronychia oblongifolia (Rutaceae), ?Pittosporum) upon which the larvae feed as leafrollers.
Remarks The species name refers to the southern distribution of this species with regard to the rest of the genus.
Oxysemaphora hacobiani, sp. nov. (Figs 25, 148, 149, 156–158)
Material examined Holotype. , ‘17.16S 145.51E, Centre Peak Mt Bellenden Ker, Q[ueensland], 1560 m, 6 Jan 1988, B. Hacobian’ (ANIC). Paratypes. Queensland: 2, Mt Lewis 13 km NW Mt Molloy, 820 m, 15.iii.1964, I. F. B. C. & M. S. U.; 3 , 3, same label data as holotype; 10 , 9, 17°16´S 145°51´E, 0.5 km WSW Mt Bellenden Ker Centre Peak, 1500–1560 m, 2–4.xi.1981, E. D. E.; 1 , 17°16´S 145°52´E, Cableway Tower 3, Mt Bellenden Ker, 1020 m, 26.x.1981, E. D. E.; 4 , 1, 17°24´S 145°49´E, Summit Camp Site Mt Bartle Frere, 1500 m, 6–7.xi.1981, E. D. E. (ANIC, BMNH, QM).
Diagnosis With their longitudinally finely striate brown wings with an irregular white band along the dorsum
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continued as a white band along thorax and head, the males of both O. hacobiani and O. notialis are quite distinct but difficult to separate. The forewings of O. hacobiani are of a darker and deeper brown and its hindwings are darker, but good material of both species is required for comparison. The female of O. hacobiani entirely lacks the fine longitudinal pattern elements present in O. notialis and has instead dark brown markings. The two well-defined dark brown marks on the dorsum are diagnostic: they are of comparable size, edged with white, and the inner mark at 369 is subrectangular and reaches to the dorsal margin without interruption, in contrast to the same mark in O. notialis that is partly separated by white scales from the dark suffusion along the dorsal margin. The male genitalia of O. hacobiani are characterised by an uncus with a gradually tapering neck and with the digits of its bifid apex long, tapering, distally pointed and at an acute angle to each other, and by a moderately narrowed valva with its ventral margin gently sinuate. The diagnostic features of the female genitalia are a sterigma with the entrance to the ostium in its posterior third and the triangular sclerotised plate anteriorly in the ductus bursae gradually tapering to the ring around the ductus bursae and with a shallow indentation in its left side.
Description Male (Fig. 148) Wingspan 18.0–23.5 mm. Head, thorax and tegulae laterally dark grey-brown sprinkled with few whitish scales, frons and inner surface of labial palpi white, a white median band along vertex and thorax, including posterior crest, tegulae with dark brown base and white tips. Forewing deep brown often with a reddish tinge, finely longitudinally striped with darker brown, some black, ochreous and red-brown and indistinctly mottled darker; a broad whitish (very rarely pale cinnamon coloured) band along dorsum to tornus with a roughly zigzagging edge of three peaks reaching to beyond 56M wing width, sprinkled with grey-brown scales throughout and with blackish brown scales scattered along dorsum with a pronounced spot at just below middle and a subtriangular mark at 6M dorsum; margin of brown wing portion suffused with black between first and second peak; costa with indistinct paired paler strigulae; a transverse pair of minute white dots above closing vein and an interrupted line of white scales along termen; fringe variably barred with white, reddish and blackish brown. Hindwings brownish grey, transversely
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mottled darker, increasingly so towards apex, no pencil apparent from base of anal margin. Abdomen paler than hindwings. Male genitalia (Figs 157, 158). Tegumen elongate ovate, weakly sclerotised. Uncus with wide, dorsally bristled base gradually tapering to bifid tip with two long, slender, distally pointed digits at an acute angle to each other. Socii long inwardly curved lobes with strong bristles in apical portion. Gnathos weak, curved lateral bands. Valva moderately and gradually narrowed below middle with ventral margin gently sinuate, large spike at base of parallel-sided cucullus sharply pointed. Juxta small, caulis hugely enlarged but weakly sclerotised, with projecting steeply bell-curveshaped lobes on each side. Aedeagus long and slender, faintly widening in apical half, sockets for numerous cornuti present. Female (Fig. 149) Wingspan 21–26 mm. Head and thorax mottled grey- and red-brown, tegulae white-tipped and usually some white-tipped scales dorsomedially on thorax. Forewings mottled brown, with dark reddish and blackish brown marks on pale brown groundcolour of mixed ochreous and red-brown scales, an indistinct outwardly oblique bar at 56M costa, a small spot at 56O and a darker apex from 6M costa to below middle of termen proximally edged with some black scales and incorporating one or two red-brown streaks between R4 and M1 and along M2; band along dorsum indicated only by mixed white and red-brown scales separating two large, white-edged blackish brown marks, the inner subrectangular at 369 and the outer more semicircular at 36: dorsum; costa with indistinct paired paler strigulae; a tranverse pair of small white dots above closing vein and an interrupted line of white scales along termen; fringe barred with light ochreous, reddish and blackish brown scales. Hindwings dark brownish grey, indistinctly mottled, especially towards apex. Abdomen as hindwings. Female genitalia (Fig. 156). Ostium in intersegmental membrane behind transversely crescent-shaped S7, with sterigma in deep semiovate emargination of hind margin of S7. Sterigma a large, weakly concave, shield-shaped plate with entrance to ostium in posterior 56:. Ductus bursae with colliculum well developed, shorter than 56O length of corpus bursae, with triangular sclerotised plate gradually tapering to ring around ductus bursae and with a deep pocket on left side. Corpus bursae roundish with two flattened, curved, hornshaped signa each from a hollow base.
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AUSTRALIAN OLETHREUTINE GENERA
Distribution Oxysemaphora hacobiani has been collected only at high altitudes in the Cairns region, in rainforest localities.
Remarks The species is named for Bart Hacobian who collected part of the type series, in appreciation of his contribution to the ANIC.
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the sclerite in the anterior half of the ductus bursae and the two hollow horn-shaped signa are characteristic for Cnecidophora. Most of these features are shared with Oxysemaphora and Dynatorhabda, but the former has much more strongly sclerotised signa surrounded by concentric wrinkles and the latter has only a single signum. The invaginated sterigma corners are found only in Cnecidophora and are diagnostic for the genus.
Description
Cnecidophora, gen. nov. (Figs 26, 159–166) Type species: Cnephasia ochroptila Meyrick, 1910, Proc. Linn. Soc. NSW 35: 279, by present designation. [Australia.]
Diagnosis Cnecidophora species are small to medium-sized grey moths with a large and conspicuous, often dark-centred whitish or reddish ochreous roundish mark occupying distal third of forewing. In some specimens the pale distal blotch is obliterated by grey scales with a red-brown tinge leaving only a small erect white mark as its proximal delineation. The forewing venation is unremarkable, with all veins present and separate, but the hindwing is quite distinct with Rs and M1 stalked, M2 at base parallel with or only slightly bent towards connate or stalked M3 and CuA1, and with a very narrow anal area with 1A+2A distally much closer to 3A than to CuP. Males of some species (C. ochroptila) have a large bundle of elongate, modified scales on underside of hindwing, originating from between CuP and 1A+2A and their tips covered by the folded anal margin. This hindwing modification, if present, is diagnostic for Cnecidophora. The male hind tibia has, at most, a loose fringe of scales along its dorsal margin. The male genitalia are exceedingly similar throughout the genus, with the slender tapering uncus, the long and narrow socii and the medially constricted and distally bifid valva with its distinctly produced ventral spike diagnostic for Cnecidophora. The male genitalia of both Oxysemaphora and Dynatorhabda are similar, but the latter has only small knob-like socii and the uncus reduced to a triangular hairy stump whereas the former has a deeply bifid and spinulose apex to its long uncus. The concave and spinulose sterigma with its deeply invaginated anterolateral corners,
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Adults (Figs 159–162) Wingspan. 11–18 mm; small to medium-sized. Head (Fig. 165). Ocelli well developed. Frons with a wide horizontal band of strongly projecting scales in upper third, remainder with very short, appressed, upwardly oriented scales. Proboscis short, about 1.5× length of labial palpus. Labial palpus moderate (1.8–2.1× diameter of eye), rather wide and sinuate; second segment curved, gradually widening towards broadly rounded apex, with long loose scales on apex and along ventral margin; terminal segment thin and quite long, but partly concealed in fringe of second segment in males of C. ochroptila, longer and more obvious in females and males of an unnamed species from New Guinea. Antenna reaching to about middle of forewing; without scales on a rather narrow band along anterior surface; cilia in male minute. Thorax. With a posterior crest. Legs practically unmodified, at most (C. ochroptila) with a fringe of long scales along dorsal margin of hind tibia in both sexes. Wings (Figs 26, 159–162). Forewing index 0.43–0.48; subtriangular, apically strongly dilated, costa slightly curved throughout, apex roundedacute, termen straight and oblique; in C. ochroptila males with very dense cover of elongate scales across discal area of underside; discal cell 0.59× length of wing, R-stem and chorda moderately developed (more weakly in male of C. ochroptila), with large accessory cell; all veins present and separate beyond discal cell, R3 much closer to R4 at base than to R2; anal loop 0.22× length of 1A+2A; with a variably developed but generically typical wing pattern of a large whitish ochreous roundish mark across terminal area in an otherwise greyish wing, pale mark usually with a darker spot in its centre (especially in females) that is sometimes expanded to cover entire pale area in wing tip, leaving only a small erect white mark as proximal delineation of the usual pale blotch (Fig. 161). Hindwing with cubital pecten usually present,
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reduced or entirely lacking (C. ochroptila) in some males, axillary tuft developed, both sexes with long hairscales from anal margin and scattered on upper surface towards 1A+2A, males of some species (C. ochroptila) with the anal margin folded under to cover a large bundle of long, modified scales originating from an elongate patch parallel to and between CuP and 1A+2A, distorting the course of CuP; R1 clearly visible as a crossvein between Sc and Rs, Rs and M1 stalked, M2 at base subparallel and widely separate from M3 (C. ochroptila) or slightly bent towards M3; M3 and CuA1 connate or
stalked, CuP moderately developed; 1A+2A usually present, but absent in males with a scale bundle, 3A halfway between 1A+2A and anal margin. Pregenital abdomen. Small transverse patch of modified, distally rounded scales along anterior margin of T4 in males of some species (present but indistinct in C. ochroptila). Sternum 8 in male with bilobed hindmargin, T8 not enlarged and posteriorly hardly projecting beyond S8. Male genitalia (Figs 163, 164). Vinculum a membranous band. Tegumen weakly sclerotised with broad pedunculi without projecting apodemes
Figs 159–162. Cnecidophora ochroptila, adults. 159–161, . 159, Bunya Mts, Q. 160, Church Point, NSW. 161, nr Mt Garnet, Q. 162, , Lamington Nat. Park, Q. Scale = 2 mm.
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AUSTRALIAN OLETHREUTINE GENERA
for insertion of muscle m4. Uncus a long and slender, curved, distally tapering and weakly sclerotised median process with scattered bristles throughout but particularly large ones in the proximal dorsal area, shape of uncus tip diagnostic. Socii long, slender, lightly inwardly curved drooping lobes covered with sparse stout bristles that run in a band to base of uncus. Gnathos two lateral bands medially fused with subscaphium, the latter with a vaulted spinulose ventral surface (above aedeagus) and two oblique and spinulose bands from its ventrolateral angles converging towards apex. Valva highly modified with bifid distal half, with large and conspicuously twobranched costal process, with basal excavation very large and reaching nearly to ventral margin, without discrete sacculus, with valva ventrally strongly constricted beyond basal excavation and widened again proximal to bifurcation into long, slender and apically rounded cucullus and a triangular to finger-shaped ventral process ending in an enormous thorn or truncate spike in a socket. Juxta small and caulis very high, anellus a
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membranous ring. Aedeagus moderate, lightly curved and tapering towards bevelled apex, base with a small ventral coecum; vesica with a bundle of several to numerous long, curved, needle-shaped deciduous cornuti. Female genitalia (Fig. 166). Ovipositor lobes subtriangular to elongate ovate. Sternum 7 rather weakly sclerotised and with moderately concave hindmargin. Sterigma a large transverse spinulose depression with a subcentral to posterior, shallow to deep and nearly parallel-sided spinulose funnel leading to ostium, with a shallow transverse fold across lamella antevaginalis with deeply invaginated lateral corners forming long, hollow, pointed lateral projections. Ductus bursae with colliculum of two small sclerites directly or at some distance below ostium, anterior half of ductus widening towards bursa and entrance of bursa often nearly encircled by the base of a large, triangular, dorsal sclerite whose apex encircles the ductus in a ring at midlength, at inception of ductus seminalis. Corpus bursae roundish, conspicuously spinulose around entrance to ductus
Figs 163–166. Cnecidophora ochroptila, Church Point, NSW, genitalia and head. 163, 164, male genitalia and aedeagus, slide T2148. 165, head, male. 166, female genitalia, slide T2149.
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and without concentric wrinkles around signa; with two signa close to base of triangular sclerite in ductus and sometimes with a curved sclerotised band above one of them, signa hollow, curved and horn-shaped, smooth and only moderately sclerotised, either slender and of roundish cross section or much larger and flattened (C. ochroptila).
Distribution New Guinea, Australia and New Caledonia (undescribed material). The Australian C. ochroptila occurs along the eastern coast from northern Queensland to southern New South Wales.
Biology Cnecidophora ochroptila has been reared from leaves of Rapanea variabilis and leaves and fruit of the mangrove Aegiceras corniculatum (both Myrsinaceae), as well as from foliage of unidentified and possibly different taxa.
Remarks Cnecidophora is doubtlessly closely related to Oxysemaphora and Dynatorhabda but each of the three taxa has a number of autapomorphies justifying their generic rank. Both the stalked Rs and M1 and the connate or stalked M3 and CuA1 in the hindwing of Cnecidophora are derived with regard to the other two genera, and this is probably also true for the narrow anal area. It has to be left open at this stage whether the course of M2 in Cnecidophora (bent only slightly towards the rather distant M3 or the two veins parallel at base) is a further apomorphy. The characteristic male hindwing modification in Cnecidophora is treated as autapomorphic at the generic level, though it is absent in one of the New Guinea species. The presumed male sex pheromone structures in the two other genera, if present, are of an entirely different nature: a long hair-pencil from the base of the hindwing anal margin in Oxysemaphora and a pencil on the underside of the forewing in Dynatorhabda. The position of the ventral valval spike on a triangularly produced process and the spinulose areas on the subscaphium are derived features in the male genitalia of Cnecidophora, the latter shared with Dynatorhabda. The deeply invaginated anterolateral corners of the sterigma of Cnecidophora are the most obvious autapomorphy in the female genitalia. Cnecidophora, Oxysemaphora and Dynatorhabda all share an essentially plesiomorphic tegumen, a
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characteristically modified valva, a simple (=plesiomorphic?) Olethreutini-type sterigma, a sclerite in the anterior part of the ductus bursae and horn-shaped hollow signa. The male hind tibia is devoid of any hair-pencils and some males lack a cubital pecten. The overall rather plesiomorphic genitalia are paralleled by a wing venation rich in ancestral character states, especially in the hindwing: a well-developed CuP reaching to the middle of the wing (in Oxysemaphora) and the presence of the crossvein R1 between Sc + R1 and Rs. The base of M2 in the hindwing is consistently distant from M3 and at most bent only moderately towards M3 or the two veins are parallel at base, but it is unclear whether this is a plesiomorphy or a derived condition. Previous unconvincing taxonomic assignments for Oxysemaphora and Dynatorhabda are discussed in the Remarks section for Oxysemaphora. The three genera form a very isolated group within the Oriental/Australian Olethreutini, and the presence of several undescribed species of Cnecidophora in New Caledonia lends support to the notion that they could be derived from the base of the Olethreutini. The horn-shaped signa, unusual for Olethreutini but present in Microcorsini, Eucosmini, Enarmoniini and Grapholitini, are regarded as groundplan condition for the Olethreutinae. According to the male genitalia, Episimoides Diakonoff from Réunion (Diakonoff 1957a) also is a close relative of Cnecidophora, but the different hindwing venation and absence of the large spike in Episimoides sufficiently differentiate the two genera. The four Cnecidophora species examined are extremely similar, in superficial appearance as well as male genitalia. Specific differences are most readily observed in the development of the male secondary sexual characters (presence or absence of hindwing scale pencil and dense scaling on forewing underside) and the shape of the uncus tip and cucullus in the male genitalia. Meyrick (1910) described C. ochroptila as a tortricine, misled by the lack of a cubital pecten in the single male holotype. The fact that C. ochroptila has been described five times in no less than four genera says much about its variable wing pattern as well as the unusual character combination of Cnecidophora.
Constituent species Several apparently undescribed species of Cnecidophora are known to occur in New Caledonia, in addition to three unnamed ones from
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AUSTRALIAN OLETHREUTINE GENERA
New Guinea and the named type species from Australia: ochroptila (Meyrick, 1910), Proc. Linn. Soc. NSW 35: 279 (Cnephasia). [NSW: Sydney.] Comb. nov. aurita (Meyrick, 1920), Exot. Microlepid. 2: 345 (Eucosma). [Qld: Brisbane.] acrocausta (Turner, 1925), Trans. R. Soc. S. Aust. 49: 59 (Laspeyresia). [Qld: Lamington Nat. Park.] ochroplaca (Turner, 1945), Trans. R. Soc. S. Aust. 69: 70 (Cnephasia). [Qld: Bunya Mts.] nephobola (Turner, 1946), Trans. R. Soc. S. Aust. 70: 217 (Argyroploce). [Qld: Brisbane.]
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and Eudemis present in the western Palaearctic region.
Biology Larvae of the Eudemis-group are leaf rollers on a wide range of plant of families, some of them polyphageous.
Remarks Some possible apomorphies with the Gnathmocerodes-group are alluded to in the Remarks for Sorolopha.
Sorolopha Lower (Figs 27, 167–179)
Eudemis-group Status remarks Falkovitsh (1962) established the tribe Eudemini as a separate entity from the tribe Olethreutini for Eudemis Hübner, Eudemopsis Falkovitsh and Choganhia Razowski, with the latter a junior synonym of Sorolopha Lower. Diakonoff (1973) established the subtribe Sorolophae for the same genera, emended to Sorolophina by Nye and Fletcher (1991). The taxon is here treated as a genus-group only, and Penthostola Diakonoff is included in the Eudemis-group.
Evidence for monophyly Falkovitsh (1962) based the Eudemini on the nature of their secondary sexual scent structures, lateral scale tufts on the abdomen rather than a tibial pencil combined with a glandular anal roll. Diakonoff (1973) defined his Sorolophae primarily on the basis of single or paired horn-shaped signa, often reduced to scobinate depressions, and slender, pendulous socii. All four genera share conspicuously elongate, slender and drooping socii. Eudemis, Eudemopsis and Penthostola, all with an undivided cucullus, have paired, horn-shaped signa, whereas it ranges from a well-developed horn to a small scobinate depression in Sorolopha.
Distribution The majority of taxa of the Eudemis-group are found in the Oriental and eastern Palaearctic regions, with Sorolopha extending into Australia
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Sorolopha Lower, 1901, Trans. R. Soc. S. Aust. 25: 73. Type species: Sorolopha cyclotoma Lower, 1901, ibid. 25: 74, by monotypy. [Australia.] Acanthothyspoda Lower, 1908, Trans. R. Soc. S. Aust. 32: 319. Type species: Acanthothyspoda elaeodes Lower, 1908, ibid. 32: 320, by original designation. [Australia.] Alypeta Turner, 1916, Trans. R. Soc. S. Aust. 40: 528. Type species: Alypeta delochlora Turner, 1916, ibid. 40: 529, by original designation. [Australia.] Choganhia Razowski, 1960, Polskie Pismo Entomol. 30: 387. Type species: Argyroploce sphaerocopa Meyrick, 1930, Ann. Soc. Entomol. Fr. 98: 719 [481], by original designation. [Vietnam.]
Diagnosis Sorolopha species are medium-sized, conspicuously coloured Olethreutini, often in shades of green, or brownish-grey with a pinkish triangle over apical area containing brown roundish subapical mark, with sinuate, porrect, stout and moderately long labial palpi. Males often with large tufts of sex scales on hindlegs, laterally on abdomen and on apical area of hindwing. The male genitalia, characterised by extremely long and thin pendulous socii bristled mainly on apex, and by lack of an apically free gnathos, are diagnostic for Sorolopha. The closely related eastern Palaearctic Eudemopsis Falkovitsh has a much broader valva, and Eudemis Hübner and the Oriental Penthostola Diakonoff possess a well-developed, apically free gnathos and
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a tuft of long bristles from the outer surface of the valva base. The signum in Sorolopha may be single as in Eudemopsis or paired as in Eudemis and Penthostola, but it is either a scobinate depression or a pointed thorn, never a large, apically rounded horn as in Eudemis and Penthostola.
Description Adults (Figs 167–170) Wingspan. 12–18 mm; medium-sized. Head (Figs 171, 172). Ocellus large to very large. Frons with projecting scales in upper part; scales on lower part small, appressed and upwardly directed. Proboscis 1.5× length of labial palpus. Labial palpus short to moderate (1.5–2.0× diameter of eye) stout, porrect, sinuate with drooping terminal segment, segmental ratio 2:3.5:1; terminal segment small, with deep subapical vom Rath’s organ to nearly base of segment; second segment lightly curved and apically little to moderately dilated by dorsal scale tuft. Antenna reaching to beyond middle of forewing; anterior surface unscaled; cilia in male at most to just over half diameter of flagellum, much shorter in female. Thorax. With conspicuous posterior crest, often fan-shaped and/or contrastingly coloured. Hind leg often strongly modified in male with tibia and sometimes first tarsal segment expanded by dorsal and ventral fringes of long, loose scales, and with various large tufts of modified scales on inner surface (in S. cyclotoma Lower also with long scales dorsally from base of tibia, as in a typical tibial pencil). Wings (Figs 27, 167–170). Forewing index 0.41–0.48. Forewing of variable shape, usually subtriangular, with distally curved costa and rounded apex in the cyclotoma-group and with oblique termen and more acute apex in elaeodesgroup; cell c. 0.65× length of wing, M-stem to between M2 and M3, chorda from halfway between R1 and R2 to between R5 and M1; with all veins separate beyond cell; with greenish or grey-brown/ pinkish wing pattern, often with apex differently coloured beyond a line running from tornus to beyond middle of costa and with a dark roundish subapical mark in cyclotoma-group. Hindwing with cubital pecten and in male often with a strongly modified anal area varying from a patch of long loose scales (S. archimedias Meyrick, S. delochlora Turner) to a large composite bundle of scales folded into a greatly expanded anal lobe (S. cyclotoma, S. leptochlora Turner), with a pad of dense scales also ventrally along anal margin in S. elaeodes; with Rs and M1 closely approximated at base; M2 clearly separate from connate M3 and CuA1 (S. cyclotoma,
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S. archimedias) or all three veins closely approximated at base (S. delochlora, S. leptochlora) or with M3 closer to M2 than to CuA1 (S. elaeodes (Lower)); A1+A2 strongly curved in some species; grey to copper brown, anal area and associated scale tufts in male often of different colour. Pregenital abdomen. Male often with conspicuous scale brushes arising laterally from segments 2–7, variously developed in different species: S. cyclotoma with an unscaled band laterally along segments 1–3, followed on each side by a strongly enlarged and dorsally converging fanlike scale tuft on segment 3; S. elaeodes with lateral tufts more prominent in distal part of abdomen; S. delochlora additionally with a small lateral pocket in membrane between segments 3 and 4. Sternum 8 in male greatly enlarged, projecting and variously modified, with medially notched (S. delochlora) or scalloped (S. cyclotoma) hindmargin. Male genitalia (Figs 173–176). Vinculum and high, narrow, triangular tegumen forming complete ring. Uncus rudimentary, a small, laterally setose, dome-shaped process (with acute apex in cyclotoma). Socii long, slender pendulous arms, apically clavate and with hairs or bristles on inner surface, developed into claw-like spines on apices. Gnathos two weak, curved lateral bands fused with sclerotised ventral surface of subscaphium. Valva with a short, blunt costal process; shape variable but nearly always with a median constriction followed by a ventral process of the cucullus; sacculus area variably setose, often enlarged into prominent lobe with band of long bristles along outer surface (S. cyclotoma); ventral process of cucullus often with strong spines, either a rounded lobe (S. cyclotoma, S. delochlora) or triangular and ending in a large spine, sometimes with scale tuft from outer surface of ventral margin. Caulis and anellus fused into broad, elongate structure. Aedeagus long, slender, hardly tapering towards apex; vesica without cornuti. Female genitalia (Figs 177–179). Ovipositor and apophyses unremarkable. Sternum 7 with caudal margin straight to deeply emarginate, often fused with sterigma. Sterigma a lightly sclerotised, short, spinulose funnel with variously developed ventral margin (lamella antevaginalis): with a simple V-shaped cleft (S. elaeodes) or with a spinulose, rounded (S. cyclotoma) or broadly flattened (S. delochlora) protuberance either side of a ventral split. Ductus bursae sclerotised in posterior part with ductus seminalis from below midlength. Corpus bursae round to inverse pearshaped, with one or two signa, often of unequal size and ranging from a minute granulate patch over a
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Figs 167–172. Sorolopha, adult males and head. 167, S. cyclotoma, Sue (Warraber) Is, Q. 168, S. delochlora, Mt Bellenden-Ker, Q [image reversed]. 169, S. elaeodes, nr Babinda, Q. 170, S. johngreeni, Mt Bellenden-Ker, Q. 171, 172, head of C. cyclotoma, male. Scale = 2 mm.
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deep granulate depression (S. cyclotoma) to an acute thorn with granulate base.
archimedias is known to be polyphageous. Cinnamomum camphora (Lauraceae) and Syzygium aromaticum (Myrtaceae) are the only reported hosts of economic importance.
Distribution Sri Lanka, India and Nepal to Taiwan, the Philippines and Japan, China, south-east Asia, Indonesian archipelago, New Guinea, Solomon Is. In Australia the genus occurs in Queensland, the Northern Territory and northernmost coastal New South Wales.
Biology There are no host records from Australia, but the information available elsewhere (Diakonoff 1973) indicates that larvae of Sorolopha are leaf rollers on a wide range of plants (Lauraceae, Myrtaceae, Magnoliaceae, Pandanaceae). Sorolopha
Remarks Diakonoff (1973) provided a comprehensive treatment of Sorolopha and related genera, figuring the genitalia of most species and subdividing the genus into five species-groups, mainly on the basis of valva shape. Kawabe (1989) reported 16 species of Sorolopha from Thailand, figuring most adults and the genitalia of six newly described species. Kuznetzov (1988c) provided genitalia illustrations of seven species found in Vietnam, one newly described. Liu and Bai (1982, 1985) named four new Chinese species and Tuck and Robinson (1994) one from Borneo and Sulawesi.
Figs 173–176. Sorolopha, male genitalia and aedeagus. 173, 174, S. johngreeni, holotype, slide T2287. 175, 176, S. delochlora, Mt Bellenden-Ker, Q, slide T2274.
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Figs 177–179. Sorolopha, female genitalia. 177, S. delochlora, nr Leidley, Q, slide T 2364. 178, S. cyclotoma, Shute Harbour, Q, slide T2362. 179, S. johngreeni, paratype, nr Tully, Q, slide ANIC 17082.
The monophyly of Sorolopha is well supported by the slender socii, hairy throughout but with stronger apical bristles, by a valva with an elbow or projection on its ventral margin and by a sterigma usually fused with S7. The diversity of secondary sexual structures found in the genus is reflected in the three generic synonyms described from the Australian fauna. Eudemopsis appears to be the most likely sister group; together with Sorolopha it lacks the large tuft of bristles from the outer valva base and the distally free gnathos present in Eudemis and Penthostola (Diakonoff 1978). The shape of the tegumen apex, the long drooping socii and the modified caulis-anellus structure could be possible synapomorphies shared with Rhectogonia, together with the modified scales laterally on the male abdomen and the female signum. The Australian S. elaeodes has suffered at the hands of taxonomists. Lower (1908) in his original description omitted any reference to the hindwing that with its yellowish to orange tinged anal area with long dark hairscales along the dorsum in the male provides the easiest means of separating S.
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elaeodes from the other three green Australian species of Sorolopha. And whilst the genitalia figured for S. elaeodes by Diakonoff (1973) are based on Australian material presumably conspecific with Lower’s type, it is doubtful whether the specimens Diakonoff mentioned from New Guinea and Sulawesi are the same species. This reservation is supported by his comment, ‘The species is rather variable’ (Diakonoff 1973: 63), an observation at odds with the homogenous series of S. elaeodes in ANIC. Both taxa treated as subspecies of S. elaeodes by Diakonoff (1973): S. elaeodes parachlora Diakonoff, 1973 and S. elaeodes temenopsis (Meyrick, 1936), are distinct species, erroneously included in S. elaeodes possibly due to the misidentification of the abovementioned material from outside Australia as S. elaeodes.
Constituent species Sixty-three named species of Sorolopha are recorded in Brown (2005). Six species occur in Australia, including one described here as new:
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cyclotoma Lower, 1901, Trans. R. Soc. S. Aust. 25: 74 (Sorolopha). [Qld: Mackay.] archimedias (Meyrick, 1912c), Exot. Microlepid. 1: 63 (Argyroploce). [China.] elaeodes (Lower, 1908), Trans. R. Soc. S. Aust. 32: 320 (Acanthothyspoda). [Qld: Cooktown and Cairns.] lasiosoma (Meyrick, 1921), Exot. Microlepid. 2: 449 (Argyroploce). [Qld: Cairns.] johngreeni, sp. nov. leptochlora (Turner, 1916), Trans. R. Soc. S. Aust. 40: 529 (Alypeta). [Qld: Cairns.] delochlora (Turner, 1916), Trans. R. Soc. S. Aust. 40: 529 (Alypeta). [Qld: Brisbane.]
Sorolopha johngreeni, sp. nov. (Figs 170, 173, 174, 179)
Material examined Holotype. . ‘17.16S 145.51E, 56O km WSW of Mt Bellenden-Ker Centre Peak, Q[ueensland], 1560 m, 3 Nov. 1981, E.D. Edwards’, ‘Genitalia slide T2287’ (ANIC). Paratypes. Queensland: 1 , 15°53´S 145°14´E, Summit Mt Misery, 13 km NW Bloomfield, 860 m, 22–23.i.1988, B. Hacobian; 1 , 8 km N Daintree R. Ferry, 6.ii.1980, B. Hacobian & N. Goodman; 1 , Windsor Tableland, NW of Mossman, 27.xii.1976, M. S. & B. J. Moulds; 17°07´0´´S 135°37´7´´E, Robson Ck, Atherton, 16.i.1996, R. L. Kitching; 1 , Mt Baldy, approx. 8 km W Atherton, 13. i.1977, M. S. & B. J. Moulds; 1 , same label data as holotype but 4.xi.1981; 4 , 17°17´S 145°38´E, Lake Eacham NP, 760 m, 23.xi.1998, E. D. E. & H. Sutrisno; 2 , 17°26´S 145°29´E, The Crater, 18 km S Atherton, 980 m, 26.xi.1998, E. D. E. & H. Sutrisno; 1 , 17°39´S 145°27´E, Millstream Falls, 5 km SW Ravenshoe, 820 m, 25.xi.1998, E. D. E. & H. Sutrisno; 1 , Palmerston NP, 27 km SW Innisfail, 21.xii.1967, R. Dobson; 2 , 1, 5 km NW by N of Tully Falls, 800 m, 15.ii.1998, 25.ii.1998 (, ANIC genitalia slide No. 17082), 22.iii.1998, E. D. E., M. Horak; 1 , 18°58´S 146°10´E, 8 km NW Paluma, 23. viii.1977, I. F. B. C. (ANIC, BMNH, QM).
Diagnosis Sorolopha johngreeni is a blue-green Sorolopha with dark grey hindwings and a characteristic wing pattern including a conspicuous snow white mark and similar costal strigulae. The male genitalia are characterised by a triangular uncus with a small posterior projection at its base, a pointed ventral cucullus lobe ending in a spine and wide and strongly curved socii arms. The female genitalia are characterised by an evenly funnel-shaped sterigma
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with a perfectly round opening ringed by a spinulose band and connected on each side through an oblique band with the corner of the excavated hindmargin of S7, and by two signa, two granulate, roundish, deep dimples of unequal size. Sorolopha johngreeni belongs to the herbifera-group of species, differing from S. herbifera (Meyrick) in wing pattern and sterigma shape, from S. autoberylla (Meyrick) and S. callichlora (Meyrick) in wing pattern and cucullus shape, and from the very closely related S. hydrargyra (Meyrick) from New Guinea and S. plumboviridis Diakonoff from the Moluccas in wing pattern and a medially more narrowed valva.
Description Male (Fig. 170), female Wingspan 16.5–18.5 mm. Head and thorax bluegreen (unless faded), pale ochreous ventrally and at base of labial palpi, tegulae and tips of large posterior crest paler, olive green. Forewings light greenish blue with three irregular, jagged, outwardly oblique deep olive green fasciae, partly edged with white and some black scales, often strongly faded, in particular the light greenish blue ground colour; wing base with some ochreous olive scales. Basal olive fascia near costa developed only along its edges as two dark bands, entire but merging into ground colour on dorsum; with inner edge an interrupted oblique black line from near base of costa halfway to dorsum and distal margin irregular, medially interrupted and bent outwardly, edged with white and black scales. Second olive fascia outwardly curved from just below 56O costa to end of dorsum, medially greatly expanded to form an often white-edged dorsal projection to beyond centre of wing, with patches of black scales on inner and distal fascia edges in middle of wing. Apical area from 6M costa to above tornus olive except for spots of light greenish blue ground colour below white strigulae along costa and along termen, and a conspicuous, inverted Yshaped white mark, partly edged with black. Costa with pairs of white strigulae, interspersed with single black ones. Fringe blue-green, with black scales on apex and middle of termen. Hindwings dark grey. Abdomen dark grey, without modified scales in male. Male genitalia (Figs 173, 174). Sternum 8 deeply bilobed. Uncus a triangular hump with a small posterior projection at base. Socii lightly hairy, rather wide and strongly curved, with large, curved spines at clavate apex. Valva strongly
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narrowed beyond nearly rectangular sacculus lobe, cucullus with triangular ventral lobe ending in single, curved, short spine, with band of bristles along ventral margin on outer surface; dorsal lobe long, slender, tapering towards rounded apex. Aedeagus sinuate, slightly tapering towards bevelled, upcurved apex. Female genitalia (Fig. 179). Hindmargin of S7 with wide subrectangular excavation. Sterigma an evenly cone-shaped, partly sclerotised funnel with a conspicuous, broad, spinulose band around projecting, perfectly round opening, laterally connected with an oblique band on each side with the corner of the excavation in S7. Ductus bursae membranous, slender, shorter than length of corpus bursae, ductus seminalis originating from close to corpus bursae, with small bulla seminalis. Corpus bursae elongate ovate, signa two granulate, roundish, deep dimples of unequal size.
Distribution Sorolopha johngreeni is restricted to rainforest areas of the Cairns-Cooktown region and on the Atherton Tableland.
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very similar and are linked by several obvious apomorphic character states (with the possible exception of states two and seven that could be plesiomorphies): 1, wing pattern; 2, apodemes for muscles m4 (tergal flexor of valva) medially projecting as distinct lobes or triangular processes from base of tegumen (from pedunculi); 3, lobeshaped and often bipartite drooping socii from close to apex of tegumen; 4, sclerotised connection between anellus and gnathos/subscaphium; the slender and relatively evenly bristled valva with 5, a hump at base of costa and 6, often with a large spine at midlength of ventral margin; 7, a large bundle of deciduous cornuti; 8, a granulate structure of the ductus bursae; 9, usually at least a lateral invagination in lamella postvaginalis. Atriscripta, gen. nov. and Demeijerella Diakonoff share four of the above apomorphies (2, 3, 7 and 8), and the tentatively included Lobesia only two (2, 3). Characters shared by subgroups within this complex, such as the lateral scale tufts at the base of the abdomen in Lobesia and Sycacantha, are discussed under the Remarks section for the genera in question.
Distribution Remarks This beautiful blue-green species is named to honour John Green, the former photographer at CSIRO Entomology, who with great skill and patience took most of the photos for this volume.
While Lobesia has a nearly world-wide distribution except for the southern half of South America the other genera are centred on the Oriental/Australian region.
Biology
Sycacantha-group Status remarks The subtribe Sycacanthae was established by Diakonoff (1973) for six obviously closely related genera, centred on Sycacantha Diakonoff. Kuznetzov and Stekolnikov (1984) considered a separate subtribe Sycacanthae unwarranted and merged it with their subtribe Olethreutina. However, the group is well characterised by several synapomorphies and its scope is here considerably expanded, though treated as a genus-group.
Evidence for monophyly Diakonoff (1973) defined his Sycacanthae primarily on their wing pattern and on horn-shaped signa often reduced to a scobinate patch. The genera initially associated with Sycacantha indeed all look
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No hosts are known for Demeijerella and Atriscripta, gen. nov., and the only information available for Sycacantha refers to two species reared from leaves of ‘kalak’ (Anonaceae) (Diakonoff 1973), suggesting that the larvae are leaf rollers. Lobesia on the other hand is reported from a wide range of host families, feeding in spun shoots, flowers, seed heads and fruit, with the larvae feeding as tiers or at least partially as borers.
Remarks Atriscripta, gen. nov. and Demeijerella are both strongly derived and are in structure and appearance so different that both groups have initially not even been recognised as Olethreutini: Demeijerella was established as a genus of the Chlidanotidae and the only previously named species of Atriscripta, gen. nov. was described as a species of Strepsicrates Meyrick. The generic
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autapomorphies have in both cases nearly obliterated the morphological evidence to reconstruct their relationships. Presence of small folds bearing modified scales at the base of the abdomen laterally to S2 in Sycacantha and, more extensively developed, in Lobesia, are probably the strongest argument for inclusion of the latter genus in the Sycacanthagroup. Such a relationship is supported by the structure of the tegumen in Lobesia, by the drooping socii as well as the apodeme for muscle m4, and though the genus is clearly strongly derived nothing in its structure would contradict a derivation from a Sycacantha-group groundplan. Deciduous cornuti are found only in a few groups of the Olethreutini and are consistently present only in the Sycacantha-group (minus Lobesia) and the Oxysemaphora-group, and otherwise reported only from Podognatha and Eubrochoneura (Zomariana-group) and from Megalota Diakonoff (Temnolopha-group).
Sycacantha Diakonoff (Figs 28, 180–193) Phaecasiophora subgenus Sycacantha Diakonoff, 1959c, Ark. Zool. 2nd Ser. 12: 181. Type species: Phaecasiophora (Sycacantha) auriflora Diakonoff, 1959c, ibid. 12: 181, by original designation and monotypy. [Java.] Sycacantha Diakonoff; Diakonoff, 1966, Zool. Verh. (Leiden) No. 85: 7, 58.
Diagnosis Sycacantha species are medium-sized broad-winged olethreutines with smallish, curved, ascending palpi and usually strongly modified male hind tibia with long, snow-white scales and two tibial pencils, one black and one pale-coloured. Most species of Sycacantha are intricately mottled throughout and large, well-defined wing markings are rare. A series of horizontal, parallel black lines near wing tip enclosed by two narrow curved silvery fasciae running one from about 36: costa to tornus and the other from the same point around apex to anterior half of termen, is typical for many Sycacantha species. The same black markings are very conspicuous in Atriscripta, gen. nov., but males of this genus lack the modified hind legs present in all those Australian Sycacantha species characterised by this pattern element. The males of nearly all
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Australian species have a conspicuously modified anal margin to their hindwing and a long hair pencil from small crescentic folds laterally in the abdominal base. The broad tegumen with its paired anterodorsal humps and deeply cleft socii, resulting in two pairs of quite dissimilar appendages, is diagnostic for Sycacantha. The closely related Phaecasiophora Grote and Psilacantha Diakonoff have extremely similar female genitalia, all characterised by a long colliculum in an otherwise conspicuously granulate ductus bursae, and by more or less developed laterally invaginated processes from the lamella postvaginalis. Because, to our knowledge, only Sycacantha is present in Australia, the usually hornshaped lateral invaginations from the sterigma can be treated as diagnostic for the genus.
Description Adults (Figs 180–183) Wingspan. 12–20 mm; medium-sized to large. Head (Fig. 192). Ocellus well developed. Vertex scales anteriorly barely protruding between antennae. Frons in upper half with a transverse band of dense, shortish and downward pointing scales; remainder of frons with very short, appressed, dorsally directed scaling. Proboscis short, c. 1.5× length of labial palpus. Labial palpus rather short (1.4–1.8× diameter of eye) and slender, curved and ascending, distally sinuate with porrect apex; second segment curved and parallel-sided, with short appressed scaling except for short but erect scales along ventral margin; terminal segment short, porrect or downward directed. Antenna reaching to just over middle of forewing; unmodified, anterior surface devoid of scaling; cilia in male minute. Thorax. With large posterior crest in most species. Legs with conspicuously modified hind tibia in males of most species: hind tibia greatly enlarged by very long, loose, usually snow-white scales along dorsal and ventral margin; with usually two parallel hairpencils from base of tibia along inner surface, the more dorsal one whitish and the more ventral one deep black in its basal portion (Fig. 193). Wings (Figs 28, 180–183). Forewing index 0.45–0.50; forewing relatively broad, widest beyond middle with costa curved throughout or bent beyond middle, distal half of costa roughly parallel to dorsum, apex obtuse and termen nearly straight and vertical; discal cell 0.65× length of wing, accessory cell short and very narrow with chorda from base of R2 or more distally to base of R4,
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chorda rarely absent, M-stem present in all Australian species but usually not a tubular vein; all veins separate beyond discal cell; anal loop 0.3× length of 1A+2A; with usually intricately mottled wing pattern of various brownish shades and often a darkened subtriangular apical area proximally followed by a curved band of horizontal, parallel, black lines laterally enclosed by two narrow curved silvery fascia running from 36: costa to tornus. Hindwing with well-developed cubital pecten and axillary tuft, with anal margin modified and raised to form a strongly sclerotised, unscaled, dorsally open groove in most species; with Rs and M1 parallel and moderately approximated at base, M2
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close to connate M3 and CuA1; 3A well developed and at considerable distance from anal margin; dark greyish. Pregenital abdomen. Males of most species with a tuft of very long hair scales originating from invaginated crescentic areas lateral to the apodemes at the base of S2. Hindmargin of S8 in male strongly bilobed, with a deep median indentation. Male genitalia (Figs 184–189). Vinculum a lightly sclerotised, narrow band. Tegumen with large, rounded, paired humps forming apex of anterolateral surface, anterior to apex of tegumen proper; with attachment point of muscle m4 (tergal flexor of valva) produced into a prominent
Figs 180–183. Sycacantha, adults. 180, 181, S. exedra. 180, , nr Cooktown, Q. 181, , Cannonvale, Q. 182, 183, S. placida, Shute Harbour, Q. 182, . 183, . Scale = 2 mm.
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triangular lobe medially projecting from base of tegumen. Uncus usually reduced, at most a small cone-shaped membranous process (S. castanicolor (Turner)). Socii deeply cleft forming two pairs of differently shaped processes; inner pair longer and usually slender and tapering to a point, often with strong spines in apical part or apex naked (S. castanicolor); outer pair smaller rounded hairy knobs or lobes. Gnathos a continuous smooth transverse sclerotised band, usually connected to anellus by an inverted Y-shaped area of sclerotisation along midline of diaphragm. Valva long and slender, constricted beyond base and usually with a pronounced elbow at about midlength; with simple costal process, a large hump at base of costa carrying a conspicuous tuft of long strong bristles, and often a pocket-shaped invagination at base of outer surface; elbow at base of cucullus variously projecting on ventral margin, often with a single or few very large spines on apex and/or basal part of cucullus with modified, flattened and bifid bristles; some species (S. exedra (Turner)) with several enlarged bristles on outer surface along basal part of ventral margin. Caulis moderately long and anellus a complete ring, wider in basal half. Aedeagus a straight or curved tube, often dorsoventrally compressed; vesica with few to a large sheaf of deciduous cornuti, varying from needle-shaped to broadly lanceolate, strongly bent beyond base in S. castanicolor. Female genitalia (Figs 190, 191). Ovipositor lobes broad, with lateral edge strongly curved. Sternum 7 triangular to trapezoidal with hindmargin medially deeply emarginate. Sterigma typically a spinulose lamella postvaginalis with two lateral, invaginated pointed processes ranging from small depressions to horn-shaped cavities (rarely entirely lacking) and a shallow, cup-shaped medial invagination or a deep pouch behind S7. Ductus bursae with a long to very long colliculum, remainder membranous but with conspicuous, evenly spaced granulation becoming less dense anteriorly but still traceable over parts of bursa; ductus either narrow throughout or gradually widening towards bursa, with ductus seminalis arising posterior to midlength. Corpus bursae from long and narrow, hardly wider than ductus, to roundish ovate, without or with one or two signa; signa ranging from depressed patches of sclerotised granulation to curved thorns.
New Guinea, the Solomon Is and Australia, with several species in Africa (Tuck personal communication). Diakonoff (1973) included Japan in the range of distribution for Sycacantha, but the genus is not mentioned in Kawabe (1982). In Australia the genus is found in coastal Queensland and northern New South Wales and in the Northern Territory on the fringes of Arnhem Land.
Biology The host records for two different species provided by Diakonoff (1973), both reared from leaves of ‘kalak’ (Anonaceae) in Java, suggest that larvae of Sycacantha are leaf rollers. No hosts are known for the Australian species, but the genus is associated with rainforest.
Remarks It is by no means a foregone conclusion that Sycacantha deserves separate generic rank, but the effort required to adequately address the problem is beyond the scope of the present study. The genus already has a rather chequered history: established as a subgenus of Phaecasiophora by Diakonoff (1959c) it was later, without any comment, treated as a distinct genus by the same author (Diakonoff 1966) and officially given new status as a separate genus only in a subsequent publication (Diakonoff [1968]). Sisona Snellen and Psilacantha Diakonoff are both extremely close to Sycacantha, with the deeply cleft socii linking all three genera. The male genitalia of Sisona are the most plesiomorphic ones within the group, with evenly bristled socii arms and an undifferentiated valva without a prominent tuft of enlarged bristles at the base of the costa. Diakonoff ([1968]) gave a comprehensive redescription of Sycacantha and pointed out that the type species, S. auriflora, is a junior synonym of S. hilarograpta Meyrick, though he did not formally establish this synonymy. Thirty-four species of Sycacantha are figured and discussed in Diakonoff (1973), and the genus is divided into two speciesgroups on the basis of socii shape. The socii of several of the Australian species are, however, intermediate between the two groups, and as there is no correlation with any other character, it appears to be a rather arbitrary division.
Constituent species Distribution Sycacantha is centred on south-east Asia and New Guinea, ranging from India to the Philippine Is,
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About 55 species are so far known world-wide (Tuck personal communication), of which c. 44 have been named (Brown 2005). Seven named and
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Figs 184–189. Sycacantha, male genitalia and aedeagus. 184, 185, S. exedra, nr Marlborough, Q, slide T2423. 186, 187, S. castanicolor, Iron Range, Q, slide T2443. 188, 189, S. niphostetha, holotype, slide T2302.
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Figs 190–193. Sycacantha, genitalia, head and hind tibia, 190, 191, female genitalia, 190, S. exedra, Cannonvale, Q, slide T2444. 191, S. placida, Minyon Falls, NSW, slide T2448. 192, head, S. exedra, male. 193, hind tibia with double pencil (arrow), S. placida, male.
at least three undescribed species occur in Australia: exedra (Turner, 1916), Trans. R. Soc. S. Aust. 40: 532 (Argyroploce). [Qld: Kuranda.] irrorea (Lower, 1916), Trans. R. Soc. S. Aust. 40: 539 (Argyroploce). [Qld: Cairns.] niphostetha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 215 (Argyroploce). [Qld: Malanda.] placida (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 268 (Argyroploce). [Qld: Brisbane.] symplecta (Turner, 1946), Trans. R. Soc. S. Aust. 70: 215 (Argyroploce). [Qld: Cape York.] sphaerocosmana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 642 (Antithesia). [NSW: Richmond R.] eurypolia (Turner, 1916), Trans. R. Soc. S. Aust. 40: 530 (Argyroploce). [Qld: Brisbane.] atactodes (Turner, 1946), Trans. R. Soc. S. Aust. 70: 216 (Argyroploce). [Qld: Cape York.]
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castanicolor (Turner, 1946), Trans. R. Soc. S. Aust. 70: 214 (Anathamna). [Qld: Cape York.]
Atriscripta, gen. nov. (Figs 29, 194–203) Types species: Argyroploce arithmetica Meyrick, 1921, Exot. Microlepid. 2: 448, by present designation.
Diagnosis Atriscripta species are relatively narrow-winged, medium-sized reddish ochreous to pinkish grey moths with few well-defined markings on forewing except very narrow and strongly oblique white strigulae on dark red-brown costa and apex and a curved band, from beyond middle of costa to
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tornus, with a number of parallel, horizontal and white-edged narrow black bars. The closely related, slightly more broad-winged genus Sycacantha Diakonoff has the same forewing pattern elements, and pinkish coloured species of Sycacantha may look deceptively similar. However, none of the known Australian species of Sycacantha combines pinkish colouring with a series of white-edged black bars in the curved subapical fascia. Frontal scaling, apex and a narrow band along ventral edge of sickle-shaped labial palpus are touched with blackish in Atriscripta. The parallel-sided, upcurved labial palpus of Sycacantha is often mottled darker but never blackened along its lower margin only. Males of Atriscripta and Sycacantha can usually be separated on their secondary sexual characters. Atriscripta males have at most slightly dilated hind tibia without any hairpencils but instead a pointed abdominal tip with closely appressed scaling and often entirely retracted genitalia with not even the tips of the usually snow-white coremata protruding. Sycacantha males have a spreading anal tuft and in all except one Australian species the hind tibia is greatly enlarged and the anal margin of the hindwing modified. The wing venation of Atriscripta is typically olethreutine with an even shorter accessory cell than Sycacantha but otherwise no distinctive feature. The male genitalia of Atriscripta are highly modified and unusual, with an apically protruding anal tube with a wellsclerotised subscaphium continuous with anellus and caulis, strongly swollen pedunculi carrying sex scales, large and weakly bristled valva with two closely adjacent spines on angle or short process of ventral margin, and strongly modified S8. The ring-shaped sterigma with a dorsal invagination, the conspicuously granulate ductus bursae and the large single horn-shaped signum with a flattened apex provide an equally unmistakable set of characters for the female genitalia.
Description Adults (Figs 194, 195) Wingspan. 16–19 mm; medium-sized. Head (Fig. 200). Ocellus well developed. Vertex scaling anteriorly barely protruding between antennae. Frons in upper part with moderately long, anteroventrally projecting scales, lower half with small, appressed, upwardly oriented scales; scales on anterior part of vertex and upper part of frons touched by dark grey or black. Maxillary palpus minute but clearly visible due to white scaling. Proboscis about 1.5× length of labial
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palpus. Labial palpus sickle-shaped, upcurved and appressed to frons, rather short (1.4–1.5× diameter of eye); second segment strongly curved, sickleshaped, widest in middle due to long loose scales along ventral margin, narrowing towards apex; terminal segment small, porrect or downwardly directed; apex of palpus and ventral margin with a band of grey or black scaling on outer surface. Antenna reaching to middle of forewing; anterior surface devoid of scales; cilia in male minute. Thorax. Smooth. Legs with male hind tibia sometimes slightly enlarged by loose scales along dorsal and ventral margins. Wings (Figs 29, 194, 195). Forewing index 0.40–0.43; forewing rather narrow, elongatesubovate with an obtuse apex, costa evenly curved, termen vertical and straight or faintly convex; discal cell 0.63× length of wing, with moderately developed M-stem to M2, with small and narrow accessory cell with chorda from halfway between R2 and R3 to between R4 and R5; all veins separate beyond discal cell; anal loop 0.29× length of 1A+2A; ground colour greyish or reddish ochreous always with at least a pinkish or vinaceous hue, with narrow dark red-brown suffusion along costa, apex and anterior part of termen, with strongly oblique very fine white costal strigulae continued as narrow silvery lines, with an outwardly curved band of conspicuous white-edged parallel and horizontal black bars running from beyond middle of costa to tornus and usually a semicircular speckled area centred on dorsum. Hindwing with cubital pecten and axillary tuft, and in males of some species with a patch of melanic sex scales on underside between CuA1 and CuP, with anal margin unmodified in male; Rs and M1 parallel at base and only moderately approximated, M3 and CuA1 connate and base of M2 close to M3, 3A well developed and at some distance from wing margin; darkish grey. Pregenital abdomen. Tip of male abdomen strongly modified, apparent even on undissected specimen: 8th segment elongated and genitalia usually entirely retracted into abdomen except for tips of coremata scales; T8 down-curved at apex, hood-like, opening displaced ventrally to a long narrowly V-shaped median split in S8, fringed with dense short scales and outlined by a U- or V-shaped median area devoid of sclerotisation; intersegmental membrane anterior to genitalia very long, striate, with ventro-lateral sclerotised areas. Male genitalia (Figs 197–199, 201, 202). Vinculum a moderate band. Tegumen high and rather narrow, with very large, club-shaped
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Figs 194, 195.
Atriscripta arithmetica, adults, Mt Bellenden-Ker, Q. 194, . 195, . Scale = 2 mm.
pedunculi with the posterior surface of their laterally protruding base densely covered by very long and often snow-white scales (coremata); with only slightly projecting attachment points for muscle m4; apical part of tegumen covered with conspicuously modified scales on dorsal and lateral surface, also outer surface of valva. Uncus absent. Socii roughly triangular, pendant or semiporrect, large, sclerotised lobes with very few isolated hairs, apex either pointed, flat and curled with some hair on inner surface (S. arithmetica Meyrick) or somewhat paw-shaped with dense stout ventral bristles. Gnathos thin lateral bands fused with very conspicuous subscaphium; anal tube with laterally compressed apex protruding beyond tegumen and with caudo-ventral surface well sclerotised, continuous with anellus and caulis as a vertical median sclerotised band, perforated by aedeagus. Valva simple and very wide, distal part of cucullus angled upward and lightly tapering towards rounded apex, with two spines next to each other on a projecting angle or short process between 36: and ¾ of ventral margin, with costal process short and knob-like and inner surface very sparsely bristled except for apex and a narrow ventral band in basal half, with outer surface densely covered by modified swollen or melanic scales, as apical part of tegumen. Caulis a smooth, concave, shield-shaped structure fused with tube-shaped anellus that in turn is connected or continuous with subscaphium. Aedeagus nearly parallel-sided, slightly curved; vesica with several to numerous, irregularly bent, long to very long, narrowly lanceolate deciduous
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cornuti (in S. arithmetica apparently part of the cornuti fused to a non-deciduous structure). Female genitalia (Figs 196, 203). Ovipositor lobes roughly sub-triangular but with clearly curved lateral margin. Sternum 7 with deep Vshaped median excavation and fused with sterigma. Sterigma a spinulose, cup-shaped structure surrounded by a rim ventrally fused with S7 and dorsally consisting of a projecting spinulose fold in front of a more or less deep transverse pocket, laterally invaginated behind S7. Ductus bursae rather short and gradually widening towards corpus bursae, with a pair of short elongate sclerites just below ostium (colliculum), and remainder of ductus conspicuously and evenly granulate, with ductus seminalis from posterior part to midlength. Corpus bursae round to ovate; with one very large apically flattened and obtuse horn-shaped signum, with its hollow base and surrounding membrane protruding in the manner of a ‘pseudocapitulum’.
Distribution Atriscripta is known from Java, New Guinea and Australia. In Australia the genus is restricted to the eastern coast of northern Queensland south to Eungella.
Biology No host plant information is available for Atriscripta, but nearly all the Australian material is from rainforest localities with a few from monsoon forest.
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Remarks The genitalia of Atriscripta, especially those of the male, are so modified that they bear at first sight no resemblance at all to those of their nearest relatives. Consequently, the first species described was
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referred to Strepsicrates Meyrick (Diakonoff 1966). The wing venation, however, is characteristic for the tribe Olethreutini and the forewing pattern strongly suggests a close relationship with Sycacantha. Several convincing synapomorphies in
Figs 196–203. Atriscripta, gen. nov., genitalia and head. 196–200, A. arithmetica. 196, female genitalia, Mt Bellenden-Ker, Q, slide T2348. 197, male genitalia, detail of socii, Mulgrave R., Q, slide T2455. 198, male genitalia, Mt Bellenden-Ker, Q, slide T2323. 199, aedeagus, slide T2455. 200, head, male. 201–203, A. sp., Papua New Guinea, 201, 202, male genitalia and aedeagus, Amazon Bay, slide T2457. 203, female genitalia, Port Moresby, slide T2458.
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the genitalia of both sexes confirm this link. In both Atriscripta and Sycacantha a quite unusual sclerotised connection has developed in the membrane between gnathos/subscaphium and anellus. In Atriscripta this has led to a continuous well-sclerotised median band incorporating caulis, anellus and subscaphium. The remainder of the male genitalia of Atriscripta can easily be seen as derived from the same ancestral form as those of Sycacantha: socii of same type as Phaecasiophora Grote (probable sister genus of Sycacantha), and a generally similar structure of tegumen and valva, though more derived in Atriscripta. The attachment point of muscle m4 at the base of the pedunculi is still recognisable but much less developed in Atriscripta. The characteristic, evenly spaced granulation in the ductus bursa, the colliculum and an invagination in the lamella postvaginalis are all synapomorphies for the female genitalia of Atriscripta and Sycacantha. The following species has been described as Strepsicrates from West Irian and is here included in Atriscripta: decorigera (Diakonoff, 1966), Zool. Verh. (Leiden) 85: 68 (Strepsicrates). [New Guinea.] Comb. nov. Sycacantha incondita Diakonoff, 1973 from Java may well be found to also belong to Atriscripta once the male is examined.
Constituent species There are at least six species of Atriscripta worldwide, two of which are named, one from Australia: arithmetica (Meyrick, 1921), Exot. Microlepid. 2: 448 (Argyroploce). [Qld: Cairns.] Comb. nov.
Demeijerella Diakonoff (Figs 30, 204–212) Demeijerella Diakonoff, 1954, Verh. K. Ned. Akad. Wet. Nat., 2 ser. 49(4): 113. Type species: Demeijerella xanthorhina Diakonoff, 1954, ibid. 49: 114, by monotypy and original designation. [New Guinea.] Parepisimia Diakonoff, 1975, Zool. Meded. (Leiden) 48: 305. Type species: Parepisimia chrysoplea Diakonoff, 1975, ibid. 48: 306, by original designation. [New Guinea: Rossel I.]
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Diagnosis Demeijerella species are small to medium-sized moths with conspicuous yellowish and brown transverse bands across forewing, often with a yellowish head. Some members of the tortricine Heterochorista Diakonoff with the same combination of transverse yellow and brown stripes with a yellow head look superficially remarkably similar, even sharing a modified male hindwing costa with Demeijerella. Absence of a cubital pecten, structure of the antennae and wing venation, however, readily identify Heterochorista as tortricines. The forewing venation of Demeijerella is characterised by stalked R3 and R4, a rare feature in olethreutines, by the close proximity of the M-stem to the base of CuA and, in most species, by CuA2 originating unusually close to angle of cell. The males of all species known so far have modified scales along costa on upper surface of hindwing, but whether the apparently glandular scale sockets along Sc and R-stem are present in all Demeijerella species remains to be confirmed. The male genitalia with the large, lobe-shaped attachment points for muscle m4 on the tegumen and the evenly bristled valva with a basal hump on costa and a conspicuous apical tuft of bristles are diagnostic for Demeijerella. A simple, ventrally split, spinulose sterigma, absence of any colliculum-like structure but presence of an area of evenly spaced granulation in ductus bursa just below ostium and a folded sclerite at entrance to corpus bursa, together with two shark-tooth-shaped signa, are characteristics of the female genitalia.
Description Adults (Figs 204, 205) Wingspan. 10–18 mm; small to medium-sized. Head (Fig. 211). Ocellus well developed. Vertex scaling anteriorly barely protruding between antennae. Frons in upper part with moderately long, forward and downward projecting scales, lower part with minute, upwardly directed and appressed scaling. Proboscis at most 1.5× length of labial palpus. Labial palpus smallish (1.6–1.8× diameter of eye), porrect and lightly sinuate; second segment gradually widened towards moderately dilated apex with longish loose scales along distal margin only; terminal segment porrect, relatively long. Antenna reaching to middle of forewing; flagellum in male entirely covered by scaling and cilia minute.
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Thorax. With a small and ephemeral posterior crest. Legs unmodified. Wings (Figs 30, 204, 205). Forewing index 0.38–0.47; forewing subrectangular, dilated towards termen, widest just before apex, costa lightly curved at base and before apex, termen straight and somewhat oblique; discal cell rather short, 0.51– 0.57× length of wing, D. chrysoplea Diakonoff with well-developed, straight M-stem to base of M3, in distal half parallel and closely approximated to base of CuA, without any trace of chorda; R3 and R4 stalked, M2, M3 and CuA1 equidistant at base with CuA1 from angle of cell, base of CuA2 nearly always close to angle of cell, in D. chrysoplea not much further from CuA1 than M3; anal loop 0.32× length of 1A+2A; with a wing pattern of two welldefined, broad, transverse, yellowish fasciae on brownish-grey to brown ground colour, the inner one just before middle of wing continuous to dorsum, the outer one in some species reduced to a large yellow triangle on costa before apex. Hindwing with cubital pecten and axillary tuft, with modified costal area in male: D. chrysoplea with wide bands of presumably glandular scale sockets along the course of Sc and M-stem and with the area between costa and Sc with very long scales lying parallel to costa on upper side, framed along costa by a row of curved scales arising from costal edge of lower surface; Rs and M1 approximate but separate at base, M2 lightly bent towards M3 at base, M3 and CuA1 connate and CuA2 from far from angle of cell, CuP a weak trace only at margin; 3A faint but distinct; greyish. Pregenital abdomen. Unmodified. Sternum 8 in D. chrysoplea male with a medially concave hindmargin and T8 with a small triangular median process on hindmargin. Male genitalia (Figs 206–209). Vinculum a moderate band. Tegumen rather weakly sclerotised, with base of pedunculi produced into large medially projecting lobes as attachment points for m4. Uncus not differentiated, a small ridge along midline with some lateral bristles. Socii moderate to large semiporrect triangular lobes, densely bristled, with their dorsal margins meeting along midline. Gnathos two weak lateral bands. Valva long and narrow, curved or bent upwardly and evenly bristled throughout except for base and apex; with straight long costal process and a large, strongly projecting angle or hump at base of costa, sacculus ill-defined, distal part of cucullus only weakly bristled except for an isolated tuft of strong and usually curved bristles from apex that is
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asymmetrically produced into a point. Caulis moderate, anellus a dorsally incomplete ring. Aedeagus small, longitudinally striate or plicate and with an apparently recurved apex; vesica reportedly with spine-shaped cornuti (Diakonoff 1954). Female genitalia (Figs 210, 212). Ovipositor lobes subtriangular. Sternum 7 unmodified. Sterigma entirely within intersegmental membrane, rather simple and little sclerotised: lamella postvaginalis membranous, finely spinulose, lamella antevaginalis consisting of two small, roughly rectangular spinulose lobes separated by a median split. Ductus bursae with an area of conspicuous, evenly spaced granulation just below ostium, remainder smooth and membranous except for an irregularly folded sclerite just before entrance into bursa, ductus seminalis from this sclerotised part of ductus bursae. Corpus bursae ovate, very finely granulate, with two large, flat shark-toothshaped signa with rounded or acute apex.
Distribution Demeijerella is known from Sri Lanka, the Andaman Is, Sulawesi, New Guinea, the Solomon Is and Australia. In Australia the genus is restricted to the eastern coast of northern Queensland south to Mission Beach.
Biology No host records are available for any of the species of Demeijerella. All the Australian localities are rainforest habitats.
Remarks The synonymy of Parepisimia with Demeijerella (Diakonoff 1981) is unquestionable. With only males with their curiously modified valva available, Diakonoff (1954) described Demeijerella in the Chlidanotinae and, later, Parepisimia in the Eucosmini (Diakonoff 1975). The structure of the female genitalia, however, places Demeijerella squarely among the Olethreutini, as does the wing venation, despite some secondary modifications. Demeijerella is characterised by numerous apomorphies, including the wing pattern, stalked R3 and R4 in the forewing, modified scales along the hindwing costa in the male, the highly derived valva with a conspuous apical tuft and long, straight costal process, and female genitalia with spinulose, medially split ventral lip to the sterigma,
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Figs 204–211. Demeijerella, adults, genitalia and head. 204, D. sp., , Mission Beach, Q. 205. D. chrysoplea, , Babinda, Q. 206–209, male genitalia and aedeagus, 206, 207, D. sp., Mission Beach, Q, slide ANIC 2443. 208, 209, D. chrysoplea, nr Mt Finnigan, Q, slide T2276. 210, D. chrysoplea, female genitalia, slide T2301. 211, D. sp., head, male. Scale = 2 mm.
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Constituent species There are five described and possibly several unnamed species of Demeijerella. In Australia the genus is represented by an unnamed species and a named one that is also present in New Guinea: chrysoplea (Diakonoff, 1975), Zool. Meded. (Leiden) 48: 306 (Parepisimia). [New Guinea: Rossel I.]
Lobesia Guenée (Figs 31, 213–234) Lobesia Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 297. Type species: Asthenia reliquana Hübner, [1825] 1816 (=Tortrix permixtana Hübner, 1811 (nec Denis & Schiffermüller, 1775)), Verz. Bekannter Schmett.: 381, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 33, 59. [Europe.] Polychrosis Ragonot, 1894, Ann. Soc. Entomol. Fr. 63: 209. Type species: Tortrix botrana [Denis & Schiffermüller], 1775, Ankündung syst. Werkes Schmett. Wienergegend: 131, by original designation (on page 208). [Europe.] Lomaschiza Lower, 1901, Trans. R. Soc. S. Aust. 25: 68. Type species: Lomaschiza physophora Lower, 1901, ibid. 25: 69, by monotypy. [Australia.]
Fig. 212. Demeijerella sp., Mission Beach, Q, female genitalia, slide ANIC 17083.
a sclerite near the entrance to the corpus bursa and two shark-tooth-shaped signa. Though of a very different appearance there is little doubt that Demeijerella is a highly derived member of the Sycacantha-group, possibly the sister group of Phaecasiophora Grote. The most convincing synapomorphies are provided by the male genitalia: 1, a strongly developed, medially projecting attachment point for muscle m4 on tegumen; 2, large, subporrect socii lobes from apex of tegumen, and 3, a simple valva with a pronounced basal hump to costa. The female genitalia are much more diversified, but presence of a very characteristic, evenly spaced granulation in at least parts of the ductus bursae is a further conspicuous synapomorphy linking the group. Demeijerella also shares several unusual characters with Lobesia, as discussed in the Remarks for that genus.
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Byrsoptera Lower, 1901, Trans. R. Soc. S. Aust. 25: 77. Type species: Byrsoptera xylistis Lower, 1901, ibid. 25: 77, by monotypy. [Australia.] A junior homonym of Byrsoptera Spinola, 1837, Essai Insectes Hémiptères: 191, – Insecta, Hemiptera. Steriphotis Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 225 (key), 259. Type species: Steriphotis peltophora Meyrick, 1911b, ibid. 36: 259, by original designation. [Australia.] Analdes Turner, 1916, Trans. R. Soc. S. Aust. 40: 533. Type species: Analdes hypolepta Turner, 1916, ibid. 40: 534 (=Grapholita extrusana Walker, 1863), by monotypy. [Australia.] Lomaschizodes Diakonoff, 1954, Zool. Verh. (Leiden) 22: 15. Type species: Grapholita extrusana Walker, 1863, List specimens Lepid. Insects Colln Br. Mus. 28: 392, by original designation. [Proposed as a subgenus of Lobesia.] [Australia.] Harmosma Diakonoff, 1963, Tijdschr. Entomol. 106: 354. Type species: Polychrosis harmonia Meyrick, 1908, Proc. Zool. Soc. Lond. 1908: 716, by
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monotypy and original designation. [Originally proposed as a subgenus of Lobesia.] [Africa.] Pirireisia Koçak, 1981, Priamus 1: 114, originally proposed as an unnecessary objective replacement name for Byrsoptera Lower, 1901. Neolobesia Bae & Komai, 1991, Tyô to Ga 42: 136. Type species: Lobesia coccophaga Falkovitsh, 1970, Vestnik Zool. 1970 (5): 62, by original designation. [Proposed as a subgenus of Lobesia.] [Russia.] Neodasyphora Bae & Liu, 1995, Japan. J. Entomol. 63: 108. Type species: Polychrosis cunninghamiacola Liu & Bai, 1977, Acta Entomol. Sin. 20: 217, by original designation. [Proposed as a subgenus of Lobesia.] [China.]
Diagnosis Lobesia species are small to medium-sized usually intricately patterned and mainly brownish moths, frequently with characteristic forewing markings consisting of roughly transverse banding on basal half ending in a transverse triangle with apex towards termen and connected with a large circular blotch centred on tornus and with three pairs of costal strigulae before apex merging into an outwardly curved white line running to termen and delineating a small round apical spot. The antennal flagellum is also scaled on its anterior surface. The strongly modified forewing venation with deflected R1 running parallel to costa and distally approximated to tip of R2 is present in most species of Lobesia and is characteristic for the genus, together with the pterostigma between R1 and costa usually present in the male. In all the Australian species examined the chorda originates between R2 and R3, and the M-stem is always absent. In the hindwing M2 and M3 are distant at base (except in males with modified venation) and M3 and CuA1 vary from quite distant to stalked. Males of Lobesia are usually characterised by a number of secondary sexual modifications: pterostigma in forewing, often a modified hindwing of triangular shape with enlarged anal area and reduced translucent scaling, tibial pencil, paired elongate pockets with modified scales laterally on the basal abdominal sternite and in a few taxa (L. extrusana (Walker), L. peltophora (Meyrick)) eversible scale pencils deeply invaginated laterally between segments 3 and 4. Of these, only the scale pockets laterally on S2 are diagnostic for Lobesia, being present at least in reduced form in all groups. The male genitalia with
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the usually hooked apodeme for muscle m4, the flap-like socii continuous with the teguminal apex and the valva with a longitudinal narrow fold or ridge along its outer surface and with several groups of spines along the basal half of the ventral edge, usually separated by one or two deep notches, are diagnostic for the genus. The female genitalia of Lobesia are very diverse, with a complex and often deeply invaginated sterigma, and primarily reflect species-groups within the genus, but inception of ductus seminalis close to ‘neck’ of corpus bursae is a diagnostic feature.
Description Adults (Figs 213–218) Wingspan. 7–19 mm; small to medium-sized. Head (Fig. 231). Ocellus well developed. Chaetosema often large. Vertex with longish, converging scales, anteriorly protruding between antennae. Frons in upper third with a transverse band of longish, anteriorly projecting scales, remainder with minute, appressed and upwardly oriented scales. Proboscis short, hardly longer than labial palpus. Labial palpus smallish to moderately long (1.3–2.1× diameter of eye), sinuate, porrect and moderately widened; second segment lightly upcurved to sinuate, moderately to quite strongly widened towards apex with long loose scales along ventral margin and apical portion of dorsal edge; terminal segment porrect, variable from rather short and stout to long and pointed, especially in female. Antenna reaching to middle of forewing; flagellum in male with scales also on anterior surface, though often of paler colour; cilia in male very small. Thorax. With large posterior crest. Legs with hind tibia in male usually with a hairpencil of variable length dorsally from base, often as long as tibia and distal portion concealed in a groove in scaling (particularly large and bipartite black/white in L. peltophora). Wings (Figs 31, 213–218). Forewing index 0.33– 0.40; forewing nearly always elongate subtriangular, sometimes longer and more slender in male and with a prominent hump at 36: costa (L. physophora (Lower)); costa usually nearly straight in basal half and lightly curved towards apex, with wing membrane especially in male variably thickened into a pterostigma between distal portion of R1 and costa, forming a protruding cylindrical structure projecting on costa if highly developed; apex rounded-acute and termen straight to weakly convex and strongly oblique; discal cell 0.61–0.64×
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Figs 213–218. Lobesia, adults, males except 214. 213, 214, L. physophora, nr Mt Garnet, Q. 213, . 214, [image reversed]. 215, L. sp., harmonia-group, nr Gogango, Q. 216, L. extrusana, Minnamurra Falls, NSW [image reversed]. 217, L. peltophora, Tewantin, Q [image reversed]. 218, L. xylistis, Townsville, Q [image reversed]. Scale = 2 mm.
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length of wing, with distal third often narrowed, with accessory cell, if present, usually very narrow with chorda varying from moderately developed to a trace only and originating from between R1 and R2 (most Palaearctic species) or between R2 and R3 (many Australian species), with M-stem at best a trace between M2 and M3; with all veins separate beyond discal cell (R4 and R5 very shortly stalked in one Palaearctic species figured by Diakonoff (1954)), with course of R1 and R2 modified in most species (in both sexes) with R1 bent and its distal portion displaced by pterostigma and running parallel to costal margin, R2 strongly sinuate with middle portion roughly parallel to costal margin and distal part often closely approximated to R3; with bases of R3, R4 and R5 moderately distant to closely approximated in species with strongly developed pterostigma, with M2, M3 and CuA1 equidistant or the latter two veins slightly closer at base; CuP present only at wing margin, base of 1A sometimes non-tubular; with very diverse wing pattern but numerous species with ‘typical’ Lobesia markings: basal half of wing with roughly transverse banding ending in a transverse triangle with apex pointing towards termen and connected with a large circular blotch centred on tornus, three pairs of costal strigulae before apex merging into an outwardly curved white line running to termen and delineating a small round apical spot. Hindwing with cubital pecten and axillary tuft present (unless sexually dimorphic) and in shape and structure in several species with a roughly triangular outline in male with narrow apex, straight or deeply concave termen, greatly enlarged anal lobe often with enlarged fringe, and a translucent sparse covering of very narrow scales except for apex (L. physophora); Rs and M1 varying from separate but closely approximated at base (more distant in modified male condition: L. physophora) to stalked (L. peltophora, L. extrusana), M2 normally distant from M3 at base and either parallel or only slightly bent towards M3, and M3 and CuA1 varying from quite distant to stalked (L. extrusana), but in modified male wing M2 and the entirely fused M3 and CuA1 may both be very short, rather close and subparallel (L. physophora); normally with CuP a weak trace and 1A+2A and 3A both present but in modified male wing CuP may be entirely reduced and 1A+2A and 3A widely separated and the former modified to a bulging crease; from snow-white to dark brownish grey, sometimes transversely marbled and apex often darker, with basal portion often translucent in male due to modified narrow scales.
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Pregenital abdomen. In male often distinctly elongated and nearly always with a pair of elongate depressions or pockets filled with modified scales laterally on S2 (Fig. 222), and in males of one species-group (L. extrusana, L. peltophora) also with two large eversible scale pencils deeply invaginated into membranous sacs laterally between segments 3 and 4 (Fig. 224); males of L. extrusana in addition with a band of persistent, modified scales along anterior margin of S4. Sternum 8 in male with faintly bilobed hindmargin. Male genitalia (Figs 219–221, 223, 225– 228). Vinculum a weakly sclerotised band. Tegumen elongate-ovate, rather weakly sclerotised, with very prominent apodemes for muscle m4 varying from semicircular lobes to large recurved and pointed hooks. Uncus reduced to a low single or bipartite hump on apex of tegumen. Socii varying from small lateral lobes to large drooping flaps weakly delineated from top of apex and carrying bristles or modified scales (L. extrusana); more rarely socii absent. Gnathos usually weakly sclerotised and curved, fused medially to form an often W-shaped band, in some groups (L. physophora) associated with a single or paired rising pointed process consisting of numerous aggregated sclerotised teeth or spinules. Valva very diverse, nearly always long and slender, with a longitudinal narrow fold along outer surface from middle of costa to below costal process, with basal half always with several groups of spines along ventral margin that are usually separated by one or two excavations in ventral edge, and with cucullus from rather short, wide and dense and evenly bristled except for an often prominent spine at its base near ventral margin to long, thin and hairy except for short spines along ventral margin in some species. Caulis from short to very long, anellus a membranous ring. Aedeagus from very short (L. peltophora) to long and slender with a slightly bulbous base, apex bevelled and in some species (L. physophora) with a few teeth on outer surface towards apex; vesica without any trace of cornuti. Female genitalia (Figs 229, 230, 232– 234). Ovipositor lobes elongate ovate, modified and slightly swollen in some species. T8 sometimes enlarged and strongly sclerotised. Sternum 7 with straight or concave hindmargin. Sterigma extremely diverse, varying from a strongly protruding spinulose hump ventrally of simple membranous ostium (in undescribed Australian species), to a
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large, raised, often roughly inverted triangular and strongly spinulose structure surrounding ostium (L. extrusana, L. peltophora) to a deeply invaginated spinulose and hour-glass-shaped cylinder (L. physophora) or smooth inverted funnel surrounding proximal portion of ductus bursae, the latter condition usually combined with paired, flat, rounded pockets invaginated behind hindmargin of S7. Ductus bursae membranous, without colliculum except for L. extrusana that has possible traces of a short bipartite colliculum, transition into corpus bursae often gradual, but if corpus bursae distinct, inception of ductus seminalis always close to bursa or from its ‘neck’. Corpus bursa varying from ovate, large and well-defined, with ductus seminalis originating from close to its ‘neck’ (L. extrusana, L. peltophora) to continuous with ductus as long, narrow, gradually widening sac with inception of ductus seminalis at about midlength (L. physophora), with intermediate forms with a very long thin ‘neck’ to corpus bursa from inception of ductus seminalis and a small inflated distal portion, often with the two anterolateral corners expanded into globular extensions; mostly without signum but with a paired spinulose depression close to ‘neck’ in an undescribed Australian species close to L. harmonia and a weakly spinulose, ill-defined and sometimes bipartite patch of sclerotisation in L. physophora and related species.
Distribution Holarctic, Africa, Asia, New Guinea, Micronesia, and Australia where the genus occurs in the coastal regions of northernmost Western Australia, the Northern Territory, Queensland and New South Wales.
Biology In the Palaearctic and Oriental regions Lobesia is found on a wide range of host families, mainly feeding in spun shoots, flowers, seedheads and fruit. The grapevine pest L. botrana (Denis & Schiffermüller) feeds in flowers and developing or ripe fruit of Vitis vinifera (Vitaceae). Many species of Lobesia are polyphagous (Diakonoff 1954; Hannemann 1961; Bae and Komai 1991). Australian host records confirm a tendency for flower and fruit feeding with members of the L. peltophora complex repeatedly reared from flowers of Melaleuca quinquenervis (Myrtaceae) (Balciunas personal communication), L. physophora reported
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from inflorescence of Litchi sinensis (Sapindaceae) and berries of Lantana sp. (Verbenaceae) and an unidentified species reared from seed pods of Buckinghamia celsissima (Proteaceae).
Remarks Due to the wide distribution of Lobesia a large body of information is available on the genus. Obraztsov (1953) and Diakonoff (1954, 1963, 1973) addressed interrelationships within the genus and its affinity to related taxa on the basis of the Holarctic respectively Oriental/Australasian fauna, providing numerous illustrations. Photos of wings and genitalia of the types of many species of Lobesia as well as species erroneously described in this genus were given by Clarke (1958). Local faunistic treatments (Heinrich 1926; Hanneman 1961; Bentinck & Diakonoff 1968; Bradley et al. 1979; Kawabe 1982; Park 1983) often also contain biological information such as host records. Bae and Komai (1991) provided an excellently illustrated account of the Japanese fauna, with a key to the subgenera. Liu and Bae (1994) first revised and illustrated the Chinese species and later (Bae and Liu 1995) described a new subgenus for one Chinese species. Lobesia is undoubtedly a monophyletic group characterised by several apomorphies such as the pockets with modified scales laterally on S2 in the male and the structure of the teguminal apex and the socii. Prominent and often hook-shaped apodemes for muscle m4 on the pedunculi, inception of ductus seminalis close to transition of ductus bursae into corpus bursae and an antenna with flagellum scaled also on its anterior surface are further characteristic features of all Lobesia, though the polarity of these character states is unclear. The typical pterostigma in the forewing of Lobesia and the resulting deflection of the course of R1 and R2 is diagnostic for the genus, if present, but these features are missing in some species with a more plesiomorphic wing venation. The large range of secondary sexual modifications present in Lobesia, together with very diverse genitalia structures in both sexes, has led to the creation of numerous genus-level taxa for the group, many of them based on Australian species. In his revisions of the Oriental, Papuan and African fauna Diakonoff (1954, 1963, 1973) accepts these groupings at subgeneric level. Bae and Komai (1991) and Bae and Liu (1995) subsequently added two more subgenera, both monotypic. The present
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revision of the rich and very diverse Australian fauna easily identifies several derived groups, but they all appear to be subordinate within Lobesia and their removal would create paraphyletic residual taxa. Characters are distributed mosaic-
like throughout the genus and some of the most obvious apomorphies for certain groups are present in a more cryptic condition in a much less derived branch. Therefore, a more informal classification based on species-groups has been adopted,
Figs 219–224. Lobesia, male genitalia and abdomen base (lateral scale tufts indicated by arrows). 219, 220, L. xylistis, male genitalia and aedeagus, Townsville, Q, slide T2159 (m4, muscle attachment). 221, 222, L. physophora, male genitalia and abdomen base, nr Mt Garnet, Q, slide T2435 (m4, muscle attachment). 223, 224, L. peltophora, male genitalia and abdomen base, Brisbane, Q, slide T2193 (va, ventral apodeme).
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Figs 225–228. Lobesia, male genitalia and aedeagus. 225, 226, L. extrusana, Mt Keira, NSW, slide T2117. 227, 228, L. sp., harmonia-group, nr Emerald, Q, slide T2318.
discussed below for those groups present in the Australian fauna. Consequently only the subgenera relevant to the Australian fauna are listed in the above synonymy of Lobesia, and these speciesgroups are equated with Diakonoff’s (1954, 1963) classification in the light of some senior synonyms. Australia has an exceptionally rich and still poorly collected fauna of Lobesia, and, given the strong sexual dimorphism impeding association of the two sexes, the identity of several named species has only been tentatively established. The otherwise African harmonia-group (Diakonoff 1963, as subgenus Harmosma Diakonoff, type species: Polychrosis harmonia Meyrick, 1908), with its underived wing venation and rather Bactra-like appearance is represented by an undescribed Australian species (Fig. 215). The genitalia of the two species are very similar. The Australian peltophora-group (Diakonoff 1954, as subgenus Lomaschizodes Diakonoff, type species Grapholita extrusana Walker, 1863) is characterised by a long thin valva usually with a deep ventral excavation, by a raised large inverted-
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triangular spinulose sterigma and in Australian species by eversible scale pencils invaginated laterally between abdominal segments 3 and 4 in the male. Diakonoff (1954) described his subgenus Lomaschizodes unaware of the fact that two generic names were already available for this group, namely Meyrick’s (1911b) Steriphotis based on peltophora Meyrick and Turner’s (1916) Analdes based on hypolepta Turner, a junior synonym of extrusana. The socii are usually large drooping flaps set with long broad scales (L. extrusana, L. paradisea Diakonoff), but in L. peltophora they are reduced to small lateral lobes with a few bristles only. Males of the peltophora-group have a well-developed pterostigma but a sexually dimorphic hindwing is found in the Papuan L. paradisea only and not in the Australian species. The corpus bursae is differentiated but lacks a signum. The physophora-group (Diakonoff 1963, as subgenus Lomaschiza Lower, 1901, type species: Lomaschiza physophora Lower) is characterised by strongly sexually dimorphic male hindwings, male genitalia with two excavations in the ventral
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Figs 229–231. Lobesia, female genitalia and head. 229, L. xylistis, Mt Keira, NSW, slide ANIC 2760. 230, L. physophora, nr Mt Garnet, Q, slide T2436. 231, head, L. extrusana, male.
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margin of the narrow valva and a gnathos with two apically free, rising processes of numerous aggregated teeth or spinules. The female genitalia have a deeply invaginated hour-glass-shaped or bulbous, conspicously spinulose sterigma that surrounds the posterior part of the ductus bursae. Corpus and ductus bursae form a narrow continuous sac without signum. Association of the two sexes is often difficult as the male, in addition to a strongly modified hindwing, also has a narrower forewing with a projecting pterostigma. The xylistis-group, with its broad and densely bristled cucullus often with a larger prominent bristle at its base in the middle of the valva, is richly represented in Australia. The female genitalia have a complex sterigma consisting of a smooth, invaginated and inverted funnel-shaped structure surrounding the posterior part of the ductus bursae and associated with two flat distally rounded pockets invaginated behind hindmargin of S7. The corpus bursae has a long slender ‘neck’ and usually a rather large irregular or slightly bipartite, weakly spinulose sclerotisation in its (?dorsal) wall, and in some species the anterolateral corners are expanded ventrally into globular expansions. Diakonoff (1954) included this group in the subgenus Lobesia, though the type species of Lobesia, Asthenia reliquana Hübner, has a spinulose and bulbous sterigma albeit combined with the characteristic pockets invaginated behind S7. Several characters of the xylistis-group point to a closer relationship with the subgenus Polychrosis, and Bae and Komai (1991) solved this problem by treating Polychrosis as a synonym of the subgenus Lobesia. Koçak (1981) unfortunately created a further synonym in describing the unnecessary replacement name Pirireisia Koçak for Lower’s (1901) Byrsoptera that is based on B. xylistis Lower and had long been synonymised with Polychrosis (Meyrick 1911b). Lobesia transtrifera (Meyrick) is also a member of the xylistis-group, judging from its female genitalia (Clarke 1958), but so far no males have been associated to confirm this relationship. Lobesia symploca (Turner) and L. arescophanes (Turner) are both known from single holotypes only and cannot be assigned to a species-group. Several species within the Australian fauna either do not fall within any of the above speciesgroups or combine characters of several groups. This supports the conclusion that whilst some of the subgenera are probably monophyletic they are all subordinate within Lobesia and their separation creates paraphyletic taxa such as the xylistis-group. Two obvious apomorphies found in this group, the
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Figs 232–234. Lobesia, female genitalia. 232, L. peltophora, Brisbane, Q, slide T2204. 233, L. extrusana, Mt Keira, NSW, slide T2118. 234, L. sp., harmonia-group, Emerald, Q, slide T2338.
prominent enlarged spine at the base of the cucullus in the centre of the valva and the expanded anterolateral corners of the corpus bursae, are further developed in the Nearctic Paralobesia viteana (Clemens). The cucullus spine has been modified into a huge spike in P. viteana and the bursa still shows a trace of the sclerotised plate but is distally tripartite with two narrow-necked globular appendices. There is little doubt that Paralobesia Obraztsov (Obraztsov 1953) is subordinate within Lobesia (Bae and Komai 1991) but the topic is well beyond the scope of this review. Lobesia is so highly derived that its affinities have been obscure. Falkovitch (1962) raised the group to tribal level on the strength of the combination of tibial pencil and abdominal scale pockets as male scent-producing organs rather than the usual tibial pencil/dorsal roll combination found in most Olethreutini. Diakonoff (1973) treats the group as a subtribe but includes several other genera without identifying convincing
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apomorphies that would link them with Lobesia and Paralobesia. Podognatha and Theorica both belong to the Zomaria-group. Didrimys is subordinate within Ophiorrhabda and its gnathos processes, deceptively similar-looking to those found in the physophora-group, are clearly convergent. Engelana Diakonoff and Xenolepis Diakonoff both lack the prominent apodemes for muscle m4 that are so characteristic for Lobesia, but a certain overall similarity in the male genitalia cannot be denied. Lobesia shares several unusual characters with Demeijerella of the Sycacantha-group. Both genera have the antennal flagellum scaled also on its anterior surface and the ductus seminalis originating from close to the corpus bursae, and they both lack a colliculum. Conspicuously developed apodemes for muscle m4 are a further trait that is found throughout the Sycacanthagroup, and the structure of the uncus/socii complex shows strong similarities with Lobesia. The small
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crescentic folds or a hairpencil at the base of the abdomen laterally from S2 in Sycacantha are paralleled by the abdominal scale pockets of Lobesia in essentially the same position, providing a strong link between the two genera. A narrow longitudinal fold along the outer surface of the basal half of the valva found in Sycacantha and most species of Lobesia could be a further synapomorphy. Consequently, as a working hypothesis, Lobesia is here treated as a member of the Sycacantha-group. Lobesia peltophora has a superficially extremely similar looking sibling species with a white hind tibia with a slender yellowish pencil in the male instead of the partly black tibia of L. peltophora with a large and conspicuous black and yellowish pencil. The genitalia in both sexes are quite different with a roughly rectangular sterigma with two protruding vertical folds in the sibling species, in contrast to the inverted-triangular sterigma of L. peltophora. Bradley’s (1961) record of Steriphotis peltophora from the Solomon Is will have to be reconsidered in view of this sibling species.
Constituent species A very cursory survey of the literature suggests at least 30 confirmed and named species of Lobesia for the Australasian and south-east Asian regions, with probably many more yet to be described. Brown (2005) listed 103 valid species worldwide. Eight named and at least as many unidentified species are known from Australia: peltophora (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 259 (Steriphotis). [Qld: Brisbane.] extrusana (Walker, 1863), List Specimens Lepid. Insects Colln Br. Mus. 28: 392 (Grapholita). [Qld: Brisbane.] discotypa (Turner, 1916), Trans. R. Soc. S. Aust. 40: 510 (Capua). [Qld: Mt Tamborine.] hypolepta (Turner, 1916), Trans. R. Soc. S. Aust. 40: 534 (Analdes). [Qld: Brisbane.] xylistis (Lower, 1901), Trans. R. Soc. S. Aust. 25: 77 (Byrsoptera). [Qld: Cooktown.] transtrifera (Meyrick, 1920), Exot. Microlepid. 2: 346 (Polychrosis). [Qld: Brisbane.] physophora (Lower, 1901), Trans. R. Soc. S. Aust. 25: 69 (Lomaschiza). [Qld: Cooktown.] arescophanes (Turner, 1945), Trans. R. Soc. S. Aust. 69: 69 (Cnephasia). [Qld: Cape York.] symploca (Turner, 1926), Trans. R. Soc. S. Aust. 50: 139 (Proschistis). [Qld: Caloundra.]
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parvulana (Walker, 1863), List Specimens Lepid. Insects Colln Br. Mus. 28: 391 (Grapholita). [Qld: Brisbane.]
Gnathmocerodes-group Status remarks Diakonoff (1973) established the subtribe Gnathmocerodides for Gnathmocerodes Diakonoff, Heteroschistis Diakonoff and Dynatorhabda Diakonoff. However, Heteroschistis is an enarmoniine genus (Horak 1981; Brown 2005) and Dynatorhabda is in this book referred to the Oxysemaphora-group. Rhectogonia Diakonoff is here added to the Gnathmocerodes-group.
Evidence for monophyly Diakonoff (1973) based his Gnathmocerodides on the single, cornute signum. Rhectogonia not only shares a single, rugose, horn-shaped signum, but also a modified gnathos with a distally bifid process, a large, sclerotised, bowl-shaped anellus, and very similar socii and valvae. Both genera have a tibial pencil and enlarged hindwing anal area at least in some species, as do many other Olethreutini, and one unnamed Western Australian species of Gnathmocerodes shares a hindwing with stalked M3 and CuA1 with Rhectogonia.
Distribution Both Gnathmocerodes and Rhectogonia are known from Sri Lanka, India, Myanmar, Thailand, New Guinea and Australia (Queensland), the former also from Madagascar, Vietnam, the Philippine Isands and the Northern Territory in Australia.
Biology Three species of Gnathmocerodes have been reared from Barringtonia acutangula (Lecythidaceae), explicitly as a leafroller where the biology was described in detail (Diakonoff 1973; Spitzer and Jaros 1985).
Remarks Rhectogonia and an undescribed Western Australian Gnathmocerodes species are among the few Olethreutini with a hindwing with stalked M3
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and CuA1. However, the tibial pencil and the genitalia of both sexes leave no doubt about their tribal position.
Gnathmocerodes Diakonoff (Figs 32, 235–243) Gnathmocerodes Diakonoff, [1968], US Nat. Mus. Bull. 257: 71. Type species: Gnathmocerodes petrifraga Diakonoff, [1968], ibid. 257: 72, by monotypy and original designation. [Philippine Is.] Agnathmacera Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 27. Type species: Gnathmocerodes (Agnathmacera) labidophora Diakonoff, 1973: ibid. 1: 28, by monotypy and original designation. [Proposed as a subgenus of Gnathmocerodes.] [New Guinea: d’Entrecasteaux Is.]
Diagnosis Gnathmocerodes species are small to medium-sized, intricately patterned, ochreous brown to brown moths with a large darker brown blotch or mark of inverted-triangular shape between 369 and 169 costa, a small pale roundish spot halfway between closing vein and termen across R5 and M1, and usually with an at least partly developed paler band zigzagging across wing from 369 costa to middle of dorsum, then back towards apex, curving around pale spot and ending in middle of termen. Gnathmocerodes has sinuate, subporrect and moderately widened labial palpi and characteristic secondary sexual modifications to hindwing and hind legs in several species. The male hind tibia and the first tarsal segment are often enlarged by scale brushes as well as a large tibial pencil, and the anal area of the hindwing may be produced into a medially projecting triangular lobe with inwardly oblique anal margin (in expanded wing), with its margin rolled under over a fringe of modified scales. A medially extended anal lobe with a strongly sinuate 1A+2A and an inwardly oblique anal margin in the expanded male hindwing is a unique feature and, if present, is diagnostic for Gnathmocerodes. The venation otherwise is unremarkable for an Olethreutini except for the unusually well-developed hindwing CuP. The male genitalia with a broad, flat, weakly sclerotised and
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dorsally hairy uncus, with drooping narrow socii, and with either medially strongly constricted or ventrally deeply excavate, usually asymmetrical valvae with a group of short spines on a digit on the margin of the basal excavation and/or an often bipartite tuft of bristles on the projecting ventral angle towards the base of the valva, are diagnostic for Gnathmocerodes. The female genitalia with a strongly raised spinulose rim encircling the entrance to the ostium, a very long tube-like colliculum and a large single signum, a granulate hollow sharp and straight horn on a sclerotised round basal plate, are equally characteristic.
Description Adults (Figs 235–237) Wingspan. 10.5–18.0 mm; small to mediumsized. Head (Figs 241). Ocellus well developed to large. Vertex scaling anteriorly barely protruding between antennae. Uppermost part of frons with a transverse band of moderately long anteriorly projecting scales, remainder with minute, appressed and upwardly oriented scales. Maxillary palpus minute but usually clearly visible due to whitish scaling. Proboscis short, hardly longer than labial palpus. Labial palpus moderately long (1.6– 1.9× diameter of eye), sinuate and subporrect, moderately widened; second segment strongly upcurved, sinuate only at tip, moderately widening towards apex and with long loose scales along ventral margin and across apex; terminal segment moderately long and porrect, longer and more pointed in female. Antenna reaching to middle of forewing; anterior surface unscaled; cilia in male very short, less than 56: diameter of flagellar segments. Thorax. With large posterior crest. Hind legs in male usually modified (unmodified in G. euplectra Lower), either with a slender tibia with a hairpencil or with tibia and first tarsal segment both greatly enlarged by densely packed long scales, usually appressed on inner and outer surface but forming a large loose crest along dorsal surfaces of tibia and first tarsal segment and on both dorsal and ventral margins of inner tibial spurs, with tibial pencil developed into a huge bundle of usually snow-white hairscales in a groove among the glossy, pale-tipped long scales of inner surface (G. ophiocosma Turner). Wings (Figs 32, 235–237). Forewing index 0.43–0.45; forewing elongate subtriangular, with
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evenly curved costa, rounded-acute apex and oblique, straight termen; discal cell 0.63–0.66× length of wing, with moderately narrow accessory cell with chorda from between R1 and R2 and with partly nontubular M-stem to base of M2; with all
Figs 235–237. Gnathmocerodes, adults. 235, G. ophiocosma, holotype, , Cairns, Q. 236, 237, G. euplectra, 236, , Mt Webb Nat. Park, Q. 237, , Townsville, Q. Scale = 2 mm.
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veins present and separate beyond discal cell, with origin of R2 varying from closer to R1 (G. tonsoria (Meyrick)) to closer to R3 (G. euplectra); intricately marbled and patterned with ochreous and brownish, always with a pale round or oval spot halfway between closing vein and termen across R5 and M1, and with a sometimes partly obliterated pale band zig-zagging across wing from a conspicuous pale bar at 369 costa to middle of dorsum, then back towards apex, curving around pale spot and ending in middle of termen, enclosing a large dark inverted triangle or blotch between 369 and 169 costa. Hindwing with well-developed cubital pecten and axillary tuft and especially in male with numerous long hairscales scattered across upper surface between CuP and anal margin, in several species (G. ophiocosma, G. tonsoria) with anal area greatly enlarged with 3A widely distant from 1A+2A and anal margin inwardly oblique in expanded wing with edge narrowly rolled under over modified curved scales fringing margin and with a fold from upper surface topographically underlying ‘dorsal roll’; with Rs and M1 parallel and closely approximated at base, M2 at base bent closely towards connate M3 and CuA1; with CuP developed to below middle; 1A+2A and 3A both well developed but base of 1A weak, 1A+2A strongly sinuate in males with enlarged anal area. Pregenital abdomen. Unmodified. Sternum 8 in male large, with medially concave hindmargin. Male genitalia (Figs 238–240). Vinculum a weakly sclerotised band. Tegumen elongate-ovate with obliquely rounded ‘shoulders’, pedunculi broad to club-shaped with broad median attachment lobes for muscle m4. Uncus a broad and flat, moderately sclerotised process varying in shape from a low triangle (G. euplectra) to a large, apically pointed pentagon (G. ophiocosma), with dense long bristles dorsally along entire margin. Socii densely bristled, drooping, elongate lobes to distally rounded bands curved toward midline. Gnathos two variably developed curved lateral bands converging at base of anal tube, often strongly sclerotised and connected with two parallel sclerotised bands rising along ventral surface of anal tube and ending in a T-shaped sclerotisation with projecting free lateral arms below apex of anal tube (G. euplectra). Valva sometimes asymmetrical and always ventrally constricted beyond large to very large basal excavation, central part of valva frequently reduced to a thin curved ‘neck’ separating cucullus (G. euplectra), with costal process small and lightly sinuate, basal half of valva
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Figs 238–241. Gnathmocerodes, male genitalia and head. 238, 239, G. ophiocosma, holotype, Cairns, Q, slide ANIC 2741. 240, G. euplectra, Rockhampton, Q, slide T1944. 241, head, G. euplectra, male.
with strongly projecting rounded-triangular ventro-distal corners often carrying asymmetrical single or double tufts of bristles, with a small group of short bristles on distal edge of basal excavation, often raised on a digit (G. ophiocosma), cucullus strongly dilated, usually broadly tongue-shaped with projecting ventro-proximal angle and varying from evenly bristled throughout with only some stronger spines along ventral margin (G. ophiocosma) or a patch of strong thorns beyond ‘neck’ (G. lecythocera (Meyrick)) to sparsely bristled with very large spines around ventro-proximal angle. Caulis rather short, anellus with dorsal portion often extended into a large, inverted V- to U-shaped sclerotisation in diaphragm reaching towards the anal tube. Aedeagus long and curved, usually slightly bulbous at the base and tapering to apex; vesica without cornuti. Female genitalia (Figs 242, 243). Ovipositor lobes elongate-oval to subtriangular. Sternum 7 often of species-specific shape, moderately to
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strongly tapering towards straight to deeply concave hindmargin. Sterigma an irregularly shaped spinulose area behind S7 with a strongly raised central rim encircling entrance to ostium, with a broader ‘upper lip’ (lamella postvaginalis) in G. euplectra and sometimes caudo-lateral angles of sterigma extended into two additional small swellings. Ductus bursae with a very long colliculum, a slender evenly sclerotised tube reaching to nearly mid-length, remainder of ductus membranous with inception of ductus seminalis anterior to midlength and last portion before corpus bursae slightly widened. Corpus bursae round to oval, weakly granular; with one large signum: a straight, sharply pointed, hollow horn with granular surface on a sclerotised round basal plate.
Distribution Madagascar, Sri Lanka, India, Andaman Is, southeast Asia to Vietnam and the Philippine Is, New
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Guinea, Australia. In Australia the genus is found in the Kimberley of Western Australia and coastal regions of the Northern Territory and Queensland south to Rockhampton.
Biology Spitzer and Jaros (1985) discussed in detail the biology of G. petrifraga Diakonoff ‘feeding in rolled or folded leaves on Barringtonia acutangula L.’ (Lecythidaceae) in Vietnam, adding a third species to the two already reared from fruit, top shoots and leaves of Barringtonia (Diakonoff 1973). One of these, G. lecythocera (Meyrick), has also been reared from a species each of Anonaceae and Malvaceae (Diakonoff 1973). The only host records from Australia are a rearing of G. euplectra from a fruit of Planchonia careya (Lecythidaceae), confirming the link with Lecythidaceae, and an undescribed species reared from Barringtonia acutangula in the Northern Territory by G. Fitt (ANIC).
Remarks Diakonoff ([1968], 1973) provided figures of adult, male genitalia and wing venation for his initial description of Gnathmocerodes (Diakonoff [1968]) and later (Diakonoff 1973) comprehensively reviewed the genus, describing or redescribing and figuring several additional species. Two species described by Meyrick were illustrated by Clarke (1958; as Olethreutes tonsoria (Meyrick) and O. lecythocera (Meyrick)). Diakonoff (1973) subdivided Gnathmocerodes into two subgenera, but as the Australian G. euplectra combines a ventrally sclerotised anal tube with two free distal processes, a feature diagnostic for the subgenus Gnathmocerodes, with a forewing venation typical of the subgenus Agnathmacera, this division cannot be maintained. As an entity, however, Gnathmocerodes is a welldefined, clearly monophyletic group with numerous autapomorphies such as the nature of the secondary sexual hindwing modification (absent in some species) and the structure of the uncus and sociicomplex, the ventrally sclerotised anal tube and the valva in the male. Diakonoff (1983c) described Gnathmocerodes (Leptocera) microchlamys Diakonoff as a new subgenus from Madagascar that will have to be reassessed once the female is known. Several apomorphies shared by Gnathmocerodes and Rhectogonia are discussed in the Remarks for the latter genus. The Australian G. ophiocosma and G. alphestis (Meyrick) are clearly very close and could be conspecific. However, given the missing abdomen of the single holotype of G. alphestis, the material currently available does not allow for a resolution of their identity. Diakonoff (1973) included a male paratype without abdomen from Australia (Geraldton near Cairns, now known as Innisfail) in the type series of G. labidophora Diakonoff that is otherwise based on New Guinea material. Since G. ophiocosma and G. labidophora are superficially virtually indistinguishable and no further material of G. labidophora has been found in Australia, the Australian paratype of G. labidophora is treated as a misidentified G. ophiocosma.
Constituent species Figs 242–243. Gnathmocerodes, female genitalia. 242, G. ophiocosma, Mt Bellenden-Ker, Q, slide ANIC 2704. 243, G. euplectra, Yeppoon, Q, slide ANIC 2752.
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There are seven named species of Gnathmocerodes, three from Australia, and at least one undescribed species is known from Australia: euplectra (Lower, 1908), Trans. R. Soc. S. Aust. 32: 322 (Tortrix). [Qld: Brisbane and Mackay.]
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alphestis (Meyrick, 1921), Exot. Microlepid. 2: 449 (Argyroploce). [Qld: Cairns.] ophiocosma (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 215 (Argyroploce). [Qld: Cairns.]
Rhectogonia Diakonoff (Figs 33, 244–250) Rhectogonia Diakonoff, 1966, Zool. Verh. (Leiden) 85: 36. Type species: Rhectogonia electrosema Diakonoff, 1966, ibid. 85: 38, figs 8–11, 53, by original designation.
Diagnosis Rhectogonia species are medium-sized to large moths with a diagnostic and sexually colour-dimorphic wing pattern of a large triangle of yellow brown, mustard colour in male and reddish brown in female over entire costa with its apex near middle of dorsum, with remainder of wing along dorsum pale grey becoming increasingly mixed with brownish in terminal third, and with a distinct white line edging brownish triangle from base to beyond its apex. Rhectogonia has sinuate, porrect, moderately widened labial palpi with a slender terminal segment and in the male a large, flat scale pencil on the hind tibia. The forewing has a narrow accessory cell, welldeveloped M-stem and all veins separate, and in the hindwing M3 and CuA1 are stalked to beyond 56:. The combination of the stalked M3 and CuA1 with a tibial pencil in the male, characteristic for Rhectogonia, is very rare in Olethreutinae. The anal area of the hindwing is conspicuously modified in the male, enlarged and covered with semierect, pale, rounded-clavate scales above and with an upcurved fold along the anal margin. In the male genitalia the strongly modified, bipartite gnathos with a medial, bi-pronged process, the characteristic bilobed cucullus and the greatly modified anellus with two antler-shaped spikes from the dorsal margin of the opening for the aedeagus are diagnostic for Rhectogonia. In the female genitalia the erect, nonverrucose, densely hairy ovipositor lobes, the complex sterigma consisting of a large, bilobed ventral pocket containing a transverse, serrate, blade-shaped structure, the ductus bursae ending in a strongly sclerotised funnel with a medial emargination in its ventral margin, and the single signum, a short, blunt, straight, slightly rugose horn of round diameter with a conical base, are diagnostic for Rhectogonia.
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Description Adults (Figs 244, 245) Wingspan. 16–19 mm; medium-sized to large. Head (Fig. 249). Ocellus well developed. Scales of upper part of frons long, anteriorly projecting, remainder of frons with minute, appressed scales. Proboscis about as long as labial palpus. Labial palpus porrect, sinuate, long (1.9–2.3× diameter of eye), strongly widened in middle, second segment subtriangular to teardrop-shaped, distally strongly widened by long, loose scales along ventral margin and in distal half of dorsal margin, terminal segment long and slender. Antennae reaching to beyond middle of wing, scaled dorsally, with minute cilia. Thorax. With very large, conspicuous posterior crest. Hind tibia in male with long, loose scales on inner surface partly concealing a broad, flat pencil of hairscales arising dorsally on its base. Wings (Figs 33, 244, 245). Forewing index 0.43–0.47. Forewing broad, widening distally, broadest just before tornus, costa curved throughout, especially in basal half, apex nearly square, hardly projecting, termen barely sinuate and not oblique; with some large, loosely appressed scales from strongly curved basal part of dorsum; discal cell with chorda parallel and close to anterior margin from base of R2 to between R4 and R5, and with M-stem well developed, to between M2 and M3; all veins separate with R4 slightly closer to R5 at base than to R3, M1 and M2 conspicuously distanced, and M3 much closer to CuA1 than to M2; CuP well developed near margin; anal loop 0.30× length of 1A+2A; mostly yellowish brown, mustard-coloured in male and deep reddish brown in female except for an irregular, light grey band along dorsum separated from brownish wing portion by a strongly sinuate narrow white line from wing base running in an even curve to near middle of dorsum where its end is sharply upcurved towards apex; with faint strigulae along costa, small dark brown spots along dorsum and an irregular series of minute black spots around ocelloid patch except along tornus. Hindwing with cubital pecten present and anal area in male greatly enlarged and modified, forming a large rounded-triangular lobe beyond 1A+2A covered on upper surface with whitish, modified, semierect, rounded-spatulate scales and with anal margin forming a broad fold on upper side; crossvein R1 between closely approximated bases of Sc+R1 and Rs present; bases of Rs and M1 parallel and approximated at base; base of M2 curved to close to base of stalk of M3
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and CuA1 that are stalked to over 56:; CuP weak and most of anterior arm of basal loop of 1A+2A missing; 3A weak, parallel and close to anal fold; grey to dark grey, darker in female. Pregenital abdomen. Male with a patch of long hairscales laterally on first segment behind each spiracle (Fig. 248) and apparently with a shallow, W-shaped, invaginated, transverse fold between T1 and T2. Sternum 8 in male with deep U-shaped medial emargination in R. electrosema, T8 with transversely triangular sclerotisation at base. Male genitalia (Figs 246, 247). Tegumen wellsclerotised, ovate to subrectangular in outline, with apical third of tegumen a particularly robust, posteriorly projecting structure ventrally delineated by gnathos, with a flat or slightly convex apex and sometimes a subapical knob that could be an uncus remnant. Socii in R. electrosema two membranous, bristled, flat, drooping, elongate-ovate lobes with their inner margins meeting at the apex of a small cone parallel and posterior to apex of tegumen, in R. ancalota (Meyrick) two widely separate, long slender, bent and tapering, hairy, sclerotised processes. Gnathos strong, apparently bipartite and consisting of two parallel structures tilted at a right angle to each other, a more ventral band of usual olethreutine shape with two medially joined lateral arms and a ventrally projecting medial lobe in R. ancalota, and a parallel but ventrally facing more dorsal portion consisting of a strongly sclerotised band laterally fused with ventral band but medially produced into either a strong, chute-shaped process ending in two long, sharply downcurved prongs (R. electrosema), or two parallel, downcurved, hairy, crescentic processes. Valva constricted beyond rounded or angled sacculus with a group of short bristles on ventral margin of narrow basal excavation and another group on apex of sacculus; cucullus with a very long, narrow dorsal lobe and a shorter, rounded to triangular ventral lobe with several strong spines or a long, finger-like process from apex, with a high transverse ridge separating the two lobes carrying stout bristles or long spines on its ventral flank; with small, slender costal process. Juxta triangular, caulis short, anellus strongly modified, a large, ovate, bowl-shaped structure with opening for aedeagus in dorsal half with a strongly sclerotised rim with a low, medial, triangular projection on its ventral margin and long, antler-shaped spikes parallel to aedeagus from its dorsal margin on each side, each a long spike with a curved, sharp point and a short, pointed lateral arm from near base in R. electrosema, the two points of more similar length in R. ancalota. Aedeagus short and slender, widest at dorso-
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ventrally flattened base and tapering toward apex; without any trace of cornuti (Diakonoff (1966) incorrectly reports cornuti). Female genitalia (Fig. 250). Ovipositor lobes modified, erect and subparallel rather than flat, very densely and finely hairy but not verrucose. Large and complex sterigma in intersegmental membrane posterior to straight hindmargin of S7. Posteriormost portion of ductus bursae forming a strongly sclerotised funnel with ventral margin of opening with a medial emargination, associated with a complex sterigma of a large, bipartite ventral pocket from which emerges a large, irregularly transverse, serrate, blade-shaped structure with an additional, parallel crest on each side. Ductus bursae long and slender, the posterior sclerotised funnel anteriorly ending in a constriction and followed by a more slender and less sclerotised portion of about equal length ending at origin of ductus seminalis, remainder of ductus bursae membranous; corpus bursae ovate with a single signum, a short, blunt, straight horn of round cross section with a conical base and rugose surface.
Distribution Rhectogonia is known from India, Sri Lanka, Myanmar, Thailand, New Guinea (Robinson et al. 1994) and Australia where it occurs in northern Queensland south to the Cairns area.
Biology No host plants are known for Rhectogonia but all Australian specimens have been collected at rainforest localities.
Remarks Diakonoff (1966) provided figures of genitalia of both sexes, head and wing venation of R. electrosema, Clarke (1958) figured adult and male genitalia of the type of R. ancalota and Robinson et al. (1994) gave a colour illustration of an adult of R. ancalota of unstated provenance. Rhectogonia shows an unusual character combination with a hindwing with stalked M3 and CuA1 together with a tibial pencil, and several features in the genitalia of both sexes pointing to a position in the Olethreutini. Meyrick (1907c) described Rhectogonia ancalota (Meyrick) as a species of Enarmonia. Diakonoff (1966) did not give a tribal assignment when he described Rhectogonia but he referred the genus to the Eucosmini in Appendix I of his south-east Asian Olethreutini
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127
Figs 244–250. Rhectogonia electrosema. 244, 245, adults, McIlwraith Range, Q. 244, . 245, . 246, 247, male genitalia and aedeagus, Tully, Q, slide ANIC 13761. 248, abdomen base, male. 249, head, male. 250, female genitalia, Cairns, Q, slide ANIC 17055. Scale = 2 mm.
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(1973). However, on describing an alleged additional species from Sumba, Diakonoff (1984a) included the genus in the Olethreutini, as did Robinson et al. (1994). The tibial pencil in the male and the genitalia in both sexes, in particular the origin of the ductus seminalis so far from the bursa and the shape of the sterigma and signum, leave no doubt that the genus belongs to the Olethreutini despite the stalked M3 and CuA1. The most closely related Australian genus, Gnathmocerodes, has M3 and CuA1 short-stalked in one unnamed northern Australian species. The monophyly of Rhectogonia is based on the stalked M3 and CuA1 and the structure of the modified anal area in the hindwing, the bipartite gnathos with its medial bipronged process and the sclerotised anellus with its two antler-shaped spikes from the dorsal margin of the opening for the aedeagus, and the modified, non-verrucose and finely hairy ovipositor lobes, the complex signum with bilobed pockets containing a transverse, blade-shaped structure and the funnel-shaped, sclerotised posterior portion of the ductus bursae. Rhectogonia shares several apomorphies with Gnathmocerodes, in particular a modified gnathos with a distally bifid process, a large, sclerotised, bowl-shaped anellus and a single, short, straight, rugose, horn-shaped signum, but also very similar socii and valvae. Both genera have a tibial pencil and enlarged hindwing anal area at least in some species, and one unnamed Western Australian species of Gnathmocerodes also has M3 and CuA1 shortstalked. Possible synapomorphies with Sorolopha are discussed in the Remarks for that genus. ‘Rhectogonia’ dyschima Diakonoff, 1984a, from Sumba, shows a character combination somewhere between Rhectogonia and Gnathmocerodes with the deeply invaginated pockets of the sterigma alluding to the former and the male genitalia closer to the latter (Diakonoff 1984a). The species does not possess the Rhectogonia apomorphies and will have to be reexamined in the context of the Oriental fauna.
Constituent species
Olethreutes-group Status remarks This group was established by Diakonoff (1973) for 28 south-east Asian genera as the subtribe Olethreutae. Kuznetzov and Stekolnikov (1984) accepted the subtribe as Olethreutina, but it is here treated as a genus-group only. Oxysemaphora Diakonoff is excluded and referred to the newly defined Oxysemaphora-group.
Evidence for monophyly Diakonoff (1973) based his Olethreutae primarily on usually two ‘cornute’ or scobinate signa, a welldeveloped, hooked uncus and the gnathos formed by a membranous, transverse band. Derivations of horn-shaped signa, most frequently paired nippleshaped patches of scobination but more rarely depressions with concentric rows of irregular teeth, and a membranous, spinulose central connection of the gnathos arms are indeed two possible apomorphies for the group.
Distribution The Olethreutes-group of genera is very widely distributed, with the highest diversity in the Oriental region, well represented in the African and Palaearctic regions and extending into the Nearctic and probably the northern Neotropical regions.
Biology Larvae of the Olethreutes-group are leaf rollers on a wide range of plant families, but the larva of Ophiorrhabda has also been reported as a borer in fruit or stem galls (Diakonoff 1973).
Remarks Diakonoff (1973) included the African Eccopsis Zeller in his Olethreutae, but Aarvik’s (2004) referral of Eccopsis to the Neopotamia-group is much more convincing.
Excluding the doubtfully assigned R. dyschima, only two species of Rhectogonia are known so far, R. ancalota described from Sri Lanka and reported also from India, Myanmar and Thailand, and R. electrosema from New Guinea and Australia:
(Figs 34, 251–267)
electrosema Diakonoff, 1966, Zool. Verh. (Leiden) 85: 38 (Rhectogonia). [New Guinea.]
Ophiorrhabda Diakonoff, 1966, Zool. Verh. (Leiden), 85: 47. Type species: Argyroploce
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Ophiorrhabda Diakonoff
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ergasima Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 271, by monotypy and original designation. [New Guinea: Woodlark I.] Cellifera Diakonoff, [1968], US Nat. Mus. Bull. 257: 40 (key), 47 [as subgenus of Hedya Hübner]. Type species: Polychrosis cellifera Meyrick, 1912b, J. Bombay Nat. Hist. Soc. 21: 869, by original designation. [Sri Lanka, India.] Didrimys Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 373 (key), 388. Type species: Platypeplus harmonica Meyrick, 1905, J. Bombay Nat. Hist. Soc. 16: 584, by original designation. [Sri Lanka.] Lasiognatha Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 395 (key), 429. Type species: Lasiognatha quartaria Diakonoff, 1973, ibid. 1: 430 (key), 432, by original designation. [Sumatra, Java.]
Diagnosis Ophiorrhabda species are small to large, usually intricately marked greyish or brownish moths, often with a reddish tinge and with a slender triangular dark mark or a band parallel to termen rising from end of dorsum, sometimes with a conspicuous large round blotch below middle of costa (dark brown in O. mormopa (Meyrick), greyish pink or greenish in O. phaeosigma (Turner)). Ophiorrhabda has moderately long sinuate labial palpi with the second segment apically at least slightly dilated, never as slender as in Dudua Walker and Trachyschistis Meyrick. The tibia and first tarsal segment in the male hind leg are greatly enlarged by scale fringes and the tibia carries a large and often composite hairpencil. A tibial pencil is present also in all the closely related genera, but the conspicuously enlarged tibia and first tarsal segment are characteristic for Ophiorrhabda. The wing venation with M3 and CuA1 in the hindwing connate or separate allows separation from the closely related Cymolomia Lederer with stalked M3 and CuA1. In males the anal margin is folded over the upper surface producing either a thickened rigid ‘anal roll’ with dense fringes of long scales from its upturned underside (O. mormopa, O. phaeosigma) or a deep fold containing a hairpencil (O. ?scaristis (Meyrick), O. ?mysterica (Turner)). The males of Dudua (Walker), Trachyschistis (Meyrick), Archilobesia Diakonoff and Euobraztsovia Diakonoff also have an anal roll, but in these genera the margin is rolled
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129
under. A pair of deep round depressions laterally on S2 in both sexes are a characteristic feature of Ophiorrhabda but are found also in Temnolopha Lower and Gatesclarkeana Diakonoff, and as a faint impression in Archilobesia Diakonoff. The male genitalia are characterised by a well-developed, ventrally bristled uncus, drooping socii, a membranous gnathos with two separate bristled pads each ending in a dorsal point (except O. ?mysterica) and a slender valva with an isolated group of short sharp spines on margin of basal excavation usually raised on a projecting corner or sinuate process. The medially separated spinulose gnathos pads are diagnostic for Ophiorrhabda when they are present, and a combination of the depressions on S2 together with a long slender valva with a sinuate process carrying a tuft of spines from the edge of the basal excavation will identify the remaining members of the genus. Dudua, Archilobesia and Trachyschistis also have a spinulose gnathos, but it is of different shape and medially continuous. Some of the most obvious apomorphies for Ophiorrhabda are found in the female genitalia: the ventrally split, rim-shaped, spinulose sterigma laterally continued into a short, angled ridge on each side and the large horn-shaped single signum with its hollow base and flattened apex are both diagnostic for the genus. Cymolomia Lederer also has only one signum, but its apex is not flattened and the genus is moreover easily recognisable by stalked M3 and CuA1 in the hindwing.
Description Adults (Figs 251–254, 256) Wingspan. 11–20 mm; small to large. Head (Fig. 255). Ocellus well developed. Vertex with moderately long scales, anteriorly barely protruding between antennae. Uppermost part of frons with a narrow transverse band of shortish porrect scales, remainder of frons with minute, appressed and upwardly oriented scales. Maxillary palpus minute. Proboscis short, little longer than labial palpus. Labial palpus moderately long (1.6–2.0× diameter of eye), sinuate, porrect to obliquely rising, slightly to moderately expanded towards apex of second segment; second segment upcurved to sinuate, distally expanded by long scales along ventral margin and apical part of dorsal edge, terminal segment drooping, rather stout. Antenna reaching to middle of forewing; anterior surface unscaled; cilia in male short, less than half diameter of flagellar segments.
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Thorax. With large posterior crest. Legs with hind tibia and first tarsal segment greatly enlarged in male, at least with a long crest of large erect scales along dorsal and ventral margins of tibia and along dorsal margin of first tarsal segment and with a hairpencil dorsally from base of tibia along its inner surface (O. phaeosigma), but usually with tibia and first tarsal segment especially dorsally with dense pads of long erect scales and with a large composite hairpencil from base of tibia along smooth inner surface of tibia (O. mormopa). Wings (Figs 34, 251–254, 256). Forewing index 0.43–0.48; subtriangular to subrectangular, distally moderately widened, with lightly and evenly curved costa and straight, slightly oblique termen; discal cell 0.66–0.68× length of wing, with moderate accessory cell with chorda from between R1 and R2 to opposite of R5 and with M-stem to between M2 and M3; with all veins present and separate beyond cell, with bases of R3, R4 and R5 equidistant or R4 closer to R5 than to R3; basal loop 0.3–0.33× length of 1A+2A; usually intricately marked and marbled, often in shades of grey or brown with reddish hues, with a dark slender triangular mark rising from end of dorsum just before tornus frequently developed into a dark band parallel to termen, and in several species with a uniformly coloured large roundish blotch below middle of costa (O. mormopa, O. phaeosigma). Hindwing with well-developed cubital pecten and axillary tuft, with scattered hairscales from upper surface, forming a deep fold often containing a pencil of hairscales arising from along 3A (O. ?scaristis, O. ?mysterica) or with the folded membrane thickened and rigid (O. mormopa, O. phaeosigma) with a dense fringe of long hairscales from the underside of the upfolded anal margin; with Rs and M1 parallel and approximated at base, M2 bent close to M3, and M3 and CuA1 connate or CuA1 from just before angle of cell; 1A+2A partly tubular but base of 1A weakly developed, 3A present but distally often covered by folded anal margin; reddish ochreous to dark grey. Pregenital abdomen. Both sexes with a pair of large deep semicircular depressions (dorsal pits) laterally on T2, less obviously developed in O. ?scaristis. Eighth segment in male long, hindmargin of S8 bilobed. Male genitalia (Figs 257–264). Vinculum a weakly sclerotised band. Tegumen high and rather narrow, with small to very conspicuous angled ‘shoulders’ and slender pedunculi without obvious muscle attachment points. Uncus a variously shaped, at least ventrally hairy process: with a distinct ‘neck’
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and distally dilated with a bipartite ventral brush of bristles (O. mormopa, O. phaeosigma), elongate-ovate with an undivided ventral brush (O. ?mysterica), or laterally compressed and apex sharply ventrally curved, ending in a tuft of strong bristles (O. ?scaristis). Socii evenly bristled, ovate, drooping lobes. Gnathos a broad transverse membranous band usually differentiated into two spinulose projecting pads culminating each in a sharp point or long cusp (O. ?scaristis), more rarely gnathos a smooth band with only weakly projecting dorsolateral corners (O. ?mysterica). Valva long, slender, lightly curved and usually with a projecting angle along ventral margin, with small costal process and conspicuous invagination from base of outer valval surface, with a group of short sharp thorns on distal part of ventral edge of basal excavation and this area usually raised as a strongly projecting and often downturned corner crowned with thorns (O. mormopa, O. phaeosigma), or as a long sinuate process apparently from base of costa (O. ?mysterica), often also with a distinct, dense patch of bristles from below distal part of basal excavation either on both sides (O. ?mysterica) or asymmetrically (O. ?scaristis), with an evenly bristled cucullus with a usually ventrally projecting basal angle and bristles along base sometimes grouped into separate tufts or raised on a wart (O. ?mysterica). Caulis moderate, anellus a wide ring. Aedeagus rather weakly sclerotised in ventral portion only, a small to moderately large, lightly bent tube; vesica without deciduous cornuti but rarely with a parallel series of sclerotised transverse ridges (O. ?mysterica). Female genitalia (Figs 265–267). Ovipositor lobes subtriangular with rounded lateral margin. Sternum 7 usually unmodified with weakly concave hindmargin, more rarely very wide at base and very short, sharply tapering towards V-shaped hindmargin (O. ?mysterica). Sterigma a projecting, finely spinulose, round to ovate and ventrally cleft rim laterally continued into downcurved spinulose ridges. Ductus bursae short to moderately long, with an ill-defined moderately long colliculum distinctly below ostium, remainder of ductus membranous and smooth, with inception of ductus seminalis close to corpus bursa and anteriormost part of ductus bursae widened and with a small lateral expansion. Corpus bursae round to very elongate-ovate; with a single horn-shaped signum with a hollow base of circular cross section and with distal part flattened into a blade, usually curved and gradually tapering (O. mormopa, O. phaeosigma), more rarely with an angled narrow base (O. ?mysterica) or signum short and straight (O. ?scaristis).
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131
Figs 251–256. Ophiorrhabda, adults and head, males except 252. 251, 252, O. mysterica, Kuranda, Q. 251, holotype . 252, [image reversed]. 253, O. phaeosigma, nr Mudginbarry Homestead, NT. 254, O. mormopa, nr Babinda, Q [image reversed]. 255, head, O. phaeosigma. 256, O. ? scaristis, Darwin, NT. Scale = 2 mm.
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Distribution Sri Lanka, India, throughout south-east Asia to the Phillippine Is and New Guinea, Solomon Is and Australia. In Australia the genus is confined to the coastal regions of the northernmost parts of Western Australia, the Northern Territory and northern Queensland south to Yeppoon.
to leaf rolling or boring in fruit or stem galls on Myrtaceae, mostly on several species of Syzygium, but also on Psidium guajava and Jambosa vulgaris. The only non-myrtaceous food plants recorded are Nephelium lappaceum (rambutan) (Sapindaceae) and Bauhinia (Caesalpiniaceae). The Australian O. phaeosigma has been reared from fruit of Syzygium armstrongii and S. eucalyptoides.
Biology Fletcher (1932) and especially Diakonoff (1973) reported numerous host records for several species of Ophiorrhabda. With few exceptions they all refer
Remarks Both Didrimys and Ophiorrhabda, as originally defined by Diakonoff (1966, 1973), are compact and
Figs 257–264. Ophiorrhabda, male genitalia and aedeagus. 257, 258, O. mysterica, nr Mt Cahill, NT, slide T2203. 259, 260, O. phaeosigma, Adelaide R., NT, slide T2256. 261, 262, O. mormopa, Mt Spec, Q, slide T2417. 263, 264, O. ? scaristis, Jim Jim Ck, NT, slide T2297.
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Figs 265–267. Ophiorrhabda, female genitalia. 265, O. mysterica, Kuranda, Q, slide ANIC 2708. 266, O. mormopa, Mt Spec, Q, slide T2418. 267, O. ? scaristis, Jim Jim Ck, NT, slide ANIC 2695.
monophyletic groups, but they both have apparently arisen from within Diakonoff’s (1973) genus Lasiognatha. No autapomorphy justifying the separate status of Lasiognatha could be found, leaving no other recourse but to synonymise the three genera. However, in its expanded state Ophiorrhabda is now defined by several obvious apomorphies in both sexes: the well-developed dorsal pits, the characteristically shaped sterigma and distally flattened signum in the female and the strongly enlarged hind tibia and first tarsal segment in the male. Diakonoff (1973) had been misled by the superficial similarity of the cuspidate gnathos in Didrimys and some species of Lobesia Guenée to include the former genus in his subtribe Lobesiae whilst at the same time drawing attention to the obvious relationship with Dudua Walker. The superficial similarity of Ophiorrhabda cellifera and Dudua anaprobola Bradley has repeatedly led to confusion. Diakonoff ([1968]) erected Cellifera Diakonoff as a subgenus of Hedya Hübner based on the female of Polychrosis cellifera Meyrick and a wrongly associated male later
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identified by Diakonoff (1973) as Dudua anaprobola Bradley. In his monograph on the South Asiatic Olethreutini, Diakonoff (1973) drew attention to this error and figured male genitalia for O. cellifera that closely correspond to those of other species with similar female genitalia, but he included O. cellifera in his new genus Lasiognatha without further mention of his earlier subgenus Cellifera. The same moths apparently also misled Clarke (1976) who, for his newly described Dudua pottsi Clarke, figured male genitalia identical to those given for O. cellifera by Diakonoff (1973), whilst those he ascribed to O. cellifera are very similar to those of O. anaprobola. This explains why he reached the conclusion that O. cellifera should be included in Dudua (Clarke 1976). As a consequence of the above discussed generic synonymies the following species, described as Platypeplus, a synonym of Dudua, is here referred to Ophiorrhabda: deceptor (Diakonoff, 1966) Zool. Verh. (Leiden) 85: 64 (Platypeplus). [New Guinea, Bismarck Is.] Comb. nov.
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Ophiorrhabda is well documented by Diakonoff (1966, 1973, as Ophiorrhabda, Didrimys and Lasiognatha), with figures of head and venation of all three genera, and genitalia of numerous species. Clarke (1958) provided photographs of wings and genitalia of O. harmonica (Meyrick) (as Argyroploce crocospila Meyrick), O. mormopa and O. cellifera. Photographs of wings and of male and female genitalia of O. dryoptycha (Meyrick) (as Olethreutes empherana Bradley) were given in Bradley’s (1961) description of O. empherana. There is some doubt whether O. favillosa Diakonoff (Diakonoff 1973) should be included in Ophiorrhabda given the very different valva and unusually modified abdomen of the male holotype, but any alternative generic assignment will have to wait until a female becomes available. Cymolomia Lederer appears to be the sister group of Ophiorrhabda with the two genera sharing such derived features as a single signum and a gnathos with a pair of raised pads. Autapomorphies for Cymolomia hartigiana (Saxesen), the type of the genus, are stalked M3 and CuA1 in the hindwing and a smooth gnathos, assuming that a spinulose gnathos is the groundplan condition for the Olethreutes-group. Ophiorrhabda ergasima (Meyrick) was described from New Guinea (Woodlark Is) and later (Diakonoff 1973) also reported from Australia. However, the specimen from Cairns referred to by Diakonoff (1973) was not found in the BMNH and no other Australian material of O. ergasima could be located. Ophiorrhabda scaristis is tentatively included in the list of Australian species. The population found in the Northern Territory and northern Queensland can at present not be differentiated from O. scaristis, given the badly damaged genitalia of the male holotype.
Constituent species Fourteen named species of Ophiorrhabda are recognised, four of which occur in Australia together with several unnamed species: mysterica (Turner, 1916), Trans. R. Soc. S. Aust. 40: 533 (Argyroploce). [Qld: Kuranda and Stannary Hills.] mormopa (Meyrick, 1906), J. Bombay Nat. Hist. Soc. 17: 136 (Platypeplus). [Sri Lanka.] melanocycla (Common, 1970), Insects of Australia: 799, 801 (Olethreutes), nomen nudum. phaeosigma (Turner, 1916), Trans. R. Soc. S. Aust. 40: 532 (Argyroploce). [NT: Darwin.]
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scaristis (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 270 (Argyroploce). [New Guinea (Woodlark Is).]
Dudua Walker (Figs 35, 268–280) Dudua Walker, 1864, List Specimens Lepid. Insects Colln Br. Mus. 30: 1000. Type species: Dudua hesperialis Walker, 1864, ibid. 30: 1000, by monotypy. [Borneo: Sarawak.] Platypeplus Walsingham, [1887] 1884–7, in Moore, Lepid. Ceylon 3: 495. Type species: Eccopsis aprobola Meyrick, 1886, Trans. Entomol. Soc. Lond. 1886: 275, by monotypy. [Tonga.] Platypeplum Walsingham, 1899, Indian Mus. Notes 4: 105. An unjustified emendation of Platypeplus Walsingham, 1887.
Diagnosis Dudua species are medium-sized to large olethreutines with rather narrow forewings and with male hindwing often either triangular with straight oblique termen or with scalloped termen and modified fringe between M2 and CuA2. Wing pattern very diverse but typically with a series of parallel horizontal black lines in a vertical band from tornus towards costa just before apex. The rather small, slender, upcurved labial palpi, appressed to the frons, are characteristic for Dudua and Euobraztsovia Diakonoff in the Australian fauna and shared in exactly the same shape otherwise only with Hedya Hübner. Slender labial palpi are also present in Trachyschistis Meyrick, but with their second segment widest at apex and with closely appressed scales along ventral margin, unlike those of Dudua with a distally tapering second segment widest in middle due to loose scales along its ventral margin. Males of Dudua have a slightly enlarged hind tibia with a hair pencil and an ‘anal roll’ with the anal margin of the hindwing broadly folded under over a clump of modified scales and entrance into this fold on lower surface covered by the crest of a second parallel fold from behind 3A. The wing venation is unremarkable for an olethreutine, with M3 and CuA1 in the hindwing originating closely together or, very rarely, stalked (D. iniqua), and with 3A adjacent to ‘anal roll’. The male genitalia with the hairy underside of their uncus continuous with the flat drooping socii, a membranous gnathos with paired and spinulose ventral humps and an elbowed valva
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Figs 268–273. Dudua, adults. 268, 269, D. aprobola. 268, , Nambour, Q. 269, , nr Yeppoon, Q. 270, D. phyllanthana, , Depot Beach, NSW [image reversed]. 271, D. iniqua, holotype [image reversed]. 272, 273, D. phyllantana. 272, , Elizabeth Beach, NSW. 273, , Forster, NSW. Scale = 2 mm.
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with a ventral band of dense long bristles fringing the ‘elbow’ at base of cucullus and with a patch of large spines below base of costa near margin of basal excavation are diagnostic for Dudua. Archilobesia Diakonoff and Trachyschistis both have very similar male genitalia, but in the former the gnathos is spinulose laterally as well as ventrally and the latter has a small roundish uncus tip with only a few strong ventral bristles. Hedya salicella (L.), the type species of Hedya, differs from Dudua in having a smooth gnathos and not distinctly elbowed valva. The structure of the female genitalia of most species of Dudua with their spinulose, ventrally split, tube-shaped sterigma and the two nipple-shaped granular signa, is identical to that of Archilobesia and Hedya salicella. However, the sterigma of D. aprobola (Meyrick) and D. phyllanthana (Meyrick) with their sharply ridged ventral expansion are unusual and readily recognisable.
Description Adults (Figs 268–273) Wingspan. 11–22 mm; medium-sized to large.
Head (Fig. 275). Ocellus well developed to large. Uppermost part of frons with a narrow transverse band of anteriorly projecting scales, remainder with minute, appressed and upwardly oriented scales. Proboscis short, at most 1.5× length of labial palpus. Labial palpus rather small (1.5– 1.7× diameter of eye), slender, strongly upcurved and appressed to frons; second segment curved throughout, with moderately long loose scales all along ventral margin and either of even width throughout or widest in middle and tapering towards apex; with terminal segment at a slight angle, short and usually pointed. Antenna reaching to just beyond middle of forewing; anterior surface unscaled; cilia in male short, less than half diameter of flagellar segments. Thorax. With large posterior crest. Legs with hind tibia in male with lengthened but appressed scales along dorsal margin and inner surface, with a long hairpencil dorsally from base concealed in groove in scaling along inner surface. Wings (Figs 35, 268–273). Forewing index 0.36–0.43; forewing elongate subovate to subtriangular, costa usually evenly curved, apex rounded-acute, termen convex and slightly to
Figs 274–277. Dudua aprobola. 274, male genitalia, East Alligator R., NT, slide ANIC 2474. 275, head, male. 276, aedeagus, slide ANIC 2474. 277, female genitalia, nr Emerald, Q, slide T2154.
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moderately oblique, tornus usually obliquely rounded; discal cell 0.65–0.69× length of wing, with accessory cell long and narrow to very narrow with chorda from roughly halfway between R1 and R2, with M-stem to between M2 and M3 (weakly developed in some species); with all veins present and separate beyond discal cell and CuA1 bent towards M3 at margin, CuA2 not sinuate; with very diverse wing pattern with pairs of strongly oblique narrow whitish strigulae in distal half of costa converging with usually silvery and red-brown lines running parallel a little below costa and usually with at least traces of a series of short parallel horizontal black lines in a vertical band from before tornus towards 169 costa. Hindwing usually with strongly developed cubital pecten, sparse axillary tuft and scattered long hairscales from upper surface of anal area, variously modified in male: all Australian species with a complicated ‘anal roll’ with thickened anal margin folded under
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over a compact clump of strongly modified scales and entrance into this fold covered by the crest of a second, parallel but dorsally concave fold invaginated just behind 3A, males often with modified, triangular wing shape with margin triangularly projecting between 1A+2A and 3A and with otherwise oblique and only faintly sinuate termen (D. aprobola) or with scalloped termen with protruding, rounded lobe between M2 and CuA2 with dense and modified fringe and with a small elongate patch of darker scales along closing vein (D. iniqua (Meyrick)); with Rs and M1 parallel and closely approximated at base, with M2 at base often non-tubular and bent closely towards M3, with CuA1 from just before angle of cell to connate or stalked (D. iniqua) with M3; with CuP often weak; 1A+2A and 3A both well developed but base of 1A weak and 3A closely adjacent to ‘anal roll’; from whitish grey to blackish grey.
Figs 278–280. Dudua. 278, D. phyllanthana, male genitalia and aedeagus, nr Cooktown, Q, slide T2293. 279, D. iniqua, holotype, male genitalia [SAMA]. 280, D. phyllantana, female genitalia, Forster, NSW, slide ANIC 2475.
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Pregenital abdomen. Unremarkable except for modified scales dorsally on male abdomen in D. tetanota (Meyrick) (Diakonoff 1973). Sternum 8 in male large, hindmargin with a triangular, semicircular or rectangular ventral excavation; T8 shorter. Male genitalia (Figs 274, 276, 278, 279). Vinculum a moderately broad lightly sclerotised band. Tegumen triangular, high and rather narrow, pedunculi tapering only at extreme base, with broad low apodemes for muscle m4. Uncus a usually roughly parallel-sided median process bent ventrally, its ventral surface nearly always densely hairy throughout, continuous with socii. Socii usually elongate, densely hairy drooping flat lateral lobes from small projecting ‘shoulders’. Gnathos a smooth, membranous transverse band with two paired downwardly directed conspicuously spinulose ventral humps. Valva roughly straight to lightly curved, from quite slender and parallel-sided to distal half dilated, with a slender, bent costal process, ventroproximal base of cucullus always a lightly projecting and densely spined ‘elbow’ often connected with an oblique band of scattered spines with the dense patch of prominent spines below base of costa on margin of basal excavation, cucullus evenly bristled with stronger spines along ventral margin and base. Caulis short, anellus a broad ring. Aedeagus short and small, parallel-sided to tapering towards apex, lightly curved; vesica without cornuti. Female genitalia (Figs 277, 280). Ovipositor lobes subtriangular to elongate-ovate. Sternum 7 with very deeply bilobed hindmargin of often diagnostic shape. Sterigma usually a slender, ventrally split, spinulose, tube-shaped structure (D. siderea (Turner), D. iniqua), more rarely with a broad ventral extension with sharp serrate ridges medially converging below ostium (D. aprobola, D. phyllanthana) and with paired spinulose protuberances above ostium (D. phyllanthana). Ductus bursae with a long, bipartite, weakly sclerotised colliculum, remainder smoothly membranous, at most slightly wrinkled in middle part near inception of ductus seminalis. Corpus bursae ovate, bursa wall granulate throughout; with two nipple-shaped signa with the hollow blunt thorns and the surrounding ring densely granular.
Distribution Madagascar, Seychelles, Amirante Is, Chagos Is, Sri Lanka, from India throughout south-east Asia to China, Taiwan, Philippine Is, New Guinea, New
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Caledonia, Vanuatu, Fiji, Tonga, Niue, Cook Is, Tahiti, Rapa, Marquesas Is and Australia (including material in the New Zealand Arthropod Collection, Auckland). The widely distributed D. aprobola is found in most of this area, ranging from the Seychelles across all of tropical Asia to Taiwan, New Guinea, Australia and across the Pacific to Micronesia and Tahiti. Clarke (1976) reports the presence of D. aprobola also in Natal. In Australia, Dudua occurs in the coastal regions of northernmost Western Australia from Broome north, the Northern Territory, Queensland and New South Wales except for the southernmost corner. D. aprobola occurs as far south as Burleigh Heads, Queensland, and D. phyllanthana south to Batemans Bay, New South Wales.
Biology Dudua aprobola is highly polyphagous, feeding on young leaves and sometimes flowers from a wide range of families (Rosaceae, Anacardiaceae, Verbenaceae, Sapindaceae, Caesalpiniaceae, Annonaceae, Lythraceae, Loranthaceae, Salicaceae (Diakonoff 1973), also on Myrtaceae (Clarke 1971) and Urticaceae (Clarke 1976)). It is found on cultivated plants such as Nephelium litchi (Sapindaceae), Psidium guajava (Myrtaceae) and frequently on Mangifera indica (Anacardiaceae), and Fletcher ([1921], 1932) describes its biology. In Australia, D. aprobola has been reared from Rosa (Rosaceae), ‘Bauhinia’ (Caesalpiniaceae), Mimosa pigra (Mimosaceae), Mangifera indica (Anacardiaceae) and Macadamia sp. (Proteaceae). Other species of Dudua with known host plants have all been reared from Glochidion (Euphorbiaceae): D. anaprobola in Fiji (Bradley 1953) and D. anisoptera Clarke in Guam from G. ramiflorum (Clarke 1986), and the Australian D. phyllanthana and D. siderea from G. ferdinandi, the former by J. Stockard and by I. F. B. Common and M. S. Upton and the latter by V. J. Robinson at Minnamurra Falls, New South Wales (ANIC).
Remarks Dudua is a large and compact group whose genitalia in both sexes are, with few exceptions, remarkably similar. The combination of the characteristic gnathos and the ventral band of dense long bristles (very rarely widely scattered, broad and flattened as in D. anaprobola) fringing the valval ‘elbow’ readily identifies species of Dudua. However, the separation from both Hedya and Archilobesia is rather tenuous in cladistic terms. Hedya salicella,
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the type species of Hedya, and Dudua share similarly derived slender labial palpi. Loss of the gnathos spinules is the only autapomorphy found for Hedya, assuming that a spinulose gnathos belongs to the ground plan of the Olethreutesgroup. A strongly developed band of bristles ventrally to the valval ‘elbow’ is the single consistent apomorphy for Dudua, but it is present as a trace in Hedya and moderately developed in Archilobesia, which, however, has an unmodified broad labial palpus. Hedya currently comprises a diverse-looking assemblage and a thorough study of its Palaearctic and Oriental taxa will be required before the phylogeny of the group can be resolved. Given the long-standing and widely accepted concept of Dudua as a discrete entity the above reservations are, for the time being, not considered sufficient reason to change the status quo. Diakonoff (1973) provided a comprehensive revision of South Asiatic Dudua, figuring genitalia of nearly all species and giving access to previous literature. Wings and genitalia of the types of many Meyrick species were figured by Clarke (1958, as Platypeplus). Additional species of Dudua were subsequently described from Micronesia (Clarke 1976) and Madagascar (Diakonoff 1981, 1983c). It is doubtful whether D. chlorohygra Diakonoff from Sikkim (Diakonoff 1973) and the Malagasy D. hemitypa Diakonoff (Diakonoff 1983c) should be included in Dudua, but a conclusive assignment will have to await discovery of the opposite sex for both species. The genitalia figured for Dudua pottsi Clarke (Clarke 1976) are identical to those of Ophiorrhabda cellifera (Meyrick) that is also treated in the same paper and has typical Dudua genitalia assigned to it. The confusion created by these two superficially similar species is discussed under Ophiorrhabda.
Constituent species Thirty one named species of Dudua are currently recognised, four of which occur in Australia: aprobola (Meyrick, 1886), Trans. Entomol. Soc. Lond. 1886: 275 (Eccopsis). [Tonga.] metallota (Lower, 1901), Trans. R. Soc. S. Aust. 25: 73 (Temnolopha). [Qld: Cooktown.] phyllanthana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 641 (Antithesia). [NSW: Sydney.] empyra (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 216 (Argyroploce). [NSW: Brunswick Heads.] auchmera (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 217 (Argyroploce). [Qld: ‘Tweed Heads’.]
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siderea (Turner, 1916), Trans. R. Soc. S. Aust. 40: 531 (Argyroploce). [Qld: Stradbroke [I.].] iniqua (Meyrick, 1921), Exot. Microlepid. 2: 449 (Argyroploce). [Qld: Cairns.]
Trachyschistis Meyrick (Figs 36, 281–286) Trachyschistis Meyrick, 1921, Exot. Microlepid. 2: 448. Type species: Trachyschistis hians Meyrick, 1921, ibid. 2: 448, by monotypy. [Australia.]
Diagnosis Specimens of Trachyschistis hians are mediumsized to large, rather broad-winged brownish olethreutines with costa strongly curved towards apex, with raised scale tufts across dorsal part of forewing, a conspicuous short oblique white line in middle of wing across closing vein and with lightly mottled hindwings. Trachyschistis has characteristic, slender, upcurved and lightly sinuate labial palpi with the second segment curved at base, straight in distal half and widest at apex. Dudua and Hedya both also have slender, upcurved labial palpi, but their second segment is evenly curved throughout and widest in middle due to long loose scales along ventral margin. The wing venation is characterised by a curiously wavy CuA2 in the forewing and in the hindwing by a very proximal origin of CuA2 and the loss of CuP. In the male the anal margin of the hindwing is thickened and narrowly turned under and the hind tibia has dorsal and ventral scale fringes and a hairpencil. The male genitalia are overall rather plesiomorphic in comparison to other genera of the Olethreutes-group, with a strongly swollen thick ‘neck’ and a small roundish ‘head’ to the uncus, inflated socii, a complex gnathos with parallel dorsal and ventral portions and a very slender, simple, sinuate and parallelsided valva. Shape of uncus, socii and gnathos, together with the absence of a large tuft of bristles from the ventral margin of the basal third of the valva, are diagnostic for the genus, with only Archilobesia sharing a dorso-ventrally bipartite gnathos of, however, different structure. The female genitalia of Trachyschistis with their large, cup-shaped sterigma, the extremely long colliculum and a strongly wrinkled anterior part of the ductus bursae and the lack of signa are unique.
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Description Adults (Figs 281, 282) Wingspan. 15–21 mm; medium-sized to large. Head (Fig. 284). Ocellus well developed. Vertex scales anteriorly barely protruding between antennae. Uppermost part of frons with a transverse band of anteriorly projecting scales, remainder with very short, appressed, dorsally oriented scales. Maxillary palpus very small, but white-scaled tip clearly visible. Proboscis short, little longer than labial palpus. Labial palpus moderately long (1.7× (male)–1.9× (female) diameter of eye) but very slender, strongly upcurved and appressed to frons, weakly sinuate with long straight middle part; second segment with curved base and straight distal half, slender with scales appressed also along ventral margin, only faintly widening towards apex; terminal segment at a slight angle, rather long and stout. Antenna reaching to just beyond middle of forewing; anterior surface unscaled and in male conspicuously serrate in distal half; cilia in male short, about 56M diameter of flagellar segments. Thorax. With very large posterior crest. Legs with hind tibia in male slightly enlarged by dorsal and ventral fringes of loose long scales and with a long slender hairpencil from dorsal base, distally concealed in a groove within scaling of inner tibia surface. Wings (Figs 36, 281, 282). Forewing index 0.43– 0.45; forewing subtriangular, strongly widened towards termen and distal half of wing faintly curved backwards; with costa strongly curved in distal part, apex nearly rectangular, termen straight, tornus very broadly rounded and dorsum lightly sinuate; with large oblique tufts of raised scales across dorsal part of wing, the most prominent between CuP and 1A+2A just beyond middle of wing; discal cell 0.63× length of wing with very narrow accessory cell with chorda from little below R2 and with a weakly developed M-stem to between M2 and M3; with all veins present and separate beyond discal cell, with R4 to costa and R5 from close to anterior angle of cell; CuP present at wing margin, forked part of 1A+2A relatively short; from ochreous ground-colour with three partially developed outwardly oblique dark grey fasciae from about one-fourth, middle and three-fourths of costa to nearly uniformly dark grey-brown, especially in female, but always with a short, very straight and inwardly oblique white streak in middle of wing across closing vein and a pale curved line rising from tornus towards three-fourths costa. Hindwing with well-developed cubital pecten and axillary tuft and with scattered long hairscales from anal area, in
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male with anal margin thickened and narrowly folded under; with Rs and M1 parallel and closely approximated at base, with M2 bent closely to M3 at base, CuA1 from just before angle of cell and CuA2 from only little beyond middle of cell; no remnant of CuP visible even in a stained wing slide, a crease associated with irregularities in scale socket pattern suggests a course distally relatively close to CuA2; 1A+2A present but base of 1A only a faint trace, 3A partly tubular and distant from dorsal margin; grey with ochreous tinge and distinctly marbled. Pregenital abdomen. Unmodified. Hindmargin of S8 bilobed with a semicircular ventral excision. Male genitalia (Figs 283, 285). Vinculum a broad weakly sclerotised band. Tegumen triangular in outline, with small but distinct ‘shoulders’ and broad short apodemes for muscle m4. Uncus a strong, ventrally curved process with a thick, swollen ‘neck’ and a much smaller rounded apex with few large bristles on underside. Socii elongate inflated pads covered with long hair. Gnathos a complicated membranous structure consisting of a medially depressed smooth transverse bulge with scattered dorsolateral spinules and of a parallel more ventral portion with two densely spinulose ventral humps above and laterally to aedeagus. Valva very long and slender, sinuate and parallelsided throughout, with slender costal process and small basal excavation with an isolated patch of shortish small spines on its distal margin, cucullus evenly bristled throughout with stronger spines towards ventral margin and base, with a band of particularly long spines running obliquely across valva towards base of costa at base of cucullus. Caulis short and anellus a mainly ventrally sclerotised broad ring. Aedeagus short and small, lightly curved and roughly parallel-sided, sclerotised ventrally only; vesica with needle-like structures but no obvious cornuti with sockets. Female genitalia (Fig. 286). Ovipositor lobes broadly elongate-ovate. Sternum 7 with concave hindmargin. Sterigma a large spinulose cup with a ventral V-shaped cleft and an irregularly excised dorsal margin. Ductus bursae with an extremely long colliculum reaching to well over middle of ductus bursae and with inception of ductus seminalis at its end, remainder of ductus bursae strongly transversely wrinkled. Corpus bursae round, without obvious surface sculpture and without signa.
Distribution Trachyschistis is endemic to Australia, occurring only in a restricted area along the eastern coast
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from the Bunya Mts in southern Queensland to Clyde Mt in southern New South Wales.
together terminal leaves of Diospyros australis (Ebenaceae).
Biology
Remarks
All the localities are rainforest habitats and one specimen has been reared from a larva binding
This monotypic genus is retained with some misgivings, but the combination of a plesiomorphic
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Figs 281–286. Trachyschistis hians. 281, 282, adults. 281, , Coraki, NSW. 282, , Bunya Mts, Q [image reversed]. 283, male genitalia, Mt Keira, NSW, slide T2257. 284, head, male. 285, aedeagus, slide T2257. 286, female genitalia, Mt Keira, NSW, slide T2258. Scale = 2 mm.
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general structure probably very close to the presumed ground plan for the Olethreutes-group contrasted with several apomorphies and conspicuous reductions (like the loss of signa and disappearance of CuP with a doubtlessly associated shift in the point of origin of CuA2) suggest considerable independent development. The complex gnathos of Trachyschistis may well be close to the ground plan condition for the Olethreutesgroup, with the dorsal portion developed into the medially attenuated gnathos of Ophiorrhabda whilst in Dudua Walker the ventral part is retained in a somewhat modified version. The shape of the labial palpi, the raised scale tufts on the forewing, the loss of CuP and a very proximal origin of CuA2 in the hindwing, the shape of uncus, socii and ductus bursae and the loss of the signa are all autapomorphies for Trachyschistis hians. The slender labial palpi, shared with Dudua, Euobraztsovia Diakonoff and Hedya Hübner, is the only derived feature pointing to possible monophyly with any of the related genera. In detail, however, there are considerable differences in the shape of the palpi between Trachyschistis and the other genera, supporting a tentative phylogeny with Trachyschistis as the sister group of Dudua, Euobraztsovia and Hedya.
Constituent species Only a single Australian species is known: hians Meyrick, 1921, Exot. Microlepid. 2: 448 (Trachyschistis). [Qld: Bulimba.]
Archilobesia Diakonoff (Figs 37, 287–292) Archilobesia Diakonoff, 1966, Zool. Verh. (Leiden) 85: 45. Type species: Argyroploce drymoptila Lower, 1920, Trans. R. Soc. S. Aust. 44: 58, by monotypy and original designation. [Australia.] Lobophora Turner, 1946, Trans. R. Soc. S. Aust. 70(2): 214. Type species: Lobophora axiologa Turner, 1946, ibid. 70(2): 214, by monotypy and original designation. [Australia.] Junior homonym of Lobophora Curtis, 1825 – Lepid., Geometridae. Enveryucelia Koçak, 1981, Priamus 1(3): 118, originally proposed as an objective replacement name for Lobophora Turner.
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Diagnosis Archilobesia species are small to medium-sized brownish moths with porrect, sinuate and broad labial palpi and with complicated wing pattern of mainly longitudinal lines and streaks in cream, brown and blackish, with a dark patch at the base of the dorsum and a cream spot on the termen below the apex, followed by a brown spot in the middle of the termen. Males with narrow forewing, conspicuously modified hindwing with deeply concave termen and dorsum and a narrowly downturned anal margin, and with slender hind tibia with a long hairpencil. The wing venation of the female is unremarkable for an Olethreutini, with M3 and CuA1 very shortly stalked, but that of the male is modified due to the modified hindwing shape: M3, CuA1 and CuA2 are very short, and CuP has disappeared together with the base of 1A. In contrast to Dudua species with a concave termen and dorsum, in the male hindwing of Archilobesia 3A is widely distant from the anal margin and never partly covered by the folded margin as in Dudua. As suggested by the name, Lobesia Guenée often has a superficially similarly modified hindwing as Archilobesia, but presence of a pterostigma in the forewing will readily identify species of Lobesia. The male genitalia of all species of Archilobesia are extremely similar and diagnostic for the genus: with a roof-shaped, ventrally bristled apex to the uncus, flat drooping socii-lobes, the gnathos a membranous transverse bulge with spinulose lateral portions and a spinulose ventral hump, the valva with very long, slender and nearly parallelsided cucullus, with a small triangular bladeshaped sacculus and a raised distal margin of the basal excavation. Dudua Walker, Trachyschistis Meyrick, Hedya Hübner, Cymolomia Lederer and Ophiorrhabda Diakonoff all have very similarlooking male genitalia, but the medially continuous gnathos of Archilobesia with spinulose areas laterally as well as ventrally together with a roofshaped apex to the uncus and a large ventral tuft of bristles from the basal part of the valva are diagnostic for the genus. Trachyschistis is the only other genus with an undivided, and dorsally as well as ventrally, spinulose gnathos, but its slender, upcurved and appressed labial palpi immediately separate it from Archilobesia. The female genitalia of Archilobesia with their simple short funnel- to tube-shaped sterigma divided into two lateral lobes and the two thorn-shaped signa each in a circle of sclerotised granulation are extremely close to those of some Dudua and Hedya species, necessitating
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examination of other structures such as the labial palpi for separation.
Description Adults (Figs 287, 288) Wingspan. 10–14(16) mm; small to mediumsized. Head (Fig. 290). Ocellus well developed. Uppermost part of frons with a narrow transverse band of porrect scales, remainder of frons with minute, appressed, upwardly oriented scales. Proboscis short, about 1.5× length of labial palpus. Labial palpus moderately long (1.7–1.8× diameter of eye), sinuate, porrect and strongly widened; second segment inverted teardrop-shaped with strongly dilated and broadly rounded apex, with long loose scales along ventral margin, apex and distal part of dorsal edge; terminal segment rather long, porrect or drooping. Antenna reaching to middle of forewing; with anterior surface unscaled; cilia in male short, less than half diameter of flagellar segments. Thorax. With large posterior crest. Legs with hind tibia in male slender, with appressed scales, with a long slender hairpencil dorsally from base of tibia. Wings (Figs 37, 287, 288). Forewing index 0.37–0.42 for males, 0.40–0.43 for females; forewing subrectangular, narrow in male, with lightly curved costa, projecting rounded-acute apex and oblique to strongly oblique convex or sinuate termen; discal cell 0.57–0.59× length of wing, accessory cell moderate with chorda from halfway between R1 and R2, M-stem to base of M3; all veins present and separate beyond discal cell; all species with a very similar intricate pattern of pale beige, brown and blackish longitudinal lines and streaks, with well-developed costal strigulae converging towards a curved pale or silvery line running to above middle of termen, with a cream spot on margin of termen below apex followed by a brown mark in middle of termen and with a dark blackish brown blotch on base of dorsum and a pale silvery cream tornal margin. Hindwing strongly sexually dimorphic, cubital pecten and axillary tuft present but both weakly developed in male, hindwing shape modified in male with termen and dorsum deeply concave and separating a roughly rectangular apical lobe from a greatly enlarged triangularly projecting anal area with 3A widely distant from anal margin that is thickened, set with a long fringe of scales and narrowly folded under; venation sexually dimorphic as a consequence of
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modified male hindwing; in female with R1 visible between base of Sc and R-stem, Rs and M1 closely approximated and parallel at base, M2 bent towards bases of connate or short-stalked M3 and CuA1; CuP only a weak trace at wing margin, both 1A+2A and 3A well developed, 1A+2A with basal fork indicated; venation similar in male except for strongly shortened M3, CuA1 and CuA2 due to convex outer margin, and with modified and enlarged anal area with CuP and forked part of 1A obliterated. Pregenital abdomen. At least in some species (A. crossoleuca (Meyrick)) with a pair of faint impressions on T2, suggesting strongly reduced ‘dorsal pits’. Sternum 8 in male with bilobed hindmargin. Vinculum a rather broad, weakly sclerotised band. Male genitalia (Figs 289, 291). Tegumen high and roughly triangular in outline, with small but distinct ‘shoulders’ and with broad short apodemes for muscle m4. Uncus with a distally tapering ‘neck’ and a usually bilobed dilated apex at right angle, roof-shaped with a median ridge and hairy beneath. Socii elongate bristled lobes. Gnathos a membranous transverse bulge with spinulose lateral portions and an often medially concave spinulose ventral hump. Valva long and very slender except at base, curved and roughly parallel-sided in distal two thirds, with slender curved costal process, sacculus developed as a small projecting triangular blade from ventral margin below lowest point of basal excavation, with various tufts of bristles from concave ventral part of outer surface between sacculus and ‘elbow’ at base of cucullus, with distal margin of basal excavation raised to form a bristled projection as termination of an oblique band of strong spines from base of more or less evenly bristled cucullus. Caulis short and anellus a broad ring. Aedeagus small, moderately to strongly tapering towards apex, straight or lightly curved. Vesica without cornuti. Female genitalia (Figs 292). Ovipositor lobes elongate-ovate. Sternum 7 with hindmargin deeply emarginate. Sterigma a short and wide spinulose tube or funnel, ventrally split and dorsally variously emarginate with the resulting lateral lobes rounded or pointed. Ductus bursae with long but rather weakly sclerotised and ill-defined colliculum, inception of ductus seminalis below colliculum and bulla seminalis small, remainder of ductus bursae membranous and smooth with only faint transverse wrinkles in middle portion, faintly widening towards corpus bursae. Corpus bursae ovate, granular throughout; with two short thorn-shaped
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signa surrounded by a ring of sclerotised fine granulation each.
Distribution Taiwan, Java, New Guinea and Australia. In Australia the genus is restricted to the northernmost coastal areas of the Northern Territory and Queensland and along the Queensland coast as far south as Yeppoon.
Biology No host records are available for the genus except a possible association with Glochidion (Euphorbiaceae) according to label data of the type series of A. crossoleuca from Java: ‘bred … from larvae feeding on leaves of Glochidion (?)’.
Remarks Diakonoff (1966, 1973) provided comprehensive information on Archilobesia and figured the genitalia of several species. Clarke (1958: pl. 249, fig. 1) figured wings and genitalia of the female lectotype of A. crossoleuca. Archilobesia is a closely knit group, all the species look exceedingly similar with only small structural differences. The modified male hindwing, the wing pattern, the shape of the gnathos and the short-stalked M3 and CuA1 in the hindwing are generic apomorphies whilst the broad, sinuate labial palpi and the female genitalia agree with the presumed groundplan condition for the Olethreutes-group. Turner (1946) erroneously claimed that R4 and R5 in Lobophora are stalked. The holotype of A. axiologa agrees entirely with the structure and wing markings of A. drymoptila. Regrettably, Koçak (1981) published a replacement name for Lobophora without any enquiry as to whether a subjective replacement name was already available. The female genitalia figured for A. drymoptila by Diakonoff (1966) belong to a specimen from Taiwan that has subsequently been described as a separate species (Diakonoff 1973). Diakonoff (1973) treated the Javanese A. crossoleuca merely as a subspecies of A. drymoptila, but the wings of A. crossoleuca are narrower and the differences in the shape of the sterigma and the hindmargin of S7 in the female are of the same order as between other species of the genus. Archilobesia crossoleuca is treated here as a separate species:
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Archilobesia crossoleuca (Meyrick, 1933), Exotic Microlepid. 4: 420 (Argyroploce). [Java.] Sp. rev. The position of Archilobesia within the Olethreutes-group is not clear. Many of the obvious similarities with Dudua are probably symplesiomorphies (structure of sterigma and signa), and the modified male hindwing found in some species of Dudua is in detail very different from that of Archilobesia. Archilobesia shares neither the derived, slender, upcurved labial palpi of Dudua, Euobraztsovia, Hedya and Trachyschistis nor the reduction to one signum characterising Cymolomia and Ophiorrhabda. Whilst the presence of traces of ‘dorsal pits’ in A. crossoleuca might indicate a closer relationship with the Ophiorrhabda lineage, the weakly developed ventral band of dense bristles below the ‘elbow’ of the valva and the possibility of Glochidion as a host point in the direction of Dudua. The link with Lobesia suggested by the generic name is purely superficial due to a similar-looking, modified male hindwing. Two species of Archilobesia are currently recognised in Australia, the northern A. drymoptila (type locality Cairns), ranging from the northern fringes of the Northern Territory to the northern part of Queensland to Mackay, with A. axiologa as a junior synonym (type locality [tip of] Cape York), and an unnamed species with material from about Townsville to Yeppoon. However, the male syntype of A. drymoptila from Cairns has a very narrow uncus neck and differs otherwise subtly from all other northern Queensland/Northern Territory males. Further material may confirm A. axiologa as a separate species, restricting A. drymoptila to the Cairns region. All Australian specimens are extremely similar superficially, but the undescribed southern species has a whitish instead of greyish hind tibia in the male and is characterised by different valva and sterigma shape.
Constituent species Four named species are known world-wide, one of which, together with an additional unnamed species, is from Australia: drymoptila (Lower, 1920), Trans. R. Soc. S. Aust. 44: 58 (Argyroploce). [Qld: Cairns.] axiologa (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 214 (Lobophora). [Qld: Cape York.]
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Figs 287–292. Archilobesia drymoptila. 287, 288, adults. 287, , nr Cooktown, Q. 288, , Cape York, Q. 289, male genitalia, nr Fogg Dam, NT, slide T2194. 290, head, male. 291, aedeagus, slide T2194. 292, female genitalia, nr Fogg Dam, NT, slide T2195. Scale = 2 mm.
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Euobraztsovia Diakonoff (Figs 38, 293–298) Euobraztsovia Diakonoff, 1966, Zool. Verh. (Leiden) 85: 39. Type species: Argyroploce chionodelta Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 274, by monotypy and original designation. [Australia.]
Diagnosis Euobraztsovia species are medium-sized brownish grey olethreutine moths with subrectangular forewings with an erect teardrop-shaped large snow-white blotch on tornus and an additional semicircular white area on base of dorsum in male. The rather small, slender, upcurved labial palpi, appressed to the frons and with the curved second segment widest in the middle, are very characteristic and shared only with Dudua in the Australian fauna. Euobraztsovia and Dudua are structurally so similar that apart from the unmistakable wing pattern of E. chionodelta only genitalia examination will discriminate between the two genera. Wing venation and secondary male characters of Euobraztsovia completely agree with those of Dudua, with the wing venation quite unremarkable for a genus of the Olethreutini. The bases of R3, R4 and R5 are rather closely approximated and CuA1 is strongly bent towards M3 at the wing margin, and M3 and CuA1 in the hindwing are connate. Males of Euobraztsovia have a slightly enlarged hind tibia with a hairpencil and a hindwing with a large ‘anal roll’ with the anal margin broadly folded under over a clump of modified scales and the entrance into this fold on the lower surface covered by the crest of a second, topographically underlying parallel fold from behind 3A. The male genitalia of Euobraztsovia with a ventrally hairy, membranous, elongaterectangular uncus laterally fused with drooping socii lobes and with highly elaborate valvae with a deeply concave central part edged by sharp ridges and blade-shaped projections set with heavy spines are diagnostic for Euobraztsovia. Podognatha has a similar, deeply concave median portion to its valva, ending in a blade-shaped ventral projection, but its uncus bears a discrete group of strong bristles and the socii are either rigid, distally naked projections or small hairy patches. The structure of the bursa copulatrix in the female genitalia with a long, weakly sclerotised bipartite colliculum and two nipple-shaped, granular signa, is exactly like that of Dudua and related genera, but the sterigma of the
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single species of Euobraztsovia is highly apomorphic and thus diagnostic for the genus.
Description Adults (Figs 293, 294) Wingspan. 13–16 mm, medium-sized. Head (Fig. 295). Ocellus well developed. Upper third of frons with a transverse band of anteriorly projecting scales, remainder with minute, upwardly oriented, appressed scales. Proboscis short, at most 1.5× length of labial palpus. Labial palpus rather small (1.5–1.7× diameter of eye), slender, strongly upcurved and appressed to frons; second segment strongly curved, with shortish but erect scales all along ventral margin and on apex, widest in middle and slightly tapering towards apex; with terminal segment at a slight angle, short and pointed. Antenna reaching to middle of forewing; anterior surface unscaled; cilia in male short, less than 56: diameter of flagellar segments. Thorax. With large posterior crest. Legs with hind tibia in male with appressed but lengthened scales along dorsal margin and inner surface, with a long hairpencil dorsally from base concealed in a groove in scaling along inner surface. Wings (Figs 38, 293, 294). Forewing index 0.40–0.43; forewing elongate, subrectangular; costa lightly curved throughout, more so towards apex, apex obtuse, termen straight to faintly concave and nearly vertical; discal cell 0.66× length of wing, accessory cell very narrow with chorda from below R2, M-stem to between M2 and M3; all veins present and separate beyond discal cell and CuA1 bent towards M3 at margin; mottled silvery grey and brown with a large snow-white teardropshaped mark centred on tornus, its pointed tip reaching nearly to costa just before apex, males with a second, semicircular white area across basal half of dorsum. Hindwing with strongly developed cubital pecten, sparse axillary tuft and scattered long hairscales from upper surface of anal area, in male with a complex ‘anal roll’ with thickened anal margin folded under over a clump of strongly modified scales and entrance into this fold covered by the crest of a second, parallel, dorsally concave fold invaginated just behind 3A; with Rs and M1 parallel and closely approximated at base, with M2 non-tubular at base and bent closely towards M3, and with M3 and CuA1 connate; with CuP present near margin, 1A+2A and 3A both well developed but base of 1A weak and 3A closely adjacent to ‘anal roll’; greyish with bronze gloss.
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Pregenital abdomen. Unmodified. Hindmargin of S8 in male weakly bilobed with shallow, Vshaped median incision. Male genitalia (Figs 296, 297). Vinculum a moderately broad lightly sclerotised band. Tegumen triangular, high and rather narrow, pedunculi very broad, club-shaped, with broad apodemes for
147
muscle m4. Uncus a membranous, elongaterectangular irregular process, ventrally densely covered with long hair and laterally continuous with rounded, drooping socii lobes covered with similar hair. Gnathos an ill-defined, faintly sclerotised transverse band, ventrally weakly spinulose, continuous with similarly sclerotised
Figs 293–298. Euobraztsovia chionodelta. 293, 294, adults. 293, , Kuranda, Q. 294, , Mossman, Q. 295, head, male. 296, 297, aedeagus and male genitalia, Cairns, Q, slide T2272. 298, female genitalia, Mt Bartle Frere, Q, slide ANIC 17048. Scale = 2 mm.
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ventral surface of projecting anal tube. Valva sinuate, broader and highly ornamented in basal half, with a sinuate costal process and a circular projecting lobe with a large group of equally spaced prominent bristles on ventrodistal margin of basal excavation, outwardly followed by a curved deep concavity running across valva from base of costa to before middle of ventral margin, distally delimited by a curved blade-shaped ridge, strongly projecting on ventral margin and with a row of heavy spines running from ventral projection to beyond middle of valva in a second parallel, curved groove that ends at base of costa in another projecting blade-like process with heavy spines on inner surface; cucullus clavate, curved, distally rounded and ‘elbowed’ at ventral margin. Caulis short and anellus a broad ring. Aedeagus moderately long, nearly parallel-sided in basal half, distal half curved; vesica without cornuti. Female genitalia (Fig. 298). Ovipositor lobes subtriangular. Sternum 8 and preceding intersegmental membrane with sterigma strongly modified: ventral intersegmental membrane strongly sclerotised and transversely deeply folded, producing a high ‘collar’ around S8 with a deep Vshaped ventral incision leading to ostium and flanked on each side by a spinulose projecting lobe that is laterally continuous with the much longer, swollen, lateral lobes of the ‘collar’. Ductus bursae with long, bipartite, weakly sclerotised colliculum followed by a membranous, lightly transversely wrinkled central portion with inception of ductus seminalis before midlength, remainder smooth and membranous. Corpus bursae ovate to pear-shaped, bursa wall weakly granulate throughout, with two nipple-shaped signa with the hollow blunt thorns and surrounding sclerotised ring densely granular.
widely distributed throughout New Guinea and its islands. Structurally the genus is obviously close to Dudua with exactly the same wing venation and male secondary sexual characters such as the tibial pencil and an ‘anal roll’ in the hindwing enclosing a clump of modified scales. The characteristic shape of the labial palpi is the most obvious snyapomorphy linking the two genera, further supported by the wing shape and apical strigulation of Euobraztsovia that conforms to that of generalised Dudua species. The genitalia in both sexes are so different, however, that separation into two genera can easily be justified with at least the males exhibiting a number of autapomorphies for each genus. Diakonoff (1966) provided a description and figures of the genitalia of both sexes of Argyroploce chionodelta and further on in the same publication gave a brief diagnosis of the newly described Euobraztsovia, together with wing venation and head figures. In the original description as well as in the subsequent treatment for the south Asiatic Olethreutini, Diakonoff (1973) stated that the male hind tibia lacked a hairpencil, a statement at variance with observations on well preserved Australian specimens.
Constituent species Only one species of Euobraztsovia is known so far, described from Australia: chionodelta (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 274 (Argyroploce). [Qld: Cairns, Innisfail.]
Eremas Turner (Figs 39, 299–306)
Distribution New Guinea and Australia. In Australia the genus has been found in northern Queensland, at Heathlands on Cape York Peninsula and in the Cairns region.
Eremas Turner, 1945, Trans. R. Soc. S. Aust. 69: 71. Type species: Eremas leucotrigona Turner, 1945, ibid. 69: 71, by monotypy. [Australia.]
Diagnosis Biology No host records are available, but all Australian localities are in rainforest areas.
Remarks Diakonoff (1966) erected the genus Euobraztsovia for a single species described from Australia and
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Eremas species are small dark greyish to blackish moths with a large pinkish tinged ochreous whitish semicircle from about 56: costa to before apex, reaching halfway across wing and bordered with a black band. The same conspicuous wing pattern is found in Statherotis Meyrick and several related genera, but with a slightly less evenly rounded pinkish mark than in Eremas and with more
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Figs 299–303. Eremas, adults, hindwing and head. 299, E. tetrarcha, , Coolangatta, Q [image reversed]. 300, E. sp., , nr Yeppoon, Q [image reversed]. 301, E. leucotrigona, , Cape York, Q. 302, male hindwing pencil, E. leucotrigona. 303, head, E. leucotrigona, male. Scale = 2 mm.
regular dark costal strigulae. Secondary sexual structures allow easy separation in males: Eremas has the margin of the forewing dorsum narrowly folded under with an associated large pencil of hair-scales on the upper side of the hindwing, features that are absent in the Statherotis-group of genera. Both sexes of Eremas lack a cubital pecten and the black sex scales present on the male hindwing of some similarlooking species of Statherotis are absent. The wing venation of Eremas is characterised by R4 in the forewing closer to R5 than to R3 (R3 and R4 approximated or stalked in many Statherotis) and by an unusually distal origin of CuA2 in the hindwing. The male genitalia of Eremas with broad lamellar lateral flanges from the tegumen with an associated tuft of long scales and with a narrow valva with a very large ventral spine are quite unique. A simple, ventrally notched funnel-shaped
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sterigma and two large slender horn-shaped signa characterise the female genitalia.
Description Adults (Figs 299–301) Wingspan. 9.5–13.0 mm; small. Head (Fig. 303). Ocellus well developed. Vertex scales anteriorly barely protruding between antennae. Frons in uppermost part with a narrow transverse band of anteriorly projecting scales, remainder with small, appressed, upwardly directed scales. Proboscis short, 1.0–1.5× length of labial palpus. Labial palpus rather short (1.5–1.8× diameter of eye) and slender, sinuate, appressed and ascending except for apex; second segment sinuate, slightly widened in distal half by loose scales along ventral margin and on dorsal margin before apex; terminal segment moderately long, porrect.
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Antenna reaching to just over middle of forewing; anterior surface unscaled; cilia in male small. Thorax. With posterior crest. Legs with modified scaling on hind femur and tibia in male: hind femur with a dense and large fan-like tuft of long scales from middle of ventroposterior surface, hind tibia with a long thin pencil of greyish hairscales from base, at rest partly concealed in a groove along inner surface. Wings (Figs 39, 299–302). Forewing index 0.43–0.48; forewing elongate subrectangular to subtriangular; costa curved in proximal half only, apex rounded-acute, termen distinctly oblique and convex, edge of dorsum in male narrowly folded over underside from beyond patch of microtrichia to about 6M dorsum, with modified opaque scales apparently within fold (i.e. on undersurface); discal cell 0.57× length of forewing, M-stem and chorda well-developed, chorda from halfway between R1 and R2 to R5; all veins present and separate beyond discal cell; anal loop 0.4× length of 1A+2A; dark brownish grey to blackish brown with a large semicircular pinkish tinged whitish ochreous mark centred on costa from about 56: costa to before apex, reaching halfway across wing, bordered with an indistinct blackish band, and with only traces of dark costal strigulae except for one larger distinct blackish spot before middle of pinkish semicircle on costa. Hindwing without cubital pecten in both sexes but with well-developed axillary tuft and in male with a large, pencil-like tuft of long hairscales on upper surface from within cell near point of divergence of Sc and base of Rs (Fig. 302), with anal margin thickened and densely scaled in male; Rs and M1 approximated at base, but not parallel; M2 bent towards connate or short-stalked M3 and CuA1 at base, CuA2 from between 36: and 6M cell length, CuP a non-tubular vestige at wing margin; vestigial 3A indicated only at base and margin; pale to dark grey. Pregenital abdomen. Unmodified. Sternum 8 in male with a faintly bilobed hindmargin, T8 with weak, inverse T-shaped median sclerotisation. Male genitalia (Figs 304, 305). Vinculum a narrow band with a very small triangular saccus. Tegumen very high and narrow anteriorly, but laterally greatly expanded, and longitudinally folded, with huge swollen, membranous ‘shoulders’ caudally paralleled by even bigger dorsally rounded wing-like flanges with large tufts of long hair-scales on each side from membrane between base of these two pairs of lateral processes. Uncus a rod- to clubshaped, weakly sclerotised curved median process with an evenly bristled rounded and sometimes
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Figs 304–306. Eremas, genitalia. 304, E. leucotrigona, male, Cape York, Q, slide T1981. 305, E. tetrarcha, male, Coolangatta, Q, slide ANIC 2461. 306, E. sp., female, Yeppoon, Q, slide T1987.
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medially notched apex. Socii moderately large, membranous, drooping and bristled ovate flaps. Gnathos two weak lateral bands fused with a large, triangular subscaphium. Valva very narrow and parallel-sided, straight or sinuate in basal half, with a large costal process and an either thin straight, finger-like erect process from base of costa with short bristles along inner edge only (E. leucotrigona) or an evenly bristled, rounded, down-curved knob (E. tetrarcha (Meyrick)), and with a huge long spine from about middle of ventral edge at base of cucullus. Caulis very short and fused with large erect diamond-shaped anellus. Aedeagus rather short and small, straight or curved with more or less bulbous base; vesica without any trace of cornuti. Female genitalia (Fig. 306). Ovipositor lobes subtriangular. Sternum 7 with evenly concave hindmargin; T7 strongly sclerotised especially along margin and of a transversely crescentic shape with concave hindmargin and produced lateral corners. Sterigma simple, a wide spinulose funnel with a large V-shaped excision in ventral edge. Ductus bursae with a very long colliculum-like sclerite below ostium extending over about 56: length of ductus to just above inception of ductus seminalis. Corpus bursae ovate, bursa wall finely granulate; with two large horn-shaped signa with an angled hollow base and sharp point.
Distribution Eremas is recorded only from Australia, from three widely separate areas along the eastern coast of Queensland: Cape York, Yeppoon, and Brisbane to Coolangatta.
Biology No biological information is available, but the locality data seem to suggest a seasonal rainforest habitat.
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absence of a cubital pecten, the lamellate lateral projections from the tegumen with associated scale tufts, the slender valva with a basal process on its inner base and a large ventral spike, and the slender, horn-shaped signa. The taxonomic position of Eremas is unclear, and no sister group can be identified at this point. The genitalia of both sexes are a curious mixture of rather plesiomorphic characters (uncus and socii in male, simple sterigma in female, possibly the slender, horn-shaped signa) with some unique apomorphies such as the lateral flanges to the tegumen and the highly derived valva. The valva with its well-developed basal process and the modified and hairy ‘shoulders’ of the tegumen recall some species of Neopotamia Diakonoff whereas the simple sterigma and the paired, slender, horn-shaped signa suggest a position in the Olethreutes-group of genera, where the genus is tentatively included for the time being. The narrow fold containing modified scales on the underside of the male forewing dorsum of Eremas and the hair-pencil on the upper side of the hindwing (Fig. 302) suggest a functional unit, a combination apparently unique among olethreutines. The dark grey hindwings of the single Eremas female available suggest that it is a specimen of E. tetrarcha, but as the locality (Yeppoon) lies between the known ranges of E. leucotrigona (Cape York) and E. tetrarcha (Brisbane, Coolangatta), the specimen is treated as Eremas species.
Constituent species leucotrigona Turner, 1945, Trans. R. Soc. S. Aust. 69: 71 (Eremas). [Qld: Cape York.] tetrarcha (Meyrick, 1920), Exot. Microlepid. 2: 349 (Argyroploce). [Qld: Brisbane.]
Zomaria-group Remarks Turner (1945) described Eremas as a tortricine, doubtlessly prompted by the absence of a cubital pecten. The single type of E. tetrarcha (figured in Clarke 1958: pl. 275, fig. 3) lacks the abdomen and the species was included in Statherotis on the basis of its wing markings (Diakonoff 1973). Eremas is a conspicuous, well-characterised genus with numerous apomorphies including the downfolded forewing dorsum and the modified scale tuft in the hindwing cell of the male, the
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Status remarks Diakonoff (1973) established the subtribe Zomariae for Zomaria Heinrich, Actinocentra Diakonoff, Eubrochoneura Diakonoff and Thysanocrepis Diakonoff. Thysanocrepis was included on the basis of a misidentified female; it clearly belongs to the Enarmoniini (see p. 292). Diakonoff (1984a) later described Zomariana Diakonoff, considering it to be intermediate between the Eucosmini and Olethreutini. Zomariana is here referred to the
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Zomaria-group. Podognatha Diakonoff also is here tentatively included in this group, as discussed in the Remarks for Podognatha.
Evidence for monophyly Diakonoff (1973) defined the Zomaria-group primarily on signum structure, which he described as ‘a stellate or substellate scobination’. He also mentioned absence of an uncus, large socii, a tendency for a strongly developed gnathos and a complex valva. These features are probably all apomorphies for the Zomaria-group, though Zomariana lacks a signum. A sterigma forming a protruding rim continuous with the sclerotised posterior part of the ductus bursae may be an additional shared derived character.
Distribution Zomaria occurs in the Nearctic and in New Guinea. Actinocentra is known from Sumatra, Eubrochoneura from Sulawesi, New Guinea and the Solomon Is, and Zomariana from Java, Sumba and Australia. The tentatively included Podognatha occurs from the Moluccan Is across New Guinea to Australia and the Solomon Is.
Biology Larvae of the Zomaria-group are leaf tiers, the Nearctic Zomaria species on Ericaceae (Heinrich 1926). Zomariana is more polyphagous, with the Australian Z. doxasticana reared from Surianaceae, Caesalpiniaceae and Mimosaceae and Z. carnicolor from Oxalidaceae in Java.
Diagnosis Zomariana species are medium-sized, greyish moths with dark grey and distally red-brown costal strigulation, variously suffused with dark grey or red-brown, and with a dark pencil of hair-scales in a groove along anal margin of hindwing. With triangular labial palpus, a pair of round deep depressions dorsally on second abdominal segment, and unmodified hind tibia in male. Wing venation typical for a member of the Olethreutini, with M2, M3 and CuA1 in hindwing all closely approximated (or the latter two very shortly stalked), but with short chorda from between R2 and R3. The forewing costa is evenly convex throughout, not concave in its central portion as in the closely related Eubrochoneura Diakonoff. The male genitalia of Zomariana, Eubrochoneura and Zomaria Heinrich are extremely similar, with very large socii, a well-developed gnathos and a subbasally constricted, membranous valva with a large bundle of long setae in a ventral indentation and with a medio-dorsally directed bristled lobe from ventral margin of basal excavation. Zomariana differs from the other two genera by particularly large socii, broadly elliptic and extending backwards, and by a series of large flattened bristles along its dorsal valva margin. The W-shaped, raised and spinulose sterigma and the long and narrow corpus bursa without signa but with inception of ductus seminalis on its neck are diagnostic for the female genitalia of Zomariana. Those of Eubrochoneura are very similar but with an uninterrupted ring-like spinulose rim as sterigma and with a signum, a scobinate crease with a group of radiating small sclerites at one end.
Description Remarks The male genitalia, the female signum, the paired depressions on T2 and the reduced tibial setae present in the Zomaria-group provide obvious links with Gatesclarkeana and related genera, and further study is needed to establish whether they constitute synapomorphies.
Zomariana Diakonoff (Figs 40, 307–317) Zomariana Diakonoff, 1984a, Entomol. Basil. 9: 411. Type species: Argyroploce carnicolor Meyrick, 1931, Exot. Microlepid. 4: 133 (as 311 in Diakonoff 1984a), by original designation and monotypy. [Java.]
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Adults (Figs 307–310) Wingspan. 12–16 mm; medium-sized. Head (Fig. 315). Ocellus well developed. Frons in uppermost part with a narrow transverse band of anteriorly projecting, shortish scales; remainder of frons with very small, appressed and upwardly oriented scales. Proboscis short, hardly longer than labial palpus. Labial palpus moderately long (1.8– 2× diameter of eye) and triangular, sinuate and widest at apex of second segment; second segment sinuate and of subtriangular outline, strongly widening towards apex and with long loose scales along ventral margin and apex; terminal segment long and thin, downwardly directed and often nearly hidden in apical scales of second segment. Antenna reaching to middle of forewing; with anterior surface unscaled; cilia in male very short, less than half diameter of flagellar segment.
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Thorax. With large posterior crest. Legs with unmodified hind tibia in male; tarsal bristles reduced to at most a single distal bristle. Wings (Figs 40, 307–312). Forewing index 0.40–0.43; forewing elongate, moderately widened distally, with lightly and evenly curved costa, oblique straight termen and broadly rounded tornus; discal cell 0.62× length of forewing, with small triangular accessory cell with chorda from between R2 and R3 to opposite R5, and with weakly developed M-stem to between M2 and M3; all veins present and separate beyond discal cell; forked part of 1A+2A 0.3× length of vein; of light greyish ground colour and genus-typical pattern restricted to dark grey costal strigulae, suffused with reddish brown in distal part of costa, but Z. doxasticana (Meyrick) with a number of very conspicuous variable markings produced by redbrown or dark grey suffusion across certain wing parts: with an ill-defined oblique darker grey fascia (Fig. 307), two large blackish grey spots on each wing in middle of disc and towards base (Fig. 309), dark grey suffusion along base of costa and across apex to entire wing uniformly dark grey, or a large red-brown semicircle over dorsum and some redbrown spots along costa (Fig. 310). Hindwing with cubital pecten and axillary tuft, and in male with an elongate patch of very long dark-based hairscales on upper surface along anal margin (Fig. 311), at rest folded as a dark pencil into a submarginal dorsal groove along 3A (Fig. 312); with Rs and M1 closely approximated and parallel at base, M2 close to M3 at base, M3 and CuA1 connate to very shortly stalked, CuA2 from just below 56: cell; 1A+2A and 3A well developed, but the course of the latter concealed in the male by the tuft of hair-scales; light grey with pinkish or coppery hue. Pregenital abdomen. Two deep round depressions or invaginations anterolaterally on T2 in both sexes (311, 312, 314), often partly concealed by broad scales from hindmargin of preceding segment. The pupa has two similar pouch-shaped invaginations laterally on the dorsum of the second abdominal segment, with a narrow and strongly sclerotised orifice. Eighth segment in Z. doxasticana male weakly sclerotised, S8 with long dense scales along hindmargin and with a V-shaped median excision, T8 with two transverse sclerites along midline, a triangle at base followed by a crescentic sclerite. Male genitalia (Figs 316, 317). Vinculum a membranous band. Tegumen short and tapering towards broadly rounded apex, rather membranous apart from conspicuous rod-like ridges along dorsal margin of pedunculi, with broad rounded-
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triangular apodemes (presumably for muscle m4) from mid-height of dorsal margin. Uncus absent. Socii two very large broadly elliptic flat lobes, in natural position extended backwards between and parallel to closed valvae, longer than high and on outer surface covered with large broad scales in dorsal part, giving way to hair-scales in ventral portion. Gnathos well developed, two lateral arms medially fused with a complicated and distally strongly sclerotised structure: a curved subtriangular plate with its apex drawn out into a lanceolate, sclerotised point, its lateral corners continued into the gnathos arms and with two spoon-shaped membranous processes from its base, on either side of the aedeagus and possibly fused with the anellus (Z. doxasticana). Valva long, of very complicated structure and highly membranous, with basal excavation and small bent costal process hidden behind a flat, large, triangular, mediodorsally directed projecting bristled lobe from inner surface of valva, with a projecting rounded or rounded-conical (Z. doxasticana) membranous sacculus, with an indentation and/or invagination in ventral margin beyond sacculus with a large bundle of long setae arising from a hump deep in its apex, cucullus broad and distally rounded, membranous and finely hairy throughout, with a row of very large flattened bristles along costa. Caulis short and triangular, anellus a narrow band. Aedeagus rather short, irregularly bent, with lightly dilated base and bevelled apex; vesica in parts of granular structure, apparently without cornuti. Female genitalia (Fig. 313). Ovipositor lobes elongate, subelliptic. Sternum 7 with a concave hindmargin. Sterigma a strongly spinulose W-shaped, dorsally cleft rim more or less continuous with tube-shaped sclerotised ductus bursae. Ductus bursae a short, straight sclerotised tube with laterally thickened walls (possible extended colliculum). Corpus bursae very long and narrow, cylindrical with rounded apex, with inception of ductus seminalis in posteriormost part of corpus bursae; without signa.
Distribution Zomariana is known from Java, Sumba and Australia. Zomariana doxasticana (Meyrick), apparently restricted to Australia, is probably the most widely distributed species of the Olethreutini in Australia, ranging across most of the continent apart from the most arid regions and the extreme south-eastern and south-western corners and Tasmania. It is self-adventive in New Zealand, on phyllodinous Acacia spp. (Dugdale et al. 2005).
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Biology The occurrence of Z. doxasticana from extremely wet rainforest habitats to areas with very low rainfall in the centre of the continent is quite remarkable. The species is polyphagous and has been reared by N. McFarland near Geraldton,
Western Australia, from foliage of Stylobasium spathulatum (Surianaceae). Other records include Senna artemisioides (Caesalpiniaceae), Acacia ligulata, A. melanoxylon, A. salicina (Mimosaceae) and one rearing from a gall on a seed pod of an unidentified Acacia sp. Diakonoff (1984a) reported
Figs 307–312. Zomariana doxasticana, adults and male hindwing fringe. 307, , nr Rockhampton, Q [image reversed]. 308, , nr Alice Springs, NT. 309, , Toowoomba, Q [image reversed]. 310, , nr Wagin, WA. 311, 312, male hindwing with anal fringe. 311, expanded; 312, enclosed in wing fold. (af, anal fringe; dd, dorsal depressions). Scale = 2 mm.
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Averrhoa carambola (Oxalidaceae) as host of Z. carnicolor in Java.
Remarks Diakonoff (1984a) provided illustrations of head, wing venation and genitalia of both sexes for his generic description of Zomariana. He figured the forewing with the M-stem ending between M1 and M2, commenting on this unusual position, and the
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hindwing with CuA2 originating from beyond 6M cell. Both these veins have a more orthodox course in Z. doxasticana, with the M-stem to between M2 and M3 and the hindwing CuA2 from below 36: cell. Diakonoff’s reference to a transtilla as ‘a slender rod with dilated oval ends’ refers to the ventral portion of the complex gnathos with its two lateral spoon-shaped processes. Zomariana and Eubrochoneura are exceedingly close and it may be debatable whether two separate
Figs 313–317. Zomariana doxasticana, genitalia, abdomen and head. 313, female genitalia, Black Mt, ACT, slide T2271. 314, abdomen base with dorsal depressions (dd). 315, head, male. 316, 317, male genitalia and aedeagus, Black Mt, ACT, slide T2268.
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genera are justified. The sinuate, medially concave costa of Eubrochoneura and the narrow, strongly elongate corpus bursa without signum in Zomariana, together with the consistently different sterigma shape, strongly support the monophyly of both groups. However, since the two genera clearly differ in their superficial appearance it seems reasonable to accept the status quo until more species become known. The complex and highly characteristic valva with a pencil of bristles and presence of an often strongly sclerotised gnathos readily identify Zomariana as a member of the Zomaria-group of genera. The male genitalia, the paired depressions on T2 and the reduced tibial setae of Zomariana provide obvious links with Gatesclarkeana and related genera, and further study is needed to establish whether they constitute synapomorphies.
Constituent species Two species of Zomariana are known, the type species from Java and one named species from Australia: doxasticana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 644 (Penthina). [NSW: Parramatta. Qld: Brisbane.] phoenicoplaca (Turner, 1945), Trans. R. Soc. S. Aust. 69: 62 (Tortrix). [Qld: Cunnamulla.] eurypolia (Turner, 1946), Trans. R. Soc. S. Aust. 70: 209 (Eucosma). [Qld: Tweed Heads (Burleigh).]
Podognatha Diakonoff (Figs 41, 318–328) Podognatha Diakonoff, 1966, Zool. Verh. (Leiden) 85: 59. Type species: Podognatha tamias Diakonoff, 1966, ibid. 85: 61, by monotypy and original designation. [New Guinea: New Ireland.] Molybdocrates Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 369. Type species: Molybdocrates opulenta Diakonoff, 1973, ibid. 1: 370, by original designation. [Moluccan Is.]
Diagnosis Podognatha species are medium-sized olethreutine moths with distally dilated forewings with the
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distal third of the costa characteristically angled or strongly curved, either pale greyish ochreous to grey with some darker speckling over basal third, with small dark grey-brown marks along costa and sometimes termen and with indistinct iridescent dots across wing (unnamed Australian species), or dark blackish brown with variably-sized ochreous blotches and metallic bluish leaden dots across wing (P. vinculata (Meyrick)). Thysanocrepis crossota (Meyrick) looks deceptively like the pale greyish species of Podognatha, in wing shape as well as markings, but absence of an ‘anal roll’ and tibial pencil in the male and a forewing with a distinctly sinuate costa, with very closely approximated R3 and R4 and without an accessory cell, readily differentiate Thysanocrepis Diakonoff from Podognatha. Podognatha has sinuate-porrect to sinuate-upcurved labial palpi, distinctly ciliate antennae and a tibial pencil in the male. The distally dilated forewing with the costa strongly curved or angled in its distal third is characteristic for Podognatha, as is the strongly modified male hindwing with the margin usually projecting as a semicircular lobe between CuP and 1A+2A, then deeply concave and projecting again as a complex narrow upfolded anal roll beyond 3A. Both features are shared with the closely related Theorica Diakonoff that has, however, a hindwing with stalked M3 and CuA1. A short, triangular accessory cell with the chorda from halfway between R2 and R3 and, in the hindwing, a very weak CuP and a short fused portion of 1A+2A with a thickened base to 1A are the most remarkable characteristics of the venation of Podognatha. The male genitalia are very diverse with a stout, strongly sclerotised tegumen with long rigid socii of diagnostic shape (reduced to hair tufts in P. vinculata) and with a very variably shaped valva with a deep oblique depression from the base of the costa to the ventral margin below the cucullus base, surrounded by several characteristic tufts of bristles or sclerotised processes. The female genitalia of Podognatha are equally diverse, with the sterigma varying from a large, well sclerotised, transversely rectangular plate, fused with the colliculum by a sclerotised tube, to a strongly spinulose, weakly sclerotised and bulging rectangular portion of intersegmental membrane. The ductus bursae often contains a partly sclerotised dilated portion and the single signum, if present, is a large convex structure consisting of a semicircular basal part of fused teeth crowned by a semicircular row of distally free teeth.
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Description Adults (Figs 318–321) Wingspan. 11.5–17.0 mm; medium-sized. Head (Fig. 326). Ocellus well developed. Uppermost part of frons with a transverse band of anteriorly oriented short to moderately long scales, remainder with very small, appressed, upwardly oriented scales. Proboscis short, hardly longer than labial palpus. Labial palpus moderate to longish (1.6–2.3× diameter of eye), sinuate-porrect to sinuate-upcurved, moderately to strongly widened; with second segment from basically upcurved and only distally sinuate, moderately widened and roughly parallel-sided in distal half (P. vinculata (Meyrick)) to strongly sinuate and distally broadly widened to inverted teardrop shape (unnamed Australian species); with third segment longish and porrect. Antenna reaching to middle of forewing; anterior surface unscaled; cilia in male moderate, nearly as long as diameter of flagellar segments, distinctly verticillate. Thorax. With prominent posterior crest. Legs with hind tibia in male with long but appressed scales along dorsal margin and inner surface, with a groove along inner surface containing a large dark hairpencil from dorsal base of tibia (bipartite in P. vinculata, with one part much shorter); tarsal bristles reduced to at most a single distal bristle. Wings (Figs 41, 318–321, 325). Forewing index 0.45–0.48; forewing elongate subovate to rhomboid, distally dilated, with costa weakly to moderately curved at base, straight to faintly convex in central portion and either strongly curved or distinctly angled in distal third, apex acute, rounded or angled, termen straight and oblique, tornus broadly rounded; discal cell 0.61– 0.67× length of wing, accessory cell triangular, short to very short and sometimes quite broad (P. opulenta), chorda from halfway between R2 and R3, M-stem only weakly sinuate, to between M2 and M3; all veins developed and separate beyond discal cell, R3, R4 and R5 usually roughly equidistant at base; either pale greyish ochreous (darker, more greyish in female), with some dark grey-brown speckling or marks over basal third and similar dark grey-brown spots along costa and sometimes along termen, and with pale grey iridescent to leaden grey metallic dots scattered across wing (P. tamias, unnamed Australian species), or brownish ochreous with a large blackish blotch over middle of costa and wing
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variously suffused with blackish brown to entirely darkened, with bluish leaden dots scattered across wing (P. opulenta, P. vinculata). Hindwing with weakly to moderately developed cubital pecten, with scattered long hairscales from upper surface of anal area and in male with highly modified tornal, dorsal and anal margin (Fig. 41): wing margin projecting as a nearly semicircular lobe between CuP and 1A+2A, then deeply concave between 1A+2A and 3A (except for one unnamed Papuan species) and sharply projecting again as a narrow anal roll with the anal margin folded or rolled over, usually containing a patch of highly modified scales (Fig. 325); Sc+R1 and Rs sometimes anastomosing beyond base, Rs and M1 closely approximated at base, M2 curved towards M3 and the two veins moderately to very close at base (P. tamias, unnamed Australian species), M3 and CuA1 connate; CuP only a weak trace, fused part of 1A+2A very short and base of 1A much thicker but interrupted, 3A well developed. Pregenital abdomen. Unmodified. Hindmargin of S8 in male moderately to strongly bilobed, hindmargin of T8 a semicircular lobe. Male genitalia (Figs 323, 324, 327). Vinculum a broad, lightly sclerotised band. Tegumen elongaterectangular to subtriangular, large, robust and well sclerotised, with prominent rounded-rectangular shoulders and broad pedunculi. Uncus a moderately long, robust, ventrally curved process with some strong apical bristles or very few curved spines from apex, distal half sometimes laterally compressed (uncus very rarely bilobed, consisting of two ventrally bristled hooded lobes (P. opulenta)). Socii typically very long, slender, porrect and strongly sclerotised arms with a bristled basal portion and more strongly sclerotised, usually flattened and often dilated apices; socii rarely reduced to a patch of bristles on shoulders of tegumen (P. vinculata). Gnathos hardly differentiated, a faint transverse band below membranous anal tube in species with porrect socii (P. tamias) and presumably fused with sclerotised ventral surface of anal tube in P. vinculata that lacks large projecting socii; some species (P. tamias, unnamed Australian species) with a sclerotised transverse bulge in membrane above aedeagus, of unclear homology. Valva very variably developed, with a small costal process and with a very diversely shaped deep depression running obliquely across valva from below base of costa and ending on ventral margin below base of cucullus usually in a tuft of long or a clump of short and sometimes
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Figs 318–321. Podognatha, adults. 318, 319, P. sp. nr rebellis, Mt Bellenden-Ker, Q. 318, [image reversed]. 319, . 320, 321, P. vinculata. 320, , Mt Bellenden-Ker, Q [image reversed]. 321, , nr Port Macquarie, NSW [image reversed]. Scale = 2 mm.
flattened bristles (P. tamias, unnamed Australian species) or in a blade-shaped process with bristles on outer surface (P. vinculata), and (except in P. vinculata) with another group of bristles between this depression and basal excavation (often raised on a blade-shaped process), and usually with a second, inwardly oriented and often bristled process extending across upper part of depression from base of cucullus below costa (absent in P. vinculata, ending in a long ladle-shaped process in unnamed Australian species), with cucullus narrower than base of valva except for P. opulenta and always with some stronger spines along ventral margin and/or across base. Caulis shortish, anellus a broad and often irregular ring. Aedeagus from very small, tapering towards apex (unnamed Australian species) to small and curved with a
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conspicuous apical thorn (P. vinculata); vesica at least in some species (P. vinculata) with a bundle of needle-like, presumably deciduous cornuti. Female genitalia. (Figs 322, 328). Ovipositor lobes rather broad. Sternum 7 weakly sclerotised, hindmargin straight to concave. Sterigma of two basic shapes: either a large, strongly sclerotised, roughly rectangular and partly spinulose plate with a subcentral opening leading to ostium, with lamella antevaginalis forming the strongly arched ventral wall of a wide tube fused with colliculum at base and with lamella postvaginalis often medially projecting as a broad flattened ridge with everted laterocaudal corners (P. vinculata), or sterigma a much less well-defined, weakly sclerotised, transversely rectangular area of bulging, strongly spinulose intersegmental membrane with a deep
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vertical fold leading to ostium (unnamed Australian species). Ductus bursae either with moderately long, lightly sclerotised bipartite colliculum and gradually widening into corpus bursa (unnamed Australian species), or with colliculum wide, modified and fused with sterigma and with a dilation with some sclerotised granulation below inception of ductus seminalis, followed by a very wide, longitudinally wrinkled membranous distal portion (P. vinculata). Corpus bursae either elongate, sac-shaped and entirely continuous with ductus bursae, without signum (unnamed Australian species), or ovate, with a large convex signum with a semicircular central part of fused teeth crowned by a semicircular row of distally free teeth (P. vinculata).
Distribution Moluccan Is, New Guinea, Solomon Is, Australia. In Australia the genus has been found in the coastal region of eastern Queensland and northern New South Wales, from north of Cooktown to Wingham.
Biology Podognatha vinculata was reared from a leafrolling larva on Alangium villosum (Alangiaceae) at Wingham, New South Wales, by J. Stockard. Diakonoff (1973) recorded label data ‘bred from Polyalthia nitidissima’ (Annonaceae) from a specimen in the British Museum from Toowong, Queensland, identified by Meyrick as ‘Argyroploce vinculata Meyr’. All Australian localities suggest rainforest habitats.
Remarks Podognatha comprises in both sexes a wide range of genitalia structures, with the two Australian species, P. vinculata and an unnamed species, illustrating extremes of the spectrum for both males and females. Added to this is a further dichotomy in forewing shape and coloration, and since intermediate forms or species with a mosaic of character states remained largely unknown, it is not surprising that Diakonoff (1973) described Molybdocrates for two dark species with bluish metallic dots including P. vinculata. However, the two groups share highly modified wing venation, a characteristic and unusual ‘anal lobe’ to the male hindwing, a male tibial pencil and similarly shaped palpi, and the distribution of genitalia characters does not support separation into two
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taxa. The distoventral strong bristles or spines from the uncus and the highly modified socii are the most obvious autapomorphies for Podognatha, and though the socii are reduced to mere hairtufts in P. vinculata, they are present in P. opulenta. The female genitalia, on the other hand, are most characteristically developed in P. vinculata and P. opulenta, with the complex structure of their ductus bursae and signum presumably representing generic autapomorphies lost in the simplified genitalia of two related unnamed species, one from Australia and one from New Guinea. Diakonoff (1966, 1973) provided comprehensive descriptions of both Podognatha and Molybdocrates, figuring head and wing venation for both taxa and genitalia of the male for the former and of both sexes for the latter (Diakonoff 1973). Theorica Diakonoff (Diakonoff 1966, 1973) is closely related to Podognatha with the two genera sharing a forewing of similar shape with scattered metallic dots and the same conspicuous modification of the male hindwing, with only minor venational differences (origin of chorda, stalked M3 and CuA1 in hindwing of Theorica). The genitalia of Theorica, though highly derived, corroborate a close relationship with Podognatha. The position of Podognatha within the Olethreutini will require further evaluation, but it is here tentatively included in the Zomaria-group. Diakonoff (1973) included Podognatha and Theorica in his subtribe Lobesiae and Molybdocrates in the subtribe Endotheniae, but both assignments are unconvincing. Several derived and unique characters of Podognatha and Theorica are found in members of the Zomaria-group of genera, and Podognatha and Zomariana share reduced tarsal bristles and a tuft of modified scales on the upper surface next to the anal margin, enfolded by the upturned anal margin in Podognatha. The position of the figures of Molybdocrates in Diakonoff’s (1973) revision among those of the subtribe Zomariae rather than the Lobesiae as indicated in the text may well suggest that he initially considered a close relationship between Zomaria and Molybdocrates. A similarly shaped forewing with a chorda originating between R2 and R3 links Zomaria frustulosa Diakonoff and Eubrochoneura Diakonoff with Podognatha (Diakonoff 1973), and the male of Z. frustulosa also has a distally projecting dorsal roll to the hindwing. Sterigma and signum of Z. frustulosa are very similar in structure to those of Podognatha opulenta and P. vinculata and provide the most convincing link between the two groups. The absence of any
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Figs 322–328. Podognatha, genitalia, hindwing and head. 322, P. vinculata, female genitalia, slide BM 24903. 323, 324, P. vinculata, aedeagus and male genitalia, Herberton, Q, slide BM 24902. 325, hindwing anal roll, P. sp. nr rebellis, male. 326, head, P. vinculata, male. 327, P. sp. nr rebellis, male genitalia, Mt Bellenden-Ker, Q, slide T2286. 328, P. sp. nr rebellis, female genitalia, Mt Bellenden-Ker, Q, slide T2291.
obvious apomorphy linking the highly derived male genitalia is not so surprising given their great diversity even within Podognatha, but structure of labial palpi and male hind tibia (with pencil present in Zomaria) as well as presence of glossy or metallic
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dots across the forewing all support tentative inclusion of Podognatha and Theorica in the Zomaria-group of genera. The description of Argyroploce rebellis Meyrick, 1931, together with the photo of the female genitalia
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of the single holotype (Diakonoff 1966), leave no doubt that the species belongs to Podognatha: rebellis (Meyrick, 1931), Exotic. Microlepid. 4: 140 (Argyroploce). [New Guinea: New Ireland.] Comb. nov. P. tamias and P. rebellis have both been described from New Ireland and could well turn out to be conspecific. The female genitalia ascribed to Thysanocrepis crossota by Diakonoff (1973) conform in every detail with the characteristic Podognatha structures as found in P. opulenta and P. vinculata and must belong to one of the pale grey Thysanocrepis lookalike species of Podognatha. This misidentification no doubt led to the inclusion of the superficially similar but totally unrelated Thysanocrepis in the subtribe Zomariae (Diakonoff 1973). Thysanocrepis is a member of the Enarmoniini and its female genitalia are figured for the first time in this volume.
Constituent species Four named and at least three unnamed species of Podognatha are known so far, of which one named and one undescribed species are from Australia: vinculata (Meyrick, 1916), Exot. Microlepid. 2: 21 (Argyroploce). [Qld: Herberton.] stilpnosticta (Turner, 1925), Trans. R. Soc. S. Aust. 49: 58 (Argyroploce). [Qld: Lamington NP.]
Neopotamia-group Status remarks Diakonoff (1973) established this group for 11 genera as the subtribe Neopotamiae, later adding Xenopotamia Diakonoff (1983c) from Madagascar. Kuznetzov and Stekolnikov (1984) accepted subtribal level for the group as Neopotamiina. Aarvik (2004) revised the African fauna, referring Eccopsis Zeller, Cosmorrhyncha Meyrick, Basigonia Diakonoff and six new genera to the Neopotamiae, which is here treated as a genus-group.
Evidence for monophyly Diakonoff (1973) defined the Neopotamia-group by possession of one of three types of signa, and by a tendency to develop a bristled process from the base of the valva. The three signa types identified by
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Diakonoff are here interpreted as derivations of an irregular patch of partly enlarged dentation, with the individual teeth increasingly larger the fewer there are, culminating in a single greatly enlarged and modified tooth of hourglass- to sickle-shape. The sterigma is a complex structure, and there is usually a process from the inner surface of the basal part of the valva (labis in Diakonoff 1973). A basal process from the inner surface of the valva is assumed to have arisen several times within the Olethreutini (also in the Zomaria-group, Sycacantha-group, Statherotis-group, and Olethreutes-group), varying from small finger-like knobs to large lobes, but the structure is particularly well developed in the Neopotamiagroup. The fixed, large cornuti found in Megalota, Geita Aarvik (2004) and some Eccopsis species may also be a derived feature.
Distribution The Neopotamia-group is well developed in the Oriental and Australian as well as the African (Diakonoff 1981, 1983c; Aarvik 2004) regions. Few taxa extend into the Palaearctic region and even fewer into North America.
Biology Species of the Neopotamia-group are leafrollers on several plant families. Megalota uncimacula (Turner) has been reared from Acacia obtusifolia (Mimosaceae) in Australia, and the African Eccopsis wahlbergiana Zeller has been repeatedly reported from Ricinus (Euphorbiaceae) (Aarvik 2004). According to Diakonoff (1973: 322), Temnolopha mosaica Lower is a ‘rather polyphagous species’, reared from Drypetes (Euphorbiaceae) in Australia.
Remarks The signa of Diakonoffiana Koçak and Costosa Diakonoff are here interpreted as derived from a patch of irregular dentation such as found in Megalota Diakonoff, with one or few particular teeth becoming increasingly enlarged to form a basally hollow sclerite. The signum of Temnolopha mosaica would in such a scenario occupy an intermediate position with a single predominant sclerite and remnants of scobination at its base. Diakonoff (1973) placed his subtribes Bactrae and Endotheniae directly after the Neopotamiagroup, and all three share a prominent basal process from the inner surface of the valva and a
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signum that could easily be viewed as pertaining to the same transformation series. Such a position of the Bactrini, i.e. as a sister group or highly derived offshoot from the Neopotamia-group rather than a separate clade, is worth considering and will probably have to be investigated with molecular methods. The very wide distribution of the Bactrini at first look suggests an early derivation within the Olethreutini, but this is contradicted by their restriction to monocotyledons as hostplants, which could well provide an alternative explanation for their wide distribution. The Gatesclarkeana-group also could be construed as derived from the Neopotamia-group on the strength of the conspicuous basal valval process and the signa shape. Furthermore, Gatesclarkeana and related genera share with Temnolopha a pair of round depressions on S2 and reduced tarsal bristles. Again, molecular studies may well help resolve the relationship between these two groups.
Temnolopha Lower (Figs 42, 329–334) Temnolopha Lower, 1901, Trans. R. Soc. S. Aust. 25: 72. Type species: Temnolopha mosaica Lower, 1901, ibid. 25: 72, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 48: 63. [Australia.]
Diagnosis Temnolopha species are medium-sized Olethreutini usually with unmistakable, contrasting and strongly sinuate wing markings in beige, grey and brown (T. mosaica) and with long palpi with a conspicuously pointed apex. Temnolopha mosaica is, together with Gatesclarkeana Diakonoff, Zomariana Diakonoff and Ophiorrhabda Diakonoff, one of only a handful of olethreutines known to possess ‘dorsal pits’, a pair of deep round depressions laterally on the second tergite. The wing venation of Temnolopha is typical for an Olethreutini, but the small accessory cell with the chorda from between R2 and R3 and not from between R1 and R2 as in Phaecadophora Walsingham distinguishes the two genera. The male genitalia are characterised by a plesiomorphic tegumen with a clavate and ventrally bristled uncus, by large, flat, drooping socii and by often asymmetrical valvae of similar structure to those of Phaecadophora and Saliciphaga Kuznetzov. The
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single signum of Temnolopha, a patch of scobination with larger central teeth and one or two greatly enlarged shark-tooth-shaped sclerites, is the diagnostic feature separating the genus from both Phaecadophora and Saliciphaga with their paired, hollow, crescentic signa without surrounding scobination.
Description Adults (Figs 329, 330) Wingspan. 12–18(24) mm; medium-sized to rarely large. Head (Figs 332). Ocellus well developed. Frons in uppermost part with two medially separate, anteriorly projecting small tufts of scales, one below each antennal base; remainder of frons with very short, appressed, dorsally oriented scaling. Proboscis short, 1–1.5× length of labial palpus. Labial palpus moderate to long (2.0–2.2× diameter of eye), porrect and strongly sinuate, rather slender and tapering to pointed apex; second segment sinuate and distally widened, but scales appressed and rather short except for some moderately loose scales along distal half of dorsal margin; terminal segment long and exposed, tapering to pointed apex. Antenna reaching to just over middle of forewing; anterior surface unscaled; cilia in male small, shorter than half diameter of flagellar segment. Thorax. With posterior crest. Legs hardly modified in male: hind tibia with somewhat longer and denser scaling than in female. Wings (Figs 42, 329–330). Forewing index 0.42–0.45; forewing elongate subrectangular, widest at 36: wing length, with lightly and evenly curved costa and nearly vertical, straight termen; discal cell 0.63× length of forewing, with small subtriangular accessory cell with chorda from halfway between R2 and R3 to R5, M-stem to between M2 and M3; all veins present and separate beyond discal cell; basal loop 0.33× length of 1A+2A; wing pattern of at least two entirely different types within genus, T. mosaica with numerous small, strongly sinuate, contrastingly coloured and well-defined markings of beige, glossy silvery grey and brown throughout wing, including an oblique-erect beige triangle over tornus and a continuous, sinuous beige band below costa from base to termen. Hindwing with well-developed cubital pecten and axillary tuft, with anal area modified in male of T. mosaica, covered with scattered very long hair-scales and with anal margin thickened and narrowly rolled over upper surface, with marginal scales on tornus enlarged, forming a dark pencil extending from dorsal roll;
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Rs and M1 closely approximated and parallel in basal part; M2 closely approximated to base of connate M3 and CuA1; 3A present; anal margin a well-sclerotised rim; dark grey, brown or purpleblack, paler towards lightly scaled and partly translucent base. Pregenital abdomen. Both sexes with two deep round dorsal depressions laterally towards base of T2, in well-preserved specimens covered by fan-like projecting scales from T1, and with a more weakly developed pair at base of T3. Sternum 8 in male lightly sclerotised, hindmargin with a median semicircular indentation. Male genitalia (Figs 331, 333). Vinculum narrow, with a small triangular saccus. Tegumen roughly triangular, with conspicuous projecting ‘shoulders’. Uncus clavate with a narrow, bent neck and with a brush of setae below apex. Socii large flat drooping lobes, densely covered with scales and long thin hair-scales. Gnathos a weakly sclerotised, ill-defined band, spinulose in T. mosaica. Valva large and wide, usually curved or bent and asymmetrical in some species (T. mosaica), with a deeply concave area separating the ill-defined sacculus and the densely bristled cucullus; with small costal process and narrow basal excavation with a sharply protruding rim and a large spinecluster above it below costa, sometimes produced into a club-shaped process crowned with cluster of spines; in T. mosaica with two asymmetrical tufts of bristles from right valva: a large tuft from sacculus and a thin tuft of very long curved bristles from ventral edge of base of cucullus, from beyond sharp ventral margin of concave subbasal portion of valva. Caulis short; anellus a wide ring with protruding laterobasal angles. Aedeagus short, sharply bent and tapering towards apex, with an asymmetrical, jagged, horizontal blade along apical half of right side; vesica without trace of cornuti. Female genitalia (Fig. 334). Ovipositor lobes subtriangular. Sternum 7 posteriorly narrowed and caudal margin deeply semicircular; hindmargin of T7 bilobed. Sterigma large and strongly sclerotised, with membrane laterally to sterigma proper also lightly sclerotised; sterigma in T. mosaica a dorsally split barrel-shaped structure continuous with the strongly sclerotised tube-shaped posterior half of the ductus bursae, with the two dorsocaudal angles (part of lamella postvaginalis) swollen into rounded and spinulose knobs. Ductus bursae strongly sclerotised in its caudal half and fused with sterigma, with ductus seminalis from short, narrow membranous central part and with ill-defined sclerotisation or a large well-defined plate (T. mosaica) in wall of basal third of ductus, which widens towards bursa. Corpus
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bursae ovate to teardrop-shaped, with only one signum consisting of a patch of scobination with larger central teeth and one or two greatly enlarged flat and erect sclerites.
Distribution South Africa, Sri Lanka, Thailand, Cambodia, Philippine Is, Japan (Ogasaware Is = Bonin Is), Indonesia, New Guinea, Australia. In Australia Temnolopha is found only in the northernmost coastal areas of the Northern Territory and in Queensland as far south as Yeppoon.
Biology Diakonoff (1973) stated that T. mosaica is a ‘rather polyphagous species’ and two collections he mentioned from central Java are ‘bred from leaves of ‘poetranjifa’ (?)’ and ‘of ‘lawas’ (? Alpinia: Zingiberaceae)’. In Australia, the species has been reared from larvae defoliating a species of Drypetes (Euphorbiaceae), possibly Drypetes lasiogyna, and nearly all localities are in areas of monsoon forest (vine thickets) rather than rainforest proper.
Remarks Diakonoff ([1968]) originally synonymised Temnolopha with Phaecadophora but later reversed his opinion, reinstating Temnolopha as a separate genus (Diakonoff 1973). Valid arguments can be provided for both decisions, and until all available species are re-examined for the characters separating the two type species it seems sensible to conserve the present nomenclatural status. The close relationship between Temnolopha, Phaecadophora, Saliciphaga and the Malagasy Cosmopoda Diakonoff (Diakonoff 1981) is indisputable, and an assessment of only their male genitalia would certainly not warrant generic rank for the four groups. However, much of this similarity, particularly in the tegumen, is due to symplesiomorphy. Apomorphies in the female genitalia suggest that Temnolopha is the sister group of Phaecadophora + Saliciphaga. Cosmopoda is probably a synonym of Temnolopha, but any formal change should deal with the entire group. Temnolopha has only a single, but rather plesiomorphic signum whereas Phaecadophora and Saliciphaga have retained two signa, but of a more derived form. Numerous structural differences between the two type species T. mosaica and P. fimbriata may well indicate further apomorphies for these two genera, but their consistency requires
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verification. Temnolopha mosaica has apically conspicuously pointed labial palpi, a feature Diakonoff (1973) mentioned in his description, though the relevant figures (Diakonoff 1973; heads in Figs G12, G13) have apparently been mixed up. A
much shorter accessory cell in the forewing and the dorsal abdominal pits further characterise T. mosaica. The male of P. fimbriata, on the other hand, has a row of long modified scales arising from between CuA1 and CuA2 on the hindwing
Figs 329–334. Temnolopha mosaica, Wessel Is, NT. 329, 330, adults. 329, . 330, . 331, male genitalia, Wessel Is, NT, slide T2420. 332, head, male. 333, aedeagus, slide T2420. 334, female genitalia, Wessel Is, NT, slide T2421. Scale = 2 mm.
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and large bundles of appressed scaling ventrally on the abdominal segments 5–7. Though the type species T. mosaica lacks a basal valval process, such a structure is well developed in the otherwise closely similar T. sponditis (Meyrick) (Diakonoff 1973) and refers the genus to the vicinity of Neopotamia Diakonoff. The shapes of the sterigma and signum support such an association. The dorsally split barrel-shaped sterigma, with its enlarged dorsocaudal angles lateral to dorsal split, is of similar structure to that of Megalota Diakonoff. Whereas the male genitalia of Temnolopha are among the least derived in the Temnolopha-group its signum appears to represent a more intermediate condition between the patch of equally-sized small erect teeth in Megalota and the regular, well-defined sclerite without basal scobination found in Phaecadophora and Neopotamia Diakonoff. Diakonoff’s (1973) remark that Temnolopha is most closely allied with Aterpia Guenée is unconvincing, especially as he then himself assigned the two genera to two different subtribes. Clarke (1958: plate 298, figs 1–1a) figured genitalia and adult of the lectotype of T. clydonias (Meyrick) from Sri Lanka, synonymised with T. mosaica by Meyrick (1909).
Constituent species Three species are currently included in Temnolopha (Diakonoff 1973), one of which was described from Australia and is present from Sri Lanka to the Philippines and Australia: mosaica Lower, 1901, Trans. R. Soc. S. Aust. 25: 72 (Temnolopha). [Qld: Cooktown, Townsville, Mackay.] clydonias (Meyrick, 1907b), J. Bombay Nat. Hist. Soc. 17: 734 (Cydia). [Sri Lanka.]
Diakonoffiana Koçak (Figs 43, 335–347) Diakonoffiana Koçak, 1981, Priamus 1: 118, established as an objective replacement name for Nyctidea Diakonoff, 1973. Nyctidea Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 295 (key), 324. Type species: Nyctidea cyanitis Diakonoff, 1973, ibid. 1: 325 (key), 327, by original designation. [Moluccan
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Is.]. A junior homonym of Nyctidea Reuter, 1903 – Insecta, Hemiptera.
Diagnosis Diakonoffiana species are small to medium-sized, usually dark-coloured olethreutine moths with a forewing with nearly rectangular apex and broadly rounded and projecting tornus, often with tufts of raised scales across forewing and a conspicuously copper-coloured wing base in female. Males of Diakonoffiana can superficially be recognised by their massively enlarged hind tibia with a large pencil of scales in a smooth glossy groove along their inner surface, in combination with long scale fringes to the first hind tarsal segment and a loose scale-tuft from its inner surface. Diakonoffiana furthermore has an apparently unique male secondary sexual structure: long hairpencils from the base of the anal margin that are inserted in ventral grooves at the base of the abdomen. The wing venation is typical for a member of the Olethreutini except for the short-stalked M3 and CuA1 in the hindwing that is, however, found in several groups throughout the tribe. The male genitalia are characterised by a single uncus, usually a long and often clavate process with a brush of bristles below the apex, by socii bristles arranged in a vertical row either along the margin of the erect-ovate socii lobes or on a vertical swelling (socii absent in D. tricolorana), and by an at least distally broad valva with a basal bristled process and a usually cone-shaped invagination below the costa at the base of the outer surface. The juxta is large in contrast to its generally small size in other genera of the Temnolopha-group. The complex sterigma of Diakonoffiana is aciculate and of basically two forms: a vertically folded median complex (D. cyanitis) usually derived from the lamella postvaginalis and often with two lateral aciculate knobs (D. vindemians (Meyrick)), or a Ushaped protruding rim from the lamella antevaginalis (D. tricolorana (Meyrick)). The laterally constricted plate-shaped signa with a hourglass-like outline found in most species of Diakonoffiana are very characteristic, though those of the type species are subquadrate and apically bilobed, and the often smaller second signum is crescentic in several species. The signa and sterigma of Apsidophora Diakonoff are extremely similar to those of Diakonoffiana, but the subtriangular forewing of the former, with its oblique termen, provides an obvious contrast to the projecting tornus of Diakonoffiana.
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Description Adults (Figs 335–339) Wingspan. 10–18 mm; small to medium-sized. Head (Figs 343). Ocellus well developed. Frons in upper part with a transverse band of moderate to very long, anteriorly projecting scales, remainder with short, upwardly oriented scales. Proboscis short, about 1.5× length of labial palpus. Labial palpus moderately long (1.7–2.0× diameter of eye), porrect to ascending, strongly sinuate; either with an apically strongly expanded subtriangular second segment and a relatively short partly concealed terminal segment (D. tricolorana) or with a sickleshaped second segment, widest in middle, and a relatively long terminal segment (D. vindemians). Antenna reaching to middle of forewing; anterior surface unscaled; cilia in male small. Thorax. With posterior crest. Male hind legs conspicuously modified: tibia greatly enlarged to an ovate shape by thick padding of closely appressed scales, with a deep smooth and shiny cavity along its inner surface filled by a large pencil of greyish (D. vindemians) or yellow (D. tricolorana) modified long scales from base of tibia (Figs 341, 340); first tarsal segment usually with long fringes of scaling and a tuft of loose scales on inner surface. Wings. (Figs 43, 335–339). Forewing index 0.42–0.45; forewing subrectangular, slightly dilated towards termen, costa slightly and evenly curved, more so near apex, termen vertical and faintly concave below apex, tornus projecting and broadly rounded; in some species with several tufts of raised scales scattered across forewing and a particularly prominent outwardly oblique band of raised scales just beyond middle of wing behind discal cell; discall cell 0.65× length of forewing, with accessory cell (absent in D. cyanitis) small and triangular with chorda to between R5 and M1, with M-stem to between M2 and M3; all veins present and separate beyond discal cell; anal loop 0.31× length of 1A+2A; wing pattern diverse, very variable in some species (D. tricolorana) and overlaid by snow-white suffusion. Hindwing with well-developed cubital pecten and axillary tuft; males with a series of extremely long hair-scales from base of anal margin that form a narrow pencil fitting into grooves at base of abdomen, and in some species (D. tricolorana) with underside of anal area thickly clothed with elongate scales and with an enlarged fringe; Rs and M1 closely approximated and parallel in basal part; M2 closely approximated at base to short-stalked M3 and CuA1; 3A well developed; greyish, sometimes grey with dense
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white fringe in anal area in male and with a conspicuous coppery ochreous wing base and fringe in the corresponding female (D. tricolorana). Pregenital abdomen. Modified in male, with patch of modified scales laterally at base of abdomen ventral to second spiracle and with two ventral grooves containing hairpencils from base of anal margin, running from lateral abdominal base towards ventral midline on S4, distally consisting of two inwardly curved crescentic folds across S3 lined with short spines or small capitate pin-like structures with enlarged sockets. Sternum 8 in male large, hindmargin with a deep, semicircular ventral excision and a band of dense scaling along edge. Male genitalia (Figs 342, 344, 347). Vinculum a moderately wide band with a small triangular, hardly projecting saccus. Tegumen from triangular and rather frail (D. tricolorana) to more robust with prominent shoulders (D. cyanitis, D. vindemians), attachment points of muscle m4 from small triangles (D. tricolorana) to large broad lobes. Uncus usually a curved and often clavate process with a brush of bristles below apex, rarely a reduced rounded prominence covered with modified scales (D. saloris Diakonoff). Socii from vertical, semiovate porrected lobes with bristles along edge to vertical rows of bristles (D. vindemians) to absent in D. tricolorana. Gnathos two more or less sclerotised lateral arms, medially fused with prominent subscaphium of often inverted-triangular shape, with two long, sharply pointed paired arms along basal margin in D. vindemians. Valva from curved elongate ovate to distinctly club-shaped in outline with dilated cucullus (D. spumans Diakonoff), with costal process often concealed behind a prominent cone-shaped invagination of outer valva surface, with a variously developed process from inner surface of valva just beyond basal excavation ranging from a small bristled hump (D. cyanitis) to a single or composite clavate or digitate bristled process; with sacculus area indistinct or with dense ventral brush of bristles or with a blade-like projecting triangular process (D. triangulana, D. vindemians) set with spines in D. saloris; base of cucullus with asymmetrically arranged pairs of pegs in D. tricolorana, close to ventral margin on right-hand valva and on inwardly directed triangular process in centre of valva on the other side. Caulis short and anellus a narrow membranous ring; juxta large and shield-shaped. Aedeagus rather small and often triangular in dorsoventral as well as lateral aspect, with bulbous base and tapering to pointed apex; vesica without any trace of cornuti.
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167
Figs 335–341. Diakonoffiana, adults and hind legs. 335–338, D. tricolorana, adults. 335, , Toowoomba, Q. 336, , Coraki, NSW. 337, , Toowoomba, Q. 338, , Rockhampton, Q [image reversed]. 339, D. vindemians, , Ravenshoe, Q. 340, 341, modified male hind legs with tibial pencils (arrows). 340, D. tricolorana. 341, D. vindemians. Scale = 2 mm.
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Female genitalia (Figs 345, 346). Ovipositor lobes elongate-ovate. Sternum 7 with faintly sinuate to deeply V-shaped hindmargin. Sterigma a variously shaped complex spinulose structure derived from either lamella antevaginalis (D. tricolorana) or postvaginalis (D. vindemians), often a vertically folded median structure above ostium associated with two dome-shaped lateral knobs, or lamella antevaginalis a projecting U-shaped spinulose rim with its distal corners extended laterally in an angled ridge (D. tricolorana). Ductus bursae with an often long, colliculum-like sclerite below ostium, inception of ductus seminalis from a blindly ending lateral expansion of ductus bursae at entrance into corpus bursae. Corpus bursae teardrop- to pear-shaped, with two often unequally sized plate-shaped signa of usually hourglass-like outline, more rarely crescentic or apically bilobed.
Distribution Sulawesi (unidentified species), Moluccan Is, New Guinea, Australia, Micronesia, with the highest diversity in New Guinea. In Australia the genus is found along the eastern coast from northern Queensland to Kiama, New South Wales, with one single record from the Northern Territory.
Biology The only host record for the genus is from Brisbane where I. F. B. Common reared several specimens each from a ‘larva joining leaves of unidentified tree’. The locality data for D. tricolorana suggest that at least some species of Diakonoffiana may exist in marginal rainforest habitats.
Remarks Diakonoffiana vindemians and a closely related species from Sulawesi provide the link between the type species of Diakonoffiana (Diakonoff 1973; as Nyctidea) and a somewhat derived but congeneric group containing D. tricolorana, included by Diakonoff (1973) in Metendothenia Diakonoff. The following species is consequently transferred to Diakonoffiana: spumans (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 452 (Metendothenia) [Papua New Guinea]. Comb. nov. The genitalia of both sexes also refer the following species from Micronesia (Eastern Caroline Is) to Diakonoffiana, as a close relative of D. vindemians:
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cyanosticha (Clarke, 1976), Insects Micronesia 9: 55 (Cymolomia) [Micronesia]. Comb. nov. Though the male genitalia of Diakonoffiana appear superficially rather diverse they are all linked by several synapomorphies: vertically arranged socii bristles, shape of invagination at base of outer surface of valva, broad cucullus and large juxta. The most convincing autapomorphies for Diakonoffiana are, however, found in the secondary sexual structures of the male. The combination of a pencil from the base of the anal margin of the hindwing inserted into apparently glandular grooves at the base of the abdomen occurs elsewhere only in Oxysemaphora in a quite different configuration. The nature of the hind tibia modification of Diakonoffiana is also quite distinct, as well as its enlarged basal hind tarsal segment. Metendothenia as conceived by Diakonoff (1973) is a conglomerate of at least three different groups, including one taxon allied to Statherotis Meyrick. Metendothenia s. str., i.e. the species congeneric with the type species M. emmilta Diakonoff, share the presence of a basal process from the inner valva surface with Diakonoffiana. Both these genera belong to the Temnolopha-group, a relationship confirmed by the shape of their signa and sterigmata. Metendothenia s.str. is characterised by a different but not less unusual set of secondary male sexual modifications. The hindlegs are unmodified but two large and very long pencils of hairscales from the thorax below the tegulae extend backwards under the wings with their tips appressed to modified scaling across the lower surface of the anal area of the hindwing. The shape of the forewing and labial palpus and the structure of the single signum in M. emmilta further differentiate Metendothenia s.str. from Diakonoffiana. Metendothenia fulvoflua Diakonoff from Madagascar (Diakonoff 1983c) has male genitalia that are very reminescent of those of Diakonoffiana, but the female genitalia are sufficiently different to advise against a recombination withouth a more thorough study. Diakonoff (1973) provided extensive descriptions of the species included in Diakonoffiana and figured their genitalia as well as the wing venation and head of the generic type. No reference was made to the abdominal modification in the male, but the pencil from the base of the hindwing dorsum was mentioned. Obvious similarity of the female genitalia points to a close relationship between Diakonoffiana and the monotypic Apsidophora Diakonoff (Diakonoff 1973). The males of both genera share a modified hind tibia but Apsidophora has strongly modified
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hind wing venation and overall more plesiomorphic male genitalia. The wing shape and genitalia of Neopotamia glyceranthes (Meyrick) (Clarke 1958: pl. 254, fig. 4, as Olethreutes glyceranthes (Meyrick)) strongly suggest a species of Diakonoffiana. Sterigma and
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signa of the unique female from the Andamans are not compatible with the species’ present assignment to Neopotamia (Diakonoff 1973). It seems sensible, however, to refrain from any new generic combination until the male becomes known.
Figs 342–347. Diakonoffiana, genitalia and head. 342–345, D. tricolorana. 342, male genitalia, Toowoomba, Q, slide T2296. 343, head, male. 344, aedeagus, slide T2296. 345, female genitalia, Maclean, NSW, slide T2280. 346, 347, D. vindemians. 346, female genitalia, Ravenshoe, Q, slide T2463. 347, male genitalia, Kuranda, Q, slide BM 24897.
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The numerous synonyms of D. tricolorana are an apt illustration of the variability of this species. It is variously mottled dark brownish grey to blackish, frequently with a snow-white spot on the costa, curved white bands across the wing or even with the distal two-thirds of the wing white (Figs 335, 338, 336). A distinct form with a red-brown costal triangle and the rest of the wing lightly powdered with white except for the base of the dorsum is found throughout the range of D. tricolorana (Fig. 337). The sexually dimorphic coloration of the hindwing – greyish with a whitish dorsal fringe in the male and with a conspicuous coppery-orange wing base and fringe in the female – further complicates the appearance of D. tricolorana. Given the strong Australian influence in the fauna and flora of the Port Moresby region it is not surprising that D. metacycla from Port Moresby is a synonym of D. tricolorana. The single holotype of D. tricolorana, with a yellow base to its hindwings, is a female though Meyrick (1881) mistook it for a male. Turner (1926) erroneously reports a costal fold for Eucosma leucatma Turner.
Constituent species Eight named species are known worldwide, two of which are from Australia: tricolorana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 638 (Helictophanes). [NSW: Sydney.] fungiferana (Meyrick, 1881), Proc. Linn, Soc. NSW 6: 640 (Helictophanes). [NSW: Parramatta.] intricata (Turner, 1916), Trans. R. Soc. S. Aust. 40: 532 (Argyroploce). [Qld: Brisbane.] leucatma (Turner, 1926), Trans. R. Soc. S. Aust. 50: 138 (Eucosma). [Qld: Yeppoon.] lychnospila (Turner, 1946), Trans. R. Soc. S. Aust. 70: 218 (Argyroploce). [Qld: Noosa.] metacycla (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 453 (Metendothenia). [Papua New Guinea.] vindemians (Meyrick, 1921), Exot. Microlepid. 2: 449 (Argyroploce). [Qld: Kuranda.]
Megalota Diakonoff (Figs 44, 348–356) Megalota Diakonoff, 1966, Zool. Verh. (Leiden) 85: 52. Type species: Polychrosis fallax Meyrick,
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Figs 348, 349. Megalota, adult, males. 348, M. helicana, nr Rockhampton, Q [image reversed]. 349, M. uncimacula, Lamington Nat. Park, Q [image reversed]. Scale = 2 mm.
1909, J. Bombay Nat. Hist. Soc. 19: 587, by original designation. [Assam.]
Diagnosis Megalota species are medium-sized brownish Olethreutini usually with an intricate wing-pattern characteristic for the genus: a large dark triangle on costa with its point drawn out and recurved towards termen and an inwardly oblique, commashaped mark in terminal area (absent in M. helicana). A dark transverse fascia including costal triangle and running to distal part of termen is developed in several species and may mask some markings. The labial palpi are rather stout and of only moderate length, and the male hind tibia in some species is enlarged by long dense scales
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Figs 350–356. Megalota, genitalia and head. 350, 351, M. helicana, male genitalia and aedeagus, Lawgi, Q, slide T2265. 352, head, M. uncimacula, male. 353, M. helicana, female genitalia, Lawgi, Q, slide T2432. 354, M. uncimacula, female genitalia, nr Gympie, Q, slide T2441. 355, 356, M. uncimacula, male genitalia and aedeagus, Lamington Nat. Park, Q, slide T2264.
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concealing a tibial pencil. The male genitalia with their large hairy uncus of two separate square or curved-triangular flat lobes and the narrow valva with a very large cone- to club-shaped basal valval process are unique and diagnostic for Megalota. A similarly intricate female sterigma that is integrated with a strongly sclerotised distal part of the ductus bursae is found in related genera such as Temnolopha Lower, Diakonoffiana Koçak and Costosa Diakonoff, but Megalota lacks a sclerotised patch in lower part of ductus bursae (present in Temnolopha mosaica Lower), and all Australian species have conspicuous invaginated triangular or horn-shaped processes in the lamella postvaginalis. Megalota has only a single signum of at least two, but usually numerous small erect shark-toothshaped sclerites.
Description Adults (Figs 348, 349) Wingspan. 11–17 mm; medium-sized. Head (Figs 352). Ocellus well developed. Frons in upper part with a transverse band of moderately long projecting scales. Proboscis short, hardly longer than labial palpus. Labial palpus moderately long (1.7–2× diameter of eye), porrect and sinuate; second segment strongly widened in distal half, with long loose scales along ventral margin and distal part of dorsal edge; terminal segment fairly long and thin. Antenna reaching to about middle of forewing; anterior surface unscaled; cilia in male minute. Thorax. With strong posterior crest. Legs with modified male hind tibia in some species: dorsal margin and inner surface with long, semi-appressed scales concealing a long hair-pencil from base of tibia. Wings (Figs 44, 348, 349). Forewing index 0.43–0.48; forewing subtriangular to subrectangular, lightly to moderately dilated towards termen; costa evenly curved; termen from straight and slightly oblique to barely sinuate and vertical; discal cell 0.62× length of wing, accessory cell small and subtriangular with chorda from halfway between R2 and R3 to R5, M-stem to between M2 and M3; all veins present and separate beyond discal cell; anal loop 0.31× length of 1A+2A; usually with complicated greyish ochreous and brownish wing markings of sinuate pattern elements, with a dark brown inverted triangle over central third of costa with its produced and outwardly curved apex in centre of wing, and usually with an oblique comma-shaped dark mark from middle of termen (absent in M. helicana
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(Meyrick)). Hindwing with well-developed cubital pecten and axillary tuft, with dense fringe of very long loose scales along anal margin in males of some species (probably correlated with modified hind tibia); Rs and M1 closely approximated at base to connate; M2, M3 and CuA1 close and equidistant at base; 3A well developed; species with enlarged and modified anal area (M. uncimacula (Turner) and others) with a wide, strongly sclerotised, crescentic rim along anal margin; dark greyish brown, Australian species not with paler wing base. Pregenital abdomen. Unmodified. Sternum 8 in male lightly sclerotised, hindmargin faintly bilobed. Male genitalia (Figs 350, 351, 355, 356). Vinculum a narrow band. Tegumen large, high and parallel-sided to widening towards uncus. Uncus two very large square or curved-triangular membranous lobes densely covered with long thick downward pointed setae on ventral surface. Socii either small hairy knobs lateral to base of anal tube, a small cluster of setae in membrane below uncus, or entirely absent. Gnathos a faint band of weak sclerotisation continuous with lightly sclerotised subscaphium. Valva small and very narrow with a very large cone- to club-shaped process from inner surface of valva beyond basal excavation or from base of costa; cucullus very long and thin and often with a ventrally strongly projecting triangular tooth at base (M. helicana, M. uncimacula), sacculus with a tuft of bristles often adjacent to a second patch of bristles at base of cucullus, basal valval process crowned by few large apical spines and covered with long hair on outer surface. Caulis long and curved and anellus a membranous ring. Aedeagus bent or curved and more or less parallel-sided, some nonAustralian species with carina and large external thorns; vesica usually with few large non-deciduous thorns. Female genitalia (Figs 353, 354). Ovipositor lobes roughly triangular. Sternum 7 strongly bilobed in the more derived species (M. helicana, M. uncimacula), with deep V-shaped median excavation in hindmargin. Sterigma a complex structure basically consisting of a wide, strongly sclerotised band ventrally around distal part of ductus bursae, continuing dorsally of ostium as lamella antevaginalis with two caudally invaginated triangular corners or short smooth horns surrounded by spinulose membrane. Ductus bursae membranous except for distal part integrated with sterigma; area between sterigma and inception of ductus seminalis below midlength in some species (M. helicana) thick-walled and set with sparse short
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stout spinules. Corpus bursae ovate, with a single signum consisting of a group of small erect sharktooth-shaped sclerites of various size and shape.
Distribution Africa, Madagascar, Sri Lanka, India, Indonesia, New Guinea, Australia. According to J. W. Brown (personal communication 2005) the genus is present in the New World from Mexico south to Argentina. In Australia the genus is restricted to coastal areas of New South Wales north of Wollongong, of southern Queensland, the Northern Territory and the Kimberley of Western Australia. Only a single specimen from Cape York Peninsula is known from northern Queensland.
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Megalota – with at least two and usually numerous small teeth grouped together – is clearly plesiomorphic. Those species with one or two teeth greatly enlarged at the cost of the rest (M. anceps (Meyrick)), M. fallax (Meyrick)) presumably represent an intermediate condition on the way to a single sclerite as in Temnolopha and Costosa. Megalota vera Diakonoff, described from New Guinea (Diakonoff 1966), is very similar to M. uncimacula and obviously closely related, but given the absence of M. uncimacula from northern Queensland the small genitalic differences justify specific status for both taxa. The specimens from Toowong near Brisbane, identified as M. vera by Diakonoff (1973), belong to M. uncimacula.
Constituent species Biology Megalota uncimacula has been reared in New South Wales from Acacia obtusifolia (Mimosaceae) at Mt Keira, Wollongong, by V. J. Robinson, and by J. Stockard from Croton verreauxii (Euphorbiaceae) at Wingham. The northern M. helicana is found in more drier or seasonal rainforest areas whereas M. uncimacula is a species of the subtropical rainforest.
Remarks The original description of the genus (Diakonoff 1966) and its comprehensive treatment in the context of the south Asiatic Olethreutini (Diakonoff 1973) provided an adequate concept of Megalota. Clarke (1958) figured the types of two species, and Diakonoff (1981) described and figured an additional species from Madagascar. Aarvik (2004) documented four African species of Megalota. Megalota is a well-defined genus and its monophyly is supported by uniquely derived genitalia with a large, bilobed, ventrally hairy uncus, a slender cone-to club-shaped process from inner valva base, few large and non-deciduous cornuti, a complex sterigma with two small posterior invaginated corners and a single signum with several enlarged teeth. The basal valval process and the structure of sterigma and signum identify Megalota as a member of the Neopotamia-group. Megalota and Costosa share a bilobed uncus and a long, slender basal valval process and could well be sister groups. Presence of only one signum is undoubtedly an apomorphy with regard to the paired signa of Costosa, but its structure in
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Twelve species are described worldwide, two of which are from Australia. The identity of two further species from the Northern Territory and northern Western Australia respectively remain to be established. The named Australian species are: helicana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 645 (Penthina). [NSW: Bulli Pass and Richmond River; Qld: Rosewood.] uncimacula (Turner, 1925), Trans. R. Soc. S. Aust. 49: 58 (Argyroploce). [Qld: Brisbane.] vera sensu Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 316 (Megalota).
Costosa Diakonoff (Figs 45, 357–361) Costosa Diakonoff, [1968], US Nat. Mus. Bull. 257: 40 (key), 80. Type species: Costosa allochroma Diakonoff, [1968], ibid. 257: 81, by monotypy and original designation. [Philippine Is.] Rhodonympha Diakonoff, 1968, Beaufortia 15: 71. Type species: Eucosma rhodantha Meyrick, 1907c, J. Bombay Nat. Hist. Soc. 18: 138, by monotypy and original designation. [Assam.]
Diagnosis Costosa species are conspicuously coloured, medium-sized to large moths with ill-defined cloudy forewing pattern of reddish to red-brown and ochreous, usually with a pale whitish ochreous apex and three groups of long, leaden strigulae with a bluish sheen along middle part of costa, the first
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group beyond 56: costa continued as an indistinct fascia of scattered leaden scales obliquely across wing. The male of Costosa has at the base of the hindwing anal margin a small projecting semicircular naked lobe folded over upper surface of wing and a hind tibia with a patch of very long and dense scaling on lower surface that is displayed as a spreading brush along the length of the tibia together with a large loose tuft from its base. The male genitalia are unique with two long naked rodlike processes arising from lateral corners of a bristled, medially fused pad-like structure across apex of tegumen (probably incorporating uncus and socii) and with a long erect slender process from a vertical bristled ridge across base of valva. A modified, spinulose T8 and a spinulose lamella antevaginalis with a pair of projecting vertical folds along midline and a spinulose lateral rim, and two unequal-sized sickle-shaped signa characterise the female genitalia (Diakonoff 1973).
Description Adults (Fig. 357) Wingspan. 14–20 mm; medium-sized to large. Head (Fig. 359). Ocellus well developed. Frons in upper part with a horizontal, ledge-like tuft of projecting scales, remainder of frons with very short, appressed, upwardly directed scales. Proboscis short, 1–1.5× length of labial palpus. Labial palpus moderately long (1.8–2× diameter of eye), porrect, gently sinuate and moderately
Fig. 357. Costosa australis, holotype [image reversed]. Scale = 2 mm.
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widened beyond middle; second segment sinuate and gradually widened towards apex, with long, loose scales along distal third of dorsal margin; terminal segment thin and downward pointing, its point hardly protruding beyond long appressed scales from ventral margin of second segment. Antenna reaching to just over middle of forewing; anterior surface unscaled; cilia in male small, c. 56M diameter of flagellar segment. Thorax. With large posterior crest. Legs with male hind tibia greatly enlarged by a thick cover of long scales especially along inner surface, appressed at rest, with a tuft of particularly long scales from base; if displayed the scales along inner surface are spread to form a brush-like structure over entire tibia, with the longest scales along dorsal and ventral margins and in basal tuft. Wings (Figs 45, 357). Forewing index 0.42– 0.45; forewing elongate subrectangular, slightly widening towards termen, with moderately curved costa and slightly oblique termen; discal cell 0.64× length of wing, with M-stem as a partly tubular vein or vestigial only, running to base of M2, chorda present or absent; all veins present and separate beyond discal cell; basal loop 0.32× length of 1A+2A; with ill-defined cloudy wing pattern of often bright red or red-brown with ochreous, usually with a pale whitish ochreous apex and three groups of long leaden strigulae with a bluish sheen along middle part of costa, the first group beyond 56: costa sometimes continued as an indistinct fascia of scattered leaden scales. Hindwing with cubital pecten and axillary tuft, with conspicuously modified anal margin in male: basal part of anal margin a small, projecting, nearly semicircular lobe folded over upper surface of wing and exposing unscaled reverse side, outlined by a large scale tuft beneath from edge of fold; with Rs and M1 closely approximate and parallel at base, M2 bent towards connate to very short-stalked M3 and CuA1; 3A developed, its distal part covered by folded anal margin. Pregenital abdomen. Unmodifed. Sternum 8 in male with a shallow median notch in hind margin. Male genitalia (Figs 358, 360). Vinculum a narrow band. Tegumen broadly triangular, wellsclerotised, with large hook-shaped attachment points for muscles m4 from pedunculi. Uncus two long and slender rod-like naked processes arising from lateral corners of a heavily bristled shieldshaped structure with convex bilobed ventral margin, possibly incorporating the socii. Gnathos two weakly sclerotised lateral bands, sometimes spinulose. Valva curved, long and narrow, with a
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very long, slender, vertically rising basal valval process, and with costal process very small and inconspicuous; sacculus blade-shaped towards margin, ending in a more or less upcurved triangular point; basal valval process arising from a vertical bristled ridge across base of valva and either parallel-sided and densely spined along entire inner side or irregular and with bristled apex and a separate, subbasal tuft on inner surface or on a side-branch of process; cucullus unmodified, densely and evenly bristled. Aedeagus rather small, straight to slightly curved, without coecum and with bevelled apex; vesica with a group of small, slender, spine-shaped deciduous cornuti. Female genitalia (Fig. 361). Ovipositor lobes subtriangular. T8 swollen, intricately folded and spinulose, in C. australis, sp. nov. trilobed with sublateral swollen triangular processes on each side separated by a deep transverse pocket from the tongue-shaped medial lobe. Sternum 7 with lightly concave emarginate hindmargin. Sterigma large, finely spinulose, a conspicuous strongly projecting
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U-shaped rim around entrance to ostium with posterior ends bent into laterally extending, horizontal spinulose ridges. Ductus bursae with a long slender colliculum below ostium with inception of ductus seminalis at its end. Corpus bursae with two sickle-shaped to reniform signa of unequal size.
Distribution Sri Lanka, India (Assam), Philippine Is, New Guinea and Australia. The few Australian localities are all in northernmost Queensland, north of Cairns.
Biology No biological information is available, but the Australian localities are all within rainforest areas.
Remarks Costosa and Rhodonympha were both described in the same year, January 1968 and May 1968
Figs 358–361. Costosa australis, genitalia and head. 358, male genitalia, paratype, Mossman Gorge, Q, slide T2082. 359, head, male. 360, aedeagus, slide T2082. 361, female genitalia, paratype, Kuranda, Q, slide BM 24895.
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(Diakonoff [1968], 1968), with figures of genitalia, wing venation and head for both taxa, including a photo of the adult for Costosa allochroma. Diakonoff (1973) later synonymised Rhodonympha with Costosa and described a third species for the genus. Photos of the adult and male genitalia of C. rhodantha (as Olethreutes rhodantha) were given by Clarke (1958: pl. 270; figs 3, 3a). Costosa is a well-defined genus and its monophyly is demonstrated by several obvious synapomorphies such as the characteristic male hind tibia, the fused and hairy uncus/socii complex with a pair of finger-like processes, the very long and slender basal valval process, and, in the female, a spinulose T8 and two crescent- or kidney-shaped signa of unequal size. Valval structure and signa shape identify Costosa as a member of the Neopotamia-group of genera. Socioplana idicopoda Diakonoff from Sumatra (Diakonoff 1983a) also has a medially continuous pad-like process across the apex of the tegumen presumably derived from the socii and/or uncus. Its similarities with Costosa include external appearance and the curiously modified male hind tibia. The shape of the valva of Socioplana points to a close relationship to Meiligma Diakonoff (Diakonoff 1973), which also has a dorsally continuous uncus/socii structure but an unmodified male hind tibia. Afrocostosa Aarvik also is a close relative, as implied by the generic name (Aarvik 2004). The modification of the male hind tibia of Costosa consists of long loose scales along its entire inner surface, longer towards dorsal and ventral margin, with a particularly large tuft from base of tibia. In display all these scales are spread to form a brush along the entire tibia rather than an isolated hairpencil from its base only. In the Australian C. australis, sp. nov., T8 is intricately folded in the female with lateral grooves and a deep dorsal pocket. Considering the projecting uncus arms and basal valval processes of Costosa males it is tempting to speculate that some sort of linking mechanism for copulation could be involved. The structure of the uncus with its ‘beak’ and the position of the deep dorsal pouch between the ovipositor lobes and T8 in certain species of the related genus Neopotamia suggest a similar function.
Constituent species Three species are described worldwide, to which a fourth from Australia is here added: australis, sp. nov.
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Costosa australis, sp. nov. (Figs 357–361)
Material examined Holotype. : ‘16.28S 145.21E, Mossman Gorge, Qld [Queensland], 11 Sept 1977, I.F.B. Common’ (ANIC). Paratypes. Queensland: 1 , 12°43´S 143°18´E, 11 km ENE Mt Tozer, 11.vii.1986, J. C. Cardale; 1 , 16°04´S 145°28´E, Pilgrim Sands, 2 km NW by N Cape Tribulation, 22.ii.1998, E. D. E.; 1 , Mossman Gorge, 2. v.1974, N. R. Badans & V. J. Robinson (ANIC genitalia slide T2082); 2, Kuranda, ?.iv.1905 (BM genitalia slide 24895), 3.xi.1904, Dodd (BMNH); (ANIC, QM).
Diagnosis The combination of the deep red-brown forewing with a yellow ochreous wing base and apex together with an evenly dark blackish grey hindwing is diagnostic for C. australis, sp. nov., in contrast to the Papuan Costosa aphenia Diakonoff with deep bronze hindwings with a yellow-golden tinge.
Description Male (Figs 357, 359), female Wingspan 15.5–18.0 mm. Head orange ochreous, scale tips above pale grey with blue iridescence (unless denuded). Thorax and tegulae ochreous variably suffused with orange, brown and pale grey. Forewing ochreous at base and apex, gradually darkening to orange ochreous and deep red-brown in centre of wing and along costa, with a more or less well-defined deep red-brown triangle extending over at least central half of costa with apex nearly to dorsum; with variably developed olive suffusion along central part of dorsum and pale red-brown suffusion across dorsal half of terminal area; several closely approximated white costal strigulae at 56: costa giving rise to wide, outwardly oblique wavy band of scattered pale blue iridescent scales, fading towards dorsum; a pair of more oblique strigulae just beyond middle of costa also starting with a few white scales each and continuing as a thin short line of iridescent pale blue scales; with one or or two shorter, less oblique white strigulae along rest of costa; iridescent pale blue scales scattered also in small streak along termen adjoining tornus and throughout olive suffusion along dorsum, sometimes delineating dorsodistal margin of redbrown triangle as an outwarly oblique short band; fringe ochreous except for a deep red-brown portion from about middle of termen to halfway round
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tornus. Hindwing dark blackish grey throughout except for pale grey band between cell and costa, with a pale grey fringe; in male upturned small lobe of anal margin shiny silvery grey and fringed by long dark grey scales. Male genitalia (Figs 358, 360). Tegumen short, robust, with broad hooks as attachment for muscles m4. Shield-shaped structure carrying uncus arms with dense short bristles except for two small curved longer tufts from medially slightly pointed dorsal margin; uncus arms long, very slender throughout, parallel and straight except for slightly recurved apex. Gnathos weakly sclerotised laterally, medially membranous, spinulose. Valva long and slender with cucullus nearly parallel-sided except near apex, sacculus ending in a flattened, upcurved, narrowly triangular point, slender basal valval process rising from a spinulose vertical ridge, laterally flattened, strongly narrowed above and below a small median expansion with a patch of bristles on inner surface, and ending in narrow, paddle-shaped, distally truncate processes with long bristles on outer surface. Juxta and caulis fused into small plate with five projections. Aedeagus short, dorso-ventrally somewhat compressed, with patch of sockets indicating cornuti. Female genitalia (Fig. 361). T8 modified, intricately folded, swollen and spinulose, trilobed with sublateral swollen triangular processes on each side separated by a deep transverse pocket from tongue-shaped median lobe. Sternum 7 with shallow V-shaped hindmargin. Sterigma a strongly projecting, spinulose, U-shaped rim distally expanded on inner side and ending on each side in a first transverse then posteriorly bent narrow band of finer, longer spinules. Ductus bursae about half as long as corpus bursae, anteriorly slightly widening, with long, slender colliculum. Corpus bursae ovate, with two unequal signa, posterior one a long, curved, nearly parallel-sided blade appearing crescent-shaped, anterior one smaller, with narrow base, broadly reniform, laterally rounded.
Gatesclarkeana-group Status remarks Diakonoff (1973) established his subtribe Gatesclarkeanae for Gatesclarkeana Diakonoff and Asymmetrarcha Diakonoff. Kawabe (1978) acknowledged a close relationship between Gatesclarkeana and his Hiroshiinoueana Kawabe without reference to subtribal position, whilst
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Kuznetzov (1981) explicitly referred Ukamenia Oku (as Aphiaris Kuznetzov) to the Gatesclarkeanae. Kuznetzov and Stekolnikov (1984) raised the group to supertribe level (Gatesclarkeanidii), comprising the two tribes Gatesclarkeanini and Endothenini and positioned at the base of the Olethreutinae. Razowski (1989) only accepted subtribal rank within the Olethreutini for the group, as Gatesclarkeanina, and he formally included Hiroshiinoueana. Horak and Brown (1991) followed Kuznetzov and Stekolnikov in including Endothenia in the group and treating it as basal within the Olethreutinae, albeit only at tribal level. Based on Dang’s (1990) convincing discussion of the synapomorphies linking Bactra Stephens and Endothenia Stephens, the recognition of a small basal excavation in Gatesclarkeana as well as the plesiomorphic nature of horn-shaped signa in olethreutines, this assignment has subsequently been modified with Endothenia removed to the Bactrini, and the Microcorsini recognised as the basal-most tribe in the subfamily (Horak 1998). Returning to Diakonoff’s (1973) concept, the Gatesclarkeana-group is here once again treated as subordinate within the Olethreutini, albeit only as a genus-group, until more light can be shed on its relationships with the Neopotamia-group and the Bactrini.
Evidence for monophyly Diakonoff (1973) defined his Gatesclarkeanae by signa ‘of scobinate-cornute type’ and by complex scent organs of two different types. The monophyly of the group is supported by several apomorphies: 1, signa scobinate patches with one or several enlarged teeth, usually pointing outwards in an at least partially circular arrangement, often resulting in a star-shape (Hiroshiinoueana Kawabe, Gatesclarkeana confracta Diakonoff); 2, sterigma a large, complex, deeply invaginated structure; 3, spiny process from sacculus region; 4, forewing with conspicuous metallic spots.
Distribution The Gatesclarkeana-group is restricted to the Oriental and Australian regions and the eastern Palaearctic
Biology Species of the Gatesclarkeana-group are leaf rollers on a wide range of plant families, often
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polyphageous, but have been found feeding as borers in mango shoots (Fletcher [1921]). Ukamenia, a polyphageous leaf roller, has been found feeding in galls of the chestnut gall wasp (Oku 1981).
Remarks The Gatesclarkeana-group has had a very chequered history, depending on which characters were considered phylogenetically most relevant. Diakonoff (1973) included the group within the Olethreutini based on signa shape and the nature of their secondary sexual structures. Kuznetzov and Stekolnikov (1984) regarded the group as the most basal one within the subfamily based on several ‘archaic’ character states: 1, retention of muscles m3 (sternal extensors of the valvae) that are absent in Microcorsini and other Olethreutini; 2, absence of a basal excavation; 3, presence of a wide vinculum; 4, retention of two signa of a ‘primitive’ structure, of lamellar dentate form. Of these, retention of muscles m3 is the only possibly valid argument for a position distinct from the rest of the Olethreutini. Careful examination of the lightly sclerotised valvae revealed that a small basal excavation is present in Gatesclarkeana, albeit hidden behind the process from the valva base, and vinculum width is very variable even within the Gatesclarkeana-group. An interpretation of ‘lamellar, dentate’ signa as a ‘primitive’ condition is no longer tenable given the horn-shaped signa of the more plesiomorphic genera in all olethreutine tribes. On the contrary, these signa composed of few enlarged teeth, often radiating from a small central circle, are a derived character possibly shared with the Neopotamiagroup of the Olethreutini, and perhaps also with the Bactrini (Endothenia). These three groups also all share a usually spiny process from the basal part of their valva and a complex, deeply invaginated sterigma in the female. Furthermore, several of these taxa are characterised by a pair of deep round depressions on T2 in both sexes and by reduced tarsal bristles. Similar paired depressions are found in the tortricine Archips Hübner and the olethreutine Zomariana Diakonoff, Ophiorrhabda Diakonoff and Temnolopha Lower. No function is known for these concavities developed already in the pupa, and their occurrence in a group of tortricines as well as three genus-groups in the Olethreutini strongly suggest a parallelism. However, the three olethreutine genera sharing dorsal depressions with Gatesclarkeana also share reduced tibial bristles and either they themselves or
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close relatives have a spiny process from the base of their valva. Only a comprehensive phylogenetic analysis, including molecular data, will establish the exact position of the Gatesclarkeana-group with regard to the remaining Olethreutini.
Gatesclarkeana Diakonoff (Figs 52, 362–376) Gatesclarkeana Diakonoff, 1966, Zool. Verh. (Leiden) 85: 48. Type species: Platypeplus erotias Meyrick, 1905, J. Bombay Nat. Hist. Soc. 16: 584, by original designation. [Sri Lanka.]
Diagnosis Species of Gatesclarkeana are small to mediumsized, rather broad-winged, brownish, often dark, intricately patterned medium-sized moths with leaden metallic streaks or spots, often with a pinkish hue at least along costa. Gatesclarkeana has moderately long, porrect, sinuate labial palpi, widened beyond middle, with a porrect, moderately long and usually exposed terminal segment. The thorax has a posterior crest except in males of species with modified white abdominal scaling dorsally at base, and the tarsal segments lack strong distal setae. Gatesclarkeana has a pair of deep round concavities at base of T2 in both sexes, visible without dissection. The male is characterised by a greatly expanded hind tibia with flat fringes of long scales from the dorsal and ventral margins, sometimes with an additional scale pencil from its inner base. The anal region of the male is usually modified, either by a dense, lengthened fringe or with the anal lobe enlarged and with modified scaling beneath. Males of some species also have ventral scale filled pockets on the abdomen on S3S5. In the forewing all veins are separate beyond the discal cell, in the hindwing Rs and M1 are parallel and moderately to very close at base, M2 is bent close to M3 at base, and M3 and CuA1 are connate to short-stalked. The male genitalia are diagnostic for the genus with large pedunculi, a hairy, distally bilobed, membranous uncus, strongly asymmetrical valvae with a sclerotised sacculus giving rise to a process (valvula) topped with blunt short spines and, on the right side only, to a sheaf of long spines enfolded between the downturned dorsal half and the ventral portion of the cucullus, and a small juxta and long caulis. The female genitalia are equally unique with an asymmetrical,
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deeply invaginated sterigma posterior to S7, consisting of a distally widening curved irregular cone with spinulose outer surface and enclosing a strongly sclerotised tube contiguous with the posteriorly flattened and sclerotised ductus bursae, with one signum formed by a scobinate, sclerotised depression, frequently with some enlarged teeth, and sometimes with a small remnant of a second signum.
Description Adults (Figs 362–367) Wingspan. 11.0–18.5 mm; small to mediumsized. Head (Fig. 373). Ocellus large. Uppermost part of frons with transverse band of long, projecting scales, remainder with very small, upwardly oriented, appressed scales. Proboscis hardly longer than labial palpus. Labial palpus moderately long (1.9–2.2× diameter of eye), sinuate, porrect, widened beyond middle; second segment sinuate and distally strongly widened by long scales in distal half of ventral and dorsal margins, terminal segment porrect, moderately long and usually exposed. Antenna not quite to middle of forewing; anterior surface devoid of scales; cilia in male from very short to nearly as long as diameter of flagellum. Thorax. With large crest except in male of one species with modified dorsal scaling at base of abdomen. Hind tibia in male often greatly expanded by a fringe of long scales along dorsal and ventral margins and flattened to nearly leaf-shaped, in one derived Australian species also with a pencil of pinkish hairscales from base of inner surface, spreading to fan-shaped if exposed. Tarsal segments distally without conspicuous setae. Wings (Figs 52, 362–367). Forewing index 0.45–53; broadly subtriangular to subovate, with costa evenly curved throughout to weakly or conspicuously angled at about 6M, apex narrowly rounded, termen straight to weakly oblique, hardly sinuate to convex; discal cell 0.61× length of forewing; chorda and M-stem well developed, chorda from below R2 to opposite R5, M-stem to between M2 and M3; all veins separate beyond discal cell; CuP well developed near margin; with usually complex, characteristic wing pattern of dark brown markings and leaden metallic streaks that are outwardly oblique and curved in distal half, ground colour often with pinkish hue especially along costa, rarely (in non-Australian species) with regular roundish spots of raised
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leaden metallic scales across entire wing. Hindwing with well-developed cubital pecten but no axillary tuft, in male anal margin modified, from a dense and enlarged fringe to a variably enlarged, projecting and down-curved anal lobe with modified dense scaling on lower surface; Rs and M1 parallel and moderately to very close at base, M2 bent close to M3 at base, M3 and CuA1 connate to very short-stalked; CuP well developed in distal half, 1A+2A at least partly tubular, 3A sometimes distally weak; medium to dark grey-brown. Pregenital abdomen. Pair of conspicuous, deep round, unscaled concavities laterally at base of T2 in both sexes; males in some species with densely scaled paired projecting lobes ventrally on S3–S5 forming scale-filled pockets, or with modified white scaling dorsally on T1–T3. Male genitalia (Figs 368–372). The two valvae are reflexed and perpendicular to the body axis rather than opposing each other and the genitalia are dorsoventrally compressed, with the valvae facing the tegumen. Vinculum short, weakly sclerotised, ventrally widened into triangular saccus-like process (bent upwardly in slides). Tegumen weakly slcerotised, narrow, triangular, a narrow band except for very large, rounded pedunculi. Uncus a hairy, distally bilobed, membranous process. Socii large, ovate, hairy lobes laterally surrounding anal tube. Gnathos usually absent, rarely present as a small median point. Valvae complex and asymmetrical with sclerotised sacculus giving rise to a dorsomedially oriented sclerotised process (valvula) topped with blunt short spines; valva constricted at end of sacculus and then expanded into large, asymmetrical, weakly sclerotised, hairy cucullus; right sacculus much larger, semicircular, with valvula slender, hook- or band-shaped, passing above aedeagus in natural position, with dorsal half of cucullus folded inwardly parallel to ventral half, enclosing a sheaf of very long spines originating from a process upcurved from ventral margin of sacculus with tips of spines extending beyond valva apex; left sacculus smaller with a shorter, more lobe-shaped valvula, and cucullus smaller without the enfolded sheaf of spines; basal excavation small and hidden behind valvula, costal process not apparent. Juxta very small, caulis very long, anellus a wide ring. Aedeagus tapering to tip, often bulbous at base, frequently with large lateral spike or teeth; no cornuti observed nor their bases but vesica scobinate. Female genitalia (Figs 374–376). Ovipositor lobes elongate ovate with acute posterior apex.
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Figs 362–367. Gatesclarkeana, adults. 362, 363, G. tenebrosa. 362, , Mt Bartle Frere, Q. 363, holotype, Kuranda, Q [image reversed]. 364, 365, G. sp. nr batianensis. 364, , Claudie R. crossing, Q. 365, , Iron Range, Q. 366, 367, G. sp. A, nr Yeppoon, Q. 366, . 367, . Scale = 2 mm.
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Hindmargin of S7 weakly to deeply concave. Sterigma a complex, strongly asymmetrical, deeply invaginated structure posterior to S7 formed as a deep invaginated membranous pocket containing a large, curved, sclerotised, posteriorly widening irregular cone with a spinulose outer surface and posteriorly continuous with innermost, nearly parallel-sided, sclerotised smooth tube turning into flattened, sclerotised posterior portion of ductus bursae; anterior portion of ductus bursae membranous, ductus seminalis originating from near corpus bursae, bulla seminalis nearly as large as corpus bursae. Corpus bursae teardrop-shaped; usually with one, rarely with very small reduced second signum; signum a depressed patch of
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sclerotised scobination, often with a few scobinations enlarged to form discrete teeth.
Distribution Gatesclarkeana has been reported from Sri Lanka and Nepal throughout south-east Asia to Taiwan, Japan, New Guinea and Australia where it has been collected along the eastern coast of Queensland from Yeppoon to Heathlands near the tip of Cape York.
Biology No host records are known from Australia, but all localities are in rainforest or monsoon forest.
Figs 368–373. Gatesclarkeana, male genitalia and head. 368, G. tenebrosa, Mt Bartle Frere, Q, slide ANIC 16860. 369, G. sp. nr batianensis, Claudie R. crossing, Q, slide ANIC 2439. 370, 371, G. sp. A, nr Yeppoon, Q, slide ANIC 2440. 372, G. sp. nr batianensis, aedeagus, slide ANIC 2439. 373, head, G. tenebrosa, male.
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Fletcher ([1921]) reports Gatesclarkeana erotias (Meyrick), a pest of litchi (Sapindaceae) in India, as a leaf roller and webber of mango leaves (Anacardiaceae), Loranthus sp. (Loranthaceae), Cynoglossum sp. (Boraginaceae) and the introduced Lantana camara (Verbenaceae), but also states that the species has been reared ‘from larvae boring mango shoots in Bombay’. Diakonoff (1973) gives numerous hosts of many plant families for G. idia (Diakonoff), with the larvae feeding on leaves, flowers and rarely fruit, and several hosts of different families for both G. domestica Diakonoff and G. senior Diakonoff, with tea (Theaceae) and Lantana reported more than once. Gatesclarkeana species seem to be polyphageous leaf rollers, possibly sometimes feeding as borers. The closely related Ukamenia Oku also usually is a leaf roller but has been found feeding in gall wasp galls (Oku 1981).
Remarks Clarke (1958) and Robinson et al. (1994) figured adults of G. erotias (Meyrick) (as Olethreutes) and G. idia Diakonoff respectively. Diakonoff (1973) provided descriptions and illustrations of genitalia of the many south-east Asian species of Gatesclarkeana. Razowski (1989) briefly reviewed the genus. Gatesclarkeana, Hiroshiinoueana Kawabe (Kawabe 1978), Ukamenia Oku (Oku 1981; with Aphiaris Kuznetzov (Kuznetzov 1981) the junior synonym), Asymmetrarcha Diakonoff (Diakonoff 1973) and some generically unassignable Australian and Oriental species are clearly all closely related but show puzzling character combinations with widely divergent male genitalia. It is doubtful that the current genera denote monophyletic groups but it would be unwise to change the status quo based
Figs 374–376. Gatesclarkeana, female genitalia. 374, G. tenebrosa, holotype, slide ANIC 16864. 375, G. sp. nr batianensis, Iron Range, Q, slide ANIC 16858. 376, G. sp. A, nr Yeppoon, Q, slide ANIC 16855.
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on this limited study. Gatesclarkeana is here accepted as defined by Diakonoff (1973), with a weakly bilobed, hairy uncus, strongly asymmetrical valvae with an enfolded spine pencil only on the right side, and with the invaginated sterigma covered with long spinules on its outer surface as possible generic apomorphies. Gatesclarkeana shares with the genera previously mentioned broad wings, a small juxta and a long caulis, a ventrally triangularly expanded but weakly sclerotised vinculum, and usually only one signum (rarely with a remnant of the second one), a depressed patch of scobination with often enlarged single teeth. Conspicuous paired concavities on T2 are present in Gatesclarkeana and figured for Ukamenia by Kuznetzov (1981), but are not mentioned in the descriptions of the other genera. In the male a modified hind tibia and ventral pockets near the base of the abdomen are found in several genera, and, if developed, are probably apomorphic for the group with their detailed structure possibly characteristic of certain subsets. Hiroshiinoueana has nearly identical male genitalia to Gatesclarkeana except for the presence of a bundle of bristles also on the left valva. Both Hiroshiinoueana and Ukamenia seem to lack spinules on the outer surface of their complex sterigma, that are otherwise exactly alike that of Gatesclarkeana. The valva of Ukamenia and a presumably highly derived Australian species (species A, Fig. 371) look radically different to those of the type species G. erotias and, unless some intermediate forms are found, are difficult to homologise.
Evidence for monophyly Diakonoff (1973) based his Statherotites on the very characteristic, paired signa, which he referred to as ‘pectinate’. These uniquely derived structures consist of several parallel rows of flattened, triangular or rounded teeth of usually increasing length, more rarely all of equal size, arranged like fish-scales. The valva is usually slender and in some groups has a spiny basal process from its inner surface. Apart from the apomorphic signum, the female genitalia have a rather simple spinulose sterigma posterior to S7 and a long, slender, wellsclerotised colliculum. There is a tendency for R3 and R4 to be closely approximated or even stalked.
Distribution The Statherotis-group is centred on the Oriental and Australian regions, with several genera found in the eastern Palaearctic, and at least Aterpia extending into the Holarctic.
Biology
Constituent species
Few host records are available for species of the Statherotis-group, but all known larvae are leaf rollers. At least two species of Statherotis feed on Nephelium lappaceum (rambutan) and N. litchi (litchi) (Sapindaceae) in south-east Asia and Japan. In Japan, the larvae of S. tonica are leaf rollers on two Elaeocarpus species (Elaeocarpaceae) (Kawabe 1982). The Nearctic Aterpia approximana (Heinrich) feeds on Lythrum (Lythraceae) (Heinrich 1926) and two Palaearctic species on Dryas (Rosacae) and Aconitum (Ranunculaceae) respectively (Hannemann 1961).
Diakonoff (1973) reported nine named species from south-east Asia, and one named and three unnamed species are known from Australia.
Remarks
tenebrosa (Turner, 1916), Trans. R. Soc. S. Aust. 40: 531 (Argyroploce). [Qld: Kuranda.]
Statherotis-group Status remarks This group was established for 21 genera by Diakonoff (1973) as the subtribe Statherotides, given as Statherotina by Nye and Fletcher (1991). Kuznetzov and Stekolnikov (1984) accepted subtribal level for the Statherotina that are here treated as a genus-group.
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The uniquely derived signum is a strong indication that the Statherotis-group is monophyletic.
Metrioglypha Diakonoff (Figs 46, 377–387) Metrioglypha Diakonoff, 1966, Zool. Verh. (Leiden) 85: 41. Type species: Grapholitha vulgana Walker, 1866, List Specimens Lepid. Insects Colln Br. Mus. 35: 1796, by original designation. [Ceram.] G. vulgana is a junior synonym of Grapholita confertana Walker, 1863, List. Lep. Br. Mus. 28: 388.
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Figs 377–380. Metrioglypha, adults. 377, 378, M. thystas, Kuranda, Q [BMNH]. 377, [image reversed]. 378, [image reversed]. 379, 380, M. phyllodes. 379, , Cooloola, Q. 380, , Brisbane, Q [image reversed]. Scale = 2 mm.
Astrobola Diakonoff, 1966, Zool. Verh. (Leiden) 85: 42. Type species: Astrobola circulata Diakonoff, 1966, ibid. 85: 44, by monotypy and original designation. [New Guinea: Sudest I.]
Diagnosis Metrioglypha species are small to medium-sized, contrastingly marked and rather colourful olethreutine moths with slender wings and without tibial pencil in the males of Australian species. Several species have a conspicuously rounded dark brown apex with white scalloped lines (M. thystas (Meyrick)), whilst the Australian M. phyllodes (Lower) has a dull bluish green forewing with
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incomplete black transverse fasciae and a pointed apex. A patch of androconial scales or a hair pencil close to the costa are present on the male hindwing of some Metrioglypha, but not the Australian species. The wing venation with all veins separate beyond the discal cell (except for the connate M3 and CuA1 in the hindwing) is typical for a member of the Olethreutini and may serve to distinguish M. phyllodes from other green-coloured Australian olethreutines such as Noduliferola neothela (Turner), Acroclita bryopa Meyrick and Rhopobota hortaria Meyrick. The three darkish green Sorolopha species can at least in the male be recognised by their strongly modified hind legs. The male genitalia of Metrioglypha with a well-developed and usually
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hairy uncus, a band-like gnathos without a free point and an often apically broadly rounded simple valva with a dense patch of short thick spines in centre of valva beyond basal excavation are diagnostic for the genus. Phaulacantha Diakonoff and Cephalophyes Diakonoff both have very similar
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male genitalia but the former has a broad and distinctly bipartite uncus and the latter a spined rather than bristled valva. A sterigma derived from a projecting spinulose fold encircling the ostium and especially the strongly transversely wrinkled ductus bursae and smooth caudal portion of the corpus
Figs 381–387. Metrioglypha, genitalia and head. 381, 382, M. thystas, male genitalia and aedeagus, Kuranda, Q, slide T2470. 383, head, M. phyllodes, male. 384, M. phyllodes, aedeagus, slide T2275. 385, M. thystas, female genitalia, Kuranda, Q, slide T2471; 386, M. phyllodes, female genitalia, Brisbane, Q, slide T2279. 387, M. phyllodes, male genitalia, Brisbane, Q, slide T2275.
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bursae are the diagnostic features of the female genitalia. The more derived Metrioglypha species have signa with rather short, triangular and very densely spaced teeth, suggesting a patch of scales.
Description Adults (Figs 377–380) Wingspan. 10–16 mm; small to medium-sized. Head (Figs 383). Ocellus well developed. Frons in uppermost part with a narrow transverse band of anteriorly projecting scales, remainder with short, appressed and upwardly oriented scales. Proboscis short, hardly longer than labial palpus. Labial palpus moderately long (1.7–2.1× diameter of eye), porrect and sinuate, with distally moderately to strongly widened second segment and with exposed third segment. Antenna reaching to middle of forewing; with anterior surface unscaled, cilia in male from minute to short, to about half diameter of flagellar segment. Thorax. With a bifid posterior crest. Legs unmodified in the known Australian species; some species elsewhere with tibial pencil in male (Diakonoff 1973). Wings (Figs 46, 377–380). Forewing index 0.36–0.42; forewing long and rather narrow, elongate-ovate to subrectangular, with evenly curved costa, slightly projecting apex, straight to sinuate and slightly oblique termen; discal cell 0.63–0.65× length of forewing, with moderately wide accessory cell with chorda from halfway between R1 and R2 to between R5 and M1, with well-developed M-stem to between M2 and M3; with all veins present and separate beyond discal cell; with diverse and intricate wing markings, Australian species either whitish to reddish ochreous with numerous dark brown, mostly longitudinally oriented spots and blotches (M. thystas) or dull bluish green with outwardly oblique incomplete black transverse fasciae (M. phyllodes). Hindwing with cubital pecten and axillary tuft, males of some species on upper side with a central patch of black androconial scales (M. confertana (Walker)) or with an expansible hair pencil close to base of costa (M. gemmarius Diakonoff) (Diakonoff 1973); with Rs and M1 parallel and closely approximated at base (stalked in Astrobola circulata Diakonoff (Diakonoff 1966)), M2 approximated to M3 at base, CuA1 from just below angle of cell seemingly connate with M3; 1A+2A and 3A well developed; dark greyish to blackish, often with paler semi-translucent base and/or bronze gloss.
Olethreutine Moths Final.indd 186
Pregenital abdomen. Unmodified. Hindmargin of S8 in male bilobed with a slight to deeply rounded median excision; T8 shorter and small. Male genitalia (Figs 381, 382, 384, 387). Vinculum a moderate band. Tegumen high and ovate in outline, hardly tapering towards pedunculi and with ill-defined broad apodemes for muscle m4. Uncus a well-developed,ventrally curved, median process, apex often dilated and nearly always hairy on dorsal (M. thystas) or ventral (M. phyllodes) surface. Socii from large drooping lobes to small groups of bristles lateral to anal tube. Gnathos a weakly sclerotised, often V-shaped transverse band. Valva curved to lightly sinuate, elongate-ovate to sickle-shaped, with short and stout costal process often concealed by humpshaped invagination from outer valva wall, with sacculus ill-defined, usually a dense patch of short stout spines in centre of valva just beyond basal excavation, often with a band of long bristles along middle of ventral margin or a dense tuft more basally, and with densely and evenly bristled cucullus often with increasingly stout spines towards ventral margin, frequently with modified and some particularly large spines just beyond ventro-proximal margin of bristled cucullus area, with a single huge spine from beyond middle of ventral valva margin in M. phyllodes. Caulis long and slender at base, anellus a moderate band, diaphragm above anellus membranous but sometimes spinulose. Aedeagus short and small; vesica without cornuti. Female genitalia (Figs 385, 386). Ovipositor lobes subtriangular. Sternum 7 with a concave hindmargin. Sterigma derived from a strongly raised spinulose fold encircling ostium, with a ventral notch and lateral portions produced into sharp points in M. thystas, with dorsal part expanded into an irregular medially depressed bulge in M. phyllodes. Ductus bursae with a long and slender bipartite colliculum below ostium followed by inception of ductus seminalis, remainder of ductus bursae with even and conspicuous transverse wrinkles throughout. Corpus bursae ovate, distinctly reticulate except towards entrance of ductus bursae; with two signa consisting of dense parallel rows of rather short triangular teeth, arranged in scale-like fashion.
Distribution Sri Lanka, India, Indonesia, New Guinea, Bismarck Is (New Hanover I.), Australia. In Australia the genus is restricted to the eastern coast of
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Queensland and northern New South Wales as far south as Lismore.
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with Metrioglypha circulata that is again recognised as a separate species: Metrioglypha circulata (Diakonoff, 1966): 42 (Astrobola). [New Guinea: Sudest I.] Sp. rev.
Biology Two rearing records from Java for M. empalinopa Diakonoff, ‘bred from leaves of ‘lunda’ (? Vitis geniculata)’ and ‘from unidentified plant’ (Diakonoff 1973) are the only host records. The locality data for the Australian species suggest rainforest habitats.
Remarks Diakonoff (1973) provided a comprehensive treatment of Metrioglypha with numerous illustrations of genitalia, synonymising Astrobola with Metrioglypha. Figures of genitalia, head and wing venation are also found in the original descriptions of Metrioglypha and Astrobola (Diakonoff 1966). Metrioglypha vulgana, the type species of the genus and subsequently recognised as a junior synonym of M. confertana (Walker) (Diakonoff 1982), lacks some of the apomorphies of more derived Metrioglypha species such as the distally broadly rounded cucullus, the patch of short thick spines in centre of valva beyond basal excavation and the shortened triangular teeth giving the signum the appearance of a patch of scales rather than the pectinate shape found in Statherotis. However, M. confertana shares the apomorphic, transversely wrinkled ductus bursae with the remainder of the genus. The type species of Astrobola is a more characteristic member of Metrioglypha, possessing the full set of apomorphies. Diakonoff (1973) synonymised Astrobola circulata Diakonoff with M. thystas and described a new species M. mellifera Diakonoff for a female from Australia (Cairns). The type series of M. thystas contains material from New Guinea (St Aignan I.) and Australia (Cairns), with a male labelled as lectotype from Cairns. Australian material, including the lectotype of M. thystas, has a distally widened valva without the strong spines at ventral base of cucullus present in the New Guinean M. circulata (figured as Metrioglypha thystas in Diakonoff (1973)). The Australian and New Guinea populations are two distinct species, with the Australian M. mellifera conspecific with the lectotype of M. thystas from Cairns. The New Guinean species comprises M. thystas ss. Diakonoff 1973 and the New Guinea material from the type series of M. thystas, all of which are conspecific
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Constituent species Eleven named species of Metrioglypha are recognised world-wide, including the two species listed below from Australia: thystas (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 276 (Argyroploce). Lectotype , ‘Cairns, Queensland, FPD, .12.[19]05’ (BMNH), hereby designated. mellifera Diakonoff, 1973, Zool. Monogr. Rijkmus. Nat. Hist. 1: 210 (Metrioglypha). [Qld: Cairns.] phyllodes (Lower, 1899), Proc. Linn. Soc. NSW 24: 93 (Dipterina(?)). [Qld: Brisbane.]
Dactylioglypha Diakonoff (Figs 47, 388–393) Dactylioglypha Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 180 (key), 188. Type species: Dactylioglypha avita Diakonoff, 1973, ibid. 1: 191, by original designation. [New Guinea.]
Diagnosis Dactylioglypha species are moderately sized greyish olethreutine moths with a wing pattern of pale and dark bands and streaks separated by whitish lines; bands either straight and broadest on dorsum and wing base and all converging towards 169 costa (D. tonica (Meyrick)), or strongly sinuate along length of wing. Species with sinuate wing pattern strongly resemble species of Asymmetrarcha Diakonoff, but D. tonica, the only species known from Australia, is immediately recognised by the greenish tinge and characteristic shape of its forewing with straight pattern elements and a conspicuously reddish fringe. The closely related Papuan D. avita lacks the greenish hue and has a greyish forewing fringe. Males of at least some species (D. tonica) have a hair pencil on their hind tibia and a lightly sclerotised patch of modified membrane with a reticulate structure laterally across the first two abdominal segments. The male genitalia with their stout, distally hairy and curved
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uncus, the usually rigid and porrect socii, the wellsclerotised gnathos with a broad paddle-shaped distal process and the sclerotised diaphragm linking gnathos and anellus are all very similar and diagnostic for the genus. The female genitalia are characterised by an either dorsally or ventrally interrupted rim-shaped spinulose sterigma, often with projecting ventro-lateral corners, by a rather broad and irregular colliculum, a smooth ductus and a corpus bursae with a granulate structure covering entire surface and with two pectinate signa with some very long truncate teeth. All the genera of the Statherotis-group have rather similar female genitalia, but the irregular colliculum of Dactylioglypha is in contrast to the usually long and thin colliculum of other taxa with a similar sterigma. The smooth ductus bursae and the long truncate teeth of the signa of Dactylioglypha allow separation from females of Metrioglypha.
Description Adults (Figs 388, 389) Wingspan. 11–15 mm; small to medium-sized. Head (Fig. 390). Ocellus well developed. Vertex scales conspicuously white-tipped, anteriorly barely projecting between antennae. Frons in uppermost part with a narrow band of long, anteriorly projecting scales; remainder of frons with minute, appressed and upwardly oriented scales. Proboscis short, not much longer than labial palpus. Labial palpus rather small (1.5–1.7× diameter of eye) and slender, upcurved and lightly sinuate towards tip; second segment evenly curved and with loose scales along ventral margin and on apex, but distally hardly dilated; terminal segment small and porrect. Antenna reaching to middle of forewing; with anterior surface unscaled; cilia in male minute. Thorax. With small posterior crest. Legs in males of at least some species (D. tonica) with a pencil of hair scales from base of inner side of tibia that is enlarged by scale fringes. Wings (Figs 47, 388, 389). Forewing index 0.43–0.50; forewing subrectangular to subtriangular (D. tonica) with seemingly truncate apex, with costa roughly straight or lightly concave to 169, thence distinctly angled, apex nearly rectangular to obtuse, termen vertical and slightly sinuate with broadly rounded and nearly projecting tornus; discal cell 0.63× length of forewing, with a small accessory cell with chorda from between R1 and R2 and often indistinct in distal portion, with partly non-tubular M-stem to between M2 and M3;
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with all veins present and separate beyond discal cell and with CuA1 strongly curved and approximated to M3 at wing margin; apex conspicuously marked by an ovate blackish spot preceded by white strigulae; wing markings in D. avita and D. tonica a number of pale and dark greyish streaks separated by whitish lines, arising from wing base and along dorsum and distally narrowing and converging towards 169 costa; remaining species with alternating pale and dark grey streaks strongly sinuate throughout. Hindwing with cubital pecten but at least in D. avita and D. tonica without axillary tuft; with Rs and M1 parallel and closely approximated at base and M2 approximated at base to connate or rarely very short-stalked M3 and CuA1; 1A+2A and 3A present, at least partly tubular, grey to blackish. Pregenital abdomen. Male of D. tonica with lateral portions of the first two segments expanded, covered with a large and lightly sclerotised ovate patch of reticulate structure. Sternum 8 in male large, with a bilobed hindmargin with a semicircular median excision; T8 much smaller and shorter. Male genitalia (Figs 391, 392). Vinculum a weakly sclerotised moderately broad band. Tegumen high and rather narrow, well-sclerotised and with prominent ‘shoulders’, pedunculi with subbasal median expansion but no well-defined apodemes for muscle m4. Uncus a strong, stout and ventrally curved median process, with rounded apical part covered by long hair. Socii usually strongly sclerotised, porrect, rigid, slender processes, sparsely haired, with upwardly curved apex in several species (D. tonica). Gnathos arms strongly sclerotised, laterally connected with socii and medially fused with an elongate sclerite in diaphragm as well as with a very large and broad paddle-shaped and strongly projecting process, forming a well sclerotised functional unit connected to anellus with base of sclerite in diaphragm. Valva long and slender, curved or lightly bent and parallel-sided, with rather slender costal process, sacculus with a dense patch of bristles at ventral edge of basal excavation, with a small hump to finger-shaped process carrying a group of bristles at margin of basal excavation in centre of valva and with cucullus usually more or less evenly bristled throughout with shorter, isolated short spines at base. Caulis moderate and anellus a broad band, its dorsal edge fused with sclerite in diaphragm. Aedeagus smallish, lightly curved; vesica without cornuti.
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Female genitalia (Fig. 393). Ovipositor lobes subtriangular to elongate-ovate. Sternum 7 with straight to concave hindmargin, rarely fused with lamella antevaginalis. Sterigma derivations of a variously developed raised spinulose fold encircling
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ostium, forming a dorsally interrupted round rim in D. tonica, more ovate with a deep ventral notch and produced into ventrolateral corners in other species. Ductus bursae with an ill-defined long colliculum, followed by inception of ductus
Figs 388–393. Dactylioglypha tonica, adults, head and genitalia. 388, 389, adults. 388, , nr Cooktown, Q. 389, . 390, head, male. 391, 392, aedeagus and male genitalia, nr Cooktown, Q, slide T2282. 393, female genitalia, Claudie R., Q, slide T2462. Scale = 2 mm.
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seminalis, remainder of ductus rather wide and smooth. Corpus bursae ovate, with granulate structure reaching to entrance of ductus; with two pectinate signa with relatively few rows of partly very long and truncate teeth.
other species of Eccopsis (Diakonoff 1981) refer the genus to the Temnolopha-group. Heinrich’s (1926) illustration of the male genitalia of D. wahlbergiana differs greatly from that given by Diakonoff (1981), but tallies with all the other species of Eccopsis figured by Diakonoff (1981), further suggesting a position of Eccopsis within the Neopotamia-group.
Distribution Sri Lanka, India, south-east Asia, Japan, New Guinea, Australia. In Australia the genus is restricted to northern Queensland north of Cooktown.
Biology In Japan, the larvae of D. tonica are leaf rollers on Elaeocarpus japonicus var. ellipticus (Elaeocarpaceae) (Kawabe 1982), with detailed biological information provided by Tominaga (1998). No host information is available from Australia, but all the Australian localities are next to or in rainforest.
Remarks Diakonoff’s (1973) comprehensive treatment of Dactylioglypha provides illustrations of genitalia of all known species and of head and wing venation of the type species. Dactylioglypha is a very closely-knit genus characterised by a number of obvious apomorphies such as the preapically bent forewing costa combined with a black apical spot, the distally bristled stout uncus and the highly derived gnathos/subscaphium-complex. The modified areas found laterally in the membrane of the first two abdominal segments in the male are present in some but not all species of Dactylioglypha (Tuck personal communication). Semniotes Diakonoff and Pomatophora Diakonoff are the only other genera in the Statherotis-group with a welldeveloped raised process from the base of the inner valva surface, but this is not paralleled by any other shared characters that could indicate a sister group relationship. Diakonoff (1981) suggested a close relationship between Dactylioglypha and the African Eccopsis Zeller, 1852. His figure of the male genitalia of Eccopsis wahlbergiana Zeller, 1852, the type species of the genus, indeed shows all the features typical for the male genitalia of Dactylioglypha. However, the male lectotype of D. wahlbergiana lacks an abdomen (Diakonoff 1957c), and the female genitalia figures for D. wahlbergiana as well as for
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Constituent species Five named species of Dactylioglypha are recognised worldwide, with D. tonica, described from Sri Lanka and India, present in Australia: tonica (Meyrick, 1909), J. Bombay Nat. Hist. Soc. 19: 606 (Argyroploce). [Sri Lanka.]
Statherotis Meyrick (Figs 48, 394–410) Statherotis Meyrick, 1909, J. Bombay Nat. Hist. Soc. 19: 591. Type species: Statherotis decorata Meyrick, 1909, ibid. 19: 591, by monotypy. [Sri Lanka.] Statheromeris Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 180 (key), 181. Type species: Statheromeris atrifracta Diakonoff, 1973, ibid. 1: 182, by original designation. [New Guinea.] Syn. nov.
Diagnosis Statherotis species are small to quite large moths, either intricately marked in shades of pale ochreous, grey or reddish brown, or with the typical Statherotis pattern of usually darkish greyish gound colour with a paler and often pinkish tinged triangle or semicircle over middle of costa, ususally bordered by a black band and sometimes entirely covered by blackish scales (Figs 397, 399). Males often have patches of black androconial scales on centre of hindwing and underside of forewing. This Statherotis-type forewing pattern with a pinkish tinged pale costal spot bordered by black is, however, also found in some related genera and characterises the genus Eremas Turner of the Olethreutes-group. Eremas males can be recognised by a large pencil of hair-scales close to costa on upper side of hindwing, and they never possess black androconial patches. The male hind tibia of Statherotis is often enlarged by scale fringes and in all Australian species carries a slender hair-pencil
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along its inner surface. The venation is characterised by a very narrow accessory cell and usually closely approximated to stalked R3 and R4. The male genitalia of Statherotis, with their slender valva with a ventral triangular process in the distal portion of the cucullus, are diagnostic for the genus. Those of the closely related Dicephalarcha can be recognised by their much longer and more slender ventral cucullus process that is devoid of bristles, at least at its base, and by the shape of the usually bifid uncus. Paired, bristled triangular to rod-shaped processes above or lateral to ostium, together with a smooth-walled ductus bursae, are the diagnostic features of the female genitalia, whilst the characteristic pectinate signa are shared with a number of related genera. Metrioglypha has very similar female genitalia but its transversely wrinkled ductus bursae affords easy separation from Statherotis.
Description Adults (Figs 394–399) Wingspan. 10–21 mm; small to quite large. Head (Figs 409, 410). Ocellus well developed. Frons in uppermost part with a narrow transverse band of anteriorly projecting scales, remainder with short, appressed and upwardly oriented scales. Proboscis short, not much longer than labial palpus. Labial palpus moderate to long (1.5–2.5× diameter of eye), porrect and sinuate, usually widened towards apex of second segment; second segment sinuate, moderately slender throughout (S. batrachodes (Meyrick)) to strongly widened before rounded apex; terminal segment from moderate to long and thin (S. euryphaea (Turner)). Antenna reaching to middle of forewing; with anterior surface unscaled; cilia in male from short to moderately long, to just below diameter of flagellar segment. Thorax. Mostly without posterior crest. Legs with hind tibia in male often enlarged by dorsal and ventral fringes of long scales and in all Australian species with a long slender pencil of hair-scales from base along inner surface. Wings (Figs 48, 394–399). Forewing index 0.40–0.45; forewing elongate-subrectangular, moderately widened distally, with lightly and evenly curved costa, straight to sinuate termen; in males of some species with an elongate patch of blackish androconial scales on lower surface along dorsal margin of discal cell; discal cell 0.6× length of forewing, with narrow accessory cell with chorda from below R2 to between R4 and R5 or R5 and
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M1, and with M-stem to between M2 and M3; all veins present and separate beyond discal cell except for R3 and R4 that may be variously approximated or stalked, with R4 running to costa; CuP present near margin; basal loop 0.3× length of 1A+2A; often dark greyish with a pale pinkish tinged triangular or semicircular mark along middle of costa, bordered by black (Fig. 396) or entirely suffused by blackish (Fig. 397), or intricately marked throughout in shades of pale ochreous, greenish or reddish brown. Hindwing with cubital pecten (except males of some species with androconia) and axillary tuft, in male frequently with variously shaped patch of blackish androconial scaling in centre of upper surface (Fig. 48) and/or with a modified anal area with dense, spatulate scales underneath and along margin; with Rs and M1 closely approximated and parallel at base, with M2 at base close to very close to connate M3 and CuA1; 1A+2A and 3A well developed; light to dark greyish, in some males with whitish anal area. Pregenital abdomen. Rarely with paired patches of modified scales on T6–8 in male. Eighth segment in male weakly sclerotised; S8 large and hindmargin bilobed with a semicircular median emargination and a band of long scales along apices of rounded lobes, T8 with an inverted V- or Tshaped sclerite at base and hindmargin usually projecting as a small median lobe, in S. batrachodes T8 greatly enlarged to a prominent subcordate lobe. Male genitalia (Figs 400–405). Vinculum a wide, hardly sclerotised band. Tegumen high and ovate in outline, with pronounced shoulders and broad, bilobed and often ill-defined apodemes for muscle m4. Uncus usually a well-developed median process, sometimes apically dilated or medially notched, at least ventral surface of apex densely hairy and often bipartite; uncus in some species (S. atrifracta (Diakonoff)) reduced to a hairy band along apex of tegumen. Socii two membranous, hairy, drooping rounded lobes of various size. Gnathos from a faintly sclerotised transverse band (S. batrachodes) to a wellsclerotised medially fused structure with a trowellike projecting apex. Valva long and very slender, costal process small and concealed by cone-todome shaped invagination from base of valva, sacculus ill-defined, with a variable hump covered with slender bristles below or just beyond basal excavation, cucullus with a strong thorn on ventral margin that is often produced into a triangular ventral process carrying enlarged spines, spines across base of cucullus often enlarged and raised on a ridge. Caulis long and slender at base, anellus
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Figs 394–399. Statherotis, males. 394, S. ? amaeboea, Eungella, Q [image reversed]. 395, S. batrachodes, Mossman, Q. 396, 397, S. pendulata. 396, regular form, nr Gympie, Q. 397, dark form, Lamington Nat. Park, Q. 398, 399, S. euryphaea. 398, regular form, upper Allyn R., NSW. 399, dark form, Mt Keira, NSW [image reversed]. Scale = 2 mm.
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a moderate band, diaphragm between anellus and subscaphium often lightly sclerotised and with vertical wrinkles. Aedeagus small and short; vesica without cornuti. Female genitalia (Figs 406–408). Ovipositor lobes subtriangular. Sternum 7 with a concave hindmargin. Sterigma either a median longitudinal spinulose bulge with ostium at its base below two variously produced paired spinulose points, or a deep short vertical groove flanked on each side by a long pointed porrect and spinulose process (S. pendulata (Meyrick)). Ductus bursae long and smooth, widening towards corpus bursa and bent or coiled, with a long straight and bipartite colliculum below ostium, followed by inception of ductus seminalis and often by a lateral dilation in ductus wall. Corpus bursae ovate, with two large signa consisting of dense parallel rows of pectination of increasing size, rarely with one of the signa reduced to a trace only (S. atrifracta).
Distribution Sri Lanka, India, Maldive Is, Seychelles, south-east Asia, China, Philippines, Taiwan, Indonesia, New Guinea, Micronesia, Solomon Is, Fiji and Australia. In Australia the genus is found in the coastal regions of the Northern Territory (Darwin), Queensland and New South Wales south to Malua Bay.
Biology At least two species have repeatedly been found as leafrollers on Nephelium lappaceum (rambutan) and N. litchi (litchi) (Sapindaceae) in south-east Asia, and there is one rearing record from Eugenia polyantha (Myrtaceae) (Diakonoff 1973). Statherotis discana is a pest on litchi in Japan (Nasu et al. 1993). In Australia, a species close to S. amaeboea (Lower) has been found webbing leaves of litchi at Babinda, Queensland, by A. Rogers. Statherotis euryphaea has been reared binding together terminal foliage of Banksia integrifolia (Proteaceae) at Jervis Bay and also on Banksia sp. at Malua Bay by I. F. B. Common, both New South Wales.
Remarks Diakonoff (1973) provided a comprehensive revision of Statherotis with numerous illustrations and extensive literature references. The monophyly of Statherotis is based on the following apomorphies: base of R3 and R4 approximated or stalked, a very slender valva with a
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broadly triangular process at about 36: ventral margin, a spinulose sterigma with a projecting point or lobe on each side and a very long, slender, well sclerotised colliculum. Many species are also characterised by patches of androconial scales below on forewing and above on hindwing, and the gnathos usually ends in a long, slender process. Statheromeris (Diakonoff 1973) is no more than an offshoot from Statherotis with a reduced uncus and one of its signa atrophied. It otherwise agrees structurally entirely with Statherotis, sharing such apomorphic features as the shape of the valva and of the sterigma. To treat Statheromeris as a separate genus would merely reduce Statherotis to a paraphyletic group. The following species are here transferred to Statherotis: atrifracta (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 182 (Statheromeris). [New Guinea.] Comb. nov. semaeophora (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 184 (Statheromeris). [Bali.] Comb. nov. Diakonoff (1973: Fig. 332) figured the wing of Statherotis atrifracta (Diakonoff) with R4 running to termen, but in both Statherotis solomonensis (Bradley) and an unnamed related species from Australia R4 runs to the costa. Dicephalarcha Diakonoff shares a valva with a tringular ventral process with Statherotis, but the valva is wider and the process longer and more slender in Dicephalarcha, and the sterigma lacks the laterally projecting lobes. Several species of Statherotis are dimorphic with regard to the pale costal spot that may be filled in by black scales (S. pendulata). In the dark form of S. euryphaea the entire costal half of the forewing is covered by dark brown scales (Fig. 399). This dimorphism is found throughout the range of the species. The identity of the Australian species S. amaeboea Lower has generally been misconstrued as it was based on a series of specimens from the Solomon I. and Sudest Is (New Guinea) identified as S. amoebaea (sic!) by Meyrick (Meyrick 1911b), and not on Lower’s type. The single female holotype unfortunately lacks the abdomen and Brisbane, the type locality, is much further south than the Cairns/ Cooktown region from where we have material of two closely related Statherotis species that could both be S. amaeboea. Neither of the two species has modified scale-tufts on S6–8 in the male, as figured in Diakonoff (1973: fig. 343) nor a deeply cleft uncus. They both are members of Diakonoff’s (1973) discana-group of species, and one of the two may be conspecific with S. discana (Felder &
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Figs 400–405. Statherotis, male genitalia and aedeagus. 400, 401, S. euryphaea, Mt Keira, NSW, slide T2260. 402, 403, S. batrachodes, Mt Bellenden-Ker, Q, slide T2298. 404, 405, S. pendulata, hr Gympie, Q, slide T2261.
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Rogenhofer). Further material will be needed to conclusively establish the identity of S. amaeboea. The similarity between the typical Statherotis forewing pattern and that of the quite distant Australian Eremas species is remarkable and has misled taxonomists: Diakonoff (1973) included Eremas tetrarcha (Meyrick) among the species of Statherotis.
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Constituent species Thirty three named species of Statherotis are recognised worldwide (Brown 2005). There are several undescribed species in Australia, and four described species: euryphaea (Turner, 1916), Trans. R. Soc. S. Aust. 40: 530 (Argyroploce). [Qld: Herberton.]
Figs 406–410. Statherotis, female genitalia and heads. 406, S. batrachodes, Mission Beach, Q, slide ANIC 1291. 407, S. pendulata, Rockhampton, Q, slide ANIC 2463. 408, S. euryphaea, Mt Keira, NSW, slide ANIC 2464. 409, 410, heads, male, 409, S. euryphaea. 410, S. pendulata (palpi unusually drooping).
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leuconephela (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 205 (Eucosma), nec (Turner, 1925) (Eucosma). [Qld: Ebor.] batrachodes (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 274 (Argyroploce). [New Guinea, Solomon Is, Qld: Duaringa.] amaeboea (Lower, 1896), Trans. R. Soc. S. Aust. 20: 161 (Dichelia (?)). [Qld: Brisbane.] pendulata (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 279 (Argyroploce). [Qld: Brisbane.] toxosema (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 216 (Argyroploce). [Qld: Yeppoon, ‘Tweed Heads’.]
Aterpia Guenée (Figs 49, 411–416) Aterpia Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 161. Type species: Aterpia andereggana Guenée, 1845, ibid. (2)3: 161, by monotypy. [Europe.] Asaphistis Meyrick, 1909, J. Bombay Nat. Hist. Soc. 19: 590. Type species: Asaphistis praeceps Meyrick, 1909, ibid. 19: 590, by monotypy and original designation. [India: Khasis.] Syn. nov. Esia Heinrich, 1926, US Nat. Mus. Bull. 132: 76 (key), 109. Type species: Olethreutes approximana Heinrich, 1919, Insecutor Inscit. Menstr. 7: 65, by original designation. [North America.] Metaschistis Diakonoff, 1953, Verh. K. Ned. Akad. Wet., Nat., 2. ser. 49(3): 94. Type species: Metaschistis sappiroflua Diakonoff, ibid. 49(3): 100, by original designation. [New Guinea.] Apeleptera Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 180 (key), 262. Type species: Argyroploce semnodryas Meyrick, 1936, Exot. Microlep. 4: 613, by original designation. [Taiwan.] Syn. nov. Leptocera Diakonoff, 1983c, Ann. Soc. Entomol. Fr. (N. S.) 19: 307. Type species: Gnathmocerodes (Leptocera) microchlamys Diakonoff, 1983c, ibid. 19: 307, by original designation. [Described as a subgenus of Gnathmocerodes.] [Madagascar.]
Diagnosis (Primarily based on the Oriental/Australian species-group.) Aterpia species are small to large, usually bluish or blackish and white olethreutine
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moths with long verticillate antennal cilia in male, often with dark forewing base, dorsum and apex, and with a white sinuate band running from before middle of costa to tornus (A. protosema (Diakonoff)). Some of the dark bluish Papuan species of Aterpia are superficially strongly reminiscent of species of Cryptaspasma Walsingham, which, however, have a shorter and stouter labial palpus, either more or less straight or sickle-shaped, but never porrect, sinuate and with loose scaling as in Aterpia. The wing venation of Oriental and Australian Aterpia is characterised by CuA1 in the hindwing originating from distinctly before angle of cell, a feature shared by only a few other olethreutine taxa such as Dynatorhabda Diakonoff, Oxysemaphora Diakonoff and Ophiorrhabda Diakonoff. The male genitalia of Aterpia are highly diagnostic with a roughly rectangular, hairy and membranous uncus, gnathos arms either fused with anal tube or produced into a median process, a strongly projecting cone-shaped sclerotised area in diaphragm between anal tube and a ventrally greatly enlarged anellus and with a simple valva with two parallel bands of dense bristles along basal half of ventral edge and usually with a triangular tooth on middle of costa. Proschistis Meyrick has a similar tegumen but with conspicuous, rigid socii. The female genitalia of Aterpia are characterised by an at least partly spinulose, plate-shaped sterigma with a median notch in its hindmargin leading to ostium, by a smooth ductus bursae with a variable portion of its posterior half lightly and evenly sclerotised, and usually by a single signum, a patch of smallish and often rather distant triangular teeth (rarely without or with two signa).
Description Adults (Figs 411, 412) Wingspan. 10.5–22.0 mm; small to large. Head (Fig. 413). Ocellus well developed. Vertex scales anteriorly barely protruding between antennae. Frons in uppermost part with a narrow transverse band of moderately long anteriorly projecting scales, remainder with minute, appressed and upwardly oriented scales. Proboscis short, little longer than labial palpus. Labial palpus rather long (1.8–2.5× diameter of eye), especially in female, porrect, sinuate and moderately widened; second segment distally moderately expanded with long loose scales along ventral margin and distal part of dorsal edge; terminal segment long. Antenna reaching to middle of forewing; with
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anterior surface unscaled; cilia in male long to very long, 1.0–1.5× diameter of flagellar segment, distinctly verticillate. Thorax. Often with large bipartite posterior crest. Legs with unmodified hind tibia in male. Wings (Figs 49, 411, 412). Forewing index 0.43–0.48; forewing subtriangular, distally moderately widened, with costa evenly curved throughout, apex rounded-acute to slightly projecting, termen straight to lightly sinuate and slightly to strongly oblique; discal cell 0.57–0.59× length of forewing, with narrow accessory cell with chorda from between R1 and R2 and with M-stem to between M2 and M3; with all veins present beyond discal cell and separate except for R3 and R4 that are usually approximated and very rarely stalked (A. praeceps); the majority of species with greyish-blackish and white pattern elements, often with dark wing base, dorsum and apex and with a sinuate white band running from before middle of costa to tornus (A. praeceps, A. protosema), with the dark areas nearly always sprinkled with at least a few deep blue iridescent scales (blue scales rarely found in A. protosema) to entire wing dark and overlaid with blue. Hindwing with cubital pecten and with long scattered hairscales from upper surface of anal area between 1A+2A and anal margin; with Rs and M1 parallel and approximated at base, M2 approximated to M3 at base, with CuA1 distinctly from below angle of cell (M3 and CuA1 connate or stalked in some non-Australian species), and CuA2 from below middle of cell; 1A+2A and 3A at least partly tubular; light to dark grey. Pregenital abdomen. Unmodified. Hindmargin of S8 bilobed in male. Male genitalia (Figs 414, 415). Vinculum a weakly sclerotised band. Tegumen high and inverted V-shaped to ovate in outline, with broad paddle-shaped pedunculi. Uncus a large and broad, rectangular to apically rounded inflated membranous process, with scattered bristles laterally and dorsally. Socii two small bristled membranous lobes laterally below uncus, often entirely reduced (A. protosema). Gnathos two well-sclerotised slender arms distally either fused with anal tube (A. protosema) or joined and produced into a sharply pointed or apically dilated median process, with a lightly sclerotised area of diaphragm between anal tube and anellus projecting as a cone-shaped evagination. Valva simple in outline, rather slender and usually roughly parallel-sided with rounded apex and nearly always with a triangular tooth from middle of costa, with straight costal process often
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concealed behind broadly rectangular invagination from outer valva surface, inner surface of valva flat to lightly concave with a strongly bristled cucullus usually with a band of particularly dense bristles along slightly raised ventral edge, with a parallel band of longer bristles from ventral part of strongly curved outer surface, usually with a patch of large spines from centre of valva beyond basal excavation. Caulis very short but anellus greatly enlarged, especially in ventral part. Aedeagus curved and slender, long to very long; vesica without cornuti. Female genitalia (Fig. 416). Ovipositor lobes elongate-ovate. Sternum 7 with concave hindmargin. Sterigma a variously shaped and at least partly spinulose, sclerotised plate (lamella antevaginalis) with anterior portion often invaginated and with a projecting ridge along its caudal margin with a median excision leading to ostium, which is invaginated behind sterigma. Ductus bursae smooth except for a very short granulate portion immediately below ostium, with an elongate ill-defined colliculum followed by inception of ductus seminalis and by a lightly and evenly sclerotised portion of ductus, remainder of ductus membranous to corpus bursae. Corpus bursae roundish to ovate; signa, if present, one or two small to moderate patches of small and usually rather distantly spaced triangular teeth.
Distribution Holarctic region, Madagascar, India, Java, Sulawesi, Philippine Is, New Guinea, Australia, with the majority of named species from New Guinea. In Australia the genus occurs only in northern Queensland south to the Cairns region.
Biology No host records are available for the Oriental/ Australian group of species, but the Australian localities suggest a rainforest habitat. The Nearctic A. approximana (Heinrich) feeds as a leafroller on Lythrum (Lythraceae) (Heinrich 1926) and two Palaearctic species on Dryas (Rosacae) and Aconitum (Ranunculaceae) respectively (Hannemann 1961).
Remarks Diakonoff (1973) provided a comprehensive treatment of Aterpia, Asaphistis and Apeleptera, with figures of genitalia of numerous species and of head
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Figs 411–416. Aterpia protosema, adults, head and genitalia. 411, 412, adults. 411, , Kuranda, Q. 412, , McIlwraith Range, Q. 413, head, male. 414, 415, aedeagus and male genitalia, Kuranda, Q, slide T2283. 416, female genitalia, Papua New Guinea, slide ANIC 2473. Scale = 2 mm.
and wing venation of sappiroflua, the type species of Metaschistis. Hannemann (1961) figured male genitalia, head and wing venation of A. andereggana, the type species of Aterpia. Photos of adult head and genitalia and a figure of the wing venation of A. praeceps, the type species of Asaphistis, are found in Clarke’s (1958) monograph of Meyrick’s types, and
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Heinrich (1926) provided a detailed description and genitalia illustrations of A. approximana (Heinrich), the type species of Esia Heinrich. Asaphistis is here synonymised with Aterpia rather reluctantly as its characteristic hindwing venation allows easy recognition of this large group in the Oriental/Australian fauna. But the origin of
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CuA1 from distinctly from before the angle of the cell in Asaphistis is probably merely a more plesiomorphic condition with respect to the connate or stalked M3 and CuA1 in Apeleptera and Aterpia. The three genera form a monophyletic group linked by very similar male genitalia with a dorsally hairy uncus, a well-developed gnathos and/or a ventrally sclerotised anal tube and a simple and rather broad valva with a band of bristles along the ventral edge of the outer surface, and by a strongly ciliate male antenna. The Holarctic species including A. andereggana form a subgroup with connate to stalked M3 and CuA1 whilst most of the Oriental/ Australian species described in Asaphistis may well constitute a monophyletic but subordinate clade with long palpi, square uncus and an additional band of bristles along the ventral margin of the inner surface of the valva. The little known and monobasic Apeleptera described from Taiwan seems to be intermediate between the two groups. In accordance with the above synonymies the following species are transferred to Aterpia: asema (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 275 (Asaphistis) [Java]. Comb. nov. catarrhactopa (Meyrick, 1938), Trans. R. Entomol. Soc. Lond. 87: 512 (Argyroploce) [New Guinea]. Comb. nov. cretata (Diakonoff, 1953), Verh. K. Ned. Akad. Wet., Nat., ser. 2, 49: 103 (Metaschistis) [New Guinea]. Comb. nov. gypsopa (Diakonoff, 1953), Verh. K. Ned. Akad. Wet., Nat., ser. 2, 49: 101 (Metaschistis) [New Guinea]. Comb. nov. haematina (Diakonoff, 1953), Verh. K. Ned. Akad. Wet., Nat., ser. 2, 49: 98 (Metaschistis) [New Guinea]. Comb. nov. hemicapnodes (Diakonoff, 1953), Verh. K. Ned. Akad. Wet., Nat., ser. 2, 49: 97 (Metaschistis) [New Guinea]. Comb. nov. hemicyclica (Diakonoff, 1953) Verh. K. Ned. Akad. Wet., Nat., ser. 2, 49: 96 (Metaschistis) [New Guinea]. Comb. nov. lucifera (Meyrick, 1909), J. Bombay. Nat. Hist. Soc. 19: 589 (Proschistis) [India]. Comb. nov. maturicolor (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 269 (Asaphistis) [New Guinea]. Comb. nov. nobilis (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 268 (Asaphistis) [New Guinea]. Comb. nov. phanerops (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 265 (Asaphistis) [Java]. Comb. nov. praeceps (Meyrick, 1909), J. Bombay Nat. Hist. Soc. 19: 590 (Asaphistis) [India: Assam]. Comb. nov.
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199
purpurascens (Diakonoff, 1953), Verh. K. Ned. Akad. Wet., Nat., ser. 2, 49: 99 (Metaschistis) [New Guinea]. Comb. nov. sappiroflua (Diakonoff, 1953), Verh. K. Ned. Akad. Wet., Nat., ser. 2, 49: 100 (Metaschistis) [New Guinea]. Comb. nov. semnodryas (Meyrick, 1936), Exot. Microlepid. 4: 613 (Argyroploce) [Taiwan]. Comb. nov. The two species A. lucifera and A. hemicyclica are only tentatively transferred to Aterpia. The signa of the female holotype of A. lucifera strongly refute inclusion in Aterpia, but an alternative assignment is not possible without male genitalia. Male specimens of A. hemicyclica or a closely related species in the ANIC suggest that A. hemicyclica and Proschistis amphibola Diakonoff are congeneric and more closely related to Cephalophyes Diakonoff than to Aterpia.
Constituent species Twenty seven described species of Aterpia are recorded worldwide and several unnamed species are known from New Guinea. One of the species named from New Guinea extends into Australia: protosema (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 274 (Asaphistis). [New Guinea]. Comb. nov.
Proschistis Meyrick (Figs 50, 417–423) Proschistis Meyrick 1907b, J. Bombay Nat. Hist. Soc. 17: 731. Type species: Proschistis zaleuta Meyrick, 1907b, ibid. 17: 731, by monotypy. [Sri Lanka.] Sporocelis Meyrick, 1907b, J. Bombay Nat. Hist. Soc. 17: 732. Type species: Sporocelis marmaropa Meyrick, 1907b, ibid. 17: 732, by monotypy. [Sri Lanka.]
Diagnosis Proschistis species are dark, rather dull, mediumsized to large moths with a leaden bluish tinge, the species found in Australia with broad, subtriangular, dark grey-brown forewings speckled with pale leaden bluish grey in basal and distal thirds. Proschistis has longish, sinuate and strongly widened labial palpi and long, verticillate antennal cilia in male. The wing venation is unremarkable, without any diagnostic features. The male of
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Figs 417–420. Proschistis polyochtha, adults, head and abdomen. 417, 418, adults. 417, , Kuranda, Q. 418, , Tully, Q [image reversed]. 419, head, male. 420, male abdomen with eversible scale tuft (arrows). Scale = 2 mm.
P. polyochtha is characterised by a unique secondary sexual character, two long membranous tubes filled with scales with enlarged sockets invaginated between the third and fourth abdominal segments and everted as fingerlike processes covered with white scales. The male genitalia with long, naked and strongly sclerotised socii, a distally dilated and rounded valva with a spinulose tumescence at its base and a variably developed bristled and raised transverse ridge beyond basal excavation, and with a characteristically modified anellus, are diagnostic for the genus. The female genitalia of Proschistis with a spinulose, concave sterigma, a ductus bursae with a granulate posteriormost portion, a variably developed colliculum and a patch of sclerotisation below midlength and with only a single scobinate signum are very similar to those of Aterpia. For the two species of the two genera so far recorded from Australia, the combination of a modified 8th segment and a very small signum will identify Proschistis polyochtha whilst presence of a well-
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developed lamella antevaginalis with a conspicuously notched posterior margin and a large signum characterises Aterpia protosema.
Description Adults (Figs 417, 418) Wingspan. 12–25 mm; medium-sized to large. Head (Fig. 419). Ocellus well developed. Chaetosema with very long bristles. Frons with shortish, anteriorly projecting scales in upper third, remainder with minute, appressed and upwardly oriented scales. Proboscis short, little longer than labial palpus. Labial palpus rather long (1.8–2.2× diameter of eye), porrect, sinuate and strongly widened; second segment roughly inverted teardrop-shaped, distally strongly widened with long loose scales ventrally and dorsally along distal third and broadly rounded apex; terminal segment long. Antenna reaching to middle of forewing; with anterior surface unscaled, cilia in male long to very
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Figs 421–423. Proschistis polyochtha, genitalia. 421, female genitalia, nr Tully, Q, slide ANIC 2732. 422, 423, aedeagus and male genitalia, Kuranda, Q, slide ANIC 2731.
long, 1–3× diameter of flagellar segment, distinctly verticillate. Thorax. With posterior crest. Legs with unmodified hind tibia in male. Wings (Figs 50, 417, 418). Forewing index 0.37– 0.52; forewing subrectangular (P. zaleuta) to broadly triangular (P. polyochtha Diakonoff) with distally bent costa (P. marmaropa (Meyrick)); costa evenly curved or bent or curved in distal third only, apex rounded-acute and lightly projecting, termen roughly straight, from nearly vertical to strongly oblique; discal cell 0.56–0.70× length of forewing, triangular in P. polyochtha, with a narrow to triangular accessory cell with chorda from between R1 and R2 to between R5 and M1 (P. polyochtha) and a well-developed M-stem; all veins present and separate beyond discal cell; R3 and R4 and R5 usually closely approximated at base (less so in P. polyochtha); dull, dark brownish or bluish greyblack, extensively marbled with whitish or leaden bluish grey). Hindwing with cubital pecten and scattered long hair scales on upper surface along 3A; with Rs and M1 parallel and closely
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approximated at base, M2 bent towards M3 at base, but not closely approximated in the broad-winged P. polyochtha and P. marmaropa, M3 and CuA1 connate from angle of cell, CuA2 from beyond middle of cell, 1A+2A and 3A well developed but not tubular in P. polyochtha, base of 1A absent in P. polyochtha; hindwing dark grey, often translucent at base. Pregenital abdomen. Male of P. polyochtha with two long, membranous eversible tubes set with scales on large modified sockets and invaginated laterally one on each side between third and fourth segment, everted as a finger-like process covered with snow-white scales (Fig. 420). Sternum 8 in male P. polyochtha with large inverted-triangular sclerotisation along midline and bilobed hindmargin with semicircular median excision; T8 with large inverted-T-shaped sclerotisation. Male genitalia (Figs 422, 423). Vinculum a broad but weakly sclerotised band. Tegumen rather broad and short, pedunculi in P. polyochtha with huge medially projecting semicircular apodemes for m4. Uncus not differentiated from apex of tegumen
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that is a large, flat, triangular to roundedrectangular process with sharp and strongly sclerotised margin in P. polyochtha and scattered hairs along rim of dorsal surface. Socii strongly sclerotised and naked processes of various shapes, in P. polyochtha rigid, porrect and strongly sinuate slender arms with ladle-shaped, enlarged and flattened tips. Gnathos two ill-defined sclerotised bands fused with sclerotised basal surface of bipartite anal tube. Valva dilated towards rounded apex and with several distinct tufts or patches of bristles; costal process long and curved, partly obliterated by an irregularly dome-shaped densely spinulose large swelling on base of valva below costa in P. polyochtha, with a variably developed raised transverse ridge across valva beyond basal excavation set with dense and often irregular spines (medially interrupted and reduced to two small irregular sharply ridged humps with scattered spines in P. polyochtha), with a dense band of long spines along ventral margin from base to beyond middle and with cucullus either evenly bristled (P. zaleuta, P. marmaropa) or with two discrete tufts and a very large circular patch of much denser bristles in basal half of cucullus (P. polyochtha). Caulis very short but anellus (in P. polyochtha) greatly enlarged with a broad, U-shaped ventral band and a darker sclerotised concave dorsal portion ending in a strongly sclerotised vertical point reaching towards base of anal tube. Aedeagus from short to moderately long, curved and sclerotised along ventral surface only (P. polyochtha), vesica without cornuti. Female genitalia (Fig. 421). Ovipositor lobes very narrow, elliptical. Eighth segment in P. polyochtha modified, swollen and strongly sclerotised, especially dorsally. Sternum 7 with a concave hindmargin. Sterigma either a strongly spinulose cup-shaped depression with several lateral processes (P. zaleuta) or a weakly spinulose membranous funnel (P. polyochtha). Ductus bursae granulate immediately below ostium and with an elongate ill-defined colliculum (only faintly indicated in P. polyochtha), remainder smooth and membranous with a faintly sclerotised patch close to inception of ductus seminalis below midlength. Corpus bursae roundish to ovate, granulate; with a single signum, a small to moderate patch of strongly sclerotised granulation or small triangular teeth.
genus has only been found in the Cairns region, in northern Queensland.
Biology No biological information is available beyond the fact that the two Australian localities are in rainforest habitats.
Remarks Taken together, Diakonoff (1973) and Clarke (1958) provided a comprehensive treatment of the genus, with the former giving extensive descriptions, references and genitalia drawings and the latter figuring wings, genitalia and wing venation of the type species of Proschistis and Sporocelis. The monophyly of Proschistis is demonstrated by several uniquely derived features in the male genitalia such as the rigid, sclerotised socii, the raised bristled ridge across the valva and probably the spinulose tumescence at its base. Proschistis and Aterpia Guenée are clearly closely related if not sister groups. They share strongly ciliate antennae in the male, similar labial palpi, a basically similar tegumen with a dorsally bristled and little differentiated uncus, gnathos arms ending in the sclerotised ventral part of a dorsoventrally bipartite anal tube and a greatly enlarged, concave anellus, and they have nearly indistinguishable female genitalia. The characteristically bipartite anal tube of Proschistis and Aterpia has given rise to some contradictory morphological descriptions with the sclerotised ventral surface (often forming a partly separate parallel tube) mistaken for the gnathos proper. Diakonoff (1973) also treats the naked rigid socii of P. polyochtha as part of a trifid gnathos but they readily fit in with the sclerotised socii of the other two species. ‘Proschistis’ amphibola Diakonoff (Diakonoff 1973), known from the female holotype only, is not congeneric with P. zaleuta. The modified 8th segment in the females of both P. amphibola and P. polyochtha is probably convergent. The possible relationship of P. amphibola is further discussed under Aterpia.
Constituent species Distribution Sri Lanka, India, Taiwan, China, Sulawesi, New Guinea, Australia and Sierra Leone. In Australia the
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Eight named species of Proschistis are recognised by Brown (2005), with one of them extending into Australia:
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polyochtha Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 282 (Proschistis). [New Guinea.]
Rhodacra Diakonoff (Figs 51, 424–429) Rhodacra Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 180 (key), 286. Type species: Argyroploce pyrrhocrossa Meyrick, 1912b, J. Bombay Nat. Hist. Soc. 21: 874, by monotypy and original designation. [Assam.] Corethrarcha Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 181 (key), 287. Type species: Argyroploce rupifera Meyrick, 1909, J. Bombay Nat. Hist. Soc. 19: 602, by monotypy and original designation. [Assam.]
Diagnosis Rhodacra species are medium-sized olethreutine moths with subrectangular to subtriangular forewing, in males with anal margin of hindwing forming a spindle-shaped roll and hind tibia with a very long hairpencil. Rhodacra pyrrhocrossa, the type species extending into Australia, is brownish and finely dusted with whitish throughout, especially on dorsum, without any well-defined markings except white costal strigulae and a conspicuous reddish fringe. Dactylioglypha Diakonoff also has a reddish fringe and overall similar colouring, but its characteristic wing shape and pattern allow easy separation. The male genitalia are diagnostic for the genus: the laterally expanded, T-shaped or trilobed uncus and a valva separated into a densely spined ventral lobe and a crescentic distal part by a deep semicircular ventral excision are both unique features. In the female Rhodacra is characterised by the two very large signa formed by a large number of triangular, scalelike flat teeth. Some species of Metrioglypha Diakonoff and Aterpia Guenée have signa of a similar structure, with dense short triangular teeth giving the appearance of a patch of scales, but they are always much smaller than those of Rhodacra and in Aterpia usually single.
Description Adults (Fig. 424) Wingspan. 12–17 mm; medium-sized.
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203
Head (Figs 425). Ocellus large. Vertex with conspicuously pale-tipped scales in R. pyrrhocrossa, barely projecting between antennae. Frons in uppermost part with shortish anteriorly projecting scales, remainder with minute, appressed and upwardly oriented scales. Proboscis short, little longer than labial palpus. Labial palpus rather small (1.7–1.8× diameter of eye), strongly sinuate, upcurved and moderately widened; second segment strongly curved and gradually widened towards apex, with long loose scales along ventral margin; terminal segment porrect and rather small, often concealed by apical scales of preceding segment. Antenna reaching to middle of forewing; with anterior surface unscaled; cilia in male minute. Thorax. With or without a slight posterior crest. Legs with hind tibia in male with long appressed scales in R. pyrrhocrossa, forming a deep groove containing a large hairpencil reaching from base to apex of tibia (Fig. 426), and with a pointed tuft of scaling from inner surface extending beyond apex of tibia, or male hind tibia smoothly shortscaled with a short hairpencil from base of tibia (R. rupifera). Wings (Figs 51, 424). Forewing index 0.38– 0.43; forewing subrectangular to subtriangular, with faintly and evenly curved costa, roundedrectangular to slightly produced apex and roughly vertical and lightly sinuate termen; discal cell with narrow accessory cell with chorda from between R1 and R2 to between R5 and M1, with R-stem to between M2 and M3; with all veins present and separate beyond discal cell, and with CuA1 strongly curved and bent towards M3 at wing margin; R. pyrrhocrossa without any distinct markings except white costal strigulae on a blackish brown costa paralleled by two slightly longer metallic silvery lines beyond middle of costa and conspicuously reddish fringe on termen and tornus, remainder of forewing brownish and finely sprinkled throughout with whitish, with several parallel horizontal blackish lines in distal part of wing, followed by an erect patch of silvery scales towards termen. Hindwing with cubital pecten and scattered long hairscales from wing base between 1A+2A and anal margin, in male with anal margin thickened and bent downward to form a large spindle-shaped roll covered with hairscales; with Rs and M1 parallel and closely approximated at base, M2 approximated at base to connate M3 and CuA1; 1A+2A and 3A present; dark greyish. Pregenital abdomen. Unmodified. Hindmargin of S8 in male only faintly bilobed in R. pyrrhocrossa.
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Figs 424–429. Rhodacra pyrrhocrossa, adult, head and genitalia. 424, adult, , Daintree, Q. 425, head, male. 426, tibial pencil, male. 427, female genitalia, Daintree, Q, slide ANIC 2471. 428, 429, male genitalia and aedeagus, Daintree, Q, slide T2304. Scale = 2 mm.
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Male genitalia (Figs 428, 429). Vinculum a moderate band. Tegumen high and ovate in outline, with pedunculi widened on inner side (? muscle apodeme). Uncus a distally dilated curved median process, with apex naked and T-shaped in R. pyrrhocrossa, hairy beneath and with two lateral and a distal lobe in R. rupifera. Socii hairy lateral knobs or lobes. Gnathos faintly sclerotised short lateral bands connected with a flat and large but weakly sclerotised subscaphium. Valva long and highly modified, with a deep ventral cleft beyond middle; with longish and sinuate costal process, with ill-defined sacculus area but with basal part of cucullus separated from hairy, crescent-shaped distal part by a deep semicircular ventral excision, with either a curved band of increasingly large spines running from edge of basal excavation to triangular ventral lobe (R. pyrrhocrossa), or ventral lobe broadly rounded with a large circular area of dense spines and bristles. Caulis long and anellus a very broad band. Aedeagus short and straight, narrowest in middle; vesica without cornuti. Female genitalia (Fig. 427). Ovipositor lobes subtriangular (R. pyrrhocrossa) to elongate-ovate (R. rupifera). Sternum 7 with straight to weakly concave hindmargin. Sterigma a large spinulose, irregularly dome-shaped area with a median split containing ostium, in R. pyrrhocrossa conspicuously reticulate and with a central erect diamond-shaped raised rim around ostium flanked by two oblique and ventrally converging sharp ridges. Ductus bursae in R. pyrrhocrossa very long, with moderately long colliculum followed by a long thin and smooth portion to inception of ductus seminalis, then abruptly widened, lightly transversely wrinkled and coiled; ductus in R. rupifera much shorter, with a longer colliculum and with anterior dilated half straight and sclerotised, with some transverse wrinkles only at entrance into bursa. Corpus bursae ovate, granulate or, towards signa, finely dentate; signa two large elongate patches of numerous parallel rows of triangular, scale-like teeth, extending over 36: of bursa length.
205
Remarks The uniquely modified valva of Rhodacra pyrrhocrossa and Rhodacra rupifera (Meyrick), together with the laterally expanded apex to their uncus, albeit hairy and with an additional distal lobe in R. rupifera, are compelling synapomorphies linking the two taxa. Given the numerous further shared derived characters, such as the spindleshaped ‘dorsal roll’ of the male hindwing and the structure of the signa, the minor differences in wing shape and female genitalia between Rhodacra and Corethrarcha cannot justify two separate monobasic genera. Diakonoff (1973) provided a comprehensive treatment of both Rhodacra and Corethrarcha, with illustrations of genitalia, head and wing venation. However, his figure of the wing venation of R. rupifera (Diakonoff 1973: fig. 447) does not agree with his description and must erroneously have been exchanged for the figure of that of Proschistis zaleuta on the same page. If the numbers are exchanged (Diakonoff 1973: figs 445 and 447) the venation figured for both species agree with the descriptions given. Photos of the types of both species of Rhodacra were given by Clarke (1958) as Olethreutes pyrrhocrossa and O. rupifera. The specimens of R. pyrrhocrossa available from Australia are smaller than the type series from India, but no other differences could be found.
Constituent species Three species of Rhodacra are known, one of these occurring in Australia: pyrrhocrossa (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 874 (Argyroploce). [India: Assam.]
BACTRINI Falkovitsh Bactrini Falkovitsh, 1962, Entomol. Obozr. 41: 884. Type genus: Bactra Stephens, 1834.
Distribution India (Assam), Thailand, Borneo, Taiwan, Australia. In Australia the genus has been collected in the McIlwraith Range and on the Daintree R. in northern Queensland.
Biology No host records are available, but the Australian localities are in rainforest.
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Status remarks Falkovitsh (1962) established the Bactrini as a tribe parallel to the Olethreutini, based on absence of any secondary sexual scent structures. This has generally been accepted, except for Diakonoff (1973), who treated the group has a subtribe within the Olethreutini and included several closely related Oriental genera. Dang (1990) provided convincing
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arguments to include the genus Endothenia in the Bactrini rather than treat it as a separate subtribe (Diakonoff 1973), tribe (Kuznetsov and Stekolnikov 1977) or include it in the Gatesclarkeanini (Horak and Brown 1991). Given the presence of a tibial pencil in Endothenia and several possible synapomorphies linking the Neopotamia-group with Bactra and Endothenia that are discussed in the Remarks for the relevant genera, the Bactrini could well be subordinate within the Olethreutini, close to the Neopotamia and Gatesclarkeana-groups. The preliminary cladistic analysis certainly suggests this, but it seems preferable to wait for molecular evidence before yet again changing the status of the group.
Evidence for monophyly Falkovitsh (1962) based the Bactrini on the lack of any secondary sexual scent structures in the male while Diakonoff (1973) defined his Bactrae primarily on the structure of the single signum. Dang (1990) described in detail four synapomorphies linking Bactra and Endothenia: 1, the apically setose and sharply recurved uncus with a depression at its dorsal base; 2, the short, downcurved tegumen; 3, the unusual position of the valva with a reduced costal process, and 4, the reduced tarsal setae. In addition, both genera have at most a single, very similar signum, the larvae are borers, and Z10–14 carbon acetate has been identified as the pheromone of Bactra verutana Zeller and as an attractant for Endothenia (Horak and Brown 1991).
Distribution Bactra is found worldwide and Endothenia is widely distributed in the Holarctic, Oriental and Australian regions, with the other genera restricted to the Oriental and Australian regions.
Biology The larvae of all three genera here discussed are borers, with Bactra living as a stem borer on Cyperaceae, Juncaceae and Poaceae and Syntozyga ephippias (Meyrick) in stems of a species of Commelinaceae (Fletcher 1932), also a monocotyledonous family. Species of Endothenia are borers in stalks, roots and seed heads of dicotyledonous plants.
considering this a primary absence rather than a secondary loss. Given the presence of a tibial pencil in the closely related Endothenia, this assumption clearly has to be rejected. Bactrini have been treated as a rather basal group due to several plesiomorphic characters, namely their small costal process, the small basal excavation and a pheromone based on a 14 carbon compound (Horak and Brown 1991). However, the boring feeding mode of the larvae is derived at both family and subfamily level, and numerous aspects of the genitalia in both sexes are clearly apomorphic. The spined process from the base of the valva ventrally to the basal excavation in the Bactrini is reminiscent of similar-looking structures in Gatesclarkeana and the Neopotamiagroup, as is the complex and deeply invaginated sterigma. The single signum of the Bactrini also is very similar to some found in the Neopotamiagroup where several genera have a single signum only. Reduced tarsal setae have been found in Olethreutini genera of several genus-groups, such as Zomariana Diakonoff, Ophiorrhabda Diakonoff and Temnolopha Lower. Examination of further character sets, in particular molecular studies, will be needed to confirm the position of the Bactrini as a separate tribe rather than a subordinate group within the Olethreutinae. Diakonoff (1973) recognised the close relationship between Bactra and Endothenia Stephens, but considered presence of male scent organs in the latter and the different facies of the two genera sufficient to establish a separate subtribe Endotheniae. Kuznetzov and Stekolnikov (1977) found the tergal extensor muscles of the valva (m2) to be absent in Endothenia marginana (Haworth) and based tribal rank for the group on this loss. Subsequently, m2 was also found to be lacking in a highly derived member of the Gatesclarkeanini (Kuznetzov and Stekolnikov 1984), and the two tribes Endotheniini and Gatesclarkeanini were joined into the supertribe Gatesclarkeanidii. This association was accepted by Horak and Brown (1991), but at tribal level with Endothenia included in the Gatesclarkeanini. However, Dang (1990) found m2 to be present in three Nearctic Endothenia species and identified convincing synapomorphies to include Endothenia in the Bactrini.
Bactra Stephens (Figs 53, 430–439)
Remarks Falkovitsh (1962) accorded the group tribal level based on the lack of any male scent organs,
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Aphelia Stephens, 1829, Nom. Br. Insects: 47. Type species: Tortrix plagana Haworth, 1811, Lepid.
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Br.: 470, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 23. [Britain.] A junior homonym of Aphelia Hübner, [1825] (Lepidoptera: Tortricidae). Bactra Stephens, 1834, Illus. Br. Entomol. (Haustellata) 4: 124. Type species: Tortrix plagana Haworth, 1811, Lepid. Br.: 470, by subsequent designation by Curtis, 1836, Br. Entomol. 13: folio 599. [Britain.] Leptia Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 169. Type species: Tortrix lancealana Hübner, [1799], Samml. Eur. Schmett. 7: pl. 13, fig. 80, by monotypy. [Europe.] A junior homonym of Leptia Dejean, 1833 (Coleoptera). Chiloides Butler, 1881, Ann. Mag. Nat. Hist. 5(7): 392. Type species: Chiloides straminea Butler, 1881, ibid. (5)7: 393, by monotypy. [Hawaii.] Noteraula Meyrick, 1892, Trans. NZ Inst. 24: 217. Type species: Noteraula straminea Meyrick, 1892, ibid. 24: 217, by monotypy. [New Zealand.] Noteraula straminea Meyrick, 1892, is a junior secondary homonym of Chiloides straminea Butler, 1881. The objective replacement name of N. straminea Meyrick is Bactra noteraula Walsingham, 1907, in Sharp, Fauna Hawaii 1(5): 689. Nannobactra Diakonoff, 1956, Zool. Verh. (Leiden) 29: 52. Type species: Bactra phaulopa Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 253, by original designation. [New Guinea: Kei Is.] Established as a subgenus of Bactra Stephens, 1834. Spinobactra Diakonoff, 1963, Tijdschr. Entomol. 106: 291. Type species: Bactra (Spinobactra) spinosa Diakonoff, 1963, ibid. 106: 291, figs 2–4, by original designation. [South Africa.] Established as a subgenus of Bactra Stephens, 1834. Protobactra Diakonoff, 1964, Zool. Verh. (Leiden) 70: 4. Type species: Bactra diachorda Meyrick, 1932, Exot. Microlepid. 4: 308, by original designation. [Brazil.] Syn. nov.
Diagnosis Bactra species are mostly medium-sized, narrowwinged, ochreous to grey-brown moths with a forewing with pointed apex, fine longitudinal striation and often a straight dark streak from base to apex or at least from a whitish dot on closing vein to apex, anteriorly sometimes enhanced by an adjoining and parallel pale streak, more rarely with
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transverse, outwardly angled pattern elements at about 56: and 36: wing or derivations thereof. Bactra has moderate to long labial palpi, sinuate, porrect and with a distally strongly widened second segment with particularly long and parallelprojecting scales from apical third of ventral margin, often concealing terminal segment. The tarsal segments have only weak distal setae. The wing venation is unremarkable for an olethreutine, with all veins separate beyond discal cell except for M3 and CuA1 in the hindwing that may be connate. The second abdominal tergite has longitudinal sclerotised rods along its lateral margins from base to beyond middle of segment, continuing the course of the strengthened lateral margins of T1. The male genitalia are diagnostic for the genus with a spatulate uncus fringed with flattened pegs, a valva deeply divided into a large, roundish sacculus lobe usually with prominent spines and a cucullus of either triangular shape with thick spines along margin or more elongate and rather evenly bristled. Often a third lobe (valvula) is present (in all Australian species), usually highly characteristic as it is fringed with similar flattened pegs as the uncus. The female genitalia are less uniform with an often complex sterigma usually consisting of a semicircular, projecting ledge from lamella postvaginalis with the ostium in a deeply invaginated, semicircular to crescentic pocket, often with a median split. The single signum of Bactra, a small reticulate or scobinate plate, invaginated to form a small pocket, is very similar to that of Endothenia.
Description Adults (Fig. 430–432) Wingspan. 9–23 mm; from small to large, mostly medium-sized. Head (Fig. 433). Ocellus well developed. Uppermost part of frons with a transverse band of long, projecting scales, pointing forward or slightly downward, remainder with minute, appressed and upwardly oriented scales. Maxillary palpus minute, appressed, hardly visible in undissected specimens, two or three-segmented in B. venosana (Zeller), with lateral bristles on first and scales on second segment. Proboscis very short, from 56O length of labial palpus to as long as palpus (Australian species). Labial palpus moderately long to very long (1.5–3.0× diameter of eye), sinuate, porrect and strongly widened; second segment sinuate and distally strongly dilated to an often triangular shape by very long scales from ventral margin and distally from dorsal edge, with scales on distal third
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of ventral margin particularly long and parallelporrect, often concealing the long, thin, drooping terminal segment. Antenna usually rather long, often reaching to over middle of forewing; anterior surface devoid of scales; cilia in male short, at most 56O width of flagellar segments. Thorax. Without crest. Legs without any modifications in male, but in both sexes with tarsal segments distally without the strong setae normally present in tortricids. Wings (Figs 53, 430–432). Forewing index 0.32– 0.37; forewing long and very slender; with evenly curved costa, acute and projecting apex, and oblique to strongly oblique straight or weakly sinuate termen; discal cell 0.62–0.63× length of forewing, with a variably sized accessory cell with a nontubular chorda (in Australian species) from between R1 to R2 to usually somewhere between R4 and R5, with M-stem only a faint trace in membrane running to between M2 and M3; with all veins present and separate beyond discal cell; CuP present at wing margin; ground colour from light yellow-ochreous to brown or dark greyish, apart from very fine longitudinal parallel striation and costal strigulation often unmarked or with several genus-specific pattern elements that may be very variably developed within a given species; a small whitish spot on closing vein posteriorly often edged by a dark halfring, a straight dark longitudinal streak from base to apex or at least from pale spot to apex, anteriorly sometimes enhanced by an adjoining parallel pale streak with wing behind streak darkened, more rarely with transverse outwardly angled pattern elements at about 56: and 36: wing-length or derivations thereof (B. blepharopis Meyrick). Hindwing with well-developed cubital pecten but weak axillary tuft, with Rs and M1 connate or stalked, M2 bent close to very close to M3 at base and CuA1 from just before angle of cell or connate with M3; CuP present at wing margin, 1A+2A and 3A at least partly tubular; whitish to grey. Pregenital abdomen. Longitudinal, sclerotised rods along lateral margins of T2 from base to beyond middle of segment, continuing the course of the strongly sclerotised lateral margins of T1; tortricoid apodemes posteriorly continuous with rod-like sclerotisations across S2 or at least vestiges of sternal rods. In male hindmargin of S8 medially concave to deeply excavated. Male genitalia (Figs 434, 435, 438, 439). Vinculum an often very broad and partly weakly sclerotised band, sometimes W-shaped and in B. venosana (Zeller) the upcurved median portion with large spines on posterior margin, B. optanias Meyrick with a T-shaped sclerotised bar along
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ventral midline of intersegmental membrane, arising from margin of vinculum and suggesting a saccus. Tegumen usually broad and dorsally strongly convex, well sclerotised. Uncus a welldeveloped, curved, spatulate process fringed with flattened pegs in distal, dilated portion, with a dorsal depression at its base. Socii lateral membranous lobes set high on apex of tegumen and covered with long scales or bristles. Gnathos at best a very weak lateral trace; ventral surface of anal tube sclerotised and very broad in some species (B. blepharopis Meyrick, punctuate in B. venosana). Valva highly modified and very diverse within genus, with very short costal process and with sacculus greatly enlarged into a complex lobe and with a third lobe (valvula) arising from boundary between sacculus and cucullus in some subgenera (Chiloides, Nannobactra); sacculus with a central punctuate area carrying one or several stout thorns (subgenus Bactra), or with its distal half upcurved, punctuate and with scattered spines (Nannobactra) or smooth except for possible hairy patches (B. optanias) but with some short to very slender spines, especially along margin (Chiloides); cucullus an elongate, usually finer bristled lobe in the subgenera Bactra and Chiloides, but much broader, subtriangular and set with huge spines along distal margin in Nannobactra (B. blepharopis); intermediate lobe from very short, triangular and with irregular marginal spines (B. capnopepla Turner) to long, distally strongly dilated and fringed with flattened pegs (valvula of Diakonoff). Juxta of triangular shape only in subgenus Bactra, in Chiloides and Nannobactra juxta increasingly wider and shorter, finally fused with caulis and anellus into an indistinctly sclerotised transverse structure (B. optanias) with aperture for aedeagus. Aedeagus a curved tube, from short and stout to long and slender, apex ventrally membranous and often with one or several thorns in membranous tip (B. capnopepla, B. optanias), and base usually with a coecum-like bulge dorsally. Female genitalia (Figs 436, 437). Ovipositor lobes diversely shaped with posterior lobes from rounded to acute. Sternum 7 often strongly modified and medially integrated into sterigma, hindmargin a membranous fold overlying invaginated parts of sterigma. Sterigma very diverse, varying from a membranous transverse fold posteriorly to funnel-shaped ostium (subgenus Bactra) to the lamella postvaginalis projecting as a transverse, semicircular ledge (spinulose in B. blepharopis) with ostium at base of deeply invaginated, often semicircular and at least partly sclerotised flat pocket, sometimes with a median
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Figs 430–433. Bactra, adults and head. 430, 431, B. blepharopis, Rockhampton, Q. 430, . 431, . 432, B. venosana, , Springsure, Q [image reversed]. 433, head, B. venosana, male. Scale = 2 mm.
split flanked by folds (B. venosana, B. testudinea Turner). Ductus bursae diverse, posteriormost part with a variably developed bipartite colliculum or evenly sclerotised (often weakly) to form a posteriorly faintly widened funnel (subgenus Bactra) or an anteriorly curved tube (B. optanias), remainder of ductus membranous, coiled or recurved at inception of ductus seminalis and gradually widening towards corpus bursae from this point. Corpus bursae ovate to roundish; with a single signum, a small scobinate or reticulate, partly invaginated semicircular to rectangular plate, forming a small pocket.
Distribution Bactra is distributed worldwide. In Australia the genus is present in all states.
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Biology All confirmed host records for Bactra refer to Cyperaceae, Juncaceae and Poaceae, particularly the former two families, with the larva living in the stem. Diakonoff (1973) reports Lavatera (Malvaceae) as foodplant of B. (Bactra) bactrana (Kennel), but this will need confirmation. Cyperus rotundus is the most important host of B. venosana.
Remarks Diakonoff (1956, 1962, 1963, 1964) revised Bactra on a world scale except for the Australian fauna, illustrating the genitalia whenever possible. Adults and genitalia of numerous types were figured by Clarke (1958), and several local fauna treatments dealt with Bactra at various levels (Heinrich 1926;
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Hannemann 1961; Bentinck and Diakonoff 1968; Zimmerman 1978; Bradley et al. 1979, Miller 1987, Razowski 1989). Bactra is a well-defined, highly autapomorphic group with unmistakable male genitalia, reduced tarsal setae and usually very characteristic external appearance. Based on male genitalia differences Diakonoff (1956) subdivided Bactra into four subgenera (Bactra, Noteraula, Chiloides and Nannobactra) in the first part of his comprehensive revision of the genus, subsequently (Diakonoff 1963) adding a fifth, the African Spinobactra. The five taxa are clearly defined, providing a useful framework for the classification of the innumerable and superficially often very similar-looking species of Bactra. It is doubtful, however, that they all represent monophyletic groups, but as only two subgenera, Chiloides and Nannobactra, are found in Australia, the present study is not able to address this question. The widely distributed subgenus Bactra is characterised by absence of a valvula and usually by a swollen sacculus with some stout spines in a punctuate central patch (Palaearctic, New World, Africa, Oriental region to New Guinea). Noteraula is restricted to New Zealand and has a row of strong spines across its valva between the sacculus and cucullus instead of a valvula with distal spines or pegs. The sterigmal area in Noteraula is tumid, somewhat mamillate and densely chitinised rather than sclerotised (Dugdale personal communication 2005). Chiloides and Nannobactra both are found in the New as well as the Old World, predominantly in tropical areas, and both are characterised by a valvula usually with a distal row of pegs. Chiloides usually has an elongate cucullus with at most some short spines alongside its marginal bristles whereas Nannobactra has short huge spines along the cucullus margin. The South African Spinobactra has a valva similar to that of Bactra but with a band of marginal spines on the sacculus instead of stout ones in its centre. Protobactra Diakonoff from Brazil is here synonymised with Bactra. The transtilla supposedly diagnostic for this taxon is merely the displaced vinculum that is conspiciously absent in the genitalia figure of the holotype of Bactra diachorda Meyrick (Diakonoff 1964: fig. 1). The following species is returned to Bactra: diachorda Meyrick, 1932, Exotic Microlepid. 4: 308 (Bactra). [Brazil.] Comb. rev. Even though Syntozyga Lower has lost the uncus there are sufficient similarities in the valva and juxta shape, the invaginated sterigma and the shared presence of only weak tarsal setae to treat Syntozyga as the sister group of Bactra.
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Furthermore, Syntozyga has been reared from stems of Commelina (Fletcher 1932), a monocotyledon. Diakonoff (1973) points to the obvious similarity of Parabactra Meyrick and Bactra in wing pattern and shape of labial palpi but concludes that the differences in the male genitalia ‘show that the relationship is only slight’. Diakonoff (1964) figured and discussed several species found in Australia, namely B. capnopepla Turner, B. venosana (Zeller) and B. optanias Meyrick. An erroneous report of the introduction of B. venosana into Australia to control nut grass (Diakonoff 1956; as Bactra truculenta Meyrick) is based on the introduction of Hawaiian material into quarantine only. The species was never released but is present in Australia. Except for B. blepharopis Meyrick, which belongs to the subgenus Nannobactra, all named Australian species are members of the subgenus Chiloides.
Constituent species According to Brown (2005), 102 species of Bactra are known worldwide, 10 of which are from Australia: capnopepla Turner, 1946, Trans. R. Soc. S. Aust. 70(2): 211 (Bactra). [Qld: Noosa.] patris Diakonoff, 1956, Zool. Verh. (Leiden) 29: 24 (Bactra (Chiloides)). [East Java.] ablabes Turner, 1946, Trans. R. Soc. S. Aust. 70(2): 212 (Bactra). [NSW: Brunswick Heads.] scalopias Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 255 (Bactra). [NT: Darwin.] anthracosema Turner, 1916, Trans. R. Soc. S. Aust. 40: 528 (Bactra). [NT: Darwin.] psammitis Turner, 1916, Trans. R. Soc. S. Aust. 40: 527 (Bactra). [SA: Adelaide.] ammopastea (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 210 (Eucosma). [NSW: Broken Hill.] venosana (Zeller, 1847), Isis Oken, Leipzig 1847: 738 (Phoxopteris). [Sicily.] truculenta Meyrick, 1909, J. Bombay Nat. Hist. Soc. 19: 586 (Bactra). [India.] scythropa Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 254 (Bactra). [Timor.] geraropa Meyrick, 1932 Exot. Microlepid. 4: 147 (Bactra). [Taiwan.] banosii Gozmany, 1960, Annls Hist.-nat. Mus. Natn. Hung. 52: 416 (Bactra). [Egypt.] testudinea Turner, 1916, Trans. R. Soc. S. Aust. 40: 528 (Bactra). [Qld: Brisbane.] optanias Meyrick, Nov. 1911b, Proc. Linn. Soc. NSW 36: 253 (Bactra), nec Meyrick July 1911b, Trans.
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Proc. NZ Inst. 43: 89 (Bactra). [Qld: Cooktown and Stanthorpe; NSW: Sydney; Vic.: Melbourne.] passercula Turner, 1916, Trans. R. Soc. S. Aust. 40: 527 (Bactra). [Qld: Brisbane, Stradbroke Is.]
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xuthochyta (Turner, 1945) , Trans. R. Soc. S. Aust. 69: 56 (Capua). [Qld: Brisbane.] blepharopis Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 255 (Bactra). [Qld: Brisbane, Duaringa.]
Figs 434–439. Bactra, genitalia. 434, 435, B. venosana, male genitalia and aedeagus, Rockhampton, Q, slide ANIC 2778. 436, B. venosana, female genitalia, Rockhampton, Q, slide ANIC 2776. 437, B. blepharopis, female genitalia, Brisbane, Q, slide ANIC 2781. 438, 439, B. blepharopis, aedeagus and male genitalia, Mt Larcom, Q, slide ANIC 2441.
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neurosticha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 211 (Eucosma). [NSW: Murrurundi.] syntaractis (Turner, 1946), Trans. R. Soc. S. Aust. 70: 205 (Eucosma). [NSW: Brunswick Heads.]
Syntozyga Lower (Figs 54, 440–450) Syntozyga Lower, 1901, Trans. R. Soc. S. Aust. 25: 70. Type species: Syntozyga psammetalla Lower, 1901, ibid. 25: 70, by monotypy. [Australia.] Eleuthodema Bradley, 1957, Nat. Hist. Rennell Island 2: 95. Type species: Polychrosis pedias Meyrick, 1920, Exot. Microlepid. 2: 347, by original designation. [India: Bengal.] Bubonoxena Diakonoff, [1968], US Nat. Mus. Bull. 257: 40 (key), 65. Type species: Bubonoxena spirographa Diakonoff, [1968], ibid. 257: 66, by original designation. [Philippine Is.] Syn. nov. Cyclacanthina Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 337 (key), 349. Type species: Cyclacanthina episema Diakonoff, 1973, ibid. 1: 352, by original designation. [Java.] Syn. nov.
Diagnosis Syntozyga species are small to medium-sized moths, intricately marked with reddish or greyish brown on beige or pale yellowish ochreous, usually with either an indistinct dark blotch, outwardly oblique bar or complete transverse fascia on costa just beyond middle, with a curved brownish line from 36: of costa to termen above tornus and with metallic scales often arranged in transverse, curved bands. Syntozyga has moderately long, slightly sinuate and porrect labial palpi, with the second segment distally widened by long, loose scales. Apart from stalked Rs and M1 in the hindwing, Syntozyga has all other veins separate beyond the discal cell except for a few species not present in Australia that have R4 and R5 stalked in the forewing and connate with R3. The M-stem is absent and the accessory cell often very small with the chorda from between R1 and R2 or R2 and R3. Syntozyga has no secondary sexual modifications in the male except the slightly lengthened antennal
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cilia in S. sedifera. Male genitalia without uncus, with ill-defined and dorsally connected socii and with a highly ornamented valva with a large basal portion set with tufts of long bristles or heavy spines and thorns on the sacculus and/or distally to basal excavation are diagnostic for Syntozyga. The female genitalia are characterised by a very diverse sterigma, often with a median lip with a row of marginal bristles laterally on each side or with a raised and strongly spinulose lamella antevaginalis, but with its anterior portion always invaginated in a semicircular or transversely crescentic membranous pocket. Syntozyga has no signum.
Description Adults (Figs 440–442) Wingspan. 8.0–14.5 mm; small to mediumsized. Head (Fig. 443). Ocellus well developed. Frons with a transverse band of long, projecting scales in uppermost part, pointing forward and slightly downward, remainder with minute, appressed and upwardly oriented scales. Proboscis weak and short, from 56O length of labial palpus to at most as long as palpus. Labial palpus moderately long to long (1.5–2.1× diameter of eye; 2.5–2.8× in female of S. anconia Meyrick), weakly sinuate, porrect or lightly ascending, moderately to strongly dilated; second segment curved and distally moderately to strongly widened with long loose scales from ventral edge and distal part of dorsal margin, in some species (S. anconia) with particularly long ventrodistal fringe, often concealing terminal segment; terminal segment long and slender, porrect. Antenna reaching to middle of forewing; anterior surface devoid of scales only in a narrow strip at base of antennae (S. psammetalla Lower) or along entire antenna (S. sedifera Meyrick); cilia in male minute to moderate, about 56O diameter of flagellum in S. sedifera. Thorax. With or without posterior crest. Legs unremarkable, without any modifications in male; mid and hind tarsus with one small apical spine/ bristle on end tarsomere. Wings (Figs 54, 440–442). Forewing index 0.36–0.43; forewing elongate subrectangular to subtriangular; with evenly curved costa, acute apex and oblique, straight to faintly sinuate termen; discal cell 0.61–0.66× length of forewing, with moderate to very small triangular accessory cell with chorda either from between R1 and R2 or R2 and R3 (often from very close to R3) to
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Figs 440–443. Syntozyga, adult males and head. 440, S. anconia, Brisbane, Q. 441, S. psammetalla, Lockerbie area, Q [image reversed]. 442, S. ? sedifera, Daintree, Q. 443, head, S. anconia. Scale = 2 mm.
between R4 and R5, without any trace of M-stem; all veins present and usually separate beyond discal cell, rarely R4 and R5 connate or stalked (and in S. episema (Diakonoff, 1973) stalked R4 and R5 connate with R3); CuP present on wing margin or absent without any trace (S. anconia); wing pattern diverse and often very intricate, mostly of beige ground colour with yellowish, reddish or greyish brown markings, characteristically with a transverse dark brown fascia (S. sedifera) or with remnants thereof; a large and indistinct blotch (S. psammetalla Lower) or an outwardly oblique dark bar (S. anconia), with an outwardly curved brown line from 36: costa to termen above tornus and variably sprinkled with metallic scales often arranged in outwardly curved transverse bands. Hindwing with cubital pecten and axillary tuft present, with
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Rs and M1 stalked, with M2 bent towards M3 at base and with CuA1 from just before angle of cell close to M3, or CuA1 and M3 connate; CuP present at wing margin, 1A+2A and 3A developed but not tubular; pale to dark greyish. Pregenital abdomen. Unmodified. Sternum 8 in male with hindmargin faintly bilobed; intersegmental membrane between 8th segment and genitalia long and longitudinally striate, lightly sclerotised in ventral half. Male genitalia (Figs 445–449). Vinculum a weakly sclerotised band, sometimes slightly Wshaped. Tegumen weakly sclerotised, dorsally tapering or rounded and rather low, pedunculi often wide and with a small triangular apodeme for muscle m4. Uncus absent. Socii two ill-defined areas of bristles laterally in membrane below apex, dorsally connected by a band of bristles across apex
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Figs 444–450. Syntozyga, genitalia. 444, S. anconia, female, Rockhampton, Q, slide T2244. 445, 446, S. anconia, aedeagus and male genitalia, Rockhampton, Q, slide T2243. 447, S. psammetalla, male genitalia, Townsville, Q, slide ANIC 2788. 448, 449, S. ? sedifera, male genitalia and aedeagus, East Alligator R., NT, slide T2192. 450, S. psammetalla, female genitalia, Endeavour R., Q, slide ANIC 16854.
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of tegumen. Gnathos reduced to a weakly sclerotised transverse band, curved and medially joined with faintly sclerotised ventral surface of anal tube. Valva very diverse and highly derived, with basal excavation large, costal process small; with basal half of valva often greatly enlarged with a prominent sacculus with a large tuft of long bristles (S. sedifera) and/or with large spines on margin of basal excavation and especially from projections beyond basal excavation; with cucullus membranous and bristled, from finger-shaped in S. anconia to a round lobe in members of Diakonoff’s Cyclacanthina. Juxta small, subtriangular and fused with often very short caulis; anellus a cup-shaped structure. Aedeagus long and curved in Australian species, often with tapering apex; vesica without cornuti. Female genitalia (Figs 444, 450). Ovipositor lobes elongate-ovate. Sternum 7 with hindmargin often concave. Sterigma very conspicuous and diverse with anterior portion always invaginated in a semicircular to transversely crescentic pocket, with lamella postvaginalis at most weakly sclerotised; entrance to ostium either behind a strongly projecting median lip with a row of marginal bristles on each side (S. psammetalla, S. anconia) or a spinulose projecting round rim (S. sedifera) often with lateral extensions, with lamella antevaginalis often well developed, varying from a smooth, curved and posteriorly projecting plate (S. anconia) to a finely spinulose transverse ridge (S. psammetalla) or an irregular densely spinulose bolster (S. sedifera). Ductus seminalis with or without an often illdefined sclerotised portion below ostium, otherwise membranous with inception of ductus seminalis from between posterior third to mid length, followed by a widening, sometimes wrinkled and coiled section towards corpus bursa. Corpus bursae ovate to roundish, without signum.
Distribution Sri Lanka, India, south-east Asia to the Philippine Is, New Guinea and Australia. In Australia the genus occurs in the coastal regions of northern Western Australia, the Northern Territory, Queensland and New South Wales south to Batemans Bay. In the tropical areas it is also found far from the coast.
Biology Fletcher’s (1932) illustrated account of the biology of Polychrosis ephippias (Meyrick) as a stem borer of Commelina benghalensis (Commelinaceae) is the only host record for the genus.
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Remarks Diakonoff (1973) provided comprehensive revisions of Syntozyga, Bubonoxena and Cyclacanthina, including the Australian species, with numerous references and illustrations of the genitalia of both sexes and head and wing venation for each genus. Figures of adult moths were given by Clarke (1958) for Lobesia pedias (Meyrick), by Diakonoff ([1968]) for Bubonoxena endaphana Diakonoff and B. spirographa Diakonoff, and by Fletcher (1932) and Diakonoff (1973: fig. 10, not Fig. 9 as stated in legend) for S. ephippias. Syntozyga, in the more inclusive interpretation chosen here, doubtlessly denotes a monophyletic group characterised by the shape of the tegumen, the modified basal portion of the valva and the anteriorly invaginated sterigma. Bubonoxena and Cyclacanthina are both synonymised as their removal from Syntozyga leaves a paraphyletic group of less derived forms. The diagnostic feature of Bubonoxena, the large tuft of long bristles on the sacculus, is also present in S. sedifera, combined with a valva otherwise typical for Syntozyga s. str., and a group or row of prominent spines on the costa (or on a projecting portion thereof) is found in S. psammetalla, the type species of Syntozyga, as well as in some species of Cyclacanthina. The wide range of female sterigmata similarly does not lend itself to separation into exclusive groups with the sterigma of S. psammetalla combining both, a medially projecting lip with lateral rows of bristles and a transverse spinulose ridge. Head structure and venation are uniform throughout the genus, with a tendency to stalking of R4 and R5 in part of Diakonoff’s Cyclacanthina. The similarities in valva form and vinculum shape, the anteriorly invaginated sterigma, the shared lack of tarsal setae and reduction of the costal process leave little doubt that Syntozyga and Bactra are closely related. This relationship is confirmed by similar labial palpi with an often conspicuously lengthened subapical fringe on the ventral surface of the second segment in both genera and by a W-shaped vinculum found in some species of both genera. It is interesting that the only species of Syntozyga of which the biology is known is a stem borer in a monocotyledonous plant, as is Bactra. There is a certain amount of variation in the degree of distal expansion of the second segment of the labial palp, but the palpus figured by Diakonoff ([1968]: fig. 103; 1973: fig. 516) for Cyclacanthina spirographa is much more slender and has much more appressed and apparently shorter scales and a much shorter terminal segment than any of the
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Australian species. The figure also does not agree with the description of the labial palpi of Cyclacanthina in the same publications. Both S. psammetalla and S. sedifera are each members of species complexes with very similar superficial appearance and male genitalia. The female sterigma appears to be the most reliable diagnostic feature, but much more material will be needed to adequately revise the group. Unfortunately the remaining syntype of S. psammetalla lacks the abdomen, and the true identity of this taxon might have to be reconsidered once a more complete inventory of the northern Queensland fauna is available. Two unidentified but clearly distinct species of the S. psammetallagroup are known from north-western Australia, in addition to the material of S. psammetalla (or a complex of species containing S. psammetalla) from north-eastern Australia. Since the male genitalia of all these taxa are very similar the reports of S. psammetalla from Indonesia (Diakonoff 1973) will have to be reconsidered since they all are males. Due to the synonymy of Bubonoxena and Cyclacanthina the following species are transferred to Syntozyga: aspersana (Kuznetzov, 1988b), Entomol. Obozr. 67: 617 (Bubonoxena). [Vietnam.] Comb. nov. endaphana (Diakonoff, [1968]), Bull US Natn. Mus. 257: 67 (Bubonoxena). [Luzon.] Comb. nov. ephippias (Meyrick, 1907a), J. Bombay Nat. Hist. Soc. 17: 731 (Chrosis). [Sri Lanka] Comb. nov. episema (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 352 (Cyclacanthina). [Java.] Comb. nov. monosema (Diakonoff, 1973), Zool. Monogr. Rijksmus. Nat. Hist. 1: 350 (Cyclacanthina). [Sri Lanka.] Comb. nov. negligens (Diakonoff, 1973) Zool. Monogr. Rijksmus. Nat. Hist. 1: 351 (Cyclacanthina). [Sulawesi.] Comb. nov. spirographa (Diakonoff, [1968]), Bull US Natn. Mus. 257: 66 (Bubonoxena). [Luzon.] Comb. nov. transversa (Diakonoff, 1973) Zool. Monogr. Rijksmus. Nat. Hist. 1: 348 (Bubonoxena). [India.] Comb. nov.
fulva (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 209 (Eucosma). [Qld: Stanthorpe.] chrysosema (Turner, 1946), Trans. R. Soc. S. Aust. 70(2): 212 (Herpystis). [Qld: Stanthorpe.] anconia (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 257 (Polychrosis). [Qld: Brisbane, Toowoomba.] pleurosema (Turner, 1916), Trans. R. Soc. S. Aust. 40: 515 (Epichorista). [Qld: Rosewood.] gonodesma (Turner, 1929), Trans. R. Soc. S. Aust. 53: 305 (Epichorista). [Qld: Lamington NP.] loxomochla (Turner, 1945), Trans. R. Soc. S. Aust. 69(1): 69 (Epichorista). [NSW: Murrurundi.] eurymochla (Turner, 1945), Trans. R. Soc. S. Aust. 69(1): 69. (Epichorista). [Qld: Stanthorpe.] sedifera (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 256 (Polychrosis). [Qld: Cairns, Brisbane, Gympie.]
Endothenia Stephens (Figs 55, 451–454) Endothenia Stephens, 1852, List Specimens Br. Anim. Colln Br. Mus. 10: 28. Type species: Tortrix gentianaeana Hübner, [1799], Samml. Eur. Schmett. 7: pl. 3, fig. 12, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 35: 35, 57. [Europe.] Alloendothenia Oku, 1963, Insecta matsum. 26: 106. Type species: Alloendothenia menthivora Oku, 1963, ibid. 26: 104, figs 1, 2, by original designation. [Japan.] Neothenia Diakonoff, 1973, Zool. Monogr. Rijksmus. Nat. Hist. 1: 364. Type species: Sciaphila hebesana Walker, 1863, List Specimens Lepid. Insects Colln Br. Mus. 28: 342, by original designation. [Proposed as a subgenus of Endothenia.] [USA.]
Diagnosis Constituent species Fifteen named species of Syntozyga are known so far, three of which are from Australia: psammetalla Lower, 1901, Trans. R. Soc. S. Aust. 25: 70 (Syntozyga). [Qld: Cooktown.]
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Australian Endothenia are small, darkish moths with intricately transversely marked subrectangular forewings with conspicuous dots and patches of metallic leaden scales. Endothenia has moderately long, sinuate, porrect labial palpi with the second segment distally strongly expanded by long, loose
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scales often concealing the moderately thin, drooping terminal segment. Males often have a scale pencil from the base of the hind tibia and a modified, narrowly upfolded anal margin. The tarsal segments lack strong distal setae. In the forewing all veins are separate beyond the discal cell, in the hindwing Rs and M1 vary from distant at base (Nearctic species) to stalked (Australian species), M2 is distant from M3 at base though curving towards it, and M3 and CuA1 are connate. The male genitalia are diagnostic for the genus with a downcurved, dorsoventrally flattened or club-shaped, usually dorsally hairy and ventrally spiny uncus with a bulge or digit at its dorsal base beyond a depression, a long, slender, upcurved, distally nearly band-shaped valva with a spiny large process arising from ventral margin of basal excavation, a small juxta without developed caulis, and a short aedeagus often with a dorsal, coecum-like bulge. The female genitalia are characterised by a variably developed sterigma posterior to S7, a very long bipartite colliculum, a dilation in corpus bursae near origin of ductus seminalis, and by a single signum, a small, invaginated scobinate plate or reductions thereof. The female genitalia of Endothenia may be similar to those of Bactra, but M2 in the hindwing is much further from M3 in Endothenia.
Description Adults (Fig. 451) Wingspan. 8–10 mm (Palaearctic species up to 22 mm); Australian species small. Head (Fig. 453). Ocellus large. Uppermost part of frons with transverse band of long, anteriorly projecting scales, remainder with minute, appressed, upwardly oriented scales. Proboscis from shorter than length of labial palpus to about 1½× its length. Labial palpus moderately long (1.6– 2.0× diameter of eye), sinuate, porrect and strongly widened distally; second segment sinuate and distally strongly widened by long scales from ventral margin and across wide apex, often concealing the moderately thin, drooping terminal segment. Antenna to middle of forewing; anterior surface devoid of scales; cilia in male short, at most 56O width of flagellar segments. Thorax. With or without (Australian species) crest. Hind tibia in male often expanded by scaling and with a scale pencil from its base, at rest concealed between scales of inner surface; tarsal segments distally without conspicuous setae. Wings (Figs 55, 451). Forewing index 0.40– 0.45; subtriangular to subrectangular, with costa
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evenly curved throughout to straight except near base and apex, apex rounded-acute, termen oblique and straight to weakly curved; discal cell 0.61× length of forewing, chorda and M-stem nontubular in Australian species, chorda from halfway between R1 and R2 to between R4 and R5, M-stem only faint, to between M2 and M3; with all veins present and separate beyond discal cell; CuP a very weak trace at wing margin; with various and intricate wing pattern based on well-developed transverse fasciae, with numerous spots and patches of metallic leaden scales, well-developed costal strigulae and often a paler to whitish band from before apex to tornus. Hindwing with welldeveloped cubital pecten but no axillary tuft, in male anal margin often modified, narrowly folded upwardly and with long fringe; with Rs and M1 from distant at base (Nearctic species: Diakonoff 1973; Miller 1987) to approximated near base to stalked (Australian species), M2 distant at base from M3 though curved towards it, M3 and CuA1 connate at base; CuP present at wing margin, 1A+2A and 3A at most partly tubular, 3A far from anal margin; pale to dark greyish or brownish. Pregenital abdomen. Unremarkable. In male hindmargin of S8 with a semicircular medial excavation. Male genitalia (Fig. 452). Vinculum a narrow band, often weakly sclerotised. Tegumen short, broad and dorsally strongly convex, i.e. downcurved. Uncus well developed, downcurved, with narrow ‘neck’, distally abruptly expanded into a dorsoventrally flattened, dorsally hairy apex with strong, short spines on ventral surface or into clubshaped process either entirely hairy tending to spiny beneath (Australian species) or with strong spines along midline only; base of uncus dorsally from a concavity ending in a ventral hump that is developed into prominent digit-shaped process in some Oriental/Australian species. Socii usually large, triangular, drooping, hairy lobes tapering from wide base to sharp point, but in some small Oriental/Australian species reduced to few hairs laterally on tegumen. Gnathos at most weak lateral remnants; anal tube not sclerotised. Valva very long and slender, with smallish to large basal excavation and with a conspicuous, large, dorsomedially directed process distally from its lower margin, tongue- to club-shaped with short spines on inner surface; with cucullus long, slender, band-shaped or slightly widened towards rounded apex, evenly hairy on inner surface; with at most rudimentary costal process. Juxta small, often narrowly crescentic, directly connected with broadly ring-
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Fig. 451–454. Endothenia polymetalla, adult, head and genitalia. 451, adult, , Cooktown, Q. 452, male genitalia, paratype, Kuranda, slide ANIC 2793. 453, head, male. 454, female genitalia, Drysdale R., WA, slide ANIC 2794. Scale = 2 mm.
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shaped anellus. Aedeagus short, often tapering from wide base, at base in some species with dorsal, coecum-like bulge; vesica with or without one to few stout, spine-shaped cornuti. Female genitalia (Fig. 454). Ovipositor lobes elongate-ovate to narrowly band-shaped. Sterigma posterior to S7, variable from a simple membranous funnel leading to ostium to a smooth sclerotised plate with a central opening surrounded by a variably projecting rim or lateral lobes. Ductus bursae with long to very long, bipartite colliculum below ostium, remainder membranous, slightly expanding towards corpus bursae and with a dilation near middle just anterior to inception of ductus seminalis. Corpus bursae teardrop-shaped, with a single signum, an invaginated trapeze-shaped small scobinate plate forming a flat pocket, often reduced to a small patch of depressed scobination.
Endothenia is widely distributed in the Holarctic, Oriental and Australian regions. In Australia it has been reported from the islands of the Torres Strait, Cape York Peninsula to Kuranda in northern Queensland, and one specimen from the Drysdale R. in the Kimberley, Western Australia.
Endothenia is a well-defined group with highly derived male genitalia. The generic apomorphies include a hindwing with M2 distant at base from M3, even if curved towards it, modified male hind tibia and anal margin of hind wing (not always present), an uncus apex that is dilated and dorsoventrally flattened or club-shaped, and a distally very slender valva with a spiny process from ventral margin of basal excavation. Dang (1990) discussed in detail the four synapomorphies in the male genitalia that link Endothenia and Bactra: 1, the apically setose and sharply recurved uncus with a depression at its dorsal base; 2, the short, downcurved tegumen; 3, the unusual position of the valva with a reduced costal process, and 4, the reduced tarsal setae. In addition, Endothenia and Bactra share a single, very similar signum, and both genera have boring larvae whereas those of Gatesclarkeana are leaf rollers. The Australian Endothenia polymetalla (Turner) is very close to Endothenia citharistis (Meyrick), initially described from India and reported from Java by Diakonoff (1973), but E. citharistis entirely lacks the orange pattern elements so typical for E. polymetalla and has different valvae. E. engone Diakonoff from Sumba (Diakonoff 1984a) also is close.
Biology
Constituent species
Endothenia species are borers in stalks, roots and seedheads of dicotyledoneous plants, but no hosts are known from Australia. Diakonoff (1973: 366) reports a biology at variance with that of Holarctic species for Endothenia citharistis (Meyrick): ‘Central Java, Telawa, teak forest, 40m, bred from leaves of ?’boenga’ (?Lagerstroemia sp. (Lythraceae))’.
According to Brown (2005) there are 47 named species of Endothenia world-wide, with one named and one unnamed species known from Australia.
Distribution
Remarks Numerous Palaearctic and Nearctic Endothenia species are described and illustrated in local fauna treatments (Heinrich 1926; Hannemann 1961; Bentinck and Diakonoff 1968; Bradley et al. 1979; Kuznetsov 1978; Miller 1987, etc.). Razowski (1989) reviewed the genus in a Palaearctic context, synonymising the Japanese Alloendothenia. Dang (1990) figured genitalia musculature and tibial setae to illustrate his convincing argument that Endothenia should be included in the tribe Bactrini, and demonstrated that Tia Heinrich, Taniva Heinrich and Hulda Heinrich are distinct genera belonging to the Olethreutini. Kuznetsov (1993) provided a key and male genitalia figures for 23 Eurasian species.
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polymetalla (Turner, 1916), Trans. R. Soc. S. Aust. 40: 535 (Laspeyresia). [Qld: Kuranda.]
ENARMONIINI Diakonoff Enarmoniini Diakonoff, 1953, Verh. K. Ned. Akad. Wet. (2)49: 89, 161. Type genus Enarmonia Hübner, [1825]. Placed on the official List of Family-group Names in Zoology by the International Commission on Zoological Nomenclature, 1955, Opin. Decl. int. Commn zool. Nom. 10 (Declaration 28): 505. Ancylisidii Pierce and Metcalfe, 1922, The genitalia of the Tortricidae: 53.
Status remark Diakonoff (1953) intended Enarmoniini for the Grapholitini including Enarmonia Hübner. Declaration 28 of the International Commission on
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Zoological Nomenclature (1955) placed the name on the official list of family-group names and it thus supersedes Ancylini. Pierce and Metcalfe (1922) had established Ancylisidii for Ancylis Hübner and Enarmonia Hübner, but the grouping was not recognised until Kuznetzov and Stekolnikov (1973) revived it as Ancylidini, including Eucosmomorpha Obraztsov, Enarmonodes Danilevsky & Kuznetzov and Semnostola Diakonoff. Despite the lack of a consistent apomorphy, the tribe has gradually gained acceptance (Razowski 1989; Horak 1998).
Evidence for monophyly Pierce and Metcalfe (1922), the first to isolate a grouping corresponding to the current Enarmoniini but comprising only Ancylis and Enarmonia, defined the group by a long valva with an angulated sacculus, a bifurcate or single uncus and the signa a pair of angulate plates. Kuznetzov and Stekolnikov (1973, 1984) defined the Ancylidini by the configuration of the male genitalic musculature, presence of prominent attachment points for muscles m4 on the tegumen and an undivided costal process. Razowski (1987) added the shape of the sterigma as a possible apomorphy for the group. There is no apomorphy known to be present in all enarmoniine genera, and in the cladistic analysis the group usually was represented as a grade between the Olethreutini and Eucosmini, albeit with a large proportion in a single clade. However, there are several derived characters present only in the Enarmoniini: socii fused across the top of the tegumen, resulting in a continuous area of hairs or bristles across tegumen apex; tuft of very long hairscales from lateral surface of tegumen near base; single, often bifid thorn from cucullus, either from its tip or ventrally from a subapical process; very large basal excavation combined with long bristles from sacculus; signum-like sclerite in neck of corpus bursae. The lateral tufts of large modified scales originate from the tegumen itself and not from a fold between tegumen and valva as in some Eucosmini (Noduliferola Kuznetzov, Gypsonoma Meyrick). Consideration of the transformation series of valva tip and cucullus spine in related genera is required to establish that a spine in a subapical position of an enarmoniine like Aglaogonia, gen. nov. is not homologous with the single ventral spine in Olethreutini like Oxysemaphora Diakonoff that is derived from a clump of sacculus spines. The enarmoniine signa
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show a transformation series from horn- or triangularly blade-shaped structures to a small depressed area of scobination, found also in the Olethreutini in a parallel development. Origin of the ductus seminalis from very close to the corpus bursae and presence of large apodemes for muscles m4 are both typical for the Enarmoniini, but the former occurs in a few taxa in other tribes and the latter is shared with most Olethreutini.
Diagnosis Enarmoniini species are small to large olethreutine moths with the forewing often with a projecting to falcate apex, a sinuate termen and with a strongly oblique silvery line from costa to termen below apex. Thorax rarely with posterior crest, often with scale tuft from ventrally curved anterior angle of tegulae enlarged, to very long and pencil-shaped. Forewing without costal fold (large upcurved scales simulating an apparent costal fold in Balbidomaga Diakonoff); R4 and R5 frequently stalked, base of R4 sometimes atrophied. Hindwing usually with cubital pecten, sometimes with lobe or ribbon from base of anal margin (Anathamna Meyrick, Cyphophanes Meyrick, Helictophanes Meyrick) or with anal roll containing a pencil from its base (Ancylis, Oriodryas Turner; Periphoeba Bradley; Thysanocrepis Diakonoff); Rs and M1 rarely distant from each other (Irianassa Diakonoff) or stalked (Anthozela Meyrick), usually parallel and closely approximated, with base of M1 often atrophied; base of M2 from distant and parallel (Anthozela, Cyphophanes, Irianassa, Loboschiza Diakonoff, Pternidora Meyrick) to bent closely to M3; M3 and CuA1 often stalked, rarely fused into a single vein (Ancylophyes Diakonoff, some Anthozela spp.) or CuA1 from below angle of cell (some Metaselena spp.); 3A present except in Balbidomaga Diakonoff. Sternum 2 of ‘tortricoid’ type, with anterolateral processes and a pair of apodemes, sometimes remnants of sternal rods present. Hind tibia in male rarely with pencil (Anathamna, Metaselena) or expandable tuft of hairscales (some Loboschiza spp.). Hindmargin of S8 in male bilobed. Male genitalia ranging from complete and plesiomorphic (Ancylis) to highly derived. Tegumen often with large apodemes for muscle m4; laterally near base often with a tuft of very long, slender scales. Uncus bifid if present, mostly absent. Socii usually prominent and large, often medially fused across top of tegumen. Gnathos variably developed. Valva often weakly sclerotised, with very large basal excavation, often with a row of long bristles from sacculus; cucullus frequently highly derived and
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often with a spine at usually downcurved apex. Caulis usually very long and often fused with anellus. Aedeagus often with a bundle of deciduous cornuti. The female genitalia are very diverse, but the ductus seminalis nearly always originates from close to the corpus bursae. Sterigma diverse, usually spinulose and posterior to S7, colliculum usually present but frequently modified, an additional signum-like sclerite sometimes present in bursa neck. Signa from horn- or blade-shaped structures, variably reduced, to small depressed areas of scobination, usually paired.
Distribution The Enarmoniini are centred on the Oriental/ Australian regions with only few genera extending into the Palaearctic region and only Ancylis, Hystrichophora Walsingham and one species of Eucosmomorpha Obraztsov into the Nearctic.
Biology The biology of relatively few Enarmoniini is known so far. Species of this tribe feed as leafrollers (most Ancylis, Cyphophanes, Eucosmogastra, Eucosmomorpha) or borers, and one Australian Ancylis species has been reared from blotch mines on Banksia (Proteaceae). Leaf rolling as well as shoot and fruit boring may be found in the same species, with Loboschiza koenigiana (Fabricius) boring in shoots when young and often rolling leaves when older, but also feeding in fruit of Meliaceae. Other shoot borers are Anthozela on Piper nigrum (Piperaceae) and Tetramoera schistaceana (Snellen) in sugar cane (Poaceae). Helictophanes species are borers in fruit of several plant families, Toonavora, gen. nov. feeds on seeds of Toona (Meliaceae), and Enarmonia formosana (Scopoli) bores under the bark of Malus and Prunus (Rosaceae). Enarmonia flammeata Kuznetzov bores into shoots of Sasa plamata (Poaceae) (Bae and Sakamaki 1995).
Remarks In Holarctic faunal treatments (Heinrich 1923; Hannemann 1961; Bentink and Diakonoff 1968; Bradley et al. 1979) the Enarmoniini have generally been included in the Eucosmini or Grapholitini, and Diakonoff’s (1973) monograph of the Olethreutini contains several South Asian genera. Pierce and Metcalfe (1922) figured the male genitalia of British species of Ancylis and
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Enarmonia. Kuznetzov (1978) provided a revision of the fauna of the European part of the former USSR, illustrated with excellent drawings, including wing venations. Razowski (1989) published a synopsis of the Palaearctic olethreutine genera with genitalia drawings of both sexes, and a comprehensive treatment of the European olethreutine fauna (Razowski 2003a), with colour photos of all species and with genitalia drawings of high quality. Kawabe (1982) provided photos of adults, in colour, and of the genitalia of some species, and Liu and Li (2002) illustrated adults and genitalia of both sexes of Chinese Enarmoniini in either the Eucosmini or Grapholitini. Apart from a few apparent sister pairs such as Anathamna and Metaselena, Eucosmogastra Diakonoff and Oriodryas, and Periphoeba and Balbidomaga, the relationships between the enarmoniine genera are not obvious at this stage. The cladistic analysis most frequently produced a long comb with very little structure. Consequently, the genera are here arranged in alphabetical sequence, without their relative position implying any relationships.
Aglaogonia, gen. nov. (Figs 56, 455–463) Type species: Ancylis eupena Turner, 1946, Trans. R. Soc. S. Aust. 70(2): 200, hereby designated. [Australia.]
Diagnosis Aglaogonia species are narrow-winged, small to very small, yellowish and grey or brown to brownish moths with silvery white costal strigulae and a projecting apex with a conspicuous and characteristic pattern of an inverted Y-shaped black mark centred on the apex and enhanced by an adjacent whitish line to just before apex, yellowish scaling on the central part of the termen and an orange ochreous fringe along the anterior half of the termen. The labial palpi are moderately long, porrect and strongly sinuate with the second segment distally greatly widened by long loose scales and the terminal segment long and slender. The wing venation, especially in combination with the apex markings, is diagnostic for the genus, with R4 and R5 stalked and M3 and CuA1 fused into one vein in the forewing, and the base of M2 curved close to the base of the stalk of M3 and
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CuA1 in the hindwing. The male genitalia are diagnostic for the genus with the tegumen teardrop-shaped in outline with a pointed apex but no uncus, with large lateral scale tufts near base, without conspicuous muscle attachments on the pedunculi and with the highly characteristic socii: two very large, long, domed lobes laterally fused with the tegumen and their pointed tips joined at the apex of the tegumen, and densely set with long, broad scales and with bristles near the rounded ventral margin. The slender valva with a rounded apical lobe and a ventral, finger-like process with a strong spine is similar to that of some Anathamna and Loboschiza species, but the large, basally bulbous aedeagus with a long, sharp point and apparently some thorns on the outer surface but no cornuti seems to be unique. The female genitalia are very similar to those of Metaselena Diakonoff with a simple spinulose sterigma with two weakly sclerotised areas with some bristles posterior to ostium and the single signum a roundish, scobinate depression.
inwardly curved band beyond 6M wing, proximally delineating ocelloid patch, and an outwardly oblique large triangle at 56: dorsum, to dense transverse strigulation or suffusion across most of wing; ocelloid patch usually indicated by a confluent series of metallic leaden dashes; costa with prominent silvery white strigulae and markings in apex characteristic for genus always
Description Adults (Figs 455–457) Wingspan. 6–13 mm; small to very small. Head (Fig. 461). Ocellus moderate. Upper part of frons with very long, anteriorly protruding and downcurved scales, remainder of frons with minute, appressed scales. Proboscis about as long as labial palpus. Labial palpus porrect, moderately long (1.7–2.0× diameter of eye), strongly sinuate, second segment distally strongly widened by long, loose scales along ventral margin and dorsally near apex, terminal segment long and slender. Antenna to beyond middle of wing, entirely scaled, no cilia visible. Thorax. Smooth, with a large scale tuft from anterior corner of tegulae. Legs unmodified. Wings (Figs 56, 455–457). Forewing index 0.31–0.33. Forewing long, slender, subrectangular, costa and dorsum roughly parallel or wing slightly widening towards termen, costa weakly curved in basal half, apex projecting, pointed to falcate, termen sinuate; M-stem and chorda absent; R4 and R5 stalked, M3 and CuA1 fused into a single vein, CuA2 from below 36: cell; CuP present only near margin; anal loop 0.18× length of 1A+2A, anterior branch very weak, hardly visible even in stained wing; wing pattern of similar elements variably expressed, with whitish ochreous to orange ochreous ground colour with reddish brown to dark grey-brown markings ranging from a dark band along costal third of wing with a transverse,
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Figs 455–457. Aglaogonia, gen. nov., adults. 455, 456, A. historica, Toowong, Q. 455, [BMNH]. 456, [BMNH]. 457, A. eupena, , Mt Edith nr Atherton, Q [image reversed]. Scale = 2 mm.
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with a conspicuous, angled line of blackish scales from apex along margin to middle of termen, usually with a second line branching off from angle running obliquely towards about 6M costa, together forming an inverted Y-shaped black mark enhanced by a parallel, adjacent, whitish line or streak running to just before apex, by a conspicuous orange ochreous fringe along anterior half of termen and by whitish or orange ochreous scaling on central part of termen between ocelloid patch and margin except for line of black scales. Hindwing with cubital pecten and axillary tuft moderately developed; Rs and M1 parallel and closely approximated at base, base of M2 curved close to base of stalked M3 and CuA1; CuP present
223
near margin only; 1A+2A and 3A both present; light to dark greyish, in A. historica (Meyrick) yellowish orange in basal half, tip dark grey-brown. Pregenital abdomen. Ventral apodemes long, slender, inwardly oblique. Sternum 8 in male with a V-shaped medial cleft in hindmargin. Male genitalia (Figs 458–460). Vinculum a short band, fused with tegumen. Tegumen little sclerotised, teardrop-shaped in outline with pointed apex, laterally near base with large tuft of ribbonshaped scales longer than tegumen from membranous bulge, with weak muscle attachments medially on pedunculi. No trace of uncus. Socii two very large lobes reaching to below middle of tegumen, laterally fused with tegumen and their
Figs 458–461. Aglaogonia, gen. nov., genitalia and head. 458, 459, A. historica, male genitalia and aedeagus, Toowong, Q, slide BM 24893. 460, A. eupena, male genitalia, McIlwraith Range, Q, slide ANIC 12959. 461, head, A. eupena, male.
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pointed tips joined at apex of tegumen, densely set with long, broad scales and with bristles near rounded ventral margin. Gnathos two weak, slender, lateral bands. Valva well-sclerotised, long, slender, with sacculus only weakly convex and valva medially faintly narrowed, distal third bipartite with a rounded apical lobe, finely bristled on inner surface, and with a long, finger-like ventral process crowned with a spine; base of valva with a shallow pocket from outer surface; costal process stout, weakly sclerotised. Juxta small, caulis moderately long, wide in lateral view. Aedeagus large, strongly sclerotised, with dorsally bulbous base tapering to
sharply pointed, dorsally membranous, long apex, in A. historica with 3–4 short thorns on outside of dorsally membranous apical portion; vesica without cornuti nor with sockets suggesting their presence. Female genitalia (Figs 462, 463). Ovipositor lobes from very slender, parallel-sided to elliptical. Ostium in membrane well posterior of weakly concave hindmargin of S7; entrance to ostium a roundish, funnel-shaped depression in spinulose membrane, ventrally delineated by a curved crease that laterally expands to an elongate patch of spinulose, weak sclerotisation on each side posterior to ostium, set with a few bristles. Ductus bursae
Figs 462, 463. Aglaogonia, gen. nov., female genitalia. 462, A. historica, Toowong, Q, slide BM 24894. 663, A. eupena, McIlwraith Range, Q, slide ANIC 17118.
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slender throughout or somewhat widening towards corpus bursae, membranous except for a weakly sclerotised, very long colliculum; ductus seminalis either from slightly widened anterior part of ductus bursae (A. historica (Meyrick)) or from posteriormost part of corpus bursae; corpus bursae teardrop-shaped, with a single signum, a rounded, domed area of scobination, with the spinules in some species widely spaced.
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Anathamna Meyrick (Figs 57, 464–471) Anathamna Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 225 (key), 261. Type species: Anathamna ostracitis Meyrick, 1911b, ibid. 36: 262, by original designation. [New Guinea.]
Diagnosis Distribution Aglaogonia is so far known only from Australia where it has been collected on the eastern coast from Wollongong in New South Wales to the tip of Cape York Peninsula in Queensland.
Biology No host records are known for Aglaogonia, but nearly all localities are in rainforest habitats.
Remarks Aglaogonia is characterised by several apomorphies of the wing venation and the male genitalia, and also by the unmistakable pattern of the forewing apex. In the forewing R4 and R5 are stalked and M3 and CuA1 are fused into a single vein, in the hindwing M3 and CuA1 are stalked. The very large, domed socii set with broad scales and the bulbous aedeagus tapering to a long, sharp point, apparently without cornuti, are both highly derived. The valva of Aglaogonia recalls those of Anathamna Meyrick and some species of Loboschiza Diakonoff, and the sterigma and signum in the female are very similar to those of Metaselena. Unlike Loboschiza, which has M2 and M3 widely separate in the hindwing, the base of M2 is curved close to M3 in Aglaogonia, as in Anathamna and Metaselena. The generic name of feminine gender (aglaos, meaning ‘splendid’ and gonia, meaning ‘angle’) refers to the pattern of the forewing apex.
Constituent species Two described and two undescribed species are known from Australia: eupena (Turner, 1946), Trans. R. Soc. S. Aust. 70: 200 (Ancylis). [Qld: Mt Tamborine.] Comb. nov. historica (Meyrick, 1920), Exotic Microlepid. 2: 343 (Acroclita). [Qld: Brisbane.] Comb. nov.
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Anathamna species are medium-sized to large, characteristic-looking, ochreous to brownish tortricids with dark greyish hindwings and subrectangular forewings with usually slightly sinuate costa and sinuate termen. They recall Homona in shape but have conspicuous strigulae along the entire costa ending in a small, dark brown, triangular mark in apex. The labial palpi are moderately long, porrect, sinuate and widened beyond middle. The veins R4 and R5 are stalked in the forewing, the hindwing has a small projecting lobe at the base of the anal region in the male, the base of M2 is curved closely towards M3, and M3 and CuA1 are connate. As in Metaselena Diakonoff, males of Anathamna have two symmetrical patches of modified scales on the posterior surface of the metathorax and two opposite on the base of the abdomen, and a scale pencil from the base of the inner surface of the hind tibia. The male genitalia are extremely similar to those of Metaselena, but in Anathamna the distal portion of the valva is usually broader, more triangular, the apical thorn is bipronged, and, if present, a tuft of bristles from a protuberance on the ventral margin towards the valva ‘neck’ is diagnostic for Anathamna. The female genitalia are very similar throughout the genus, and the combination of a very large sclerite with one or two large horns at the entrance of the corpus bursae and two horn-shaped signa, both in the centre of the corpus bursae, are diagnostic for Anathamna. Females of Cyphophanes Meyrick also have a conspicuous sclerite at the entrance to the corpus bursae and horn-shaped signa, but in Cyphophanes the sclerite is blade-shaped and the signa are of unequal size.
Description Adults (Figs 464, 466) Wingspan. 15–22 mm; medium-sized to large. Head (Fig. 467). Ocellus moderate. Upper part of frons with long, anteriorly projecting scales, remainder of frons with minute, appressed scales.
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Figs 464–467. Anathamna, adults, abdomen and head. 464, A. plana, , Port Darwin, NT [BMNH]. 465, A. sp., abdomen base with modified scales on S2, male (va, ventral apodeme; ap, anterolateral process). 466, A. sp., , Iron Range, Q. 467, head, A. plana, male. Scale = 2 mm.
Proboscis longer than labial palpus. Labial palpus porrect, sinuate, moderately long (1.8–2.2× diameter of eye), second segment sinuate, moderately to strongly widened beyond middle by long, often loose scales ventrally and especially dorsally towards apex, terminal segment moderately long, slender and short-scaled. Antenna short, hardly reaching middle of wing, anterior surface unscaled, cilia minute. Thorax. In some species with small crest. Posterior surface of metathorax in male with two symmetrical ovate patches of dense, modified scaling, opposite similar patches at abdomen base. Hind tibia in male with a scale pencil from base of inner surface, often concealed by long, loose scaling. Wings (Figs 57, 464, 466). Forewing index 0.38–0.41. Forewing elongate subrectangular with
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costa strongly curved in basal fourth, then straight or slightly sinuate; termen sinuate and small apex sometimes projecting no more than middle of termen, dorsum straight except towards base; Mstem to M3, chorda absent in the four Australian and Papuan species examined but a very short chorda from beyond R3 to M1 figured for A. ostracitis (Diakonoff 1966); R4 and R5 stalked nearly to middle, all other veins separate beyond cell; R2, R3 and R4 roughly parallel, R3 not swollen distally and its base closer to R4 than to R2; CuP well developed near margin; anal loop 0.27× length of 1A+2A in A. plana Meyrick; yellowish ochreous to brownish grey with very conspicuous costal strigulae of dark brown, yellowish ochreous and silvery leaden along entire costa ending in conspicuous brown triangle in apex, with a brown spot or oblique bar below apex
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near middle of termen, and sometimes with a clearly defined dark brown wing base at least in anterior half and/or with a variable dark suffusion across dorsal half of wing or as an inwardly curved streak at 36: dorsum. Hindwing with large cubital pecten and axillary tuft; base of anal area with a small, projecting lobe with a thickened inner margin in male; crossvein between Sc+R1 and Rs present; Rs and M1 parallel but clearly separate at base, M2 curved to close to M3 at base, M3 and CuA1 connate from angle; CuP present towards margin, 1A+2A and 3A well developed; rather dark brownish grey. Pregenital abdomen. Sternum 2 with two large ovate patches of dense modified scales (Fig. 465). In male hindmargin of S8 deeply bilobed. Male genitalia (Figs 468, 469). Vinculum short, a moderate band fused with tegumen to form a
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ring. Tegumen high, subtriangular, a weakly sclerotised band (shape usually distorted in slide preparations). Apex of tegumen usually rounded but in one undescribed Papuan species with a conical hairy process (uncus?). Socii two ventrolateral, curved, densely bristled pendulous lobes from just below apex of tegumen, continuous with two patches of dense long bristles on dorsal surface of apex of tegumen. Gnathos two curved to triangular lateral bands from posterior margin of tegumen. Valva long and very slender, strongly sclerotised and with a very conspicuous triangular pocket-shaped invagination at base on outer surface below very large, straight costal process, widest in basal half with a very large basal excavation fringed along ventral margin with a band of bristles, with a more or less narrow ‘neck’ at 36: valva length often with a tuft of bristles from a small protuberance on
Figs 468–471. Anathamna, genitalia. 468, A. plana, male, Darwin, NT, slide BM 20961. 469, A. sp., male, Mt Webb Nat. Park, Q, slide ANIC 12917. 470, A. plana, female, near Cahills Crossing, East Alligator R., NT, slide T2245. 471, A. sp., female, Bamaga, Q, slide ANIC 17049.
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ventral margin, and with a narrowly to broadly triangular distal portion with dense bristles on inner surface, rounded dorsodistally and ending in a ventrodistal point carrying a large, bipronged thorn. Juxta small, caulis a very long, weakly sclerotised, wide band. Aedeagus long and slender, curved to sinuate, tapering towards apex; vesica with a large bundle of lanceolate cornuti. Female genitalia (Figs 470, 471). Ovipositor lobes subelliptic. Ostium in membrane posterior to usually evenly curved-concave hindmargin of S7 (in one Papuan species hindmargin of S7 with a deep W-shaped emargination); sterigma usually a simple, spinulose, membranous cup-shaped depression leading to ostium (surrounded by a U-shaped raised ridge in A. ostracitis and the above-mentioned undescribed Papuan species). Ductus bursae narrow and roughly parallel-sided, shorter than corpus bursae, membranous except for indistinct colliculum; corpus bursae ovate with a short narrowed posterior extension leading to ductus bursae and giving rise to ductus seminalis, with 1–2 sclerites in this region, a large hollow structure consisting of a single or two large horns at entrance to corpus bursae and sometimes a small transverse bar next to origin of ductus seminalis. Two hollow, curved, horn-shaped signa of roughly equal size, surrounded by a ring of scobination.
Distribution Anathamna is known from Papua New Guinea and Australia, where the genus occurs in the northernmost part of the Northern Territory, including Darwin (A. plana) and Kakadu NP, and on Cape York Peninsula in Queensland south nearly to Cooktown (unidentified species).
Biology No host records are known for Anathamna. In Australia the genus has been collected from localities with monsoon forest or at least some monsoon forest plants, but not from typically wet rainforest areas.
Metaselena and Anathamna are closely related, sharing numerous unique apomorphies discussed below. Whereas there is no doubt about the monophyly of the highly derived Metaselena with a string of unique features, especially in wing venation and female genitalia, Anathamna is overall much less derived and shows the plesiomorphic condition in most features. However, the presence of a large sclerite bearing one or two horn-shaped projections at the entrance to the corpus bursae is a conspicuous synapomorphy supporting the monophyly of Anathamna. Metaselena sometimes has a small sclerite near the entrance to the corpus bursae associated with the origin of the ductus seminalis, but this seems to be homologous with the second very small transverse sclerite in Anathamna in the same position. Cyphophanes also has a large sclerite at the entrance to the corpus bursae, but the male genitalia are quite different from those of Anathamna. The small projecting lobe with a thickened inner margin at the base of the anal region of Anathamna may be another apomorphy for the genus. Metaselena undoubtedly is the sister group of Anathamna, with the two genera sharing a series of apomorphies such as the stalked R4 and R5, the very similar male genitalia, the tibial pencil on the hindleg, and especially the unique paired patches of modified scaling posteriorly on the metathorax and opposite on the base of the abdomen. Their superficial appearance is similar apart from the modified wing shape of Metaselena, and the very conspicuous strigulae along the entire costa ending in a small dark triangle in the apex are probably a synapomorphy. Heteroschistis Diakonoff from the Solomon Is (Diakonoff 1966) and the Sri Lankan Aemulatrix Diakonoff (Diakonoff 1982) are closely related to Anathamna with very similar male genitalia, but R4 and R5 are separate in both genera. Heteroschistis also differs in details of the valva structure and in Aemulatrix the socii are fused rather as in Cyphophanes. Embolostoma Diakonoff, established in the Chlidanotinae (Diakonoff 1977), is also a close relative with R4 and R5 separate, but with long, slender, sinuate labial palpi and valvae of very different shape.
Remarks Anathamna was initially described for three species (Meyrick 1911b), one each from Australia, New Guinea and the Solomon Is, and Meyrick (1928) later added a fourth species from New Ireland off New Guinea. Diakonoff (1966) reviewed the genus, figuring the genitalia.
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Constituent species Seven species of Anathamna have been described from the Australian/Papuan region (Brown 2005), with one described and one unidentified species known from Australia. The generic identity of A. chionopyra Diakonoff from New Guinea needs
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confirmation, A. spermatophaga Diakonoff & Bradley is referred below to Toonavora, gen. nov., and A. megalozona Meyrick from Sri Lanka (Clarke 1958: pl. 142) is not congeneric with A. ostracitis. plana Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 261 (Anathamna). [NT: Port Darwin.]
Ancylis Hübner (Figs 58, 472–486) Ancylis Hübner, [1825] 1816, Verz. Bekannter Schmett: 376. Type species: Tortrix harpana Hübner, [1799], Samml. Eur. Schmett. 7: pl. 13, fig. 77, by subsequent designation by Fletcher, 1929, Mem. Dep. Agric. India (Entomol.) 11: 14. [Europe.] Epicharis Hübner, [1825] 1816, Verz. Bekannter Schmett: 376. Type species: Tortrix derasana Hübner, [1813], Samml. Eur. Schmett. 7: pl. 32, fig. 206, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 7, 58. A junior homonym of Epicharis Klug, 1807, Magazin Insektenk. (Illiger) 6: 197. [Europe.] Anchylopera Stephens, 1829 [June], Nom. Br. Insects: 47. Type species: Pyralis lundana Fabricius, 1777, Genera Insect.: 294, by subsequent designation by Curtis, 1831, Br. Entomol. 8: folio 376. [Germany.] Phoxopteris Treitschke, 1829, in Ochsenheimer, Schmett. Eur. 7: 232. Type species: Tortrix siculana Hübner, [1799], Samml. eur. Schmett. 7: pl. 13, fig. 79, by subsequent designation by Duponchel, 1834, in Godard & Duponchel, Hist. Nat. Lépid. Papillons Fr. 9: 22. [Europe.] Anticlea Stephens, 1834, Illust. Br. Entomol. (Haustellata) 4: 113. Type species: Tortrix harpana Hübner, [1799], Samml. eur. Schmett. 7: pl. 13, fig. 77, by subsequent designation (for Philalcea Stephens, 1835) by Curtis, 1836, Br. Entomol. 13: folio: 583 (but cited as ramella Linnaeus). A junior homonym of Anticlea Stephens, 1831, Illust. Br. Entomol. (Haustellata) 3: 236. [Europe.] Philalcea Stephens, 1835, Illust. Br. Entomol. (Haustellata) 4: 396. Type species: Tortrix harpana Hübner, [1799], Samml. eur. Schmett. 7: pl. 13, fig. 77, by subsequent designation by Curtis, 1836, Br. Entomol. 13: folio 583 (but cited as ramella Linnaeus). Established as an objective
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replacement name for Anticlea Stephens, 1834. [Europe.] Phoxopteryx Sodoffsky, 1837, Bull. Soc. Imp. Nat. Moscou 1837(6): 93. An unjustified emendation of Phoxopteris Treitschke, 1829. Sideria Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 156. Type species: Tortrix achatana [Denis & Schiffermüller], 1775, Ankündung syst. Werkes Schmett. Wienergegend: 131, by monotypy. [Europe.] Ancylopera Agassiz, 1847: Nomencl. zool. (Index univl.): 21, 22. An unjustified emendation of Anchylopera Stephens, 1829. Palaeobia Meyrick, 1881, Proc. Linn. Soc. NSW 6: 635 (key), 660. Type species: Palaeobia hibbertiana Meyrick, 1881, ibid. 6: 662 (key), 665, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 43, 61. [Australia.] Lamyrodes Meyrick, 1910, Proc. Linn. Soc. NSW 35: 162 (key), 182. Type species: Lamyrodes phileris Meyrick, 1910, ibid. 35: 182, by monotypy. [Australia.]
Diagnosis Ancylis species are small to medium-sized olethreutines with projecting and sometimes falcate apex, a sinuate termen usually partly edged with a line of black scales and an often characteristic wing pattern including well-developed dark interstrigular spaces along the entire costa, in distal half often with the white strigulae prominent and giving rise to some strongly oblique metallic leaden lines. The labial palpi are long, porrect and sinuate with the second segment distally strongly widened by very long, loose scales often concealing the long, slender terminal segment. The antenna in the male is entirely scaled without any visible cilia. Nearly all Australian Ancylis species have R4 and R5 stalked, the hindwing venation varies with M2 from quite distant from M3 to bent close to M3 at base, and with M3 and CuA1 mostly stalked but entirely fused in a few species. Males of at least some Australian species have a modified anal margin folded upwards over a scale pencil from a small lobe at the base of the hindwing. The genitalia in both sexes are usually diagnostic, in particular those of the female. In the Australian fauna the male is characterised by the distally bifid uncus (if present) of two parallel, ventrally curved hooks, the large, pendant, densely hairy socii, a sacculus with a strongly projecting and
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often produced ventrodistal angle and a very large basal excavation reaching nearly to its ventral margin, a long and broad caulis strongly curved in lateral view, and by a large number of deciduous, lanceolate cornuti. The female genitalia are characterised by a sclerotised funnel-shaped sterigma with a projecting ventral lip of diverse shape (lamella antevaginalis), a ductus seminalis arising from ‘neck’ of corpus bursae, and, except in A. carpalima, by two large, flattened, triangular, blade-shaped signa with compressed, hollow bases arising from triangular or T-shaped sclerotised areas dorsally and ventrally near middle of corpus bursa.
Description Adults (Figs 472–476) Wingpan. 7–17 mm; small to medium-sized. Head (Fig. 477). Ocellus moderate. Upper part of frons with a very long, flattened tuft of anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis longer than labial palpus. Labial palpus porrect, sinuate, long to very long (1.6–2.7× diameter of eye), second segment distally rounded and greatly widened by very long loose scales along entire ventral margin and distal half of dorsal margin, terminal segment long to very long, slender, downwardly inclined, often concealed in apical scales of second segment. Antenna to middle of costa or beyond, entirely scaled except for narrow line on anterior side, cilia not visible. Thorax. Smooth. Legs not modified. Tegulae with short scale pencil from anteroventral corner. Wings (Figs 58, 472–476). Forewing index 0.30–0.37. Forewing long and slender, subrectangular, costa and dorsum usually roughly parallel except in basal third, more rarely wing distally slightly widening, apex projecting to falcate, termen strongly sinuate except in A. longestriata (Durrant) with a straight, strongly oblique termen; chorda and M-stem from well developed to absent, chorda from below R2 to R5 or between R5 and M1, M-stem usually to between M2 and M3 but as a trace to between M3 and CuA1 in A. carpalima (Meyrick); R4 and R5 separate (Holarctic species), connate (A. carpalima) or stalked to 56O (all other Australian species), M2, M3 and CuA1 usually equidistant in Australian species except A. carpalima where base of M2 is obsolete, M3 from angle of cell and bent close to M2 at margin and CuA1 originating from before angle of cell; CuP present at margin; anal loop 0.27–0.29 length of
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1A+2A, anterior arm weak in A. carpalima; wing pattern extremely diverse in Australian species but except for A. longestriata all with well-developed dark interstrigular spaces along entire costa, in distal half often with white strigulae prominent and giving rise to some strongly oblique metallic leaden lines, nearly always with a line of blackish scales along part of strongly sinuate termen, ocelloid patch often indicated, remainder of pattern very diverse, frequently sexually dimorphic; A. longestriata unlike any other Ancylis, with very long, elongate-ovate forewing with strongly oblique, straight to weakly convex termen, of reddish ochreous ground colour fading to brownish grey towards margin, with a broad, lightly curved, longitudinal white streak from base to near end of cell and with radiating white stripes between the veins from R1 to M2. Hindwing with welldeveloped cubital pecten and axillary tuft, in male with anal margin modified and narrowly folded upward and, at least in some Australian species including A. erythrana Meyrick, A. anguillana Meyrick and A. carpalima, enclosing a scale pencil from small lobe at base of hindwing, in A. carpalima anal angle acute and projecting; Rs and M1 closely approximated and parallel at base, M2 from quite distant to bent close to M3 at base, M3 and CuA1 stalked or entirely fused (A. phileris (Meyrick), A. synomotis (Meyrick), A. biscissana (Meyrick), A. carpalima); CuP present at margin; 1A+2A present but not tubular, 3A present and distant from anal margin, at most partly tubular; usually greyish, often with a coppery tinge or orange-grey. Pregenital abdomen. Apodemes long and slender, often continued as indications of sternal rods. Male genitalia (Figs 478–483). Vinculum a broad band. Tegumen from elongate ovate to roundish, rather delicate and moderately sclerotised, dorsally with strong bristles near base of uncus, often with pronounced muscle attachment medially at base of pedunculi. Uncus usually well developed, distally bifid, two parallel and ventrally curved hooks, often long and slender, or uncus reduced to two small tubercles or entirely absent and top of tegumen broadly rounded-triangular (A. carpalima). Socii well developed and densely hairy, large, pendant lobes from ‘shoulders’ of tegumen below uncus or from apex if uncus absent. Gnathos two weak, hardly sclerotised, curved, lateral bands. Valva long and slender, nearly always with ventral emargination separating sacculus from cucullus; sacculus with strongly projecting ventrodistal angle,
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often produced into a long point, margin of basal excavation reaching nearly to ventral margin of sacculus; cucullus elongate-ovate to sickle-shaped with bristled inner surface and often with short
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spines towards ventral margin, or shape variously modified, very slender and distally tapering in A. synomotis, bifurcate in many Palaearctic species or subtriangular and distally downcurved to a
Figs 472–477. Ancylis, males except 476. 472, A. erythrana, nr Forster, NSW. 473, A. longestriata, Mt Gingera, ACT [image reversed]. 474, A. synomotis, nr Yeppoon, Q [image reversed]. 475, A. carpalima, Mossman, Q [image reversed]. 476, A. hibbertiana, female, nr Bateman’s Bay, NSW [image reversed]. 477, head, A. erythrana. Scale = 2 mm.
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Figs 478–483. Ancylis, male genitalia (attachment points for m4: arrows). 478, A. synomotis, Yungaburra, Q, slide ANIC 12949. 479, A. erythrana, Canberra, ACT, slide ANIC 12939. 480, A. hibbertiana, nr Ulladulla, NSW, slide ANIC 12951. 481, A. segetana, nr Mt Coree, ACT, slide T2230. 482, A. carpalima, McIlwraith Range, Q, slide ANIC 12954. 483, aedeagus, A. segetana, slide T2230.
naked point in A. carpalima; costal process large, sometimes bipartite. Juxta rather small, plateshaped, caulis long, broad and strongly curved in lateral view, anellus cup-shaped and fused with aedeagus. Aedeagus moderately large to large, usually lightly curved, tapering towards apex, in A. carpalima and related species aedeagus very long and slender, strongly curved; vesica usually with very large number of deciduous, lanceolate
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cornuti, apparently fewer in A. carpalima and related species. Female genitalia (Figs 484–486). Ovipositor lobes long and narrow, elongate-ovate, rarely margins pressed together (A. erythrana Meyrick). Ostium well posterior of S7. Sterigma a lightly sclerotised funnel with a projecting ventral lip of diverse shape, funnel in most Australian species parallel-sided and anteriorly narrowed and
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separated from rest of slender, membranous ductus bursae by short colliculum of two small, lateral sclerites; lamella postvaginalis spinulose and often with several large bristles on each side; corpus bursae teardrop-shaped with ductus seminalis arising from transition from corpus to ductus bursae; with two large, flattened, triangular, blade-shaped signa with compressed, hollow bases arising from triangular or T-shaped sclerotised areas dorsally and ventrally near middle of corpus bursa; signa surrounded by minute granulation and often a third patch of granulation between the two signa; A. carpalima with an accessory bursa on a narrow ‘neck’ from next to origin of ductus bursae and with only one signum, a long triangular blade with a very narrow, rounded, hollow base.
Distribution Ancylis is widely distributed and species rich across the Holarctic, Oriental and Australian regions. In Australia it is found in all states and territories.
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Biology Larvae of Ancylis are mostly leaf rollers on a large number of host families. Kuznetzov (1978) lists Ericaceae, Salicaceae, Fagales, Betulaceae, Rhamnaceae and Rosaceae, with the young larva sometimes mining a leaf before producing a shelter, few damaging soft fruit such as strawberries, and some Australian species producing large blotch mines. Biological information can be found in Heinrich (1923) and Miller (1987) for Nearctic species and Swatschek (1958), Hannemann (1961), Bentinck and Diakonoff (1968), Kuznetzov (1978) and in great detail in Bradley et al. (1979), among many others, for European species. In Australia, A. anguillana (Meyrick) has been reared from Correa reflexa (as Correa speciosa) (Rutaceae) and Pomaderris aspera and P. sp. (Rhamnaceae), A. artifica (Meyrick) from Alphitonia excelsa (Rhamnaceae), A. erythrana (Meyrick) from Hibbertia obtusifolia (Dilleniaceae), all feeding in spun-up shoots, and the following species from blotch-mines: A. hibbertiana (Meyrick) on Hibbertia scandens, H. volubilis and H. denta
Figs 484–486. Ancylis, female genitalia. 484, A. anguillana, nr Mt Coree, ACT, slide T2135. 485, A. erythrana, Canberra, ACT, slide ANIC 12950. 486, A. carpalima, near Ravenshoe, Q, slide T2139.
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(Dilleniaceae), A. synomotis on Banksia grandis, B. integrifolia and B. sp. (Proteaceae). In a note stating that A. carpalima is an Australian species that had been erroneously reported from India, Fletcher (1932: 20) said: ‘Indian species, formerly identified as A. carpalima, were presumably A. lutescens, Meyr., whose larva feeds on leaves of Zizyphus jujuba [Rhamnaceae]’.
Remarks Despite the unmistakably Ancylis-like forewing shape and pattern of most Australian species, Meyrick (1881, 1911b) only referred carpalima to that genus as all the other species have R4 and R5 stalked, a character not found in Holarctic species. Meyrick (1881) therefore described the genus Palaeobia for several Australian species, discussing their similarity with Ancylis (as Phoxopteryx) but stating that they also lacked a properly falcate apex. Lamyrodes (Meyrick, 1910), a genus described in the Tortricinae, was based on a single male of an unusual-looking species with hardly any wing pattern, a non-sinuate termen and with M3 and CuA1 in the hindwing entirely fused. However, the genitalia of all these Australian taxa leave no doubt that they are perfectly good Ancylis species. Traditionally, Ancylis has often been separated into subgenera based on hindwing venation, Ancylis with M3 and CuA1 stalked and Anchylopera Stephens with M3 and CuA1 fused into one vein. Even in the Holarctic region, groupings based on venation are not concordant with groupings based on genitalic structure, and the Australian taxa make it even more obvious that fusion of M3 and CuA1 has occurred several times within Ancylis, hence the configuration of these veins does not denote monophyletic groups. Given the wide distribution of the genus there is much information about Ancylis available from the many reviews of Holarctic Olethreutinae, among others in Heinrich (1923) and Miller (1987) for Nearctic species and Swatschek (1958), Hannemann (1961), Bentinck and Diakonoff (1968), Kuznetzov (1978), Bradley et al. (1979), Kawabe (1982), Park (1983) and Razowski (1989, 2003a) for the Palaearctic fauna. For the Oriental region one still has to rely largely on the original descriptions in combination with Clarke’s (1958) type illustrations, apart from the treatment of the Chinese fauna by Liu and Li (2002). The monophyly of Ancylis is supported by the two large, flattened, triangular, blade-shaped signa with compressed, hollow bases arising from triangular or T-shaped sclerotised areas, and
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possibly also by the sclerotised, funnel-shaped sterigma with a ventrally protruding lip. Razowski (1989) mentions the projecting and often falcate apex as a further apomorphy for Ancylis, but in the context of the other Oriental and Australian Enarmoniini, many with a projecting apex, this character carries little weight. Ancylophyes Diakonoff, with similar wing shape, has one round, horn-shaped signum but the second one flattened and very similar to those of Ancylis, albeit much longer, and the two genera also share a funnelshaped sterigma. There is no doubt that they are very close, with Ancylophyes generally much more derived, with M3 and CuA1 fused into one vein in both wings, the tips of a greatly elongate pencil from the anteroventral corner of the tegula inserted in the folded anal margin rather than a scale pencil from the base of the hindwing, with the top of the tegumen modified into an onion-shaped, bristled dome probably incorporating the socii and without any trace of an uncus, the ostium within S7 and the ovipositor greatly modified in all species examined. The strongly developed gnathos of Ancylophyes, medially fused with the anal tube, in conjunction with stalked R4 and R5, is an indication that both groups are monophyletic since the groundplan condition for Ancylis would be R4 and R5 separate, as in all Holarctic species, and the gnathos strongly reduced. The expanded anal area with a pronounced angle posterior to 3A is possibly a further apomorphy for Ancylis. The characteristic male genitalia of Ancylis are in large measure due to plesiomorphies, in particular the combination of the bifid uncus and the large, pendant, hairy socii. This explains the striking similarity to some Epinotia genitalia. The most plesiomorphic male genitalia of the two genera differ only in the enarmoniine apomorphies, the very large basal excavation and the elongate and dorsoventrally curved and widened caulis of Ancylis, and perhaps a stout and more strongly sclerotised tegumen in Epinotia without pronounced muscle attachments on the pedunculi. Judging from the male genitalia, Ancylis is the least derived genus of the Enarmoniini, and it probably is the sister group to the rest of the tribe, though this is not corroborated by the preliminary cladistic analysis. If confirmed, this would imply that the genus must be of considerable age, an assumption supported by its wide distribution across the Old and New World, its unusually diverse biology including leaf rollers and species producing large blotch mines, and the great morphological diversity in wing venation and genitalic structures. The Australian Ancylis species, with their stalked R4
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and R5, could well be part of a subordinate monophyletic group. Ancylis carpalima differs conspicuously from the remaining Australian Ancylis species in wing shape, venation and genitalia of both sexes, probably most significantly by the single signum of quite different shape. However, together with several Oriental species of similar appearance and genitalia, A. carpalima is no more than a very derived Ancylis and to treat the group as a separate genus would leave the rest paraphyletic. The Indian Ancylis aromatias Meyrick (Clarke 1958: figs 3–3a, 4–4b) shares exactly the same wing shape and pattern and has similarly modified female genitalia with a large accessory bursa arising from the posterior part of the corpus bursae, but its two Ancylis-type signa still provide the link with the remainder of the genus. Pseudacroclita hapalaspis (Meyrick), the type species of a monobasic genus (Oku 1979), has a tegumen and valva structure similar to that of A. carpalima, but given the differences in wing shape, venation and especially female genitalia this must be due to parallel reduction rather than to synapomorphy.
Constituent species Brown (2005) listed 132 species of Ancylis worldwide, including those of Ancylophyes that is treated as a synonym, with Razowski (1989) reporting over 50 species from the Palaearctic region. Twenty four named species are known from Australia, and several undescribed ones are present in the ANIC. artifica (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 237 (Acroclita). [Qld: Brisbane.] erythrana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 664 (Palaeobia). [NSW: Sydney.] hibbertiana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 665 (Palaeobia). [NSW: Sydney, Bulli Pass.] himerodana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 666 (Palaeobia). [NSW: Murrurundi.] stilpna (Turner, 1925), Trans. R. Soc. S. Aust. 49: 55 (Acroclita). [Tas.: Cradle Mt] synomotis (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 238 (Acroclita). [NSW: Sydney.] fidana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 667 (Palaeobia). [NSW: Sydney.] acromochla (Turner, 1946), Trans. R. Soc. S. Aust. 70: 199 (Acroclita). [WA: Waroona, Perth.] pseustis (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 239 (Acroclita). [SA: Adelaide; WA: Perth, Geraldton.] coronopa (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 237 (Acroclita). [Vic.: Beaconsfield.]
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mesoscia (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 235 (Acroclita). [Vic.: Mt Macedon.] segetana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 669 (Palaeobia). [Vic.: Warragul.] argillacea (Turner, 1916), Trans. R. Soc. S. Aust. 40: 505 (Lamyrodes). [NSW: Glen Innes.] anguillana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 662 (Palaeobia). [NSW: near Parramatta.] volutana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 663 (Palaeobia). [Vic.: Warragul.] erythrosema (Turner, 1945), Trans. R. Soc. S. Aust. 69: 57 (Capua). [WA: Albany, Bunbury.] erythrotypa (Turner, 1946), Trans. R. Soc. S. Aust. 70: 198 (Acroclita). [WA: Albany.] ochrophara (Turner, 1946), Trans. R. Soc. S. Aust. 70: 198 (Acroclita). [WA: Albany, Busselton.] sciodelta (Meyrick, 1921), Exotic Microlepid. 2: 446 (Acroclita). [WA: Yallingup, Cap Naturaliste.] glycypolia (Turner, 1945), Trans. R. Soc. S. Aust. 69: 58 (Capua). [WA: Margaret R.] rubrisignis (Turner, 1946), Trans. R. Soc. S. Aust. 70: 198 (Acroclita). [WA: Busselton.] phileris (Meyrick, 1910), Proc. Linn. Soc. NSW 35: 182 (Lamyrodes). [SA: Mt Lofty.] phileris (Lower, 1893), Victorian Naturalist 12: 71 (Proselena), nomen nudum. colonota (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 238 (Acroclita). [WA: Albany, Perth.] hemiochra (Turner, 1946), Trans. R. Soc. S. Aust. 70: 197 (Acroclita). [WA: Denmark.] infectana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 663 (Palaeobia). [NSW: Sydney, Mittagong.] longestriata (Durrant, 1891), Proc. Linn. Soc. NSW 6(1): 17 (Palaeobia). [NSW: Mt Kosciusko and Tumut.] biscissana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 674 (Holocola). [NSW: Sydney, Bulli Pass.] ochropepla (Turner, 1926), Trans. R. Soc. S. Aust. 50: 138 (Acroclita). [Qld: Byfield, near Yeppoon.] Comb. nov. carpalima Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 244 (Ancylis). [Qld: Cairns.]
Ancylophyes Diakonoff, stat. nov. (Figs 59, 487–492) Ancylophyes Diakonoff, 1988a, Entomol. Ber. (Amst.) 48: 195 (replacement name for Ancyloides Diakonoff, 1982).
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Ancyloides Diakonoff, 1982, Zool. Verh. (Leiden) 193: 63 (established as subgenus of Ancylis; a junior homonym of Ancyloides Kuznetzov, 1964, Entomol. Obozr. 43: 882, – Lepidoptera, Tortricidae). Type species: Ancylis (Ancyloides) stenampyx Diakonoff, 1982, ibid. 193: 64. [Sri Lanka.] Diakonoffiella Kuznetzov, 1997a, Entomol. Obozr. 76: 195 (unnecessary replacement name for Ancyloides Diakonoff, 1982), syn. nov.
Diagnosis Ancylophyes species are narrow-winged, small moths with subrectangular forewings with a strongly produced, pointed apex and deeply sinuate termen and a reduced wing pattern of few blackish brown markings, including well-developed costal strigulae with a dark costa in distal half, and variable reddish ochreous suffusion sprinkled with blackish brown scales on pale ochreous ground. The genus can be recognised superficially by the combination of its wing shape, long, sinuate labial palpi, distally dark-banded antennae and the derived wing venation with M4 and M5 stalked and with M3 and CuA1 entirely fused in the forewing and long-stalked or entirely fused in the hindwing. Furthermore, the males are externally recognisable by the long scale pencil from the anteroventral corner of the tegula with its tip inserted in the downcurved anal margin of the hindwing, and the female by the diagnostic, flattened, sclerotised ovipositor lobes and the entrance to the ostium within S7. The male genitalia are highly diagnostic for the genus with an onion-shaped, laterally hairy top to the tegumen and a remarkably long anal tube projecting far beyond the apex of the tegumen and strengthened throughout by two sclerotised bands arising from the medial portion of the gnathos, and by a ventrally deeply emarginate valva with an ovate cucullus and triangularly enlarged sacculus with a straight and inward rolled ventral margin with a row of strong bristles. The female genitalia are equally unique with an extensible ovipositor with flattened, strongly sclerotised triangular or crescentic ovipositor lobes, the ostium on S7 and two unequal signa, one a small hollow horn and the second a much larger, sharply triangular blade. Superficially, Ancylophyes looks somewhat similar to the equally narrow-winged Ancylis carpalima Meyrick, but A. carpalima has totally different venation with R4 and R5 separate and M3 and CuA1 both present and separate in the forewing,
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and a hindwing with a triangularly expanded anal area with the anal margin folded upwardly and containing a scale pencil originating from the base of the anal area.
Description Adults (Figs 487, 488) Wingspan. 8.5–11.0 mm; small. Head (Fig. 489). Ocellus moderate. Upper part of frons with long anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis about as long as labial palpus. Labial palpus porrect, long (2.2–2.5× diameter of eye), strongly sinuate, slender at base and apex but strongly widened in middle, second segment slender at base but with very long, loose scales on dorsal and ventral margins beyond middle, terminal segment very long and thin with appressed scales, partly concealed by long scales from second segment. Antennae to beyond middle of wing, entirely scaled, especially in male with conspicuous alternating dark and pale banding in distal half, no cilia visible. Thorax. Without posterior crest, but with a very long scale pencil from anteroventral angle of tegulae, crossing forewing beneath base and its tip inserted in narrow ventral fold along anal margin of hindwing. Legs unmodified. Wings (Figs 59, 487, 488). Forewing index 0.32–0.36. Forewing long and slender, subrectangular, widest in basal third, with costa strongly curved in basal third then straight to slightly concave, apex pointed and strongly produced, termen deeply sinuate with posterior half projecting nearly as much as apex; M-stem well developed, distally closely approximated and parallel to hindmargin of cell, no trace of chorda, closing vein rudimentary; R3 closely approximated at base to long stalk of R4 and R5, bases of M1, M2 and entirely fused M3 and CuA1 equidistant, fused M3 and CuA1 distally curved towards costa, CuA2 from 36: cell; CuP visible only near margin; anal loop 0.33 length of 1A+2A, 3A present; with strongly reduced wing pattern based on costal strigulae and variable reddish ochreous and blackish brown suffusion on pale ochreous ground colour; basal half of costa with about seven small, strongly oblique, blackish brown strigulae, distal half of costa edged with blackish brown including apex, especially just beyond middle and towards apex, with strigulae apparent as white lines that are strongly oblique except that before apex; with a blackish brown row of scales along termen; with
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reddish ochreous suffusion bordering dark brown band along distal part of costa and extended to varying degree across wing, often as a large subrectangular patch along basal third of dorsum extending to beyond middle of wing, and often also with scattered blackish brown scales across wing,
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especially in an indistinct broad oblique band from beyond middle of costa to base of dorsum. Hindwing long and narrow, costa strongly sinuate, with moderate cubital pecten and axillary tuft; in male anal margin thickened just beyond base and narrowly folded under, holding tips of long, slender
Figs 487–492. Ancylophyes monochroa, adult, head, abdomen and genitalia. 487, adult, , Byfield, Q. 488, , nr Cooktown, Q. 489, head, male. 490, abdomen tip, 8th segment, male. 491, male genitalia, Cape Hillsborough, Q, slide ANIC 12927. 492, female genitalia, nr Bowen, Q, slide T2430. Scale = 2 mm.
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scale pencil originating from tegula; Rs and M1 parallel and closely approximated at base, base of M2 curved close to base of long-stalked (A. stenampyx Diakonoff) or entirely fused (A. monochroa Diakonoff) M3 and CuA1, CuA2 from 36: cell, closing vein rudimentary; no trace of CuP even at margin; 1A+2A with basal loop but as 3A only partly tubular; pale brownish grey. Pregenital abdomen (Fig. 490). Ventral apodemes very long and slender, distally angled and hollow only at base, with rudiments of ventral rods present. Sternum 8 very large in male, its hindmargin with two strongly projecting rectangular lobes separated by a medial cleft (Fig. 490); T8 with a transverse, dumbbell-shaped sclerite at base and a small, subrectangular medial sclerite projecting on hindmargin. Male genitalia (Fig. 491). Vinculum well sclerotised, rather short, fused with tegumen and with valvae. Tegumen with moderately sclerotised ovate frame with small lobe-shaped muscle attachments, a ventrally split, onion-shaped top with apex (?uncus) blunt and nipple-shaped (A. monochroa) or produced into a sharp point (A. stenampyx), and with socii fused with tegumen as lateral patches of long bristles. Gnathos with a very prominent X-shaped central portion with the two dorsal arms very long sclerotised bands fused with the hugely enlarged and basally bulbous anal tube that in length far exceeds the tip of the tegumen. Valva with a very large, nearly circular emargination in ventral margin beyond middle separating a large, ovate, variably down-curved cucullus that is densely hairy on inner surface by a narrow neck from large, triangularly expanded sacculus with straight, inwardly rolled ventral margin set with row of strong bristles and distally projecting; costal process small. Caulis moderately long, anellus forming a large, cup-shaped sheath. Aedeagus long and slender, tapering towards apex, straight or slightly downcurved at base; vesica without cornuti nor with sockets suggesting presence of cornuti. Female genitalia (Fig. 492). Ovipositor radically modified, extensible with long apophyses and with flattened, strongly sclerotised, narrowly triangular to crescentic ovipositor lobes with only few bristles along margin. Ostium distally on S7, sterigma a narrow, sclerotised funnel fused to a circular opening in S7. Ductus bursae narrow, membranous, without trace of colliculum, with ductus seminalis arising close to corpus bursae; corpus bursae teardrop-shaped with two unequal signa, one a small, curved, hollow horn and the
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second more anterior one much larger, a long triangular blade.
Distribution Ancylophyes is so far known from Sri Lanka, Sumba and Australia, where it occurs in northeastern Queensland between Yeppoon and Cooktown, with one specimen from Darwin, Northern Territory.
Biology An unidentified species of Ancylophyes has been reared from Ficus erythrosperma (Moraceae) at Ohu near Madang, Papua New Guinea (morphospecies XXXX114 collected by the Papua New Guinea insect ecology project (Miller et al. 2003)) The localities for the few Australian specimens are mostly associated with monsoon forest. The curiously modified and extensible ovipositor suggests an unusual oviposition behaviour.
Remarks Ancylophyes was initially described as Ancyloides, a subgenus of Ancylis Hübner (Diakonoff 1982), but that name is a junior homonym of Ancyloides Kuznetzov, described as a subgenus of Eucosmomorpha Obraztsov (Kuznetzov 1964). Diakonoff (1988a) later proposed the replacement name Ancylophyes. Commenting on the unusual ovipositor Diakonoff (1982) assumed it to be an adaption merely on the species level and decided on the strength of the Ancylis-like signa to treat the taxon as a subgenus of Ancylis, figuring the genitalia of both sexes. Australian females of A. monochroa (Diakonoff), described as an Anchylopera from a single male from Sumba (Diakonoff 1984a), have exactly the same derived ovipositor and this modification is here regarded as a genus-level apomorphy and Ancylophyes treated as a genus. Ancylophyes is characterised by a number of apomorphies, some of them unique. In the forewing R4 and R5 are stalked and M3 and CuA1 fused into a single vein, in the hindwing M3 and CuA1 are at least long-stalked if not fused into a single vein and the anal margin in the male is folded under to hold the tips of a long scale pencil from the tegula. Apomorphies in the genitalia include the onion-shaped tegumen top, the greatly enlarged and partly sclerotised anal tube and the characteristic valva shape in the male, and in the
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female the modified ovipositor and the derived position of the ostium on S7. Ancylis and Ancylophyes share several apomorphies such as a similar wing shape, similar valva and a tubeshaped sterigma. The flattened signum in Ancylophyes is of the same basic structure as those of Ancylis. While there is no doubt that Ancylophyes is a monophyletic group the arguments for regarding also Ancylis as monophyletic and thus a separate taxon are discussed in the Remarks for the latter genus.
Constituent species Apart from two unidentified species from Papua New Guinea (Miller personal communication 2003), Ancylophyes is known from two named species, the generic type A. stenampyx from Sri Lanka and the species occurring in Australia, described from Sumba as Anchylopera. monochroa (Diakonoff, 1984a), Entomol. Basil. 9: 406 (Ancylis (Anchylopera)). [Sumba.] Comb. nov.
Anthozela Meyrick (Figs 60, 493–500) Anthozela Meyrick, 1913, Ann. Transv. Mus. 3: 280. Type species Anthozela chrysoxantha Meyrick, 1913, ibid. 3: 280, by monotypy. [South Africa.] Gephyroneura Obraztsov, 1968, J. New York Entomol. Soc. 76: 181. Type species: Laspeyresia hemidoxa Meyrick, 1907c, J. Bombay Nat. Hist. Soc. 18: 145, by original designation. [India.] A junior homonym of Gephyroneura Turner, 1919, Trans. Entomol. Soc. Lond. 1919: 417, – Lepid. Anthelidae. Mehteria Koçak 1981, Priamus 1: 117, replacement name for Gephyroneura Obraztsov, syn. nov.
Diagnosis Species of Anthozela look very alike superficially, so much so that most were regarded as one single, widely distributed species. They are small moths with bright yellow head, thorax and basal half of forewing, either speckled with black or suffused with orange, and the remainder of the forewing, separated from the yellow base by a leaden transverse band not quite reaching to costa, is of blackish ground colour
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with an intricate pattern of sinuous, bright orange to red lines, enclosing leaden spots and bands. The labial palpi are short, porrect and slightly sinuate. The hind wing venation is characterised by M2 distant at base from M3, Rs and M1 are connate to short-stalked and M3 and CuA1 vary from connate to fused into a single vein. Both male and female genitalia are diagnostic for the genus, and in the male the hood-shaped gnathos processes with pegs along the dorsal and lateral margin are unique. The female of A. chrysoxantha has not been examined, but the other species are characterised by a flimsy anterior appendix or expansion to the corpus bursae, and share with Eucosmogastra Diakonoff and Oriodryas Turner a posterior appendix giving rising to both ductus bursae and ductus seminalis, and two thorn- or horn-shaped signa.
Description Adults (Figs 493, 494) Wingspan. 10–12 mm; small. Head (Fig. 500). Ocellus large. Upper part of frons with anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis slightly longer than labial palpus. Labial palpus porrect, short (1.4–1.6× diameter of eye) and rather slender, lightly sinuate, second segment quite slender throughout and distal segment rather thick (A. hemidoxa, A. hilaris Turner) or second segment distally slightly widened and terminal segment slender, clearly angled forward (A. chrysoxantha). Antenna short, not reaching middle of wing, stout and scaled dorsally only in male, with minute dense cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 60, 493, 494). Forewing index 0.43–0.51. Forewing broadly triangular to subrectangular, with costa evenly curved, apex rounded and termen oblique (A. chrysoxantha, A. hemidoxa) to apex faintly produced and anterior half of termen roughly at right angle to costa (A. hilaris); chorda from below R2 to between R4 and R5, M-stem to between M2 and M3; all veins separate beyond discal cell; CuP well developed near margin; anal loop 0.33 length of 1A+2A; basal half of wing bright yellow, either evenly dotted with clusters of blackish to black scales (A. hilaris) or suffused with orange, especially along costa (A. chrysoxantha); posterior half of wing blackish with intricate pattern formed of numerous, strongly sinuous and partly parallel, bright orange to red lines enclosing shiny leaden scales in loops and in a strongly oblique band
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Figs 493–495. Anthozela, adults and male genitalia. 493, A. hilaris, holotype , [image reversed]. 494, A. sp., , Papua New Guinea [image reversed]. 495, A. chrysoxantha, male genitalia, South Africa, slide BM 24915. Scale = 2 mm.
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running from middle of costa towards termen below apex; costal strigulae well developed. Hindwing with sparse cubital pecten and axillary tuft; in A. chrysoxantha with anal margin narrowly rolled over upper surface, apparently concealing scale pencil; Rs and M1 connate (A. hemidoxa) to short-stalked (A. hilaris), M2 widely separate from M3 or stalk of M3 and CuA1 at base; M3 and CuA1 varying from connate (A. hilaris) to stalked (A. hemidoxa) to fused into one vein (A. chrysoxantha); CuP present towards margin; 1A+2A and 3A well developed; varying from grey to dark grey with an orange gloss to vividly orange in basal portion or entire wing except for dark mark in apex. Pregenital abdomen. Ventral apodemes broad at base, triangular. Sternum 8 in male with a subrectangular to shield-shaped sclerotisation with an emarginate to V-shaped hindmargin with, in A. chrysoxantha, a medially interrupted row of long stiff bristles (Fig. 499); T8 with transverse sclerotised bar or narrow crescent along anterior margin. Male genitalia (Figs 495–497). Vinculum a moderately wide band fused with tegumen to form a ring; triangular or semicircular membranous lobes carrying long scales associated with vinculum in some species (A. hemidoxa, A. bathysema Diakonoff). Tegumen rather wide and low, with apex either depressed and truncate (A. chrysoxantha) or a large rounded to subtriangular hump (A. hilaris). No trace of uncus in A. chrysoxantha and A. hilaris, but in A. hemidoxa according to Obraztsov (1968) uncus a pair of papillae. Socii two small lateral patches of bristles, hidden behind gnathos. Gnathos strongly modified, a weakly sclerotised U- to V-shaped medial band below socii giving rise to two raised large lobes together forming a hood-shaped structure, with a row of strong pegs along dorsal and lateral margin of each lobe. Valva rather membranous, usually elongate-subrectangular, in A. bathysema distally widened; sacculus somewhat swollen, its rounded angle crowned with a short blade or large point; distal part of valva simple, tongue-shaped and bristled ventrally on inner surface, only in A. chrysoxantha base of cucullus ventrally projecting in a rounded angle set with stronger bristles; costal process very large and long in A. chrysoxantha and A. hemidoxa, very small in A. hilaris. Juxta reduced to transverse, dumb-bellshaped or curved bar connecting both valvae and forming base of a wide, flat band with lateral flanges formed by caulis, nearly twice the width of the aedeagus. Aedeagus small, short, usually tapering
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towards apex; vesica with a sheaf of small, deciduous, aciculate or sinuous cornuti. Female genitalia (Fig. 498). Ovipositor lobes rather narrow. Ostium posterior to lightly V-shaped hindmargin of S7; sterigma a strongly sclerotised smooth ring- to funnel-shaped structure with thick ventral rim either ventrally projecting or in slide preparations often tipped backwards and concealing sockets of two scale patches in membrane behind ostium. Ductus bursae long and slender, colliculum well developed in A. hilaris as two lateral sclerites; corpus bursae complex, posteriorly ending in a narrowed appendix with a sclerotised apex giving rise to both ductus bursae and ductus seminalis, anteriorly with a second appendix of very thin membrane either nearly continuous with rest of corpus bursae or separated as an ovate accessory sac by a thin neck; no bulla seminalis. One or two signa, large, straight or curved horns.
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Distribution Anthozela has been recorded from South Africa, Sri Lanka, India, Indonesia, Taiwan, Japan, Papua New Guinea and Australia, where only a few specimens have been collected in rainforest habitats in northern Queensland, at Kuranda and at Cedar Bay near Cooktown.
Biology Anthozela hemidoxa has been reared from shoots of pepper vine in India (Fletcher [1921]). In Indonesia Enarmonia hemidoxa is figured as the pepper shoot borer and is considered a minor pest on black pepper (Piper nigrum) (Piperaceae), ‘eating irregular holes in top leaves, the young petioles and the shoots’ (Kalshoven 1981: 226). As discussed by Diakonoff (1984a), these records clearly refer to more than one species. Komai (personal
Figs 496–500. Anthozela, genitalia, abdomen and head. 496, A. hilaris, male genitalia, Kuranda, Q, slide BM 24911. 497, A. hemidoxa, male genitalia, Papua New Guinea, slide BM 24921. 498, A. hilaris, female genitalia, Kuranda, Q, slide BM 24910. 499, 8th abdominal segment, male, A. chrysoxantha. 500. head, male, A. chrysoxantha.
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communication 2005) observed larvae of A. hemidoxa sensu Kawabe boring into shoots of Piper kadsura on Iriomote I., Ryukyu Is.
Remarks Clarke (1958) figured the adult and genitalia of the male lectotype of Laspeyresia hemidoxa. Further information about Anthozela is available under Gephyroneura, with Obraztsov (1968) providing drawings of head, wing venation and genitalia of both sexes of A. hemidoxa, and Diakonoff (1984a) photos of the genitalia of both sexes of A. bathysema from Sumba. The monophyly of Anthozela is based on the male genitalia with raised gnathos processes carrying peg-shaped, modified bristles along the dorsal and lateral margins. Further apomorphies for the genus are the conspicuous wing pattern, the course of M2 distant from M3 in the hindwing, a blade- or point-like projection from the sacculus, and possibly, if confirmed for A. hemidoxa and A. chrysoxantha, the absence of a bulla seminalis and the presence of an accessory sac of very thin membrane at the apex of the corpus bursae. The course of M2 in the hindwing has in the past been regarded as an unequivocal grapholitine trait, with A. hemidoxa and A. hilaris originally described as Laspeyresia and Meyrick (1913: 280) stating ‘related to early forms of Laspeyresia’ with regard to Anthozela. Obraztsov (1968), while describing Gephyroneura in the Laspeyresiini, drew attention to the links with the enarmoniine Pseudophiaris Obraztsov and Eucosmomorpha Obraztsov. The remarkably modified male genitalia and the wing pattern are so similar in chrysoxantha, the type species of Anthozela, and A. hemidoxa, the type species of Gephyroneura, that the differences in wing venation cannot justify two separate genera. Oriodryas has very similar genitalia in both sexes and is probably the sister taxon. In particular, it shares the highly modified juxta/caulis complex fused into a broad band with wide lateral flanges, with the original juxta shape unrecognisable. Very little material of Anthozela is available in collections, but the observations by Diakonoff (1984a) and the fact that a handful of specimens from Australia and Papua New Guinea represent two if not three different species, all different from A. hemidoxa, suggest that it will be worthwhile looking at so-called A. hemidoxa from India to Japan. Of the species originally included in Gephyroneura, the generic position of the Chinese Gephyroneura perdigna Kuznetzov remains
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unresolved as it lacks pegs on the socii (Kuznetzov 1976), one of the apomorphies of Anthozela. In addition to the South African type species A. chrysoxantha, two non-Australian species are here included in Anthozela: hemidoxa (Meyrick, 1907c), J. Bombay Nat. Hist. Soc. 18: 145 (Laspeyresia). [India.] Comb. nov. bathysema (Diakonoff, 1984a), Entomol. Basil. 9: 407 (Gephyroneura). [Sumba.] Comb. nov.
Constituent species Brown (2005) lists six species of Mehteria, including two Australian ones that are referred to Loboschiza. One species of Anthozela occurs in Australia, represented by very few species from the Cairns/ Cooktown region in northern Queensland: hilaris (Turner, 1916), Trans. R. Soc. S. Aust. 40: 536 (Laspeyresia). [Qld: Kuranda, near Cairns.] Comb. nov.
Balbidomaga Diakonoff (Figs 61, 501–511) Balbidomaga Diakonoff, 1983a, Zool. Verh. (Leiden) 204: 64. Type species: Balbidomaga dorophora Diakonoff, 1983a, ibid. 204: 65, figs 40, 44; pl. 8, fig. 39, by original designation. [Sumatra.]
Diagnosis Balbidomaga species are small to medium-sized greyish ochreous moths with a projecting redbrown apex followed by white scales enhancing the impression of a strongly sinuate termen, and often with a small, dark, triangular mark on the middle of the costa. The labial palpi are long and sinuate with a distally widened second segment with a conspicuous horizontal to forward oblique whitish band across anterior half above middle. Males of Balbidomaga are characterised by a fake costal fold, a semicircular flap of large, glossy scales at 569 costa covering a pencil of hairscales, and by a very narrow anal area with a band of long, modified scales between anal margin and 1A+2A. Balbidomaga and Periphoeba Bradley have a very similar wing venation without M-stem, a very short chorda adjacent to the radius (chorda absent in Periphoeba), R3 adjacent and parallel to R4 nearly to margin, R4 and R5 connate or short-stalked, and bases of M3 and CuA1 closely approximated and parallel for
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some distance at base in the forewing, and the hindwing with M2 bent very close to M3 at base, M3 and CuA1 connate to stalked, CuA2 from just beyond middle of cell, 1A+2A close to anal margin and 3A absent in Balbidomaga. The males of the two genera are easily separated by their secondary sexual characters, the fake costal fold in Balbidomaga and the large scale pencil in the anal margin in Periphoeba, while separation of the females has to be based on wing pattern or genitalia. The genitalia in both sexes are diagnostic. The male is characterised by a large, capitate uncus and a well-sclerotised, large and complex gnathos, both unique in structure, and by a valva with a blade-shaped process on its ventral margin, either as a straight blade or an angled flange if the valva is ventrally emarginate. Males of Balbidomaga share with those of Periphoeba long, band-like scales laterally on the
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basal half of the tegumen, small socii (if present), an overall similar valva and sometimes a conspicuous, single row of very large bristles on the sacculus. The female genitalia of Balbidomaga are characterised by a spinulose sterigma usually sitting in a deep, often key-hole shaped emargination of S7, except in B. uptoni, sp. nov. where the sterigma is a U-shaped, spinulose rim posterior to the hindmargin of S7, by an entirely membranous ductus bursae and by an area of regular, minute, ovoid granulation in the corpus bursae ranging from a curved band to half the corpus bursae.
Description Adults (Figs 501–503) Wingspan. 8.5–12.5 mm (up to 19 mm in nonAustralian species); small to medium-sized.
Figs 501–505. Balbidomaga, adults and head. 501, 502, B. uptoni. 501, , Paluma, Q [image reversed]. 502, , nr Atherton, Q [image reversed]. 503, B. sp., , Papua New Guinea. 504, scale tuft on costa in male, B. uptoni. 505, head, male, B. uptoni.
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Head (Fig. 505). Ocellus moderate. Upper part of frons with long, strongly projecting scales, remainder of frons with minute, appressed scales. Proboscis barely longer than labial palpus. Labial palpus porrect, sinuate, very long (2.3–2.8× diameter of eye), second segment sinuate and distally strongly widened by long, loose scales along ventral and dorsal margins; terminal segment long, slender; second segment conspicuously bicoloured, darker in basal half and ventrodistally and pale whitish to whitish grey dorsodistally, the two areas separated by a horizontal or slightly outwardly inclined straight line, sometimes with fringe of apical scales dark, terminal segment with dark base and pale tip. Antenna to middle of wing, entirely scaled, with cilia not visible. Thorax. Smooth, with short scale tuft from tegula. Legs not modified. Wings (Figs 61, 501–504). Forewing index 0.34–0.40. Forewing elongate subrectangular, slightly widening distally, broadest before tornus, with costa lightly curved in basal half, straight distally, apex strongly projecting, termen sinuate; upperside in male at about 569 costa with a semicircular flap of large, broad, outwardly oblique, glossy scales suggesting a costal fold but consisting of recurved scales originating from costal margin or adjacent underside and concealing a tuft of hairscales extending outwardly parallel to costa (Fig. 504); M-stem indicated by a shadow in wing membrane only, chorda very short and close to radius; R2 from beyond )6D cell, parallel and close to R3 to margin, R3 from just before angle of cell, adjacent and parallel to R4 nearly to wing margin, R4 and R5 connate to short-stalked, M2 and M3 converging but only just separate at base, bases of M3 and CuA1 closely approximated and parallel for some distance from angle, CuA2 from beyond 6M cell; CuP present at margin; anal loop 0.25× length of 1A+2A; ground colour glossy whitish grey to pale greyish brown, with costal strigulae ending in a conspicuous, roundish, redbrown spot in apex followed by white scales on termen below apex, often with a variably developed, triangular, brown mark on middle of costa, pattern on dorsal half of wing sometimes only ill-defined, dark, transverse strigulation but often with an outwardly oblique bar, triangle or indistinct suffusion at 569 dorsum and with a row of dark dots along dorsum, rarely with a dark longitudinal line above middle of wing in distal 36: or costal part of wing up to this line suffused with dark brown; costal scale flap in male elongate and silvery grey in B. dorophora and the Australian B. uptoni, sp. nov.,
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smaller and dark brown in several New Guinea species. Hindwing with moderately developed cubital pecten and axillary tuft, in male with anal lobe truncate by long, straight anal margin lined on upperside with dense long hairscales from enlarged sockets from narrow area between 1A+2A and margin, on underside same marginal area with a band of minute scaling; cell without closing vein; Rs and M1 parallel and closely approximated at base, M2 bent very close to M3 at base, M3 and CuA1 connate to short-stalked, CuA2 from just beyond middle of cell; CuP present near margin, 1A+2A parallel and close to anal margin, no trace of 3A (in slide of male wing); pale to dark brownish grey, in B. uptoni, sp. nov. wing semi-translucent with very small, sparse scales. Pregenital abdomen. Long, slender, curved apodemes; in male with a large tuft of long scales with medially converging tips on T5 (Fig. 508). Male genitalia (Figs 506, 507, 511). Hindmargin of S8 concave. Vinculum short. Tegumen elongateovate, robust and well sclerotised, laterally in basal half with a band of long, band-like scales. Uncus conspicuous, an often large, posteriorly projecting, complex, capitate structure separated by a narrow ‘neck’ from tegumen, in B. uptoni, sp. nov. with two lateral horn-shaped processes and median capitate portion with bristles. Socii two small, lightly sclerotised bristled patches in B. dorophora and B. uptoni, sp. nov., absent in several New Guinea species. Gnathos a large, conspicuous structure, from moderately sclerotised and clearly bipartite with lateral arms dorsally extended into broad lamella and with angled, rising, paired, spoonshaped distal portions (B. dorophora, B. uptoni, sp. nov.) to a well-sclerotised, rigid, somewhat barrelshaped structure with the two dorsodistal lobes dorsally overlapping. Valva well-sclerotised, long and slender with distal half bristled on inner surface; either elongate-ovate with a single row of 5–6 conspicuous long bristles ventrally on sacculus and a blade-shaped projection along median third of ventral margin (B. dorophora, B. uptoni, sp. nov.) or valva constricted before middle by a semicircular ventral excavation, with only scattered small bristles on sacculus and ventroproximal angle of cucullus with an angled flange (several unnamed New Guinea species); costal process small. Juxta and caulis not modified, caulis moderately long. Aedeagus moderately long, rather slender, tapering towards apex and lightly sinuate in lateral view; no trace of cornuti or their sockets. Female genitalia (Figs 509, 510). Ovipositor lobes reniform, transversely ribbed by enlarged
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bristle bases, with jagged lateral margins. Ostium either caudally of S7 and hindmargin of S7 weakly concave (B. uptoni, sp. nov.) or ostium in a usually key-hole shaped emargination in hindmargin of roundish S7, often with two shearing lines with jagged ridges running obliquely forwards on both sides anterior and parallel to the obliquely curved lateral margins of S7. Sterigma a U-shaped, narrow,
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spinulose raised rim surrounded by spinulose membrane in B. uptoni, sp. nov., a similar spinulose structure in other species sitting within emargination of S7, often with an inner, parallel, at least partly sclerotised ring. Ductus bursae membranous, without colliculum, very slender in distal 36: to inception of ductus seminalis, anterior third widening towards teardrop-shaped corpus
Figs 506–511. Balbidomaga, genitalia. 506, 507, B. uptoni, male, Eungella, Q, slide T2142. 508, B. sp., abdomen, modified dorsal scales on T5, male, Papua New Guinea, slide GP164. 509, B. uptoni, female, Eungella, Q, slide T2143. 510, B. sp. nr rhacodyta, female, Papua New Guinea, slide GP296. 511, B. sp., male, Papua New Guinea, slide GP164.
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bursae; corpus bursae with a variable area of regular, minute, ovoid granulation (signum?), in B. uptoni, sp. nov. a curved band and in other species covering half the bursa.
Distribution Balbidomaga is known from Sumatra, New Guinea and Australia, where it occurs in Queensland in the Eungella NP, just west of Mackay, and in the Cairns and Atherton Tableland area.
Biology No host is known for Balbidomaga. Nearly all the Australian localities are upland rainforest, and the several undescribed New Guinea species have mostly been collected above 2000 m in rainforest with Nothofagus, with only one species from lower localities at 1300–1500 m. The type species originates from an altitude of 1430 m in Atjeh, Sumatra.
Remarks Balbidomaga dorophora was described by Diakonoff (1983a) from males only, with figures of adult, male genitalia, wing venation and head, and with a discussion of the unusual character combination and the superficial similarity of the species with some Chlidanotini. The species is figured in colour by Robinson et al. (1994). Clarke (1958) figured the adult and the genitalia of one of the two female syntypes of the Papuan Ancylis rhacodyta Meyrick, which is here referred to Balbidomaga: rhacodyta (Meyrick, 1938), Trans. R. Entomol. Soc. Lond. 87: 510 (Ancylis). [Papua New Guinea.] Comb. nov. The monophyly of Balbidomaga is supported by numerous apomorphies, some unique. The fake costal fold in the male consisting of a flap of large, glossy scales originating from the costa and the adjacent underside and concealing a scale pencil parallel to the costa is unique in the Olethreutinae. The truncate, modified anal area with reduced 3A and a band of long, modified scales between anal margin and 1A+2A in the male is a further apomorphy for the genus. The highly derived and complex uncus and gnathos are unique to Balbidomaga, and the extended areas of conspicuous granulation in the corpus bursae appear to be a further apomorphy in contrast to the small round patches in related genera. Balbidomaga shares with Periphoeba an unusual and strongly derived wing venation as well as
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similar forewing shape, and overall similar male genitalia with modified scales laterally on the tegumen, a well-developed uncus, similar socii and valva of the same general shape often with a conspicuous row of large bristles on the sacculus. The signa in both genera are patches of granulation. The two genera clearly form a sister group but no other close relatives have been identified. Vein 3A is well developed, very close to the margin in Periphoeba, but absent in Balbidomaga uptoni, sp. nov. with its modified and truncate anal region. The Australian B. uptoni, sp. nov. shows a curious combination of the capitate central part to the uncus with a lateral horn on each side of exactly the same configuration as the horns of the bifid uncus of Periphoeba. Assuming that the bifid uncus of Ancylis, Epinotia and Periphoeba denotes the olethreutine groundplan condition the lateral horns in the uncus of B. uptoni, sp. nov. would have to be interpreted as plesiomorphies within Balbidomaga. In a parallel development the socii, present in shape similar to those of Periphoeba in B. uptoni, sp. nov., are absent in species with a more highly derived uncus. The Sumatran B. dorophora, the Australian B. uptoni, sp. nov. and a very similar-looking species in New Guinea are the most plesiomorphic group in the genus with less derived male genitalia and the ostium in the female well posterior to S7. However, Balbidomaga is essentially a Papuan group with a number of highly derived species confined to the mountains of New Guinea.
Constituent species Two named species of Balbidomaga are known from beyond Australia, the type species B. dorophora from Sumatra and the Papuan B. rhacodyta. The Australian B. uptoni, sp. nov. is described below, and at least seven unnamed species are known from Papua New Guinea. uptoni, sp. nov.
Balbidomaga uptoni, sp. nov. (Figs 61, 501, 502, 504, 506, 507, 509)
Material examined Holotype. , ‘Eungella, Q[ueensland], 2400ft, 7 May 1955, I.F.B. Common’, ‘Genitalia slide No. T2142’ (ANIC). Paratypes. Queensland: 1 , 16°30´S 145°16´E, 9 km N of Mt Lewis (33.5 road km from Julatten turn-off), 30.
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i.1988, D. C. F. Rentz; 1 , Mt Edith 29 km SE Atherton, 1036 m, 18.iii.1964, I. F. B. C. & M.S.U.; 2 , 17°16´S 145°53´E, Lower slopes Mt Bellenden-Ker, 500 m, 20. x.1981, E. D. E.; 1, 17°16´S 145°23´E, Cableway Tower 3, 1020 m, Mt Bellenden-Ker, 28.x.1981, E. D. E.; 1, 17°26´S 145°29´S, The Crater, 18 km S Atherton, 980 m, 24. xi.1998, E. D. E. & H. Sutrisno; 1, 27.4 km S Atherton, 915 m, 19.iii.1964, I. F. B. C.; 2 , 2, 18°57´S 146°11´E, Mt Spec, 884 m, 792 m, 5.v.1955, 4.iii.1964, I. F. B. C., I. F. B. C. & M.S.U.; 1 , 18°58´S 146°10´E, 8 km NW Paluma, 23.viii.1977, I. F. B. C.; 1 , 19°00´S 146°13´E, Paluma, 19. ix.1985, I. F. B. C.; 1 , 6.4 km W of Paluma, 915 m, 13. iv.1969, I. F. B. C. & M.S.U.; 2 , 1, same locality as holotype, 7.v.1955, 1.iii.1964, 4.iii.1964, I. F. B. C., I. F. B. C. & M. S. U. (ANIC, BMNH, QM).
Diagnosis As the only Balbidomaga species known in Australia, the male of B. uptoni is immediately diagnosed by its fake costal fold somewhat distant from the wing base, and even the female can be easily recognised by wing shape and pattern. Balbidomaga uptoni is much smaller than most other know Balbidomaga species, though there is an equally small and very similar-looking unnamed species from Papua New Guinea in the ANIC, with quite different genitalia. The male genitalia of B. uptoni are characterised by an uncus with a swollen, ovate median portion and two short, sharp, lightly curved lateral horns, an elongate ovate valva with a large blade-shaped ventral projection, and a subbasally constricted aedeagus, medially widened before tapering to pointed apex. The female genitalia are characterised by the modified ovipositor lobes, a sterigma posterior to S7 rather than within a keyhole-shaped emargination of S7, and a well-defined band of granulation rather than an indistinct patch.
Description Male (Fig. 501) Wingspan 8.5–12.0 mm. Frons and inner surface of labial palpi whitish ochreous, rest of head and thorax mottled ochreous or greyish ochreous, outer surface of second palpus segment grey near base and on apex with longitudinal orange ochreous dash in between (Fig. 505), vertex scaling distally grey-brown. Forewing with strongly projecting apex, sinuate termen and a large, conspicuous flap of recurved, large scales covering a depression with a scale pencil at 56M costa, suggesting a costal fold (Fig. 504); R3 close and parallel to R4, R4 and R5 connate; ground colour glossy whitish ochreous,
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costal flap with some dark grey scales distally and on costa, rest of costa with a series of white strigulae separated by orange brown streaks, with a dark brown mark just beyond costal flap and an orange brown spot in apex; two silvery lines converging near termen below apex, one from 36: costa, the second from just before apex; a broad, obliquely transverse band of interrupted dark brown pattern elements from 56: costa to 56: dorsum; distal half of wing with ochreous scales along veins and near dorsum, and with some long, sinuous dark brown lines across wing, an indistinct dark brown subtornal patch and a whitish ochreous line around tornus to middle of termen; fringe basally white just below apex, barred with ochreous and dark grey. Hindwing dark grey-brown with a coppery gloss, sparsely scaled and translucent between small, modified scales. Abdomen with a paired patch of modified scales on T5; whitish ochreous at base, rest grey. Male genitalia (Figs 506, 507). Modified tegumen scales long, band-like near base; uncus complex, with very large, ovate, medioventral portion with some lateral bristles, flanked by a short, sharp, lightly curved horn on each side; socii small bristled patches; gnathos large, two long parallel arms each dorsally expanded into a broad lamella. Valva elongate ovate, medially not strongly constricted, but with a large blade-shaped projection in middle of ventral margin; with few strong bristles below basal excavation and cucullus bristles arranged nearly in a semicircle. Aedeagus constricted subbasally in lateral view, then strongly widened before tapering to pointed apex; no trace of cornuti. Female (Fig. 502) Wingspan 11–13 mm. Similar to male, but without costal flap and wing pattern except in wingtip simplified: costa with small dark strigulae along basal third and with a conspicuous, dark brown subtriangular mark in middle, rest as in male; dorsodistal parts of wing more evenly dark grey than in male, with some indistinct dark transverse lines across dorsum and reddish ochreous bands along veins. Hindwing greyish ochreous with coppery gloss. Female genitalia (Fig. 509). Ovipositor lobes modified, reniform, transversely ribbed by enlarged bristle bases. Sterigma posterior to weakly concave hindmargin of S7, a U-shaped, narrowly raised, spinulose rim within a transverse, triangular area of spinulose membrane. Ductus bursae shorter than corpus bursae, entirely membranous, slender, with short portion between corpus bursae and origin of
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ductus seminalis slightly wider. Corpus bursae ovate with a well-defined, oblique band of conspicuous granulation from middle of bursae to laterally close to its ‘neck’.
Distribution and biology Balbidomaga uptoni occurs along the Queensland coastal region from Eungella to somewhat north of Cairns, in rainforest localities usually at somewhat higher altitudes. No host information is available.
Remarks The species is named after Murray S. Upton, who collected a good proportion of the Lepidoptera in the ANIC, including some of the type series of this exciting species.
Cyphophanes Meyrick (Figs 62, 512–518) Cyphophanes Meyrick, 1937 Exot. Microlepid. 5: 160. Type species: Cyphophanes dyscheranta Meyrick, 1937, ibid. 5: 160, by monotypy. [Java.]
Diagnosis Species of Cyphophanes are very small to mediumsized, characterised by subrectangular forewings, often with a rounded apex, of grey-brown to
blackish brown ground colour sprinkled with metallic blue scales but with little pattern otherwise except a few white costal strigulae. The males, except for one New Guinea species, have a long, narrow, scaled lobe at the base of the anal area separated from the rest of the hindwing by a deep cleft. The labial palpi are basically porrect and sinuate, but they can be very short. The wing venation is characterised by R5 running to costa and a strongly oblique closing vein in the forewing, and by the base of M2 widely distant from M3 in the hindwing. The male genitalia are diagnostic with a narrow, high tegumen with well-developed ‘shoulders’, narrow and slender valvae with an apical thorn and a long, tapering aedeagus. In the female genitalia the unusual position of the simple U- to ring-shaped sterigma between the posterior margin of S7 and the ventrally extended posterolateral corners of T7 is diagnostic, as are the three sclerites in the corpus bursa, the two pimpleto thorn-shaped signa (one sometimes minute) and the similar-looking but more blade-shaped sclerite at the entrance to the corpus bursae.
Description Adults (Figs 512, 513) Wingspan. 7.5–15.0 mm; very small to medium-sized. Head (Fig. 515). Ocellus large. Anteriorly projecting tuft of scales from upper part of frons; remainder of frons with minute, appressed scales. Proboscis not much longer than labial palpus.
Figs 512, 513. Cyphophanes, males. 512, C. gracilivalva, holotype, Sue (Warraber) Is, Q. 513, C. sp., Papua New Guinea. Scale = 2 mm.
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Labial palpus porrect and sinuate, variably long (1.33–2.00× diameter of eye), short and slender with often very small terminal segment in small species but longer with distally widened subtriangular second segment and with long thin third segment in larger species. Antenna not reaching middle of forewing; anterior surface without scales, cilia in male short. Thorax. With a posterior tuft of slightly raised scales. Legs unmodified. Wings (Figs 62, 512, 513). Forewing index 0.41–0.48. Forewing elongate subrectangular with apex weakly pointed to broadly rounded; costa curved throughout, more so towards apex; closing vein usually strongly oblique, accessory cell sometimes very narrow and M-stem to just before M3; with all veins separate beyond cell, R4 and R5 closely approximated and parallel at base in some species (C. dyscheranta) and R5 to before apex, M2 widely distant from M3 at base; anal loop 0.28 length of anal vein in Australian species near C. dyscheranta; grey-brown to blackish brown with little obvious pattern but with conspicuous dots and patches of appressed bluish metallic scales scattered across wing; ground colour grey brown to blackish brown, in some species with a paler golden brown streak from apex or with two indistinct, outwardly angled paler fasciae in distal half of wing; the few white costal strigulae very oblique and short: usually 2–3 less distinct ones at about 56: costa and always 4–5 from middle of costa to before apex, the first giving rise to a long, strongly oblique metallic blue line connected to a roughly transverse broad band of scattered metallic scales beyond 6M wing; further concentrations of metallic blue scales in an elongate mark before termen and in an indistinct, outwardly angled band from strigulae at 56: costa to middle of dorsum. Hindwing with cubital pecten and axillary tuft moderately developed; in all except one New Guinea species basal part of anal area separated as a ribbon-shaped, scaled lobe by a deep and narrow emargination from remainder of wing; with Rs and M1 closely approximated and parallel in proximal third, M2 widely distant at base from M3 (C. dyscheranta) or base curved towards stalk of M3 and CuA1 (unidentified Papuan species), M3 and CuA1 connate to short-stalked; CuP very weak even towards margin; 3A very short and to just beyond emargination separating scaled lobe at base of anal area. Pregenital abdomen. Unmodified. Sternum 8 in male with hindmargin weakly bilobed; T8 with rounded hindmargin.
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Male genitalia (Figs 514, 518). Vinculum small, fused with tegumen. Tegumen high and narrow with well-defined ‘shoulders’, weakly sclerotised, formed by two broad, parallel bands, broad also at base. Apex of tegumen either a broad, rounded lobe or weakly to strongly bilobed, dorsally and sometimes laterally with dense long hair-like bristles or scales. Socii present in some species (C. dyscheranta) as flimsy membranous lobes laterally to anal tube, covered with long, hairlike bristles extending to ‘shoulders’ of tegumen. Gnathos usually absent, only in C. dyscheranta apparent as a weak sclerotised streak laterally on tegumen; anal tube spinulose below in C. dyscheranta and related Australian species. Valva simple and moderately sclerotised, narrow to very narrow, roughly parallel-sided or tapering towards often curved apex, with double or single apical thorn and long bristles below basal excavation and on apex; costal process long, straight and slender. Caulis moderately long, straight. Aedeagus long, straight, evenly tapering to much narrower apex, often with one to several lanceolate cornuti. Female genitalia (Figs 516, 517). Ovipositor lobes moderately broad. Hindmargin of S7 anterior to hindmargin of T7 and posterior corners of T7 extended ventrally, sometimes as pronounced, sclerotised lobes, to frame sterigma. Sterigma usually a spinulose, weakly sclerotised U- to ringshaped structure in membrane, in one derived New Guinea species a small sclerotised cup behind margin of S7. Ductus bursae membranous except for a colliculum of two narrow, elongate sclerites; anterior end of ductus bursae, beyond inception of ductus seminalis, granular and widening towards corpus bursae. Corpus bursae ovate, with two signa and a third signum-like sclerite at entrance to bursae, all three (except rarely the smaller signum) associated with a strongly sclerotised, flat, roundish base and varying in shape from a minute pimple to a large, hollow thorn, with the sclerite at entrance of corpus bursa a blade-shaped structure and always the largest of the three.
Distribution Cyphophanes occurs in Sri Lanka (Diakonoff 1982), India (Meyrick 1916), Indonesia, New Guinea and Australia, where it is found in the north-eastern coastal region and the northern tips of the continent from near Maryborough to the islands off Cape York in Queensland and the north-eastern corner of Arnhem Land in the Northern Territory.
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Biology In his description of C. dyscheranta, Meyrick (1937: 160) reported that the species has been bred in January ‘from larvae feeding on leaves of Murraya (Rutaceae) (Dr L. G. Kalshoven)’ in Telawa, Java. In Australia the genus is found in diverse associations, from subtropical woodland to tropical rainforest.
Remarks Clarke (1958) figured head, wings, wing venation and male genitalia of the lectotype of C. dyscheranta and wings and male genitalia of that of Helictophanes dryocoma (Meyrick), and Diakonoff (1982) provided a photo of the male genitalia of the latter from Sri Lanka. Apomorphies for Cyphophanes include the metallic blue scales scattered across the forewing, the narrow, scaled, ribbon-shaped lobe at the base of the anal area in the male hindwing (absent in one New Guinea species), the narrow tegumen with very broad pedunculi, the simple, slender valva hardly dilated at base, and the signum-like sclerite at the entrance to the corpus bursae. As stated by Meyrick (1937) in his original description, Helictophanes seems to be the most closely related genus, sharing very similar forewing shape and a narrow lobe from the base of the anal area in the male hindwing. The sclerite at the entrance to the corpus bursae provides a link with Anathamna. Argyroptocha Diakonoff from the Philippines has a similar tegumen and equally narrow valvae, but differs in having a bifurcate valva apex, dense, strong, short bristles on the end of the sacculus, and a hindwing without a free anal lobe and with M2 closely approximated to M3 at base. Articolla myriolychna Turner was referred to Cyphophanes in Horak et al. (1996), but dissection of the female holotype, still the only specimen of this species, revealed that it is a species of Helictophanes. However, the following species from India is here referred to Cyphophanes: dryocoma (Meyrick, 1916), Exotic Microlepid. 2: 21 (Argyroploce) [India.] Comb. nov.
species near C. dyscheranta and the new species described below: gracilivalva, sp. nov.
Cyphophanes gracilivalva, sp. nov. (Figs 512, 514, 516)
Material examined Holotype. , ‘10.12S 142.49E, Sue (Warraber) I., Q[ueenslan]d, 1 Dec 1977, E.D. Edwards’ (ANIC). Paratypes. Northern Territory: 1 , 11°01´S 136°45´E, Rimbija Is, Wessel Is, 19.i.1977, E. D. E. Queensland: 3 , 2, same data as holotype, 1–8.xii.1977; 2, 6.4 km SSE Yeppoon, 2 & 4.ii.1970, I. F. B. C. (ANIC, QM).
Diagnosis From the first of the two unnamed Cyphophanes species present in Australia, C. gracilivalva can be separated by its darker colour, smaller size and short, straight palpi. The second unnamed Australian species, known from a single male from Iron Range, is similarly sized and similarly dark as C. gracilivalva but has more defined orange-brown pattern elements, in particular two parallel, welldeveloped, orange-brown lines from costa, angled to follow termen. The length of the ribbon-shaped anal lobe is diagnostic, reaching nearly to the anal angle in C. gracilivalva but not much further than halfway in the male from Iron Range. The male genitalia of C. gracilivalva are characterised by a deeply bilobed tegumen apex with the two lobes parallel rather than diverging, by an elongate subtriangular valva with a hardly projecting sacculus, by a small, curved cucullus tapering to a single apical spine, and by an only weakly curved aedeagus. The female genitalia are characterised by a ring-shaped sterigma, an elongate-bulbous thickening below ostium, a triangular, toothshaped sclerite in bursa ‘neck’and two small pimple-shaped signa.
Description Constituent species In addition to the two named Cyphophanes species from the Oriental region there are two unidentified species from Papua New Guinea in the ANIC. Three species occur in Australia, probably one of the two New Guinea species, a second unnamed
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Male (Fig. 512) Wingspan 8.0–9.5 mm. Head, thorax and tegulae dark grey-brown, with blue iridescence on tegulae; palpi and lower part of frons ochreous brown, paler than rest of head; palpi very short, nearly straight, only weakly widened and terminal
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segment very short. Forewing with R4 and R5 distant at base; ground colour dark grey-brown to blackish brown, with four indistinct, transverse and outwardly curved bands of large patches of glossy
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bluish black scales, the first near base, the second in middle of wing, the third from ¾ costa to tornus and the fourth across apex; intervening areas partially scattered with narrowly pale-tipped brown
Figs 514–518. Cyphophanes, genitalia and head. 514, C. gracilivalva, paratype, male genitalia, Sue (Warraber) I., Q, slide T2410. 515, head, C. gracilivalva, male. 516, C. gracilivalva, paratype, female genitalia, Sue (Warraber) I., Q, slide T2428. 517, C. dyscheranta, female genitalia, Java, slide T2426. 518, C. dyscheranta, male genitalia with aedeagus, Java, slide T2425.
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scales, especially on middle of dorsum, in a large triangular area before tornus reaching to centre of wing, and in a small dense patch across R5 some distance from the apex; costal strigulae often indistinct in basal third and reduced to 4–5 small whitish spots along costa in distal half, rarely some confluent along costa; with few variably developed, short, oblique, orange brown spots or short oblique lines, one on costa from before middle, the next usually longer and from before 36: costa, and often a few groups or a line of orange brown scales along costal half of termen; fringe dark grey-brown, concolorous with wing. Hindwing with M2 distant and nearly parallel to very short stalk of M3 and CuA1, dark grey-brown, distinctly paler towards base; the lobe from base of anal margin ribbonshaped, reaching nearly to anal angle. Male genitalia (Fig. 514). Tegumen narrower at apex, deeply bilobed, the two distally rounded lobes parallel and with long, narrow scales laterally and across apex. Socii absent. Anal tube not spinulose. Valva elongate subtriangular, widest at base, sacculus hardly projecting, with a few bristles below basal excavation and a line of persistent long scales from outer surface near middle; bristled cucullus short, curved, slightly widened beyond ‘neck’ then tapering to single apical spine. Juxta small, caulis rather long. Aedeagus weakly curved in basal half, gradually tapering from wide base to pointed apex; no trace of cornuti. Female Wingspan 9.5–10.5 mm. As for male, but palpi not paler than rest of head, and without anal lobe in hindwing. Female genitalia (Fig. 516). Posterior corners of T8 extended ventrally as triangular lobes. Sterigma a moderately sclerotised, smooth large ring behind weakly concave hindmargin of S7. Ductus bursae as long as corpus bursae, membranous, gradually widening towards corpus bursae, elongate-bulbous and of thicker membrane below ostium with two narrow lens-shaped structures (colliculum?) anteriorly; anterior third from inception of ductus seminalis scobinate and more strongly widening. Corpus bursae ovate, with a small, triangular, tooth-shaped, signum-like sclerite in bursa ‘neck’, slightly larger than signa; with two small signa, pimple-shaped, one subapical and the second not far from bursa ‘neck’.
Distribution and biology Cyphophanes gracilivalva is known from islands off the north-eastern tip of Arnhem Land (Northern
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Territory) and off Cape York, and from Yeppoon (Queensland).
Remarks The bilobed tegumen apex and valva shape link C. gracilivalva much more closely to C. dryocoma than to C. dyscheranta, but the shape of the valva, as indicated by the specific name, and the weakly curved aedeagus leave no doubt that it is a separate species.
Eucosmogastra Diakonoff (Figs 63, 519–534) Eucosmogastra Diakonoff, 1975, Zool. Meded. (Leiden) 48: 307. Type species: Eucosmogastra anthochroa Diakonoff, 1975, ibid. 48: 308, textfig. 3, pl. 4, figs 14, 15, by original designation. [New Guinea.] Nenomoshia Clarke, 1976, Insects of Micronesia 9(1): 11 (key), 99. Type species: Argyroploce poetica Meyrick, 1909, J. Bombay Nat. Hist. Soc. 19: 437, by original designation. [New Guinea.]
Diagnosis Eucosmogastra species are unusual-looking, medium-sized olethreutine moths with characteristic blackish hindwings with yellowish or, more rarely, whitish patches in their basal half and with diversely patterned, elongate-subrectangular to distally slightly dilated forewings often marked with iridescent bluish or purplish blotches or with a pale grey irregular band along the basal half of the dorsum ending in an anteriorly projecting triangle. The labial palpi are either short, upcurved, rather slender and of equal width throughout or long, porrect with the second segment ventrodorsally strongly dilated to form a well-defined triangle with the long and slender terminal segment optically a continuation of the pale dorso-apical margin of the second segment. Eucosmogastra has chorda and M-stem well developed and all veins present and separate beyond the cell in the forewing, and a hindwing with Rs and M1 not very closely approximated at base, M2 bent towards M3 at base, M3 and CuA1 connate, and with long hairscales on both sides of 3A on the upper side. Males of some species (E. anthochroa, E. pyrrhopa (Lower), E. miltographa (Meyrick)) are further characterised by a modified and upwardly rolled
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anal margin concealing a scale pencil from the base of the wing. The genitalia in both sexes are diagnostic. Males of Eucosmogastra are characterised by a variably developed, ventrally bristled uncus, the conspicuous, uniquely modified socii set with flattened, hand-shaped, scale-like structures rather than bristles, a weakly sclerotised, band-shaped gnathos with medial spines, bristles or scobination, and by a slender valva with a sacculus ending in a broad tooth or an often serrate short blade. Females of Eucosmogastra are characterised by a sterigma consisting of a large, projecting, wellsclerotised and at least partially spinulose lip of diverse shape and ventral to the entrance to the ostium, by two equally large, slender, strongly curved, horn-shaped hollow signa widening near their circular base, and usually by a longitudinal band of needle-like spinules anteriorly in the corpus bursae. Oriodryas Turner has very similar female genitalia, but its sterigma is a sclerotised plate surrounding the ostium rather than only a ventrally projecting lip as in Eucosmogastra.
Description Adults (519–521) Wingspan. 12.5–18.5 mm (Australian species, one unidentified New Guinea species over 20 mm); medium-sized to large. Head (Figs 532, 533). Ocellus and chaetosema large. Vertex with moderately long to long scales, upper part of frons with anteriorly projecting scales, shortish in species with slender, upcurved labial palpi or with a long, horizontally projecting flat tuft in species with sinuate, triangular labial palpi, remainder of frons with minute, appressed scales. Maxillary palpus not visible on undissected head. Proboscis longer than labial palpus. Labial palpus of two radically different shapes: in E. anthochroa and E. poetica short (1.5–1.8× diameter of eye), upcurved, slender and of roughly same width throughout, with second segment of same width throughout or widest in middle and with moderately long, loose scales along entire ventral margin and around apex, and with terminal segment short, stout and only weakly inclined; in E. pyrrhopa and E. miltographa longer (1.9–2.4× diameter of eye), dorsoventrally strongly widened and conspicuously triangular, with second segment a triangle formed by ventral and dorsal margins and the conspicuously paler scale tips of the distally hugely widened apical tuft ending in a perfectly straight margin, the terminal segment long and slender, anteriorly inclined so that its pale scaling looks like an extension of the pale dorso-apical margin of the
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Figs 519–521. Eucosmogastra, adults, 519, 520, E. poetica. 519, , Bamaga, Q [image reversed]. 520, , nr Babinda, Q. 521, E. pyrrhopa, , Mt Webb, Q [image reversed]. Scale = 2 mm.
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second segment. Antenna to middle of wing, anterior surface without scales, with short cilia. Thorax. With very long, bipartite posterior crest; with a loose tuft of long scales from anteroventral angle of tegulae reaching to below base of hindwing. Legs not modified. Wings (Figs 63, 519–521). Forewing index 0.40–0.49. Retinaculum in male a hook from subcosta, in female a row of strong scales behind cubitus. Forewing elongate subrectangular or distally slightly dilated, costa curved at base and before apex with remainder straight to very faintly curved, apex pointed, slightly projecting in E. pyrrhopa, termen oblique and straight to sinuate; chorda forming a long, narrow accessory cell, Mstem well developed, lightly sinuate, to between M2 and M3; all veins present and separate beyond discal cell; CuP well developed near margin; anal loop 0.33–0.37× length of 1A+2A; wing pattern and colouring very diverse, usually complex and often with iridescent bluish to purplish blotches or markings (E. poetica, E. anthochroa); E. pyrrhopa and E. miltographa both with very different pattern with an irregular pale greyish band along basal half of dorsum ending in a triangle in middle of dorsum, E. pyrrhopa with pale greyish brown ground colour and little pattern except brown costal strigulae and scattered transverse strigulae across wing and dark brown suffusion in dorsal half of wing, E. miltographa with short orange red sinuate marks across entire wing on olive ochreous ground colour, recalling some Loboschiza species. Hindwing with a well-developed cubital pecten; axillary tuft not developed but upperside with dense, long hairscales on both sides of 3A particularly in male, and in males of some species (E. anthochroa, E. pyrrhopa, E. miltographa) anal margin modified, rolled upwards and enclosing a small to large scale pencil from base of hindwing, fringed with long hairscales and distal portion raised or triangularly folded over; Rs and M1 not very closely approximated at base, M2 bent close to M3 at base, M3 and CuA1 connate from angle of cell, origin of CuA2 variable, from not much beyond middle of cell to 36: cell; CuP present at margin, 1A+2A and 3A well developed; blackish with variably developed yellowish ochreous to orange yellow areas in basal half of wing (whitish and translucent in E. poetica), and sometimes with a yellow band along termen. Pregenital abdomen. Ventral apodemes short and stout in E. anthochroa, long and slender in distal 36: in remaining species dissected; anterior margin of S2 between apodemes deeply V-shaped and variably thickened. Male of E. anthochroa with
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T5–T7 with modified scaling (Fig. 534), particularly on T6 and T7 with various large tufts of long, strongly modified scales arising from lateral membranous and partly sclerotised lobes on both segments, in E. miltographa modified scales from two lateral membranous lobes next to spiracle on T7 only. In male hindmargin of S8 bilobed with a medial V-shaped emargination; hindmargin of T8 usually a projecting round or triangular lobe. Male genitalia (Figs 522–528). Vinculum very short. Tegumen ovate, tapering to subtriangular apex, with very broad, weakly sclerotised pedunculi, in some species (E. anthochroa) with a large bundle of very long scales dorsally from base of tegumen. Tegumen apex usually triangular (two small paired knobs in E. miltographa), with few bristles on dorsal surface and with a dense patch of strong, stout, short spines on apex or ventrally just below apex or on a second lower knob ventral to tegumen apex. Socii two large, pendant, swollen lobes often with a narrowed and even stalk-like base and with large, hand-shaped, scale-like structures instead of bristles. Gnathos weakly sclerotised, a broad, indistinct, U-shaped transverse band with a bipartite central clump of heavy spines in E. poetica, two more separated bunches of bristles in E. anthochroa, or with only medial scobination in the remaining species dissected. Valva well-sclerotised, long and slender to very slender, the often rectangularly projecting sacculus distally ending in a large tooth or blade, often serrate, remainder of valva from broadly crescentic and evenly bristled on inner surface with increasingly strong spines towards ventral margin (E. poetica) to very slender and constricted beyond middle, with bristles on inner surface often separated into a medial and a distal patch (E. anthochroa); costal process large, often bifid. Juxta small, caulis moderately long with lateral flanges. Aedeagus short, tapering towards oblique apex; vesica with a large bundle of lanceolate cornuti. Female genitalia (Figs 529–531). Ovipositor lobes from reniform to very slender elongate-ovate. Sterigma well posterior to weakly concave hindmargin of S7, a large, well-sclerotised, projecting and at least partially spinulose lip of diverse shape ventral to entrance to ostium, membrane dorsally to ostium spinulose but not sclerotised. Ductus seminalis narrow and membranous except for a lightly sclerotised lateral constriction just below ostium (colliculum); corpus bursae teardrop-shaped or ovate with a small posterior appendix giving rise to ductus bursae and ductus seminalis with an often lightly
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sclerotised granulose patch just posterior to origin of ductus seminalis, corpus bursae anteriorly usually with a longitudinal band of needle-like spinules and with two equally large, slender, strongly curved, horn-shaped hollow signa widening near circular base.
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Distribution Eucosmogastra is known from Sri Lanka, India, Southern Mariana Is (Guam), Papua New Guinea, and Australia where it has been recorded from Brisbane to Bamaga at the tip of the Cape York Peninsula in Queensland, from Darwin in the
Figs 522–528. Eucosmogastra, male genitalia and aedeagus. 522, 523, E. miltographa, Claudie R., Q, slide ANIC 2739. 524, E. anthochroa, Papua New Guinea, slide BM 24918. 525, 526, E. poetica, Bamaga, Q, slide T2478. 527, 528, E. pyrrhopa, nr Sarina, Q, slide T2267.
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Figs 529–534. Eucosmogastra, female genitalia, head and male abdomen tip. 529, E. pyrrhopa, Yeppoon, Q, slide T2281. 530, E. poetica, Bamaga, Q, slide T2479. 531, E. anthochroa, Papua New Guinea, slide BM 24919. 532, head, E. poetica, male. 533, head, E. pyrrhopa, male. 534, modified male abdomen tip, E. anthochroa.
Northern Territory and from the northernmost part of Western Australia in the Kimberley. Clarke (1976) assumed that the presence of E. poetica in Guam is due to an accidental introduction.
Biology Fletcher ([1921]) reported that E. poetica was reared from larvae rolling leaves of Polyalthia longifolia (Annonaceae) at Pusa, India.
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Remarks In the original descriptions of Eucosmogastra and its junior synonym Nenomoshia both authors (Diakonoff 1975; Clarke 1976) provided illustrations of male genitalia, head and wing venation, and Clarke (1976) also gave a photo of the adult. The male of E. anthochroa has very conspicuously modified scaling on T5–T7, which is absent in E. poetica, but there is no doubt that the two species are congeneric, with E. poetica possessing all the apomorphies of Eucosmogastra.
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Clarke (1958) figured adult and genitalia of the male types of E. miltographa and E. poetica (both from Sri Lanka and India), and of E. aeraria (Meyrick) and E. callicratis (Meyrick) (both from Khasi Hills, India), all as Olethreutes. The monophyly of Eucosmogastra is supported by several apomorphies in the genitalia of both sexes, but also by the blackish hindwing with yellowish or more rarely whitish patches in its basal half and with dense, long hairscales along 3A on its upper surface. Apomorphies in the male genitalia include the conspicuous, uniquely modified socii, pendant and often somewhat swollen lobes set with flattened, hand-shaped, scale-like structures rather than bristles and a slender valva with a sacculus ending in a broad tooth or an often serrate short blade. A large sterigma consisting of a projecting, well-sclerotised and at least partially spinulose lip of diverse shape ventral to the entrance to the ostium and a longitudinal band of needle-like spinules anteriorly in the corpus bursae are apomorphies for the female genitalia of Eucosmogastra. Judging from the various character states within Eucosmogastra a ventrally bristled uncus, a gnathos with medial spines or bristles and a valva with a rectangularly projecting sacculus and a crescentic distal portion probably are the groundplan conditions for the genus. All these features are also found in Oriodryas Turner. Eucosmogastra and Oriodryas furthermore share very broad pedunculi, a modified sacculus tip, a similarly shaped aedeagus, similar signa and a modified anal margin in the hindwing concealing a scale pencil from the base of the wing, strongly suggesting that the two genera are sister groups. Given the bristled tegumen apex of E. miltographa, the patch of bristles ventrally on the apex in E. pyrrhopa and then the tuft (E. poetica) or ventral patch of bristles (E. anthochroa) on a second hump or knob ventral to and considerably lower than the apex of the tegumen, it is assumed that these bristles represent the uncus, moved ventrally below the apex of the tegumen in the latter two taxa. Meyrick (1911b) considered the population of E. miltographa occurring from eastern Indonesia (Halmahera) to Papua New Guinea and to Australia to be sufficiently distinct to be called var. miltoxantha, based on the orange hindwing patches. However, this character is variable even between material from Sri Lanka and India.
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Constituent species Brown (2005) listed seven named species of Eucosmogastra, three of which occur in Australia, including E. poetica with a distribution including Sri Lanka, India, Guam, New Guinea and Australia and E. miltographa named from material from India and Sri Lanka. Several undescribed species are known from New Guinea. poetica (Meyrick, 1909), J. Bombay Nat. Hist. Soc. 19: 437 (Argyroploce), replacement name for mosaica Meyrick. mosaica (Meyrick, 1907c), J. Bombay Nat. Hist. Soc. 18: 138 (Eucosma), nec (Lower, 1901) (Temnolopha). [India.] pyrrhopa (Lower, 1896), Trans. R. Soc. S. Aust. 20: 161 (Tortrix (?)). [Qld: Mackay.] miltographa (Meyrick, 1907c), J. Bombay Nat. Hist. Soc. 18: 139 (Eucosma). [Sri Lanka, India.] miltoxantha (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 284 (Argyroploce), established as miltographa var. [Indonesia, New Guinea, Australia: Cairns.]
Helictophanes Meyrick (Figs 64, 65, 535–551) Helictophanes Meyrick, 1881, Proc. Linn. Soc. NSW 6: 637. Type species: Helictophanes uberana Meyrick, 1881, ibid. 6: 639, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 43, 61. [Australia.] Artiphanes Diakonoff, 1966, Zool. Verh. (Leiden) 85: 65. Type species: Articolla prospera Meyrick, 1909, J. Bombay Nat. Hist. Soc. 19: 591. [India.] Syn. nov. Ruthita Clarke, 1976, Insects Micronesia 9: 24. Type species: Ruthita argillacea Clarke, ibid. 9: 26, text fig. 6, pl. 2, fig. c, by original designation. [Southern Mariana Is.]
Diagnosis Species of Helictophanes are small to medium-sized and have a triangular or subrectangular forewing that, together with the strongly oblique costal strigulae and the orange and silver lines curving around the apex (Fig. 550), gives them a characteristic appearance. The forewing shape of H. prospera with the costa angled at 6M is unique,
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and males of those species with a ribbon-shaped process from the base of the anal area in the hindwing can be recognised easily. The long, sinuate palpi with a yellowish lateral spot usually surrounded by iridescent blue-grey scales on the second segment are typical for many enarmoniine genera. The venation of Helictophanes is characterised by a weak basal part of R4 (Figs 64, 65) and an often strongly oblique distal margin to the discal cell, and in H. prospera by a very thick R3 connate with R4. The male genitalia with their swollen cucullus and the sinuate or long, rod-like sclerites ventrally along the anal tube are diagnostic for Helictophanes. The female genitalia are characterised by their signa, two large, curved blades tapering to sharp points, arising from Tshaped bases.
Description Adults (Figs 535–540) Wingspan. 9–16 mm; small to medium-sized. Head (Fig. 551). Ocellus moderate. Projecting tuft of long scales from upper part of frons; remainder of frons with minute, appressed scales. Proboscis slightly longer than labial palpus. Labial palpus strongly sinuate, long (2.0–2.6× diameter of eye), second segment apically strongly widened by long scales along dorsal and ventral margin and often with characteristic colouring: laterally dark grey with blue iridescence and with central orangeyellow patch, terminal segment long and thin. Antenna from less than half length of forewing to beyond middle, with conspicuously paler scales on lower surface, entirely scaled or with a narrow anterior band with short cilia. Thorax. With a long, pointed posterior crest of hardly raised scales; with small scale tuft from anteroventral angle of tegula. Legs unmodified. Wings (Figs 64, 65, 535–540, 550). Forewing index 0.40–0.41. Retinaculum in male a hook from subcosta, in female a row of strong scales behind cubitus. Forewing in Australian species subtriangular, with costa curved distally or sharply angled at 6M (H. prospera), in non-Australian species forewing often subrectangular; apex acute to nearly rectangular, always broadly rounded, termen more or less oblique; all veins separate beyond cell; cell often with strongly oblique distal margin, with chorda, if present, from below R2 to R5, and Mstem to between M2 and M3; R3 variably approximated to base of R4 (Figs 64, 65), sinuate, very thick and connate with R4 in H. prospera; basal part of R4 only a weak trace from angle of cell; M3
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either closer to M2 or to CuA1; anal loop 0.29 length of anal vein in H. prospera; smooth-scaled, with three short, parallel, oblique yellowish and silvery lines from before middle of costa followed by a dark mark at or beyond middle and by three parallel white, silvery and yellow lines curving from before 56M costa to below apex, with four silver strigulae separated by orange scales along apical fourth of costa (Fig. 550). Hindwing with cubital pecten and axillary scale tuft well developed, in males of some species (H. prospera, H. scambodes (Meyrick)) with long, scaled or naked, ribbon-like process from base of anal margin; Rs and M1 parallel and closely approximated in basal half or the two veins connate or short-stalked; M2 bent towards stalk of M3 and CuA1 at base; CuP well developed in apical third; anal veins well developed. Pregenital abdomen. Unmodified. Sternum 8 and T8 in male weakly sclerotised, T8 a narrow triangle in H. prospera. Male genitalia (Figs 541–547). Vinculum wide and fused with tegumen, forming complete ring. Tegumen weakly sclerotised except for rim-like anterior margin often with two pointed muscle attachment points; with folds of scale-bearing, modified membrane between tegumen and valva. No developed uncus; apex of tegumen a variably projecting transverse band or triangular lobe with two to four prominent dorsal setae. Socii variable, from two large, elongate lobes projecting beyond apex of tegumen (H. uberana) to ill-defined, hairy lateral patches (H. prospera). Gnathos at most two very weak lateral arms, each connected to S-shaped sclerites along ventral surface of prominent anal tube (H. prospera), often distally lengthened and reaching beyond apex of tegumen (H. uberana). Valva weakly sclerotised, with large basal excavation, abruptly constricted beyond ventrally projecting sacculus area, costal process sometimes large; cucullus swollen, often club-shaped with deeply rounded outer and densely long-haired inner surface, tapering to single or bifid apical thorn, or flattened with thorn subapical. Caulis straight and moderately long, anellus a wide membranous ring. Aedeagus short, weakly sclerotised except ventrally towards apex, vesica with numerous sinuate, spindle-shaped cornuti. Female genitalia (Figs 548, 549). Ovipositor lobes elongate and narrowed to moderately broad. Sternum 7 subtriangular, weakly sclerotised, posterior margin sometimes with a small emargination with sharp, sclerotised edge. Sterigma a pronounced ventral lip delineating a membranous
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Figs 535–540. Helictophanes, adults. 535, 536, H. prospera. 535, , nr Ravenshoe, Q. 536, , Base Cableway, Mt Bellenden-Ker, Q [image reversed]. 537, 538, H. uberana, , Richmond R., NSW. 539, 540, H. scambodes, Kuranda, Q [BMNH]. 539, . 540, [image reversed]. Scale = 2 mm.
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Figs 541–547. Helictophanes, male genitalia and aedeagus. 541, 542, H. prospera, Mt Bellenden-Ker, Q, slide T2324. 543, H. scambodes, Kuranda, Q, slide BM 24892. 544, 545, H. uberana, Richmond R., NSW, slide T2468. 546, 547, H. sp., Mt Keira, NSW, slide T2248.
cup, membranous lamella postvaginalis densely spinulose. Ductus bursae moderately long to long, with ductus seminalis from below mid-length; membranous and smooth except for a wide ring of scobination in basal third below inception of ductus seminalis and a small folded sclerite (colliculum) below ostium (often weakly developed). Corpus bursae roundish, finely granulated, with two large signa, one dorsally close to entrance of ductus and one ventrally; signa two
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large curved blades tapering to sharp points, arising from T-shaped bases.
Distribution Helictophanes has been recorded from India (Assam), New Guinea, Micronesia and Australia, where it is restricted to rainforest areas on the eastern coast of New South Wales and Queensland, from Mt Dromedary near Narooma to the Cairns region.
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Figs 548–551. Helictophanes, female genitalia, forewing apex and head. 548, H. prospera, female genitalia, Brisbane, Q, slide T2374. 549, H. uberana, female genitalia, Mt Keira, NSW, slide T2473. 550, detail wing pattern, forewing apex, H. sp., Papua New Guinea. 551, head, H. prospera, male.
Biology A specimen of H. uberana in the ANIC has been reared from fruit of Alyxia ruscifolia (Apocynaceae) by A. W. S. May. Turner (1946) reported rearing a series of H. prospera from the fruit of Omphalea (as Omphaletis) queenslandiae (Euphorbiaceae) and the holotype of H. myriolychna (Turner) from fruit of Hearnia sapindina (Meliaceae).
Remarks The synonymy of Ruthita and Helictophanes is obvious with the two type species sharing very similar genitalia in both sexes, even though Rs and M1 in the hindwing are connate in H. uberana and separate in H. argillacea. Helictophanes prospera, the type species of Artiphanes, looks superficially quite distinct with an angled costa and a conspicuous, ribbon-like process from the base of the anal area of the hindwing, both derived features absent in H. uberana. However, an undescribed Australian
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species with subtriangular wings combines a short, scaled, ribbon-like process from the base of the anal area with elongate socii projecting beyond the apex of the tegumen and with apically straight and elongate ventral sclerites to the anal tube, the two derived traits characterising H. uberana and H. argillacea. The subtriangular wings of the undescribed species, with a barely oblique distal margin to the forewing cell, are probably plesiomorphic with respect to the uniquely derived wing shape and venation of H. prospera. Articolla cyclidias Meyrick from Sri Lanka (Meyrick 1907b) with exactly the same venation as H. prospera is probably congeneric as well, but given its reduced tegumen, study of the female genitalia will be required to confirm this. The monophyly of Helictophanes is based on the swollen and enlarged cucullus and the S- or rodshaped sclerites ventrally along the anal tube in the male, and the curved, blade-shaped signa with a T-shaped base in the female. The ribbon-like process
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from the base of the hindwing margin in H. prospera and some other species provides a link to Cyphophanes that has an elongate lobe in the same position. The prominent R3 in H. prospera recalls the modified venation of Metaselena Diakonoff. Helictophanes uberana has an extremely variable wing pattern, with H. uberana based on ochreous brownish forms and the junior synonym H. metallocosma on a mixed series of two species. Both syntypes of H. metallocosma lack the abdomen and are in very bad condition, with the specimen from Brisbane a dark grey specimen of H. uberana with considerable white scaling and the specimen from Mt Tamborine apparently a specimen of an undescribed species of Helictophanes with narrower wings and a more oblique termen, with further material from upland New South Wales rainforest areas in the ANIC. To ensure nomenclatural stability, the Brisbane specimen is here formally designated as the lectotype. The genitalia of H. prospera show considerable variation in the shape of the apex of the tegumen, the aedeagus and the signa even within the Australian population that at present is considered conspecific with the type from Assam. Only the female holotype is known of H. myriolychna, but wing pattern, shape and venation and the female genitalia refer the species to Helictophanes. Kawabe (1989) figured H. prospera from Thailand, erroneously identified as Enarmonia mixanthes (Meyrick), a species that could possibly also belong to Helictophanes. Cyphophanes dryocoma (Meyrick) was included in Helictophanes by Clarke (1958), but is transferred to Cyphophanes above.
Constituent species Five species of Helictophanes are known from the Oriental/Australian regions, with four of the named and one undescribed species found in Australia. uberana Meyrick, 1881, Proc. Linn. Soc. NSW 6: 637 (Helictophanes). [NSW: Sydney.] metallocosma Turner, 1925, Trans. R. Soc. S. Aust. 49: 57 (Helictophanes). Lectotype ‘Brisbane, 28–2–23 [Turner]’, ‘Helictophanes metallocosma Turn. Type’, hereby designated. [Qld: Brisbane.] prospera (Meyrick, 1909), J. Bombay Nat. Hist. Soc. 19: 591 (Articolla). [India.] Comb. nov. scambodes (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 277 (Argyroploce). [Qld: Cairns.] Comb. nov. myriolychna (Turner, 1946), Trans. R. Soc. S. Aust. 70: 218 (Articolla). [Qld: Mossman.] Comb. nov.
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Irianassa Meyrick (Figs 66, 552–560) Irianassa Meyrick, 1905, J. Bombay Nat. Hist. Soc. 16: 609. Type species: Irianassa sapphiropa Meyrick, 1905, ibid. 16: 609, by monotypy. [Sri Lanka.]
Diagnosis Irianassa species are medium-sized, conspicuous and unmistakable moths with iridescent blue, broadly triangular, blackish forewings with a bluntly rounded apex preceded by two large white strigulae, with a terminal notch some distance from the apex accented by a triangular tuft of white fringe scales, and usually with a few curved silvery or metallic blue lines originating from the costa, one running from about 36: of costa towards the terminal notch. The hindwings are brownish black and have nearly always an unscaled, transparent area along the hindmargin of the cell. Irianassa has large ocelli and short, slender, upcurved labial palpi with a relatively large, pointed terminal segment. All veins are widely separate in the forewing with CuA1 from far below the angle of the cell and both the M-stem and chorda well developed. The hindwings lack a cubital pecten, have hairscales along 1A+2A, a frenulum of only two bristles in the female, Rs and M1 widely separate at base, and M2 distant from M3 that is connate with CuA1 from the angle of the cell. The male genitalia are highly diagnostic with long scales laterally on the tegumen with its lateral margins sclerotised, projecting and sometimes modified, without uncus, with long, slender, strongly sinuous gnathos arms originating from near base of tegumen and fused at base of anal tube (sometimes partially reduced), with a rather simple, elongate valva with a very large basal excavation, a truncate apex with a ventral spine and a variably developed dorsal lobe, with a small juxta, and with caulis and anellus forming a hoodshaped structure, and with a very long, slender aedeagus. The female genitalia are unique and have long, parallel-sided, modified ovipositor lobes with strongly projecting posterior lobes, a smooth, short, chute-shaped sterigma in the intersegmental membrane, a ductus bursae with a modified colliculum and an oblique sclerotised ring near corpus bursae, and the corpus bursae partially lined with composite spines with star-shaped bases.
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Description Adults (Figs 552, 553) Wingspan. 12–17 mm; medium-sized. Head (Fig. 554). Ocellus large, conspicuous. Upper part of frons with shortish, anteriorly oriented scales, remainder with minute, appressed scales. Maxillary palpus not visible on undissected head. Proboscis slightly longer than labial palpus. Labial palpus small (1.4–1.7× diameter of eye), slender, upcurved, with appressed scales, second segment curved, narrowly crescentic, third segment large, nearly 56O length of second segment, pointed. Antennae rather short and stout, to middle of wing, dorsally scaled and ventrally with very short, dense cilia. Thorax. Without posterior crest; blackish with strong bluish iridescence as rest of body. Legs unmodified. Wings (Figs 66, 552, 553). Forewing index 0.46– 0.49. Forewing broadly triangular with a conspicuous notch in termen far from apex, with costa evenly curved, apex blunt, broadly rounded, termen vertical and strongly sinuate with notch at M1, enhanced by a V-shaped tuft of whitish fringe scales, tornus broadly rounded; M-stem well developed, straight, to just behind M2, chorda from halfway between R1 and R2 to base of R5; all veins present and widely separate beyond cell; bases of R3, R4 and R5 nearly equidistant, M1 slightly closer to R5 than to M2, M3 from angle of cell, CuA1 from far below angle and CuA2 from about middle of cell; CuP present at wing margin; anal loop 0.22 length of 1A+2A; blackish brown with bluish iridescence across entire wing, with two very large, nearly vertical, white strigulae before apex and a series of metallic bluish or silvery bands originating along costa, sometimes from remnants of white strigulae, with a partially fused and otherwise roughly parallel pair from just below middle of costa strongly outwardly curved and running to beyond middle of dorsum, followed by a short, oblique dash at about 69 costa, then by a curved bluish line connecting to end of apical strigula and a further short, oblique dash close to white strigulae; termen with a notch at M1 enhanced by a tuft of whitish fringe scales and with a subparallel, curved silvery band along its dorsal 36:. Hindwing usually partially unscaled and translucent, without cubital pecten but in both sexes with long hairscales along 1A+2A, frenulum in female with two bristles only; Rs distant from Sc+R1 and extending only about 56: below angle of cell before fading, bases of Rs and M1 conspicuously distant, M2 distant from and subparallel with M3,
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M3 and CuA1 connate from angle of cell, CuA2 from about 69 cell; distal half of CuP well developed, halfway between CuA2 and 1A+2A; 1A+2A with dense scale sockets, but not tubular and without basal loop, 3A present; blackish brown, usually with
Figs 552–554. Irianassa, adults and head. 552, I. sp., , McIlwraith Range, Q [image reversed]. 553, I. aetheria, , Darwin, NT [image reversed]. 554, head, I. sp., male. Scale = 2 mm.
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species-specific portion of dorsal half unscaled and translucent. Pregenital abdomen (Fig. 557). Unmodified; anterolateral process of S2 reduced. In male hindmargin of S8 weakly bilobed, membrane between genitalia and S8 very long and sometimes ventrally sclerotised. Male genitalia (Figs 556, 559). Vinculum wide and short, fused with valva. Tegumen robust, with ventroposterior margin sclerotised and projecting, laterally with dense, very long scales, muscle attachment on pedunculi small. No trace of uncus, anal tube passing between ventrally projecting lateral tegumen margins (sometimes with a hooklike process) and socii and reaching to beyond apex of tegumen. Socii from large, densely hairy, projecting lobes to small, drooping remnants. Gnathos arms extremely long, slender, strongly sinuous bands originating from tegumen margin near base, fused at base of anal tube and extending as a sclerotisation along its ventral surface; in derived genitalia only base of arms present and hook-like sclerotisations at base of anal tube. Valva with very large basal excavation to well beyond middle with dense, long bristles along its ventral margin, widened towards truncate apex with a ventral spine and a variably developed dorsal lobe; inner surface distally variably bristled, sometimes with blunt spines and a dorsal keel beyond basal excavation; with large, club-shaped costal process. Juxta very small, fused with elongate, hood-shaped structure formed by caulis and anellus. Aedeagus very long and slender, lightly to strongly sinuate, apex dorsally membranous and spinulose; no cornuti nor their sockets visible. Female genitalia (Figs 555, 560). Ovipositor lobes modified, long, straight, parallel-sided, with posterior lobes strongly projecting, sometimes with bristles arranged in transverse rows and/or some papillae enlarged and peg-shaped towards apex. Hindmargin of S7 concave. Sterigma a smooth, short, chute-shaped sclerite forming projecting lamella antevaginalis in intersegmental membrane. Ductus bursae gradually widening anteriorly, membranous except for short sclerotised portion just below ostium (modified colliculum?) and an oblique sclerotised ring near corpus bursae, with ductus seminalis originating in anterior third; corpus bursae ovate, with composite spines with star-shaped base in a ring around subapical entrance to corpus bursae and forming large patch on opposite bursa wall.
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Distribution Irianassa has been reported from Sri Lanka, India, New Guinea, Samoa and Australia, where it has been collected from the tip of Cape York to the Cairns region in Queensland and in the vicinity of Darwin in the Northern Territory.
Biology No host record is known for Irianassa. The Australian specimens have been collected in rainforest localities.
Remarks Meyrick (1905: 609) described Irianassa in the Plutellidae, with the remark ‘Allied to Hilarographa’. Fletcher (1929) included the genus in the Glyphipterigidae and Heppner (1982) transferred it to the Chlidanotinae (Tortricidae). A forewing pattern with pronounced costal strigulae, together with the notched termen and short labial palpi, give Irianassa some superficial similarity to hilarographine Chlidanotinae, but the structure of the genitalia and antenna refer the genus to the Enarmoniini despite the lack of a cubital pecten. The widely distant Rs and M1 in the hindwing probably led Turner to describe the Australian I. aetheria Turner in Laspeyresia. Clarke (1969) figured adult, genitalia and wing venation of the male holotypes of I. sapphiropa and I. poecilaspis Meyrick. Based on these figures the two species do not look congeneric, and further study is clearly required. The number of unique apomorphies characterising Irianassa is well illustrated by the diverse family assignments for the genus and its striking appearance. Given only the highly derived genitalia of I. sapphiropa and related species with modified, partially translucent hindwings, it was difficult to find convincing synapomorphies to refer Irianassa to any of the olethreutine tribes, though forewing shape and pattern and the extremely large basal excavation did strongly suggest a position in the Enarmoniini. An undescribed Australian species with entirely scaled hindwings and more plesiomorphic male genitalia including large socii and a distally bifid valva with a large ventral thorn squarely fits into the Enarmoniini, but no synapomorphies pointing to a sister group have been identified. The most conspicuous apomorphies of Irianassa are a
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Figs 555–560. Irianassa, genitalia and abdomen. 555, I. sp., female genitalia, Mt Bartle Frere, Q, slide T2170. 556, I. sp., male genitalia, Papua New Guinea, slide ANIC 13808. 557, male abdomen, 8th segment, I. sp., Papua New Guinea. 558, detail of corpus bursae of I. aetheria. 559, I. sp., male genitalia, Papua New Guinea, slide T2168. 560, I. aetheria, female genitalia, nr Darwin, NT, slide ANIC 13807.
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forewing with iridescent scaling, a unique wing pattern, a notch on the termen and CuA1 from well before the angle of the cell, a hindwing without cubital pecten and with Rs and M1 widely distant at base, small and upcurved labial palpi, male genitalia with gnathos arms apparently from near the base of the tegumen and with caulis and anellus fused to form a hood-shaped structure, and entirely unique female genitalia with modified anal lobes, a chute-shaped sterigma and the corpus bursae set with numerous composite spines.
Constituent species Heppner (1982) and Brown (2005) listed four species of Irianassa from outside of Australia, but the generic relationships of I. poecilaspis from Assam will have to be reassessed (Clarke 1969). One named and two unidentified species are known from Australia. aetheria (Turner, 1946), Trans. R. Soc. S. Aust. 70: 219 (Laspeyresia). [Qld: Cape York.]
Loboschiza Diakonoff (Figs 67, 561–575) Loboschiza Diakonoff, [31 January 1968], US Nat. Mus. Bull. 257: 40 (key), 93. Type species: Argyroploce clytocarpa Meyrick, 1920, Exot. Microlepid. 2: 349, by original designation. [Philippines.] Rhadinoscolops Obraztsov, September 1968, J. New York Entomol. Soc. 76: 187. Type species: Pyralis koenigiana Fabricius, 1775, Syst. Entomol.: 653, by original designation. [‘India orient.’]
Diagnosis Given their conspicuous pattern and colour, species of Loboschiza can often be superficially recognised. They are small to medium-sized and usually have fairly long, sinuate labial palpi, seldom they are slender and upcurved. The male antennae has short to moderately long cilia. The forewing pattern is usually bipartite with the basal half to two-thirds either bright yellow, orange or with bright red bands on brownish ground, and the darker distal portion, separated by a transverse leaden band, brownish or blackish, often with bright orange sinuous scribbles. More rarely, the entire forewing
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is bright red with obliquely transverse leaden lines and black scribbles, or dark brown with a yellow transverse band before middle and an ochreous band along termen. In the forewing the M-stem and sometimes also the chorda are usually present, in the hindwing M2 and M3 are nearly always widely separated and roughly parallel, M3 and CuA1 are connate or stalked. In the male genitalia the valva has a bipartite cucullus with a ventral process usually ending in a thorn, and/or a large tuft of long bristles from the ventral margin beyond the sacculus, the socii are two sublateral, bristled lobes with free apex, and the aedeagus has a large bundle of cornuti. The female genitalia are quite variable, with a colliculum and usually with a variably developed sclerotised area in the ductus bursae associated with the ductus seminalis, and with either one or two signa, usually concave scobinate areas, rarely with a small thorn in the centre.
Description Adults (561–563, 565, 566) Wingspan. 8–14 mm; small to medium-sized. Head (Fig. 564). Ocellus well developed to large. Projecting tuft of long scales from upper part of frons, remainder with minute, appressed scales. Proboscis slightly longer than labial palpus. Labial palpus usually sinuate, porrect to slightly oblique, moderately long (1.5–2.5× diameter of eye), with second segment distally strongly widened above and below, terminal segment long, thin and slightly drooping; labial palpus rarely small, slender and upcurved, with terminal segment continuing curve of second segment (L. furiosa (Meyrick)). Antenna short, hardly to middle of forewing, usually with short but dense cilia in male, rarely with longer cilia (56O width of flagellum in L. furiosa). Thorax. Smooth. Hindlegs in some species (L. hemicosma (Lower)) with an expandable tuft of hairscales dorsally on base of tibia. Wings (Figs 67, 561–563, 565, 566). Forewing index 0.42–0.49. Forewing from subtriangular to subrectangular; with all veins separate beyond cell, R4 and R5 rarely connate; chorda, if present, from between R1 and R2 or between R2 and R3 to between R5 and M1, and M-stem to M2 or between M2 and M3; in L. exemplaris (Meyrick) and L. hemicosma R4, R5 and M1 all closely approximated at angle of cell and accessory cell small and narrow; anal loop 0.29 length of anal vein in L. koenigiana (Fabricius); smooth-scaled, with very diverse, unusual and conspicuous,
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Figs 561–566. Loboschiza, adults and head, males. 561, L. koenigiana, Byfield, Q [image reversed]. 562, L. exemplaris, Mt Webb Nat. Park, Q. 563, L. sp., Lockerbie, Q [image reversed]. 564, head, L. koenigiana. 565, L. delomilta, Kuranda, Q [BMNH]. 566, L. furiosa, Cairns, Q [image reversed]. Scale = 2 mm.
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usually bipartite wing pattern with basal half to 36: either bright yellow, orange or with bright red bands on brownish ground, and distal portion darker, brownish or blackish, often with bright orange sinuous scribbles; distal portion separated from rest of wing by a transverse leaden band; costa with strigulae throughout; forewing of L. furiosa and L. delomilta (Turner) not bipartite but bright red throughout, with obliquely transverse leaden lines and black scribbles across wing, and an undescribed Australian species dark brown with a transverse yellow band before middle and an ochreous band along termen. Hindwing with cubital pecten and axillary tuft weak to well developed; Rs and M1 closely approximated and parallel near base; M2 distant from stalk of M3 and CuA1 at base with M2 and M3 roughly parallel,
only in one undescribed species base of M2 curved towards M3; M3 and CuA1 connate to stalked to about 56M; CuP developed in apical third, anal veins well developed. Pregenital abdomen. Some species (L. koenigiana, L. hemicosma) with a tuft of modified scales from enlarged sockets between ventral apodemes; in others (unidentified species from Woodlark Is, Papua New Guinea) dorsally in the membrane between segments 3 and 4 with a pair of expandable scale tufts each next to a patch of modified, short and dense scales with large sockets and on segment 7 with laterally projecting triangular lobes; L. clytocarpa with a lateral, longitudinal groove with modified scales below tergites on segments 3–5. Sternum 8 and T8 in male weakly sclerotised, hindmargin of S8 bilobed, T8
Figs 567–572. Loboschiza, male genitalia. 567, 568, L. koenigiana, nr Marlborough, Q, slide T2110. 569, 570, L. martia, Shute Harbour, Q, slide T2306. 571, L. exemplaris, nr Cooktown, Q, slide ANIC 2713. 572, L. delomilta, Kuranda, Q, slide BM 24913.
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usually with a sclerotised, transverse bar at base and a narrow triangle of sclerotisation extending to hindmargin. Male genitalia (Figs 567–572). Vinculum a moderate band, fused with tegumen to form a complete ring. Tegumen moderately sclerotised, often with obvious lobe projecting from base of anterior margin. No developed uncus. Socii two sublateral, distally free, somewhat pendulous bristled lobes, with bristles sometimes extending to back of tegumen; in L. furiosa and L. delomilta socii forming a dorsally bristled, dome-shaped structure crowning the triangular tegumen tip. Gnathos two often weak, curved lateral arms arising from posterior margin of tegumen and medially fused with anal tube. Valva moderately sclerotised, usually somewhat narrowed beyond ventrally projecting sacculus region and distal half usually bipartite and inner surface densely hairy; sacculus small to very elongate, bristled, often followed by a dense patch of long bristles from ventral margin; cucullus usually with a rounded distal lobe, rarely crescentic and/or swollen, and with a subapical, triangular, ventral lobe nearly always ending in an often bipronged thorn; cucullus in L. furiosa and L. delomilta not bipartite but with an oblique ridge across inner surface, thorn ventrally on rounded apex; costal process long. Caulis very long except in L. furiosa and L. delomilta, anellus weakly developed. Aedeagus long and slender, often slightly curved at base and tapering towards apex; vesica with a very large bundle of curved, lanceolate, deciduous cornuti, often filling apical 36: of aedeagus. Female genitalia (Figs 573–575). Ovipositor lobes fairly slender. Position of ostium variable, usually posterior to hindmargin of S7 in shallow, cup-shaped depression often surrounded by ring of spinulose membrane (L. hemicosma, L. halysideta (Walsingham), L. martia (Meyrick)), sometimes below a triangular medial projection of the hindmargin of S7 (L. koenigiana) or at the bottom of a sclerotised tube fused to S7 (L. furiosa). Ductus bursae about as long as length of corpus bursae, narrow and widening towards corpus bursae, nearly always with two sclerotised regions, the colliculum often only two lateral, minute, crescentic sclerites, the second sclerotisation variably developed near middle of ductus bursae at origin of ductus seminalis, in one undescribed Australian species together with origin of ductus seminalis at junction with corpus bursae on ventral surface; in some species (L. koenigiana) the entire ductus bursae sclerotised with only a narrow membranous ring
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below colliculum. Corpus bursae globose to ovate, with dimpled surface and in some species with a conspicuous ring-like crease at junction of ductus and corpus bursae; with one or two signa, usually a somewhat depressed area of scobination only, often inconspicuous, but rarely with a small thorn (L. halysideta) or a remnant thereof in its centre.
Distribution Loboschiza is widely distributed across the IndoAustralian region, from Sri Lanka, India and China to Japan, the Philippines, Papua New Guinea and Australia, including an endemic species on Christmas I. In Australia it occurs in riverine vegetation (Drysdale R.) and rainforest and marginal rainforest locations from the Kimberley in Western Australia, across the Northern Territory and Queensland to Wingham and the Allyn R. near Gloucester, New South Wales.
Biology Fletcher (1932: 33–34) described the biology of L. koenigiana from Pusa, India, with the larvae ‘nibbling the surfaces’ of ‘rolled or superimposed leaves’ of Melia azedarach (Meliaceae), adding that ‘when young, the larva bores the tender topshoots of its foodplant’, and ‘sometimes, the larva continues as a shoot-borer until it is full-grown, but most larvae seem to be leaf-feeders’. Yasuda (1969) figured the larva of L. koenigiana from Japan, feeding in spun leaves of M. azedarach. I. F. B. Common reared L. koenigiana in August/ September from fruit of Owenia venosa (Meliaceae) from Mt Larcom, Queensland, and the single holotype of L. deloxantha was reared in July by A. J. Turner from fruit of Aglaia (as Hearnia) sapindina (Meliaceae) from Mossman, Queensland. Unlike his undoubted references to rearings of L. koenigiana from Melia, Fletcher ([1921]: 62) states cautiously that ‘at Surat it [L. koenigiana] is said to have bred from larvae on mogra (Jasminum sambac)’ (Oleaceae). The host plants listed for Loboschiza (Robinson et al. 2001) are predominantly Meliaceae, with two references to Jasminum sambac (Oleaceae) and one to Hibiscus rosa-sinensis (Malvaceae).
Remarks Clarke (1958) figured adult, abdomen and male genitalia of the holotype of L. clytocarpa. Drawings of head, wing venation and male genitalia of
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L. clytocarpa were provided by Diakonoff ([1968]) and of head, wing venation and genitalia of both sexes of L. koenigiana by Obraztsov (1968). No universally valid apomorphy has been identified for Loboschiza, but except for an undescribed Australian species they all have M2 distant from and parallel to M3 in the hindwing. Both male and female genitalia are remarkably diverse throughout the genus, but the extremes are linked by intermediate states. Several species have a patch of modified scales at the base of the abdomen between the ventral apodemes. The two species L. furiosa and L. delomilta both differ from the rest of the genus in not having a bilobed valva tip, different wing pattern and labial palpus and, in L. furiosa, the only female available, with the posterior part of the ductus bursae fused with S7. But the presence of a tuft of long bristles from the ventral margin of the valva beyond the sacculus provides a link with the majority of Loboschiza species, and the two taxa are included for the time being. Loboschiza and Rhadinoscolops were described in the same year, and later synonymised by Diakonoff
(1982). The features separating the two taxa: different secondary sexual structures in the male and a different course of M3 and CuA1 in the hindwing (stalked in Loboschiza and connate in Rhadinoscolops), are variable throughout the genus. Confusion about the position of the Enarmoniini is reflected in Diakonoff’s ([1968]) comment ‘with eucosmine affinities’ on Loboschiza and Obraztsov’s (1968) inclusion of Rhadinoscolops in the ‘Laspeyresiini’ while comparing it with Eucosmomorpha Obraztsov, Pseudophiaris Obraztsov and Cimeliomorpha Obraztsov, all three now regarded as Enarmoniini. Anthozela shares the course of M2 widely distant from M3 in the hindwing with the majority of Loboschiza, also the bipartite wing pattern, and is possibly the sister taxon. The modified scales beneath at the base of the abdomen in some species provide a link with Anathamna and Metaselena. Species of Loboschiza are often collected as singletons, and the smaller species look very similar superficially, so the genus will probably turn out to be very rich in species. Given the course of M3 and
Figs 573–575. Loboschiza, female genitalia. 573, L. koenigiana, Mt Larcom, Q, slide T2111. 574, L. thoenarcha, Kuranda, Q, slide BM 24912. 575, L. sp., Annan R. crossing, Q, slide ANIC 12962.
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CuA1 most of the species here recognised as Loboschiza were initially described as species of Laspeyresia. The endemic Christmas I. species L. halysideta (Walsingham) had been provisionally, but quite correctly, transferred to Loboschiza by Tuck (1988), after Meyrick (1912a) had earlier referred it to Polemograptis on the basis of the red forewing markings. The following non-Australian species are here transferred to Loboschiza: cinnabaritis (Meyrick, 1928), Exotic Microlepid. 3: 451 (Laspeyresia). [St Matthias Is.] Comb. nov. gratulata (Meyrick, 1916), Exotic Microlepid. 2: 23 (Laspeyresia). [Philippines, Mindanao.] Comb. nov. mediana (Walker, 1866), List Specimens Lepid. Insects Colln. Br. Mus. 35: 1798 (Tospitis) [New Guinea.] Comb. nov. turifera (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 876 (Laspeyresia). [India.] Comb. nov. An undescribed Australian species with dark brown forewings with a yellow, transverse band before middle and an ochreous band along termen is probably the sister group to the remaining species as it has the base of M2 in the hindwing bent towards M3, large socii, and the sclerotised region of the ductus bursae associated with the origin of the ductus seminalis close to the corpus bursae rather than halfway between it and the ostium.
Constituent species Including L. halysideta from Christmas I. and L. koenigiana, we are aware of seven species of Loboschiza from outside mainland Australia. It is doubtful whether L. vulnerata (Walker) is indeed a junior synonym of L. koenigiana, but only study of the entire south-east Asian fauna of Loboschiza will resolve this question. Eight named and three unnamed species are known from Australia: koenigiana (Fabricius, 1775), Systema Entomologiae: 653 (Pyralis). [‘India orient.’] koenigana (Fabricius, 1787), Mantissa Insectorum 2: 237 (Pyralis), misspelling of koenigiana. aurantiana (Pryer, 1877), Cist. Entomol. 2: 235 (Hemerosia). [China.] vulnerata (Walsingham, 1900), in Swinhoe, Cat. East. and Aust. Lepid. Heterocera 2: 571 (Eucelis). [Myanmar.] martia (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 291 (Laspeyresia). [Qld: Cairns.] Comb. nov. hemicosma (Lower, 1908), Trans. R. Soc. S. Aust. 32: 321 (Leptarthra). [Qld: Kuranda.] Comb. nov.
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exemplaris (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 296 (Laspeyresia). [NT: Port Darwin.] Comb. nov. deloxantha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 219 (Laspeyresia). [Qld: Mossman.] Comb. nov. thoenarcha (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 291 (Laspeyresia). [Qld: Cairns.] Comb. nov. furiosa (Meyrick, 1921), Exotic Microlepid. 2: 450 (Laspeyresia). [Qld: Cairns.] Comb. nov. delomilta (Turner, 1946), Trans. R. Soc. S. Aust. 70: 220 (Laspeyresia). [Qld: Cape York.] Comb. nov.
Metaselena Diakonoff (Figs 68, 576–583) Metaselena Diakonoff, 1939, Zool. Meded. (Leiden) 21: 126 (key), 156. Type species: Metaselena alboatra Diakonoff, 1939, ibid. 21: 158, figs 5O– P, 6E, by original designation. [New Guinea.]
Diagnosis Metaselena species are very small to medium-sized olethreutine moths with subrectangular to subovate forewings, with the termen protruding further than the often obtuse apex and nearly always with a unique and very reduced wing pattern of mainly prominent brownish costal strigulae on a pale cream-coloured to greyish yellow ground. The labial palpi are rather short, sinuate, porrect or curved upward. The forewing venation is unique and diagnostic for the genus with R3 distally swollen and its often very slender base much closer to R2 than to the stalk of R4 and R5, sometimes R2 and R3 on a common stalk or R2 also slightly swollen distally. In the hindwing the cubital pecten is very weakly developed or absent, the anal area in the male is expanded to form a projecting triangular lobe usually with a thickened anal margin, and the venation very variable with either M2, M3 and CuA1 all rather close together, sometimes with M3 and CuA1 stalked, or with CuA1 distant from M3, halfway between M3 and CuA2. Metaselena males share with those of Anathamna a tibial pencil on the hindleg and two symmetrical patches of modified scales on the posterior surface of the metathorax and two opposite on the base of the abdomen. The male genitalia are very similar to those of Anathamna, but the distal portion of the valva is much more slender in Metaselena, at most slightly swollen before the apex but never expanded into a triangle,
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and the apical thorn is simple and not bipronged as in Anathamna. The female genitalia of Metaselena are characterised by: a single signum, a rounded, deeply concave, strongly granulose patch; a small sclerite in the ductus bursae posterior to origin of the ductus seminalis, and by a sterigma forming a spinulose ridge on either side of the ostium connected with a narrow ventral rim.
strigulae along entire costa ending in a triangle in apex, and sometimes a suffusion or mark at 56: dorsum (two undescribed Papuan species differ with additionally either an outwardly curved dark brown band from middle of costa to dorsum before apex with its inner margin ill-defined and slowly
Description Adults (Figs 576, 577) Wingspan. 7–15 mm; very small to mediumsized. Head (Fig. 578). Ocellus moderate. Upper part of frons with anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis hardly longer than labial palpus. Labial palpus sinuate, porrect or curved-ascending, short (1.3– 1.7× diameter of eye), either with second segment sinuate and distally widened by long loose scales dorsally and ventrally and terminal segment pointing forward, or with second segment evenly curved and slender throughout and terminal segment only weakly angled forward. Antenna to middle of wing, with short cilia. Thorax. Smooth. Posterior surface of metathorax in male with two symmetrical ovate patches of dense modified scaling, opposite two similar patches at abdomen base. Hind tibia in male with a scale pencil from base of inner surface, often concealed in long scaling. Wings (Figs 68, 576, 577). Forewing index 0.43–0.50. Forewing elongate subrectangular to elongate ovate, costa curved in basal half then slightly sinuate or straight, apex from rectangular to strongly obtuse, termen straight to outwardly oblique with protruding tornus; M-stem diverging from anterior margin of cell opposite R1 and ending between M2 and M3, chorda not developed; R2 and R3 variably modified in both sexes with R3 always much closer to R2 than to R4 at base and ending well before costa in a more or less pronounced swelling of vesicular appearance, with base of R3 often very slender, sometimes arising near base of R2 or occasionally R2 and R3 connate or short-stalked and stalk of R4 and R5 deflected dorsally, rarely also R2 swollen distally; R4 and R5 stalked, remaining veins separate beyond discal cell; CuP weakly developed near margin only; anal loop 0.4× length of 1A+2A in M. lepta Horak & Sauter; nearly always with strongly reduced wing pattern of few red-brown to blackish brown markings on uniformly cream-coloured to pale greyish yellow ground consisting of conspicuous
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Figs 576–578. Metaselena, adults and head, males. 576, M. allophlebodes, holotype, Mission Beach, Q [image reversed]. 577, M. lepta, paratype, Sue (Warraber) Is, Q [image reversed]. 578, head, M. sp., Papua New Guinea. Scale = 2 mm.
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AUSTRALIAN OLETHREUTINE GENERA
fading towards base, or with a clear-cut large greybrown semicircle over basal 36: of wing with its margin below and roughly parallel to costa to middle of wing then curving to dorsum before tornus). Hindwing with cubital pecten very weakly developed or absent; anal area in male enlarged, forming a projecting triangular lobe with 3A far distant from often strongly modified, thickened anal margin; Rs and M1 parallel but clearly separate at base, position of M2, M3, CuA1 and CuA2 variable, in some species M2, M3 and CuA1 all quite close to each other, in others (M. lepta) M2 and M3 close or connate but CuA1 distant and halfway between M3 and CuA2, and in M. allophlebodes Horak & Sauter M3 and CuA1 stalked; CuP present towards margin, 1A+2A and 3A well developed; cream to grey. Pregenital abdomen. Sternum 2 with two large ovate patches of dense modified scales (Fig. 580). Hindmargin of S8 in male bilobed. Male genitalia (Figs 579, 583). Vinculum short, a weakly sclerotised band fused with tegumen to form a ring. Tegumen high, subtriangular, a weakly sclerotised band (shape usually distorted in slide preparations). Apex of tegumen usually rounded or with small, cone-shaped hump, rarely (M. allophlebodes) produced into long, slender, naked and distally bifid point (uncus?). Socii two ventrolateral, densely bristled pendulous lobes from just below apex of tegumen continuous with dense long bristles on dorsal apex of tegumen. Gnathos vestigial, two weakly developed lateral bands below socii. Valva very long, strongly sclerotised, abruptly narrowed medially with costa produced into long, slender, curved process usually somewhat widened subapically and with a simple apical thorn; large ventral lobe on basal half of valva with dense long bristles along ventral margin of large basal excavation; very conspicuous triangular, pocketshaped invagination at base of valva on outer surface below long and very slender costal process. Juxta small, caulis a long wide weakly sclerotised band. Aedeagus long and slender, straight to lightly curved, tapering towards apex; vesica with a large bundle of sinuate, lanceolate cornuti. Female genitalia (Figs 581, 582). Ovipositor lobes narrowly to broadly subelliptic. Apophyses usually fairly short but especially anterior pair greatly lengthened in M. allophlebodes with its extensible 8th segment. Ostium in membrane posterior to often deeply concave and strongly sclerotised hindmargin of S7; sterigma two posteriorly widening spinulose ridges on either side of cup-shaped depression leading to ostium, sometimes with one or several large bristles on each
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ridge and usually ventrally connected by a narrow lip to form a U-shaped spinulose rim around entrance to ostium. Ductus bursae narrow, roughly parallel-sided, no colliculum apparent, with a small sclerite just posterior to origin of ductus seminalis at entrance to corpus bursae and sometimes sclerotised near ostium or with a sclerotised band along entire length; ductus seminalis from close to corpus bursae; corpus bursae globose to ovate with one signum dorsally close to entrance of ductus bursae. Signum a rounded, deeply concave, strongly granulose patch.
Distribution Metaselena is known only from Papua New Guinea and Australia, where the genus occurs on the north-eastern coast of Queensland north of Dunk I.
Biology No host record is known for Metaselena. Nearly all material consists of singletons, and the material of M. lepta Horak & Sauter from Sue (or Warraber) I. off Cape York is the only series available. The vegetation on Sue I. is dry monsoon forest on coral sand.
Remarks Metaselena was originally described from a single female as a genus in the Tortricinae (Diakonoff 1939), no doubt due to the lack of a cubital pecten and the single scobinate signum. The unusual genus caught the author’s attention early in her work on Papuan Tortricidae, resulting in a revision of all available material (Horak and Sauter 1981) and the recognition of the many related genera in the Oriental/Australian region. Metaselena is highly derived and characterised by a number of apomorphies. The forewing venation is unique, with R3 modified in both sexes, distally swollen and its often very slender base approximated towards R2 or, rarely, the two veins are even stalked. The hindwing has a strongly reduced cubital pecten and the anal region in the male is greatly enlarged to a projecting triangular lobe usually with a thickened inner margin. The male genitalia are very similar to those of Anathamna, but the long and extremely slender distal portion of the valva in Metaselena with a simple apical thorn represents a more derived condition. The single signum in Metaselena, a concave patch of scobination close to the origin of the ductus bursae, is a further apomorphy for the genus.
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Figs 579–583. Metaselena, genitalia and abdomen. 579–581, M. lepta, paratypes, Sue (Warraber) Is, Q. 579, male genitalia, slide T1719. 580, abdomen base with modified scales on S2, male. 581, female genitalia, slide T1716. 582, 583, M. allophlebodes. 582, female genitalia, paratype, Kuranda, Q, slide BM 20957. 583, male genitalia, holotype, Mission Beach, Q, slide T1724.
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AUSTRALIAN OLETHREUTINE GENERA
Metaselena and Anathamna are sister groups and share a number of conspicuous synapomorphies discussed in the Remarks for Anathamna. The male genitalia of the Sri Lankan Aemulatrix Diakonoff are very similar to those of Metaselena, especially the shape of the valva with a very slender distal process with a simple thorn, but Aemulatrix has R4 and R5 separate, a hindwing with M2 distant from the stalk of M3 and CuA1 and with a small projecting lobe from the base of the anal area, and the socii fused rather like Cyphophanes. Heteroschistis has very similar genitalia to Metaselena, especially the female with a V-shaped, raised, spinulose sterigma and a single signum only, a concave scobinate patch. The wing venation of Heteroschistis is plesiomorphic with all veins separate beyond the cell and with the base of M2 bent close to M3 in the hindwing, but the shape of the valva with a bipronged subapical spine on the dorsal lobe is here treated as an apomorphy for Heteroschistis. Cimeliomorpha Diakonoff and Anoecophysis Diakonoff also have a single, concave, scobinate signum, but the former genus is characterised by conspicuous metallic scales on the forewing, distally upcurved pointed labial palpi, very different male genitalia with short caulis and aedeagus and a hindwing with M2 widely distant from M3. Anoecophysis has an extremely slender valva throughout with the distal portion bipartite and the thorn on a triangular ventral process, and the structure of the ductus bursae is very different from that of Metaselena.
Constituent species Nine species of Metaselena have been described from Papua New Guinea and Australia, with two described and one unnamed species known from Australia. lepta Horak & Sauter, 1981, Aust. J. Zool. 29: 240 (Metaselena). [Qld: Sue (Warraber) I.] allophlebodes Horak & Sauter, 1981, Aust. J. Zool. 29: 264 (Metaselena). [Qld: Mission Beach.]
Oriodryas Turner
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Diagnosis The male of Oriodryas olbophora is unmistakable, a medium-sized olethreutine moth with subtriangular forewings of white ground colour with a sparse pattern in silvery white, grey and blackish, and with rather dark grey hindwings with a greatly enlarged anal area with a modified margin rolled upwardly and containing a large scale pencil from the base of the anal margin. The female is darker, its forewing suffused with brownish, and it looks less unusual but can, like the male, be identified by the characteristic forewing venation with a distally narrowed cubital cell with a sinuate anterior margin and a very oblique accessory cell, and with M2 and M3 both strongly sinuate and widely separated at base but curving towards each other nearer to margin. The labial palpi are only moderately long, evenly upcurved close to the head and sinuate only at the very tip, of even width throughout with long loose scales along entire ventral margin of second segment. The genitalia in both sexes are diagnostic, also the laterally projecting lobe-like folds between T1 and T2 and S2 in the male. The male genitalia are characterised by a stout, well-sclerotised tegumen, a small, lobeshaped uncus with ventral bristles, pendulous socii, a highly derived gnathos with paired, paddleshaped distal lobes with heavy bristles on their outer surface, with a slender valva with a flap-like process associated with the costal process, and with juxta and caulis fused into a broad, flat structure with wide lateral flanges. The female genitalia are characterised by a large sterigma, a flat, mostly smooth, roundish plate with the central opening surrounded by a partially raised spinulose rim and posteriorly by a spinulose hump, and by two hornlike signa. Eucosmogastra Diakonoff and Anthozela Meyrick have similar genitalia in both sexes to Oriodryas. Eucosmogastra can be differentiated in the male by highly derived socii with hand-shaped scale-like structures instead of bristles and a much smaller gnathos with at best a small clump of bristles, and in the female by a sterigma with a ventrally projecting lip or groove rather than a flat plate. Anthozela has large raised socii with pegs in the male and a flimsy anterior expansion of the corpus bursae.
(Figs 69, 584–589)
Description Oriodryas Turner, 1925, Trans. R. Soc. S. Aust. 49: 59. Type species: Oriodryas olbophora Turner, 1925, ibid. 49: 59, by monotypy. [Australia.]
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Adults (584, 585) Wingspan. 14–16 mm (see Remarks); mediumsized.
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Head (Fig. 586). Ocellus moderate. Upper part of frons with anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis longer than labial palpus. Labial palpus moderately long (1.6–1.8× diameter of eye), sinuate only
towards tip otherwise evenly upcurved close to head, of equal width throughout, second segment strongly upcurved and sinuate only at apex, with long, loose scales of similar length along entire ventral margin and a small apical tuft dorsally,
Figs 584–589. Oriodryas olbophora, adults, head, hindwings and genitalia. 584, , Dorrigo Nat. Park, NSW [image reversed]. 585, , The Crater, nr Atherton, Q. 586, head, male. 587, hindwing pencils, male. 588, male genitalia, Mt Tamborine, Q, slide ANIC 12937. 589, female genitalia, Herberton, Q, slide BM 24901. Scale = 2 mm.
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AUSTRALIAN OLETHREUTINE GENERA
terminal segment moderately long, pointing forward. Antenna to middle of wing, anterior surface without scales in male, with short cilia. Thorax. With posterior crest and with a long scale tuft from anteroventral corner of tegulae reaching to below base of hindwing. Legs unmodified. Wings (Figs 69, 584, 585, 587). Forewing index 0.41. Forewing subtriangular, widest just before tornus, with costa straight and curved only near apex, apex rounded-acute, termen oblique; discal cell with sinuate anterior margin and widest at 36: then narrowing towards oblique, weakly developed closing vein, accessory cell triangular with chorda from below R2 to opposite R5; all veins present and separate beyond cell with course of R2 and R3 modified with both veins strongly sinuate and widely separated at base but curving towards each other nearer to margin, bases of R3, R4 and R5 all rather close and M2, M3 and CuA1 equidistant at base; CuP present at margin; anal loop 0.35× length of 1A+2A, 3A present; white with variously developed, indistinct, scattered markings in pale greyish ochreous, silvery white and grey to black, in female markings more brown and entire wing much darker, suffused with brownish; costa with short blackish strigulae throughout, with a short transverse black dash or small triangle at 56: costa, an often incomplete oblique grey to blackish narrow band from below 6M costa to termen above tornus, dorsum with three small triangular black marks in middle third and a variably developed grey suffusion along basal half extending towards centre of wing at about 569 and as a long triangle just before middle; with a series of short, partly parallel, curved black lines around apex and along termen; with indistinct silvery white transverse lines especially in distal half. Hindwing with welldeveloped cubital pecten and axillary tuft, in male with anal area triangularly expanded with a thickened margin rolled upwardly and containing a large scale pencil from base of anal margin (Fig. 587); with Rs and M1 closely approximated and parallel at base, base of M2 curved towards M3 at base, M3 from angle of cell and CuA1 very close to but not connate with M3, CuA2 from just beyond middle of cell; CuP present at margin but not tubular; 1A+2A and 3A well developed, 3A widely distant from anal margin; grey, brownish grey in female. Pregenital abdomen. Ventral apodemes long, slender and curved; in lateral membrane of first two segments a longitudinal, lobe-like deep fold, naked on outer surface but with dense sockets on
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lateral inner surfaces, its apex projecting. In male hindmargin of S8 bilobed, with medial V-shaped emargination. Male genitalia (Fig. 588). Vinculum short, a moderately wide band. Tegumen robust, broad and well-sclerotised. Uncus a small, elongate, distally rounded lobe with stout bristles on ventral surface. Socii two membranous, ovate, pendulous, sparsely hairy lobes, obscured by gnathos. Gnathos prominent and well-sclerotised, arms short and very broad at base and each with an upwardly angled, large, subtriangular distal lobe with strong, rather short and backward oriented bristles on lateral, outer surface and their smooth inner surfaces facing each other. Valva well-sclerotised, slender with nearly straight costa, rounded apex with bristles on inner surface and a sinuate ventral margin set with short strong bristles especially in the two convex regions just before apex and below middle, with sacculus strongly projecting as a rectangle with a flattened, blade-shaped angle; costal process large and with a flap-like process at base below costa; inner surface with a longitudinal rib running roughly parallel to costa towards apex, outer surface with a conspicuous, angled line of weakness running from costa near costal process to base of sacculus. Juxta and caulis entirely fused into a broad flat structure with a transverse band as base (juxta) and an ovate distal portion with a broad, lamellar lateral margin (caulis), with a lamellar, hourglass-shaped dorsal sclerotisation to anal tube. Aedeagus small and very short, widest at base; vesica with a very large bundle of lanceolate cornuti. Female genitalia (Fig. 589). Ovipositor lobes broadly elliptic. Hindmargin of S7 concave. Sterigma a very large, roundish, sclerotised plate with central round opening surrounded by a partially raised spinulose rim and posteriorly by a large spinulose hump, ventral and lateral portions of sterigma smooth. With long, narrow colliculum below ostium, rest of ductus bursae membranous, joining teardrop-shaped corpus bursae at apex of slightly produced posterior end that also gives rise to ductus seminalis. Corpus bursae finely dimpled and with two slender, horn-shaped signa of roughly equal size.
Distribution Oriodryas is known from Australia only, where it occurs from the Dorrigo NP behind Coffs Harbour in New South Wales to the Atherton Tableland behind Cairns in northern Queensland.
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Biology No host is known for Oriodryas. All the material has been collected in rainforest, in lowland and upland subtropical rainforest in northern New South Wales and southern Queensland and in upland rainforest in northern Queensland.
Periphoeba Bradley (Figs 70, 590–597) Periphoeba Bradley, 1957, Nat. Hist. Rennell Isl. 2: 96. Type species: Periphoeba adluminana Bradley, 1957, ibid. 2: 96, pl. 1, fig. 8; pl. 5, figs 52–54, by original designation. [Rennell I.]
Remarks Turner (1925) described Oriodryas olbophora based on a single male from the Lamington NP near Brisbane. Recently collected material from northern Queensland confirmed that the single, much darker female from Huberton (misspelling for Herberton) Hills in the BMNH, collected by F. P. Dodd, is indeed O. olbophora. Turner (1925) gave a Meyrick-type ‘expanse’ of 19 mm for the holotype, which is the measurement from centre of thorax to apex times two, hence the size divergence. Oriodryas is a highly derived genus with numerous apomorphies including the shape of the labial palpi, the distally narrowed discal cell and the unusual course of R2 and R3 in the forewing, the modified anal area of the hindwing in the male containing a scale pencil (shared with Thysanocrepis Diakonoff), the lateral lobe-like folds at the base of the male abdomen, the ventrally bristled uncus, the complex gnathos with large bristled distal processes, and the large plate-shaped sterigma. Both Eucosmogastra and Anthozela are closely related to Oriodryas, with similar, but less derived venation in the first two and with several synapomorphies in the genitalia of both sexes. The three genera share a very similar male valva, but are differentiated by socii and gnathos structures. The female genitalia also are similar in all three genera with large, well-sclerotised sterigma separated from S7 and a similarly shaped corpus bursae with a produced posterior end from which originates the ductus seminalis, and with two horn-shaped signa. Anthozela also shares the fused and flattened juxta/ caulis complex with Oriodryas, and could well be the sister group.
Constituent species Only one named Australian species is known: olbophora Turner, 1925, Trans. R. Soc. S. Aust 49: 59 (Oriodryas). [Qld: [Lamington] NP.]
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Diagnosis Periphoeba species are small to medium-sized, greyish ochreous olethreutines with a strongly projecting reddish brown forewing apex, a sinuate termen and an unmistakable wing pattern with pronounced costal strigulae giving rise to metallic leaden lines in distal half and with remainder of wing densely transversely strigulated with dark brown wavy lines. The labial palpi are very long and sinuate with the second segment distally greatly widened and the third segment long and slender, and of very characteristic colouring with a yellowish ochreous band distally across the second segment followed by a grey dorsoanterior margin with a purplish sheen. Males of Periphoeba are characterised by a truncate hindwing anal area with the anal margin narrowly folded upwards over a large scale pencil from the base of the wing. Periphoeba and Balbidomaga Diakonoff have very similar wing venation without M-stem and chorda (very short chorda present in Balbidomaga), R3 adjacent and parallel to R4 nearly to margin, R4 and R5 connate in Periphoeba and connate or shortstalked in Balbidomaga, and bases of M3 and CuA1 closely approximated and parallel for some distance at base in the forewing, and the hindwing with M2 bent very close to M3 at base, M3 and CuA1 stalked, CuA2 from just beyond middle of cell, 1A+2A rather close to anal margin, and 3A next to anal margin in Periphoeba and absent in Balbidomaga. The males of the two genera are easily separated by their secondary sexual characters, the fake costal fold in Balbidomaga and the large scale pencil along the anal margin in Periphoeba, while separation of the females has to be based on wing pattern or genitalia. The genitalia in both sexes of Periphoeba are diagnostic. The male is characterised by the strongly sclerotised, dorsoventrally flattened, bipronged uncus, a gnathos reduced to lightly sclerotised lateral arms, a ventrally deeply emarginate valva with a tuft of bristles from the tip of the sacculus and with several long spines from the outer surface of cucullus, a very narrow caulis and an aedeagus wide at base and with a lateral
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AUSTRALIAN OLETHREUTINE GENERA
thumb-like process near middle and unusual, somewhat boomerang-shaped cornuti. Males of Periphoeba share with those of Balbidomaga long, band-like scales laterally on the basal half of the tegumen, lightly sclerotised socii lobes (lost in some Balbidomaga species), an overall similar valva with a conspicuous single row of very large bristles on the sacculus (absent in some Balbidomaga species). The female genitalia of Periphoeba are characterised by the unique position of the ostium in a slightly sclerotised, spinulose, ledge-shaped band fused with the hindmargin of S7 at a 90° angle, by a ductus bursae widened and lightly sclerotised in its posterior 36:, and by usually two signa formed by roundish, depressed areas of scobination.
Description Adults (590, 591) Wingspan. 7.0–13.5 mm; small to mediumsized. Head (Fig. 594). Ocellus moderate. Upper part of frons with a very long, flattened tuft of anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis shorter than labial palpus. Labial palpus porrect, sinuate, very long (2.1–2.6× diameter of eye) and relatively slender, second segment strongly sinuate and distally greatly widened by long scales along distal half of dorsal margin with their dark tips forming purplish tinged, nearly straight dorso-anterior edge of the neatly subtriangular segment, with a parallel yellowish ochreous band below, third segment very long and slender, downward inclined in
Figs 590, 591. Scale = 2 mm.
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continuation of dorsoanterior edge of second segment, grey with purplish sheen above, ochreous below. Antenna to beyond middle of wing, entirely scaled, cilia not visible. Thorax. Smooth, with long scale pencil from anteroventral corner of tegula. Legs not modified. Wings (Figs 70, 590, 591). Forewing index 0.34– 0.36 (to 0.41 in non-Australian species). Forewing elongate subrectangular, slightly widening distally, broadest before tornus, with costa lightly curved throughout but more so in basal half, apex strongly projecting, termen sinuate; chorda and M-stem absent; R2 from about 6E cell, R3 from close to angle of cell, very close to and parallel with R4 nearly to wing margin, R4 and R5 connate from angle of cell, M2 and M3 close together or connate at base, M2 straight but M3 bent at base, bases of M3 and CuA1 close and parallel for some distance, CuA2 from just below 6M cell; CuP present at margin, close to 1A+2A; anal loop short, 0.19× length of 1A+2A, anterior arm weak; forewing pattern very similar in all known species with ochreous to reddish ochreous ground colour transversely strigulated with numerous, narrow, wavy, dark brown lines throughout wing, entire costa with oblique, dark brown strigulae mixed with white ones in distal half giving rise to several oblique, thin, metallic leaden lines, one particularly prominent at 56O costa and one at 6M recurving to costa just before apex, with distal half of costa and apex reddish brown between strigulae; ocelloid patch variably indicated by a series of blackish spots between veins bordered proximally and distally with indistinct, transverse, wavy, leaden bands;
Periphoeba, males. 590, P. trepida, Eungella, Q [image reversed]. 591, P. sp., Sulawesi.
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margin of termen with a line of black scales interrupted by a few white scales in concavity below apex. Hindwing with strongly sinuate costa, with moderately developed cubital pecten and axillary tuft, in male with anal lobe truncated by a modified anal margin narrowly folded upwardly and enclosing a large hair pencil from base of wing; base of M1 weak and closely approximated to Rs, M2 bent very close to M3 at base, M3 and CuA1 stalked, CuA2 from below 36: cell; CuP present near margin; 1A+2A strongly curved, 3A present but parallel and very close to margin, the two veins converging towards margin; light grey with coppery gloss to dark grey, semitranslucent with small, sparse scaling towards base. Pregenital abdomen. Unmodified. Hindmargin of S8 in male weakly concave. Male genitalia (Figs 592, 593, 597). Vinculum short. Tegumen ovate, little sclerotised except for sclerotised apex, laterally at base with a dense bundle of very long scales nearly twice the length of the tegumen, dorsally below uncus with two patches of large bristles. Uncus strongly sclerotised, a pair of long points curved ventrally and forming a bipronged fork, with sclerotised lateral ‘shoulders’ giving rise to moderately large, pendant, lobeshaped socii distally lightly sclerotised and sparsely bristled except along ventral margin. Gnathos weakly sclerotised lateral bands variously widened medially. Valva long, moderately sclerotised, with nearly straight costa but with a deep semicircular ventral excavation just beyond middle separating rounded-triangular cucullus by a narrow ‘neck’ from base of valva; cucullus distally densely bristled on inner surface with short strong spines towards ventrodistal margin and with 3–7 strong, long spines on outer surface; basal part of valva projecting rectangularly on ventral margin with a conspicuous tuft of bristles from angle, a row of long bristles on narrow band between basal excavation and ventral margin, and a long, slender costal process. Juxta and caulis entirely fused, caulis very slender. Aedeagus broad and roughly rectangular at base in ventral view, with a lateral thumb-like process just below middle and much narrower beyond; vesica with a large bundle of long, sinuate, somewhat boomerang-shaped cornuti. Female genitalia (Figs 595, 596). Ovipositor lobes subelliptical, rather narrow, with jagged lateral margin. Ostium in a ledge-shaped, dorsoventrally oriented posterior extension of S7, a spinulose band fused at a 90° angle with hindmargin of S7. Sterigma not conspicuous, a simple round opening in lightly sclerotised,
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spinulose band. Ductus bursae about as long as corpus bursae, lightly sclerotised and wider in posterior 36:, colliculum not apparent, with ductus seminalis originating from close to corpus bursae. Corpus bursae ovate or pear-shaped with a small rounded posterior prominence next to origin of ductus bursae; with usually two signa, rarely one only, depressed areas of conspicuous, granulate scobination, one next to origin of ductus bursae, the second slightly smaller one on the opposite side about 56: from origin of ductus bursae.
Distribution Periphoeba is known from Sri Lanka, Sulawesi, the Solomon Is, New Guinea and Australia, where it occurs in Queensland from just north of Mackay in the Eungella NP to the tip of Cape York.
Biology No host is known for Periphoeba. The Australian localities are in rainforest, rainforest margin or monsoon forest from sea level to 750 m.
Remarks Periphoeba was described from a single female (Bradley 1957) from the Solomon Is, with illustrations of the adult, head, wing venation and female genitalia. Periphoeba palmodes (Meyrick) from Sri Lanka, with adult and female genitalia figured by Clarke (1958), is a senior synonym of the type species of Periphoeba according to Brown (2005). The monophyly of Periphoeba is supported by several apomorphies. Whilst a scale pencil along the anal margin of the hindwing is present in males of other Enarmoniini such as Ancylis and Oriodryas, the detailed structure of the anal region in Periphoeba with the close approximation of 3A to the anal margin is quite unique. Apomorphies in the male genitalia include the strongly sclerotised, dorsoventrally flattened, bipronged uncus, a gnathos reduced to lightly sclerotised lateral arms, a tuft of bristles from the tip of the sacculus, long spines on the outer surface of the cucullus, the very narrow caulis, the shape of the aedeagus with a lateral thumb-like process near its middle and the unusual, somewhat boomerangshaped cornuti. Apomorphies of the female are the unique position of the ostium in a slightly sclerotised, spinulose, ledge-shaped band fused with the hindmargin of S7 at a 90° angle and a
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AUSTRALIAN OLETHREUTINE GENERA
ductus bursae widened and lightly sclerotised in its posterior 36:. Periphoeba shares with its sister genus Balbidomaga an unusual and strongly derived wing venation as well as similar forewing shape, a truncate
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anal area in the hindwing of the male and overall similar male genitalia with modified scales laterally on the tegumen, a well-developed uncus, similar socii and valva of the same general shape often with a conspicuous row of large bristles on the sacculus.
Figs 592–597. Periphoeba, genitalia and head. 592, 593, P. trepida, male genitalia and aedeagus, Eungella, Q, slide ANIC 1290. 594, head, P. trepida, male. 595, P. trepida, female genitalia, nr Hope Vale Mission, Q, slide T2328. 596, P. sp., Woodlark Is, Papua New Guinea, female genitalia, slide T2460. 597, P. sp., Sulawesi, male genitalia, slide T2459.
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Both genera have ovipositor lobes with jagged lateral margins and patches of granulation as signa. Tokuana Kawabe and Neoanathamna Kawabe both have similar-looking male genitalia, probably due to symplesiomorphies as the female genitalia of both genera are quite different from those of Periphoeba. The type of P. trepida (Meyrick), a single specimen from Cairns, has not been found, and the species was listed among the unplaced Eucosmini in Horak et al. (1996). However, there can be no doubt that Meyrick’s description refers to a Periphoeba species. Periphoeba species look extremely alike superficially and show only small differences in their genitalia, with the Australian P. trepida the only named species represented by males. Undescribed material from New Guinea and Sulawesi, the latter including males, is very close superficially and in genitalia structure except for one single, strongly abraded female from Sulawesi with dark grey hindwings, a radically modified, triangular S7 and only one signum. However, there are slight variations in wing shape, the shape of the sclerotised portion of the ductus bursae and subtle differences in the male genitalia, demanding a study beyond the scope of the present review to confirm that P. palmodes and P. adluminana are indeed synonyms.
Constituent species Three species of Periphoeba have been named, one from Sri Lanka, one from the Solomon Is and one from Australia, with the first two treated as synonyms (Brown 2005). Unidentified material is known from Sulawesi and New Guinea, containing at least one very distinct species from Sulawesi where two species have been collected at the same locality. trepida (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 249 (Eucosma). [Qld: Cairns.]
and brown with a long, falcate apex with a redbrown spot followed by a slender, Y-shaped blackish mark on white along the deep, rectangular indentation on the termen behind the apex. In the male the basal half of the hindwing is translucent with distantly spaced, small, black scales, and the anal lobe is very small. The labial palpi are only moderately long, porrect and sinuate with the second segment distally strongly widened by long, loose scales and the terminal segment long and slender, white on inner surface and ventrodistally, otherwise brownish. The veins R4 and R5 are longstalked and M3 and CuA1 separate at base in the forewing and in the hindwing M2 is quite distant from the stalk of M3 and CuA1. The genitalia of both sexes are diagnostic for the genus. The male genitalia are characterised by a broadly rounded tegumen with or without a nipple-shaped uncus remnant and with long, hairy, subcrescentic socii dorsally and laterally attached to the tegumen and usually with a projecting, short, band-like process from the middle of the median margin, and by a long and medially narrow valva with a strongly projecting sacculus that may be produced into a finger-shaped process, followed by a very deep, roundish, ventral emargination with the cucullus gradually widening to a broad apex consisting of a rounded dorsal lobe and a triangular ventral projection ending in a small spine. The female genitalia are characterised by a conspicuous sterigma, a prominent, lightly sclerotised, bandshaped ventral lip surrounded by spinulose membrane, an indistinct, ring-like sclerotisation anterior to the ostium in the otherwise membranous ductus bursae, a single, distally flattened, curved, dagger-shaped signum with a serrate inner edge, surrounded by scobination, and a scobinate, domed area next to the origin of the ductus bursae.
Description
Pseudancylis, gen. nov. (Figs 71, 598–601) Type species: Ancylis acrogypsa Turner, 1916, Trans. R. Soc. S. Aust. 40: 524, hereby designated. [Australia.]
Diagnosis Pseudancylis species are narrow-winged, small olethreutines contrastingly patterned in whitish
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Adults (Fig. 598) Wingspan. 9–12 mm; small. Head (Fig. 599). Ocellus moderate. Vertex with long scales, medially converging and anteriorly protruding between antennae, upper part of frons with very long, anteriorly protruding and downcurved scales, remainder of frons with minute, appressed scales. Maxillary palpi minute, scaled, often not visible on undissected head. Labial palpus porrect, moderately long (1.5–1.7× diameter of eye), strongly sinuate, second segment distally strongly widened by long, loose scales along entire ventral margin and dorsally towards apex, terminal
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segment long and slender, concealed among scales from second segment; white on inner surface and ventrodistally, remainder dark brownish. Antenna to beyond middle of wing, entirely scaled, no cilia visible. Thorax. Smooth, with long scale pencil from anteroventral angle of tegula. Legs unmodified. Wings (Figs 71, 598). Forewing index 0.27–0.35 (P. acrogypsa 0.34–0.35). Forewing long and slender, widest before middle with costa strongly curved at base and faintly concave beyond middle, apex long and falcate, termen strongly sinuate but not oblique, with indentation behind apex deep and rectangular; M-stem and chorda absent; R4 and R5 stalked to beyond middle, M3 and CuA1 separate at base; CuP present at margin, 1A+2A with 1A at base much weaker; contrastingly marked with whitish and red-brown to blackish brown with silvery leaden pattern elements; with alternating whitish and blackish costal strigulae and with apex
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red-brown, with a large whitish blotch at 56M costa or along entire basal third of costa, with an irregular brownish triangle based on costa from before middle to before apex reaching to middle of wing or to dorsum and sprinkled with leaden scales, with a whitish band along termen containing an inwardly oblique blackish line on costa, a Y-shaped black mark along angle of termen behind apex and several longitudinal, parallel, leaden marks in tornal region, and with remainder of dorsal half of wing brownish with variable transverse whitish bands, especially in basal half; fringe on termen with a cinnamon hue. Hindwing narrow, anal region small, cubital pecten and axillary tuft weakly developed; Rs and M1 parallel and closely approximated at base, base of M2 curved towards stalk of M3 and CuA1, but not very close; CuP present at margin, 1A+2A and 3A both present; pale brownish grey, especially in male transparent in
Figs 598–601. Pseudancylis acrogypsa, adult, head and genitalia. 598, adult, , Mt Edith nr Atherton, Q. 599, head, male. 600, male genitalia, Kuranda, Q, slide ANIC 12957. 601, female genitalia, nr Ravenshoe, Q, slide ANIC 12958. Scale = 2 mm.
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basal half with small, black (modified?), distantly spaced scales above and below. Pregenital abdomen. Ventral apodemes long, slender. In male hindmargin of S8 weakly concave, T8 with a long, narrow, weakly sclerotised triangle. Male genitalia (Fig. 600). Vinculum a short, wide, weakly sclerotised band fused with tegumen. Tegumen weakly sclerotised, broadly rounded, apex rounded or pointed, pedunculi without obvious muscle attachments. Uncus entirely reduced or possibly present as a nipple-shaped remnant (P. rostrifera (Meyrick)). Socii long, hairy, somewhat crescentic lobes, dorsally and laterally attached to tegumen and at least sometimes with a projecting, short, band-like process from middle of medial margin (P. acrogypsa, P. rostrifera). No trace of gnathos. Valva well sclerotised, long and very narrow in central section, with a strongly sinuate costa and ventral margin with a deep, roundish emargination beyond pointed, strongly projecting sacculus angle that may be produced into a fingershaped process, base of cucullus a narrow neck gradually widening to broad apical third, bristled on inner surface and consisting of a rounded dorsal lobe and a triangular ventral projection ending in a small spine; outer surface of valva with a shallow pocket at base; costal process straight. Caulis rather wide in lateral view. Aedeagus moderately long, often wider at base; vesica with a bundle of lanceolate cornuti. Female genitalia (Fig. 601). Ovipositor lobes subreniform, posteriorly with several very prominent tubercules in P. acrogypsa. Ostium in membrane well posterior of weakly concave hindmargin of S7. Sterigma a prominent, lightly sclerotised, smooth, band-shaped ventral lip, short and wide in P. acrogypsa, long, semicircular and narrow in P. rostrifera sensu Diakonoff from Sumba (Diakonoff 1984a), surrounded by spinulose membrane. Ductus bursae membranous except for an indistinct, ring-like sclerotisation anterior to ostium, without colliculum, ductus seminalis originating close to corpus bursae; corpus bursae teardrop-shaped, with a single, distally flattened, curved, dagger-shaped signum with a serrate inner edge and a hollow base, surrounded by scobination, and with a scobinate, domed area next to origin of ductus bursae.
Distribution Sri Lanka, Java, Borneo, Sumba and Australia where it occurs on the Atherton Tableland and the hills south of Cooktown in northern Queensland.
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Biology No host records are known for Pseudancylis, but all Australian localities are in upland rainforest.
Remarks Though Pseudancylis superficially looks like Ancylis the two are only distantly related, and Pseudancylis is characterised by several apomorphies in wing structure and the genitalia of both sexes. The veins R4 and R5 in the forewing are stalked, in the hindwing the base of M2 is quite distant from the stalk of M3 and CuA1 and in the male the scaling in the basal half is modified with the scales very small, much darker and distantly spaced. In the genitalia, the long, hairy socii with a medial, band-like process are an apomorphy for Pseudancylis, as are the shape of the valva with the very pronounced sacculus tip, the band-shaped sterigma forming a ventral lip and the single, dagger-shaped signum. No sister group can be identified for Pseudancylis, but possible synapomorphies link the genus with Eucosmogastra Diakonoff and Thysanocrepis Diakonoff. Pseudancylis shares with these two genera a ventral, lip-shaped sterigma and a small sclerotisation in the otherwise membranous ductus bursae anterior to the ostium, and with Thysanocrepis also a projecting sacculus tip. Eucosmogastra has two horn-shaped signa in contrast to the single, more flattened, daggershaped signum of Pseudancylis. Diakonoff (1984a) treated material from Sumba as conspecific with P. rostrifera (Meyrick) from Sri Lanka, but the male genitalia of the very narrowwinged P. rostrifera (Clarke 1958: pl. 146, figs 2, 2A) have a much shorter sacculus tip than that figured for P. rostrifera sensu Diakonoff from Sumba (Diakonoff 1984a: fig. 28). The Australian P. acrogypsa (Turner) has similar wing pattern and genitalia in both sexes as the other two species, but its forewings are wider than those of P. rostrifera and with differences in pattern detail, the male genitalia are unique through the absence of any uncus remnant but have a similarly drawn out sacculus tip as P. rostrifera sensu Diakonoff, and the sterigma in P. acrogypsa is shorter and wider than in P. rostrifera sensu Diakonoff and the signum more strongly curved. The three taxa are here considered to be three distinct species, and the identity of further material from Java and Borneo referred to by Diakonoff (1984a) will have to be established. The following two Oriental species are here referred to Pseudancylis:
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rostrifera (Meyrick, 1912b), J. Bombay. Nat. Hist. Soc. 21: 862 (Ancylis). [Sri Lanka.] Comb. nov. percnobathra (Meyrick, 1933), Exotic. Microlepid. 4: 417 (Ancylis). [Sumatra.] Comb. nov.
Constituent species Two named non-Australian species are known. Genitalia differences suggest that P. rostrifera sensu Diakonoff from Sumba (Diakonoff 1984a) is a third species, and Diakonoff (1984a) referred to further, unstudied material from Java and Borneo. The one named Australian species of Pseudancylis, listed as a synonym of Eucosma symploca (Turner) by Brown (2005) among the unplaced Eucosmini, is designated as the type species of the genus: acrogypsa (Turner, 1916), Trans. R. Soc. S. Aust. 40: 524 (Ancylis). [Qld: Kuranda.] Comb. nov.
Pternidora Meyrick (Figs 72, 602–606) Pternidora Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 225 (key), 285. Type species: Pternidora phloeotis Meyrick, 1911b, ibid. 36: 286, by monotypy. [Australia.]
Diagnosis Pternidora species are small to very small moths with subtriangular forewings indistinctly transversely banded, with a greyish ochreous basal half and a much darker, blackish distal portion usually separated along an oblique line from before apex to middle of dorsum, and with two scale tufts triangularly projecting beyond margin in central third of dorsum. The labial palpi are only moderately long, sinuate and rather stout with a short terminal segment. In the forewing, Pternidora has M-stem and chorda present and all veins separate with M3 distant from CuA1 at base, in the hindwing Rs and M1 are parallel but not adjacent at base, M2 is widely distant from M3 at base and M3 and CuA1 are connate from angle. The genitalia in both sexes are diagnostic. The male is characterised by a tegumen of onion-shaped outline with dense, large scale sockets laterally in basal half, a dorsally bristled narrowed apex and gnathos arms distally continued as sinuate slender bands along anal tube, and by a highly derived, membranous and flat valva of roughly elongate-ovate outline with ventral
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margin projecting in a small elbow set with short strong bristles, and with a band of conspicuous scale sockets on outer surface along ventral margin. The female genitalia are characterised by a cup-shaped sterigma fused with S7 in a deep emargination, by a ductus bursae with a colliculum of two small lateral crescentic sclerites in posterior half, followed by an oblique ring of sclerotisation and with a large Lshaped sclerite and several thorn-shaped structures arranged in a spiral in anterior third, and by two slender, horn-shaped signa.
Description Adults (Figs 602, 603) Wingspan. 7–10 mm; small to very small. Head (Fig. 604). Ocellus moderate. Vertex with shortish, appressed scales, upper part of frons with anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis barely longer than labial palpus. Labial palpus porrect, sinuate, stout and only moderately long (1.6–1.7× diameter of eye), second segment sinuate and distally strongly widened by moderately long scales along ventral and dorsal margins; terminal segment short and rather stout. Antenna to middle of wing, stout in male, anterior surface without scales, with minute cilia. Thorax. Smooth. Legs not modified. Wings (Figs 72, 602, 603). Forewing index 0.42–0.52. Forewing subtriangular, widest just before tornus, with costa nearly straight and curved only just before apex, termen oblique, dorsum in middle with two conspicuous, triangular tufts of long scales projecting beyond margin; M-stem well developed, nearly straight, to between M2 and M3, chorda from between R1 and R2 to between R4 and R5; all veins present and separate beyond discal cell, M3 distant from CuA1 at base; CuP well developed near margin; anal loop 0.39× length of 1A+2A; with a greyish ochreous basal half and a much darker, blackish distal portion, with a grapholitine-type wing pattern with numerous indistinct, parallel, transverse lines slightly angled below costa in grey, red-brown or blackish brown on very pale grey ground colour, closer and darker towards wingtip and often wing abruptly very dark beyond a line from before apex to middle of dorsum, with a conspicuous transverse leaden band from end of dorsum reaching 36: to costa and a second shorter one at 36: of dorsum, and with an interrupted blackish row of scales around apex and along termen. Hindwing with moderately developed cubital pecten and axillary tuft, with tuft
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Figs 602–606. Pternidora, adults, head and genitalia. 602, P. ? phloeotis, , Toowoomba, Q [image reversed]. 603. P. sp., , Mt Spec, Q [image reversed]. 604. head, P. sp., male. 605. P. sp., male genitalia, Mt Bartle Frere, Q, slide T2313. 606. P. sp., female genitalia, Cooktown, Q, slide ANIC 12928. Scale = 2 mm.
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of narrow, modified scales above on base of costa; with Rs and M1 parallel but not adjacent at base, M2 widely distant at base from M3 and the two veins roughly parallel, M3 and CuA1 connate from angle of cell; CuP present near margin, 1A+2A and 3A well developed; dark grey-brown with a reddish brown sheen, darker at apex. Pregenital abdomen. Ventral apodemes short and hollow apparently to tip, anterolateral processes absent. Hindmargin of S8 in male with a medial, V-shaped emargination; 8th segment dorsally with an inverted T-shaped sclerotisation. Male genitalia (Fig. 605). Vinculum very short. Tegumen of teardrop-shaped outline, broad near base and tapering towards tip, a weakly sclerotised band moderately wide near base and on its lateral surface in basal half with large, conspicuous scale sockets; tip of tegumen narrow, with a paddleshaped or truncate apex with few to many hair-like dorsal bristles. Socii absent. Gnathos two weak, curved lateral arms medially connected and both continuing as two narrow sinuate bands extending posterodorsally along anal tube that does not exceed tip of tegumen in length. Valva very membranous, short, wide and flat, with shape somewhat distorted in slide preparation but essentially elongate-ovate with base of costa strongly curved, with a rounded-triangular apex and the ventral margin projecting in a small elbow set with dense, short, strong bristles at 36: that extend as thinner bristles along ventral margin to apex; with a patch of long hair-like bristles in centre of valva on inner surface and a band of conspicuous scale sockets on outer surface between ventral margin and a straight, rib-shaped sclerite on outer surface from base of valva towards ventral margin before apex; base of valva on outer surface with a rounded pocket-shaped invagination; costal process very long, narrow and straight. Juxta short and wide, caulis long and anellus inconspicuous. Aedeagus long, sinuate and slender with a bulbous base; vesica with 2–3 very large, nail-shaped cornuti and numerous much smaller spine-shaped ones. Female genitalia (Figs 606). Ovipositor lobes elongate ovate. Ostium in deep U- or V-shaped emargination in S7. Sterigma a ventrally sclerotised, smooth, shallow to moderately deep cup fused with S7. Ductus bursae mostly membranous, tapering from corpus bursae to ostium, with a colliculum of two small, crescentic lateral sclerites about 56: from ostium followed by an oblique, irregularly sclerotised ring-shaped area near middle of ductus bursae, just posterior to origin of ductus seminalis;
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anterior third of ductus bursae greatly widened, containing a large irregularly curved L-shaped sclerite, with a ventral bulge set with several large, complex thorn-shaped structures arranged in a spiral; corpus bursae with a dimpled structure throughout and with two slender, horn-shaped signa of equal size.
Distribution Pternidora is so far known from Papua New Guinea and Australia only, from single to at most three specimens of several species ranging from the Border Ranges NP in New South Wales and from Brisbane to the Claudie R. just south of Cape York in Queensland and from Darwin to the East Alligator R. in the Northern Territory.
Biology No host is known for Pternidora. The localities vary from wet rainforest to monsoon forest, and two specimens have been taken in gardens, one in Toowoomba, Queensland, and one in Bulolo, Papua New Guinea.
Remarks Meyrick (1911b) described Pternidora phloeotis based on two specimens caught by T. Bacheler in Brisbane, a male in the BMNH without abdomen and a second male in the ANIC with an abdomen. A perfect male from Toowoomba near Brisbane agrees superficially in every respect with the type material and is figured here. The monophyly of Pternidora is supported by several apomorphies in the genitalia of both sexes, but also by the wing pattern and the two projecting scale tufts on the dorsum of the forewing. Apomorphies of the male genitalia include the shape of the ventrally bulbous and dorsally narrowed tegumen with modified scale sockets laterally in the basal half and the membranous, wide and flat valva with the costa strongly curved at base, a triangular apex, an elbow-shaped projection on the ventral margin set with short, dense bristles and a band of modified scale sockets along the ventral margin on the outer surface. The cupshaped sterigma fused with S7 in a deep emargination, the ductus bursae with a posterior constriction, an oblique ring of sclerotisation and a large L-shaped sclerite and several thorn-shaped structures arranged in a spiral are apomorphies of the female genitalia.
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No obvious sister taxon of Pternidora can be identified. The rather membranous valva with a longitudinal rib along its outer surface is shared by Helictophanes Meyrick, but female genitalia and wing venation are quite different in the two genera. A hindwing venation with M2 and M3 parallel and widely separate at base is shared by Anthozela Meyrick and Irianassa Diakonoff and nearly all Loboschiza Diakonoff and Cyphophanes Meyrick. Thysanocrepis Diakonoff shares the long scales laterally on the tegumen, and Cyphophanes and Anathamna Meyrick an irregular sclerite in the bursa neck. Only 14 specimens of Pternidora are available representing some six species. Two species are present as pairs, but no female of P. phloeotis has been collected so far. Consequently, female genitalia and wing venation not from the type species of the genus have been figured.
Constituent species One named and several unnamed species are known from Australia, and a single specimen of probably yet another unnamed species is known from Papua New Guinea. phloeotis Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 286 (Pternidora). [Qld: Brisbane.]
Tetramoera Diakonoff (Figs 73, 607–615) Tetramoera Diakonoff, [1968], US Nat. Mus. Bull. 257 (1967): 39 (key), 68. Type species: Grapholitha schistaceana Snellen, 1890, Ber. VersStn. ZuckRohr. W. Java (Meded. Proefstn SuikRiet W. Java “Kagok”) 1: 96, pl. 2, figs 5–7, by original designation. [Indonesia.]
Diagnosis Tetramoera species are superficially quite unmistakable, small to medium-sized brownish moths with elongate-subrectangular forewings with a blunt apex, a strongly sinuate termen and an intricate wing pattern of mostly longitudinal, fine, parallel, cream, grey and brown lines interrupted beyond middle of wing by a transverse series of irregular dark marks, and with conspicuous costal strigulae and a prominent leaden line running from beyond middle of costa to below apex then curved
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along termen to tornus. The labial palpi are long, porrect and sinuate, with the second segment distally strongly widened and with a transverse ochreous streak surrounded by dark scaling. Species of Bactra Stephens and Pammenopsis Kuznetzov also have longitudinally striate forewings, but the forewing shape is very different in both Bactra and Pammenopsis, with a pointed apex and a strongly oblique termen, and in Pammenopsis the males have a conspicuous fringe of black hair scales along the anal margin of the hindwing and the ovipositor in the female is modified, smooth and strongly sclerotised. Tetramoera has all veins separate in the forewing and M3 and CuA1 variable from separate to stalked in the hindwing. The male genitalia are highly characteristic with a uniquely modified, deeply bilobed, flat and densely hairy valva with the dorsal lobe without a thorn but with a flap-shaped process from its ventral margin. The female genitalia are characterised by a single signum varying from a long crease with a small blade along central ridge to a flat, sclerotised ribbon or an ovate concave plate with a small thorn on central ridge.
Description Adults (Figs 607, 608) Wingspan. 9–14 mm (Australian species); small to medium-sized. Head (Fig. 609). Ocellus moderate. Upper part of frons with long, anteriorly projecting scales, remainder of frons with minute, appressed scales. Maxillary palpus not visible on undissected head, in T. gracilistria (Turner) minute, naked, apparently 2segmented remnant. Proboscis longer than labial palpus. Labial palpus porrect, sinuate, long (1.9–2.3× diameter of eye), second segment sinuate and distally strongly widened, often to triangular shape, by very long, loose scales, especially ventrally, often concealing terminal segment; terminal segment short to moderately long, slender, with short scales, downwardly inclined; second segment with a horizontal ochreous streak in distal half, surrounded by dark scaling. Antenna to middle of wing, stout in male, entirely scaled, with minute cilia. Thorax. Smooth. Legs not modified. Wings (Figs 73, 607, 608). Forewing index 0.35–0.42. Forewing elongate subrectangular except for narrowed base, with costa lightly curved throughout or distally straight, termen sinuate with apex and dorsal part of termen equally protruding; M-stem present (not a tubular vein in T. gracilistria) and running to M3, chorda from below R2 to
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beyond R5; all veins separate beyond cell, R3 and R4 adjacent at base in T. gracilistria; CuP present near margin; anal loop 0.29× length of 1A+2A in T. gracilistria; with complex wing pattern of mainly longitudinal, fine parallel lines of cream, grey and brown, interrupted by a transverse series of
Figs 607–609. Tetramoera gracilistria, adults and head. 607, , Stannery Hills, Q. 608, , nr Hope Vale Mission, Q. 609, head, male. Scale = 2 mm.
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irregular small dark marks at about 36: wing, and with well-developed costal strigulae giving rise beyond middle to a sinuous leaden line running obliquely to below apex then curved to tornus along termen. Hindwing with well-developed cubital pecten and axillary tuft; Rs and M1 close but clearly separate at base, M2 curved close to M3 at base, M3 and CuA1 close but separate (T. gracilistria, T. calligrapha (Meyrick)) to connate (T. schistaceana) to stalked (T. isogramma (Meyrick)); CuP present towards margin, 1A+2A and 3A well developed; brownish grey. Pregenital abdomen. Unmodified. Hindmargin of S8 in male bilobed. Male genitalia (Figs 610–612, 615). Vinculum short, fused with valvae and tegumen.Tegumen high, ovate, a weakly sclerotised band. Apex of tegumen usually produced into narrow, elongateovate process, sometimes medially notched, covered dorsally and laterally with dense, long, hair-like bristles that ventrolaterally extend along a narrow band; in T. calligrapha apex produced into a pair of finger-shaped processes crowned with hairpencils. Socii very small drooping lobes with only a few bristles. Gnathos two variably developed curved lateral bands. Valva large and rather weakly sclerotised, deeply bilobed and densely hairy, with ventral sacculus area developed into a large hairy lobe nearly as long as dorsal lobe in some species, dorsal lobe without apical thorn and of variable shape, crescent-shaped to elongate triangular and widest at apex but always with a ventrally projecting lobe on inner surface varying from an inconspicuous blade-shaped process in T. schistaceana to a quadrate or band-shaped lobe; with a well-sclerotised, small, triangular, pocketshaped invagination at base on outer surface and with costal process straight and moderately long. Juxta very small, caulis short, anellus a broad band around aedeagus. Aedeagus stout, narrower towards apex, base bulbous and upcurved, with one or two dorsal, finger-like appendices; vesica usually lined with barb-like spinules, and with two to many needle-shaped cornuti. Female genitalia (Figs 613, 614). Ovipositor lobes rather broad, with large triangular posterior lobe. Ostium in membrane posterior to curvedconcave hindmargin of S7. Sterigma simple, a spinulose depression surrounding ostium, sometimes with a prominent bristle on each side, and in T. schistaceana with a wide and shallow pouch-shaped invagination leading to ostium. Ductus bursae membranous, without colliculum, at least as long as corpus bursae, slender but slightly
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widening towards corpus bursae with ductus seminalis originating from near middle. Corpus bursae ovate, with a single signum of somewhat variable shape even within a species, varying from a long sclerotised crease with a blade-shaped crest along ridge to a flat sclerotised ribbon or an elliptic, sclerotised depression with a small ridge or thorn on central ridge (T. schistaceana), and with either entire bursa finely granulate (T. schistaceana) or at least a patch of granular membrane opposite signum (T. gracilistria).
Distribution Tetramoera is widely distributed from South Africa, Madagascar, Mauritius, Sri Lanka, India, Myanmar, throughout south-east Asia, China, Japan (Ryukyu Is), Taiwan, the Philippines, Indonesia, Papua New Guinea to Australia, where the genus is found along the eastern coast of Queensland from Yeppoon to Cape York, and in the northernmost parts of the Northern Territory and in the Kimberley of Western Australia.
Biology Tetramoera schistaceana is a serious pest of sugar cane (Saccharum officinarum) (Poaceae), usually as a borer in the young shoots (Kalshoven 1981). The species was initially described from Java and Diakonoff ([1968]) provided references to its biology from Indonesia. No host is known from Australia, but the wing pattern of Tetramoera is typically that of a graminaceous feeder. The collecting localities in Australia are all somewhat drier localities, mostly monsoon forest, rarely dry woodland with possibly a few monsoon forest species, but not the typically wet rainforest. The sex pheromone of T. schistaceana has been identified as: Z9–12Ac and E9,11–12Ac in a ratio 100:1 (Kinjo et al. 1996).
genitalia of the types of T. isogramma and T. calligrapha, and the latter is here transferred to Tetramoera: calligrapha (Meyrick, 1912b), J. Bombay. Nat. Hist. Soc. 21: 865 (Eucosma). [India, Myanmar.] Comb. nov. The monophyly of Tetramoera is supported by several apomorphies, especially the uniquely modified, deeply bilobed flat and hairy valva without an apical thorn but with a usually flap-like ventral lobe from the ventral margin of the dorsal lobe. Further apomorphies are the single, elongate signum, usually a sclerotised crease with a narrow blade or small thorn on its central ridge, sometimes reduced to just a ribbon of sclerotisation, and the derived forewing shape and pattern. Given the overall plesiomorphic wing venation and the highly modified genitalia it is difficult to identify the sister group of Tetramoera. Overall similarities in male genitalia with Toonavora, gen. nov., such as a weakly sclerotised valva and short caulis and aedeagus, together with the possible homology of similar flap-like processes from the inner surface of the dorsal lobe in Tetramoera and next to the apical thorn in Toonavora, gen. nov., could indicate a closer relationship between the two genera. The finger-like aedeagus process is shared by Pternidora Meyrick. Superficially the Australian T. gracilistria and the lectotype of T. isogramma from Sri Lanka look extremely similar, but the former has much darker hindwings and two membranous, finger-like aedeagus processes, the latter only one. It is not clear whether all Tetramoera in Australia are T. gracilistria as there is some variation in the genitalia of both sexes, in uncus and valva shape as well as in the development of the signum. Tentatively, the Northern Territory population is considered to be distinct from T. gracilistria, based on a dorsal valva lobe with a rounded rather than straight apical margin and with a basally narrower flap-like process, and a signum with a keel.
Remarks Diakonoff ([1968]) described Tetramoera for T. schistaceana, based on material from Java, and T. isogramma, based on a series from Sri Lanka and South Africa, with a Sri Lankan specimen selected as the lectotype by Clarke (1958). Diakonoff figured adult, head and venation, and genitalia of both sexes for T. schistaceana (Diakonoff [1968]) and adults and male and female genitalia for T. leptalea Diakonoff from Madagascar (Diakonoff 1988c). Clarke (1958) provided photos of adults and male
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Constituent species Seven named species of Tetramoera are currently known from South Africa to Australia, including one Australian species, and there seems to be an unnamed Australian species from the Northern Territory closely related to T. gracilistria. It has to be confirmed that T. flavescens Kuznetzov from Vietnam (Kuznetzov 1988b) and T. flammeata Kuznetzov from the Kuril Is, described as a species of Enarmonia (Kuznetzov 1971), do indeed belong
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Figs 610–615. Tetramoera, genitalia. 610, 611, T. gracilistria, male genitalia and aedeagus, nr Hope Vale Mission, Q, slide ANIC 12924. 612, T. schistaceana, aedeagus, slide GP742. 613, T. gracilistria, female, nr Cooktown, Q, slide T2429. 614, T. schistaceana, female, Java, slide GP743. 615, T. schistaceana, male genitalia, Java, slide GP742.
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to Tetramoera (Kuznetzov 2001), given the very different signa in both species. gracilistria (Turner, 1946), Trans. R. Soc. S. Aust. 70: 206 (Eucosma). [Qld: Palm I.]
Thysanocrepis Diakonoff (Figs 74, 616–620) Thysanocrepis Diakonoff, 1966, Zool. Verh. (Leiden) 85: 45. Type species: Argyroploce crossota Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 266, by original designation. [Solomon Is.]
Diagnosis Thysanocrepis specimens are broad-winged, medium-sized to large olethreutine moths with long, sinuate labial palpi with a distally widened second segment. They are easily recognised by the shape and pattern of their variously suffused greyish ochreous to grey forewing characterised by a strongly sinuate costa with a conspicuous blackish triangle in its concave central portion as part of a series of small blackish marks along the costa. In the forewing R4 and R5 are adjacent and parallel at base or short-stalked, in the hindwing the bases of M2, M3 and CuA1 are all very close with M3 and CuA1 connate in some specimens, and the male has the anal margin rolled upwardly concealing a long scale pencil from a small lobe at the base of the hindwing. The genitalia in both sexes are diagnostic for Thysanocrepis. The male genitalia are characterised by a hairy, cone-shaped and distally notched uncus in combination with elongate, pendulous socii and without any trace of a gnathos, and by uniquely derived narrow valvae with a twice deeply emarginate ventral margin with a sclerotised, distally projecting, bristled sacculus followed by a bristled, tongue-shaped medial process with a blunt apical thorn and a widened, truncate apex to the valva, and by a short, wide aedeagus with an oblique apex and a large bunch of minute cornuti. The female genitalia are characterised by a spinulose sterigma ventrally to the ostium, a transverse sclerite with a deep Vshaped emargination in its hindmargin, and by two unequal-sized signa of unique shape, the smaller one with a cone-shaped base and a crescentic top with a jagged inner surface, the larger one with a more cylindrical base and the top an open semicylinder widening towards a deeply jagged rim.
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Description Adults (Figs 616, 617) Wingspan. 16.5–19.0 mm; medium-sized to large. Head (Fig. 618). Ocellus moderate. Upper part of frons with moderately long, anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis about as long as labial palpus. Labial palpus long (2.0–2.1× diameter of eye), porrect, strongly sinuate, second segment distally strongly widened by long, loose scales along dorsal and ventral margin, terminal segment long and slender; pale ochreous with four blackish marks laterally on second segment, the first one small, ventrally at base of segment, the second a large obliquely transverse streak just before middle, the third a strongly bluish iridescent ovate mark on loose dorsal scales near apex and the fourth comprising the ventral scales near apex, terminal segment touched with blackish ventrally and laterally at base. Antenna to middle of wing, scape anterodistally with long projecting blackish scales, anterior surface of flagellum without scales, cilia in male small. Thorax. Smooth, in male with a large, triangular tuft of scales from anteroventral corner of tegula. Hind tibia in male with long and rather loose scaling. Wings (Figs 74, 616, 617). Forewing index 0.43–0.48. Forewing widening towards apex and broadest before apex, with a conspicuously sinuate costa in both sexes with subbasal and subapical convex portions separated by concave central part, apex pointed, termen moderately oblique and nearly straight; anterior margin of cell evenly curved, M-stem present, chorda absent; all veins present and separate beyond cell except for R4 and R5 that are adjacent and parallel at base or shortstalked; CuP present at anal margin; wing greyish ochreous to grey fading to whitish ochreous in old material, variably tinged with reddish ochreous and grey, with markings consisting only of a series of small, blackish brown spots, bars and triangles along costa, with the fifth dark mark from base a prominent large triangle on concave middle part of costa, followed by a minute dot, then three about equally spaced bars or triangles and a last roundish spot in apex; several very fine, parallel, grey lines curving from before apex to lower part of termen and costal half of wing sprinkled with silvery white scales; base of costa and tegulae and distal third of wing from R5 to dorsum often with red-brown suffusion. Hindwing with well-developed cubital pecten and axillary tuft, in male with anal margin
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Figs 616–620. Thysanocrepis crossota, adults, head and genitalia. 616, 617, adults. 616, , New Britain, Papua New Guinea [image reversed]. 617, , Papua New Guinea. 618, head, male. 619, male genitalia, Cedar Bay, Q, slide BM 24922. 620, female genitalia, Papua New Guinea, slide ANIC 12921. Scale = 2 mm.
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rolled upwardly concealing a long scale pencil originating from small lobe at base of hindwing; with Rs and M1 closely approximated and parallel at base, bases of M2, M3 and CuA1 all very close and M3 and CuA1 connate in some specimens; CuP present at margin; 1A+2A and 3A well developed, 3A distant from anal margin; dark brownish grey with coppery gloss. Pregenital abdomen. Ventral apodemes long, triangular at base, slender towards tip. In male hindmargin of S8 with shallow, V-shaped, medial emargination; hindmargin of T8 rounded-convex. Male genitalia (Fig. 619). Vinculum a narrow, weakly sclerotised band. Tegumen weakly sclerotised, short and wide, shape distorted in slide preparation, laterally along entire length with very long, dense scales from large sockets. Uncus a hairy, membranous cone with a cleft apex like two paired nipples. Socii membranous, distally rounded, narrow, pendulous, hairy lobes. No gnathos remnants apparent. Valva small, narrow, moderately sclerotised, with a nearly straight costa but ventral margin with two deep, rounded emarginations, the deeper more basal one separating the sclerotised, distally projecting and densely bristled sacculus from a tongue-shaped, densely bristled medial process with a blunt thorn at apex, and the second, longer and more shallow emargination between this medial process and rounded, distally widened truncate apex; costal process long and straight. Juxta small, caulis rather short. Aedeagus short, wide, with oblique apex; vesica with a large bunch of minute cornuti. Female genitalia (Fig. 620). Ovipositor lobes elongate-ovate. Hindmargin of S7 curved-concave. Sterigma a strongly spinulose transverse sclerite ventrally to ostium (lamella antevaginalis) with a deep V-shaped medial emargination in posterior margin leading to ostium, lamella postvaginalis not developed except for some spinules in membrane. Ductus bursae slender and membranous except for well-developed colliculum of two small, lateral, elliptic sclerites; corpus bursae globular with a small, nipple-shaped lateral extension where ductus bursae and ductus seminalis originate, with two large, complex signa of roughly similar shape but different height, the smaller one ventrally, close to origin of ductus bursae, with a low, cone-shaped base and a crescentic top with a jagged inner surface, the larger, dorsal one with a long, more cylindrical base and a semi-cylindrical top widening towards strongly jagged rim. Bulla seminalis very small with a long connection to corpus bursae.
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Distribution Thysanocrepis has been reported from Papua New Guinea, Indonesia (western Sulawesi) and Australia, and the BMNH has material from India (Assam), Thailand, Malaysia, Brunei and the Solomon Is. In Australia it has been collected in northern Queensland, in lowland rainforest near Cairns to Cedar Bay south of Cooktown.
Biology No foodplant is known for Thysanocrepis. The Australian specimens have all been collected close to sea level, from rainforest or rainforest margins. The New Guinea specimens comprise both older material from close to sea level and more recent material collected further inland on New Britain and mainland New Guinea at up to 1450 m by W. W. Brandt.
Remarks In his description of Thysanocrepis, Diakonoff (1966) provided drawings of head, venation and male genitalia of T. crossota, and later (Diakonoff 1975) of the female genitalia of T. celebensis Diakonoff. Diakonoff (1966) remarked on the unusual combination of olethreutine neuration with eucosmine genitalia and suggested the genus might belong somewhere near ‘Articolla’ prospera Meyrick. Later (Diakonoff 1973), misled by the genitalia of a superficially similar-looking but unrelated female, discounted his initial correct assignment and included Thysanocrepis in the subtribe Zomariae of the Olethreutini. The description of a second species based on a correctly identified female from Sulawesi (Diakonoff 1975) did not result in a reassessment of the taxonomic position of the genus. Thysanocrepis is characterised by the forewing shape with a strongly sinuate costa and with R4 and R5 adjacent and parallel at base or short-stalked. Apomorphies in the male genitalia include a totally reduced gnathos, the moderately sclerotised, slender valva with a bristled, distally sclerotised and projecting sacculus, a tongue-shaped, bristled, ventral cucullus process with a blunt apical thorn, and a rounded, distally widened apex. The shape of the transverse, sclerotised and densely spinulose sterigma and the unique signa are apomorphies in the female genitalia. No sister taxon to Thysanocrepis has been identified, but several derived features are shared with other enarmoniine genera. A scale pencil
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originating from the hindwing base and concealed by an uprolled anal margin and the dense, long scales with large sockets laterally on the tegumen are both shared by several enarmoniine genera. The paired, papillate and hairy tegumen tip of Thysanocrepis, together with the well-developed socii, suggests that a hairy and somewhat bilobed uncus may be an apomorphy of a subordinate group within the Enarmoniini. A similar hairy and bilobed uncus, together with distinct socii, is found in Helictophanes uberana Meyrick and Cyphophanes dyscheranta Meyrick. Cursory examination has found no consistent specific differences across the wide range of Thysanocrepis, but more detailed study is needed to synonymise T. celebensis with T. crossota.
Constituent species Two named species are known, one from Sulawesi and one from Papua New Guinea, the latter extending into northern Australia. crossota (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 266 (Argyroploce). [Solomon Is., New Guinea.]
Toonavora, gen. nov. (Figs 75, 621–626) Type species: Eucosma aellaea Turner, 1916, Trans. R. Soc. S. Aust. 40: 526, hereby designated. [Australia.]
Diagnosis Toonavora species are small to medium-sized, vividly marked moths with a complex wing pattern of orange to reddish brown, cream and leaden on dark blackish brown, incorporating all the typical grapholitine pattern elements of conspicuous strigulae, a well-developed ocelloid spot and paired outwardly curved mid-dorsal streaks that are often expanded into a conspicuous whitish mark on middorsum. They have sinuate, moderately long labial palpi with a yellowish ochreous patch distally on the second segment. In the forewing all veins are separate beyond the cell and in the hindwing M3 and CuA1 are stalked. The strongly modified male hindwing venation is unique and diagnostic for Toonavora, with M3 and CuA1 stalked to over 6M or entirely fused into a single vein, and with CuA2 branching from hindmargin of cell at a very narrow
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angle and with a course unusually close to CuA1 right to wing margin. The genitalia in both sexes also are diagnostic for the genus. The male is characterised by a cone-shaped tegumen tip covered with long bristles, by an elongate-ovate valva with sinuate dorsal and ventral margins with a ventral, subapical thorn and a hairy flap from inner surface next to thorn, by a short caulis/ anellus complex modified into a large cup-shaped structure with a triangular dorsal lobe, and by a short aedeagus with a downcurved ventral point on apex. The female is even more unusual with a strongly extensible ovipositor, the ostium in an inverted T-shaped suture between S7 and the downcurved posterior corners of T7, with sclerotisations along the margins, and with two triangular, blade-shaped signa from a flat base.
Description Adults (Figs 621, 622) Wingspan. 10.5–17.0 mm; small to mediumsized. Head (Fig. 623). Ocellus large. Upper part of frons with long, anteriorly projecting scales, remainder of frons with minute, appressed scales. Proboscis little longer than labial palpus. Labial palpus porrect, sinuate, moderately long (1.6–1.9× diameter of eye), second segment sinuate and strongly widened in distal half by long, loose scales ventrally and especially dorsally towards apex, often nearly concealing terminal segment, terminal segment moderately long, short-scaled, pointing obliquely downward; colouration of labial palpus typically enarmoniine with a dorsolateral yellowish ochreous patch distally on second segment usually surrounded by blackish scaling, but at least with a blackish area across middle of second segment and a blackish terminal segment. Antenna to middle of wing, ventral band without scales, cilia small. Thorax. Smooth. Legs unmodified. Wings (Figs 75, 621, 622). Forewing index 0.39–0.43. Forewing elongate, slightly dilated towards termen, with costa evenly curved, apex rounded-pointed and projecting, termen oblique and weakly sinuate; M-stem to between M2 and M3, chorda from halfway between R1 and R2 to R5; all veins separate beyond discal cell, base of M3 closer to CuA1 than to M2 and the two veins relatively close right to margin; CuP well developed near margin; anal loop 0.33× length of 1A+2A in T. aellaea; with a vivid and complex pattern of reddish brown, cream and leaden on blackish brown ground, with a series of distally cream-
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Figs 621–623. Toonavora aellaea, adults and head. 621, 622, adults. 621, , Allyn R., NSW [image reversed]. 622, , Mt Keira, NSW. 623, head, male. Scale = 2 mm.
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coloured strigulae along costa mostly giving rise to oblique leaden lines edged with reddish brown on both sides, with a series of longitudinal, parallel blackish bars in the tornal area separated by reddish brown lines and basally and distally flanked by irregular leaden bands, with dorsal region to beyond middle speckled with irregular dots of reddish brown scales and with a pair of outwardly curved narrow loops on middle of dorsum, leaden in centre and surrounded by mixed cream and reddish brown scales, often both marks confluent or suffused by white to form a large whitish mark on middle of dorsum. Hindwing with moderate cubital pecten and anal tuft, sexually dimorphic, narrower in male with modified venation; Rs and M1 parallel but clearly separate at base; in female M2 curved towards M3 at base but not closely approximate, M3 and CuA1 stalked to about 56:, and CuA2 at a normal angle from beyond 36: cell with a course distant to CuA1; in male M2 nearly straight and moderately distant from M3 at base, M3 and CuA1 stalked to over 6M or entirely fused into one vein (the two conditions found on one wing each of the same specimen), and CuA2 at a very small angle from beyond 36: of cell, parallel and close to hindmargin of cell and stalk of M3 and CuA1; CuP only a weak trace at margin, 1A+2A with anal loop reduced, 3A present; blackish brown with coppery gloss. Pregenital abdomen. Unmodified. Hindmargin of S8 bilobed in male. Male genitalia (Figs 624, 625). Vinculum entirely fused with valvae and with tegumen. Tegumen very weakly sclerotised, a narrow band (shape distorted in microscopic slides). Apex of tegumen a membranous, narrow cone with dense, long, hair-like bristles extending ventrolaterally, presumably formed of uncus and socii. Gnathos two lateral, medially upcurved arms, barely recognisable. Valva rather weakly sclerotised, elongate-ovate with both ventral and dorsal margins sinuate and a strong, single or bifid thorn on ventral margin before rounded, bristled apex, and with a hairy flap-like process next to thorn from an angled ridge running on inner surface from thorn towards base of valva; with a large tuft of long bristles before middle of ventral margin, and ventral margin of basal excavation ending in a knob-shaped process at base near juxta; costal process short and distally tapering. Juxta quite large, caulis short, fused together with anellus to form a large cup-shaped structure with dorsal margin projecting as a large triangle and with a small, well-sclerotised, second triangular process
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Figs 624–626. Toonavora aellaea, genitalia. 624, 625, male genitalia and aedeagus, Mt Bellenden-Ker, Q, slide T2237. 626, female genitalia, Mt Webb Nat. Park, Q, slide T2373.
below aedeagus. Aedeagus small, straight, distally expanding with apex forming a ventral, strongly sclerotised, downcurved blunt point; vesica with a sheaf of long, slender, sinuate cornuti. Female genitalia (Fig. 626). Ovipositor modified, slender and greatly extensible, with ovipositor lobes narrowly elliptic, erect, smooth on inner surface. Apophyses, especially posterior pair, greatly lengthened. T7 longer than S7 and posterolateral corners of T7 ventrally extended to nearly meet in ventral midline posterior to ostium, forming an inverted T-shaped fissure between hindmargin of S7 and the two posterolateral corners of T7, with ostium at point of intersection; downcurved posterolateral margins of T7 sclerotised parallel to hindmargin of S7 and forming a medially interrupted transverse dark band posterior to ostium, and with a small bent nose-shaped sclerite below ostium (lamella antevaginalis), sometimes flanked on each side by a strong bristle. Ductus bursae narrow, membranous, without colliculum, with ductus seminalis originating at 36: from ostium, anterior third
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slightly widening and joining corpus bursae laterally. Corpus bursae globose, with two unequal signa, long sharply pointed triangular blades, the shorter one laterally from a flat base and the larger one medially from a ridged base.
Distribution Toonavora is known from Papua New Guinea and Australia, where the genus occurs along the eastern coast from Kiama (south of Wollongong in New South Wales) to just north of Cooktown in Queensland.
Biology The Papuan T. spermatophaga (Diakonoff & Bradley) has been described from specimens reared by John Dobunaba from seeds of Toona sureni (Meliaceae) in Bulolo, Papua New Guinea. In Australia, T. aellaea is frequently reared from seeds of Toona ciliata, and the collecting localities of T. aellaea exactly coincide with the distribution of
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Toona ciliata. There is no doubt that Toonavora larvae feed on seeds of Toona spp.
Remarks The Papuan T. spermatophaga was described as an Anathamna with the genitalia figured (Diakonoff and Bradley 1976), but venation and genitalia in both sexes are distinct and unlike Anathamna, and the species is here transferred to Toonavora: spermatophaga (Diakonoff & Bradley, 1976), Bull. Entomol. Res. 66: 401 (Anathamna). [New Guinea.] Comb. nov. The monophyly of Toonavora is based on several unique apomorphies in wing venation and the genitalia of both sexes. M3 and CuA1 in the hindwing are stalked in both sexes, but the male hindwing is further modified with M3 and CuA1 stalked nearly to margin or entirely fused, and with CuA2 throughout closely approximated to CuA1. Apomorphies in the male genitalia include a caulis/ anellus complex modified to form a large, cup-like structure with a triangular dorsal lobe, and an aedeagus with the apex ventrally produced into a downcurved point. The female genitalia are equally derived with a modified and strongly extensible ovipositor and an usual position of the ostium. This has tentatively been interpreted as an inverted Tshaped fissure between S7 and the posterior corners of T7 rather than an incision in S7. Toonavora shares a more or less ventral hairy process from the cucullus with Tetramoera and Thysanocrepis. The forewing markings of Toonavora with all the elements of a typically grapholitine wing pattern may well be close to the groundplan wing pattern for the Enarmoniini. Helictophanes shares the stalked M3 and CuA1 in the hindwing, the subapical ventral thorn on the valva, a short caulis and two large blade-shaped signa, and T. aellaea has sometimes been mistaken for Helictophanes scambodes (Meyrick).
Constituent species Two species of Toonavora are known to date, one from Papua New Guinea and one from Australia: aellaea (Turner, 1916), Trans. R. Soc. S. Aust. 40: 526 (Eucosma). [Qld: Beerburrum near Nambour.] Comb. nov.
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EUCOSMINI Meyrick Eucosmidae Meyrick, 1909, J. Bombay Nat. Hist. Soc. 19: 582. Type genus: Eucosma Hübner, 1823. Eucosminae; Heinrich, 1923, US Nat. Mus. Bull. 123: 1. Eucosmini; Obraztsov, 1946, Z. Wien. Entomol. Ges. 30(1945): 23.
Status remark Heinrich (1923) established the Eucosminae as a subfamily, in accordance with the American tradition with Olethreutidae as a separate family. In a European context, Obraztsov (1946) treated the same group at tribal level. More recently, the Enarmoniini have been removed as a separate group, with only a few genera including Ancylis Hübner in the European fauna, but much more diverse in the Oriental/Australian region. Some authors considered the Enarmoniini and Eucosmini to be sister taxa, both at subtribal level within a composite tribe Eucosmini (Razowski 1989), but more recently the Eucosmini have generally been accepted as a distinct tribe (Horak 1998; Razowski 2001; Kuznetzov and Stekolnikov 2001).
Evidence for monophyly Traditionally, the Eucosmini were defined by a hindwing venation with M2 at its base curved towards and approximated to the stalk of M3 and CuA1 (Heinrich 1923; Obraztsov 1946), in contrast to the Olethreutini with separate or at most connate M3 and CuA1 and the Grapholitini with M2 at its base distant from and parallel to M3. This definition serves with very few exceptions to separate the three tribes in the Nearctic and European fauna, much less so in the Oriental/ Australian region. Furthermore, enarmoniine hindwing venation falls across the entire range of this spectrum. There is no apomorphy known to be present in all eucosmine genera, but several derived characters occur only in the Eucosmini or elsewhere only in such a distant position in the Tortricinae that an independent multiple origin is a reasonable assumption: presence of a notch at base of antenna, presence of a costal fold, raised scales on the forewing dorsum, absence of 3A in the hindwing, the sterigma smooth or only very finely spinulose, horn-shaped signa often reduced and entire bursa with spines. Stalked R4 and R5 and presence of a
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scale pencil within the hindwing anal roll are both frequently found in Eucosmini and Enarmoniini, but never in Olethreutini. Most Eucosmini are further characterised by a thorax without posterior crest, absence of apodemes for muscle attachments on the pedunculi, usually by presence of deciduous lanceolate cornuti, and a ductus bursae often sclerotised between corpus bursae and origin of ductus seminalis. These four character states contrast those found in nearly all Olethreutini. The hindwing in males of many Eucosmini has modified, very dark and small scales that appear to be more widely spaced with translucent wing membrane visible in between. Given the lack of a universal synapomorphy for the Eucosmini, it had become accepted wisdom that the group may not be monophyletic, and in particular, may be paraphyletic in terms of the Grapholitini (Horak and Brown 1991; Horak 1998; Razowski 2003a). However, in the very preliminary cladistic analysis (see p. 17), the eucosmine genera frequently formed a clade at the top of the tree, as a remarkably resilient grouping even if various characters were excluded. The consistent absence of 3A in the eucosmine hindwing except in the overall probably plesiomorphic genera Rhopobota Lederer, Epiblema Hübner and Crocidosema Zeller, is particularly interesting since 3A is generally present in grapholitine genera. On balance the current evidence seems rather to favour the possibility of monophyletic Eucosmini, but only a cladistic analysis based on a re-scored matrix including molecular data will settle the issue.
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Meyrick) or entirely fused into a single vein (Eccoptocera Walsingham, some species of Spilonota Stephens, Holocola Meyrick, Crocidosema and others); 3A at most a trace or entirely lost in most genera. S2 of ‘tortricoid type’, with antrolateral processes and paired apodemes. Tegumen never with conspicuous apodemes for muscles m4. Uncus frequently small and paired or absent, never ventrally hairy. Socii generally large, often sclerotised or otherwise modified. Valva with well-developed cucullus usually delimited by ‘neck’, in Acroclitagroup with flattened bristles ventrally on outer surface. Aedeagus with bundle of deciduous cornuti. Female genitalia with sterigma often well-sclerotised, smooth or only very finely spinulose, in emargination of S7 or fused with S7. Ductus bursae often with a sclerotised ring near inception of ductus seminalis to variably sclerotised in anterior half. Signa nearly always paired, horn- to thorn-shaped, if lost then wall of corpus bursae spiny.
Distribution The tribe is present world-wide, including New Zealand, but the genus-groups are not evenly distributed across the faunal regions. Epinotia Hübner, Eucosma Hübner and Epiblema are particularly well represented in the Nearctic and Palaearctic regions, whereas the genera related to Spilonota have their greatest diversity in Australia and the genera associated with Acroclita are most numerous in the Oriental region.
Biology Diagnosis Species of the Eucosmini are very small to large olethreutine moths, often narrow-winged and with usually well-developed olethreutine wing pattern of costal strigulae and ocelloid patch. Antenna in male often with a notch near base. Thorax without posterior crest except in some Noduliferola species. Legs unmodified. Forewing often with costal fold in male and in some groups with raised scale patches on dorsum; R4 and R5 frequently stalked. Hindwing often with expanded anal region with anal roll containing scale pencil from base, sometimes with modified scales along anterior margin of cell; Rs and M1 often stalked, M2 nearly always curved closely to M3 at base (not close in some Strepsicrates Meyrick and sexually dimorphic Acroclita males), M3 and CuA1 nearly always stalked (separate only in sexually dimorphic males of Acroclita Lederer and Tritopterna Meyrick, widely distant in Herpystis
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The Eucosmini include leaf rollers and leaf tiers as well as borers in flower heads, shoots, stems and roots, more rarely they mine conifer needles or produce a blotch mine (Epinotia nemorivaga Tengström (Hannemann 1961)). The overall more plesiomorphic genera appear to be leaf rollers, but external and internal feeding may be found in the same genus, as in Eucosma Hübner (Hannemann 1961). Many Holarctic Eucosmini are restricted to conifers and the group comprises serious forestry pests. The Australia-centred Spilonota-group has a strong association with Myrtaceae and has possibly tracked the radiation of this family in response to increasing aridity.
Remarks There are numerous chapters on the Eucosmini in geographically localised publications. The more
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comprehensive treatments range from the still very relevant monograph by Heinrich (1923) on the Nearctic Eucosminae, Hannemann’s (1961) diagnostically illustrated fauna of Germany, Obraztsov’s (1964, 1965, 1967) incomplete revision of the European genera, to Bentinck and Diakonoff’s (1968) fauna of the Netherlands and the volume by Bradley et al. (1979) detailing the biology of British species with exquisite drawings. More recently, Kuznetzov (1978) provided a revision of the fauna of the European part of the former USSR, illustrated with excellent drawings, including wing venations, and Miller (1987) a diagnostic pictorial guide to the species of midland North America. Razowski (1989) published a synopsis of the Palaearctic Olethreutinae with genitalia drawings of both sexes and a comprehensive treatment of the European olethreutine fauna (Razowski 2003a), with colour photos of all species and genitalia drawings of high quality. Kawabe (1982) provided photos of adults, in colour, and of the genitalia of some species, and Liu and Li (2002) illustrated adults and genitalia of both sexes for the Chinese fauna. While Pierce and Metcalfe (1922) erected two groups based on signa shape for the European Eucosmini, Heinrich (1923) recognised ‘three main lines of development’ for the Nearctic fauna, without elaborating on characters. Obraztsov (1946) did not indicate any subdivisions within the Eucosmini, but roughly followed Heinrich’s (1923) sequence in his arrangement. Kuznetzov (1978) and Razowski (1987) each presented the eucosmine genera in a different sequence, but with Epinotia near the beginning in both cases. Kuznetzov (1978), without any comments, located Spilonota at the beginning and Rhopobota Lederer and Acroclita at the very end. In the introduction to the generic descriptions, Razowski (1987) acknowledged that eucosmine phylogeny is still not understood, but outlined six tentative lineages: Epinotia/Zeiraphera/ Crocidosema, Eucosma/Phaneta/Pelochrista, Epiblema to Rhyacionia, Eriopsela/Thiodia, Gibberifera/Gypsonoma, and Rhopobota/Spilonota. While revising the Australian eucosmine genera, unique synapomorphies were found to support several genus-groups, even though they were not always present in all species. However, if they were present they were diagnostic for the particular grouping. Two groups are particularly prominent in the Australian fauna, the genera around Strepsicrates Meyrick and Spilonota, usually with myrtaceous hosts and often with an antennal notch in the male, and a large group including Acroclita and Rhopobota, characterised at least in some
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species in each genus by large, flattened bristles from the outer surface of the valva. These two groupings frequently were recovered as clades in the cladistic analysis and are here treated as informal genus-groups. Epiblema and Coenobiodes Kuznetzov on the one hand and Icelita Bradley and Eucosmophyes Diakonoff on the other consistently appeared as sister pairs in the cladistic analysis. The first two genera, together with the absconditanagroup, share several obvious synapomorphies and are referred to as the Epiblema-group, corresponding to one of Razowski’s (1987) groupings. Likewise, Crocidosema Zeller belongs to one of the lineages defined by Razowski (1987) here treated as the Epinotia-group. Elucidation of the position and relationships of Icelita, Eucosmophyes, Herpystis Meyrick, Whittenella, gen. nov., ‘Acroclita’ tothastis Meyrick and several other yet to be named Oriental/Australian eucosmine genera has to await a more rigorous cladistic analysis, including molecular data. Their present position at the end of the tribe, unassigned to genus-groups, is rather arbitrary and not based on phylogenetic considerations. Some of the derived features of these taxa may simply be a result of their small size.
Epinotia-group Status remarks This is merely a convenient informal genus-group, corresponding to Razowski’s (1987) tentative Epinotia/Zeiraphera/Crocidosema lineage.
Evidence for monophyly The Epinotia-group is overall plesiomorphic and may well be a basal grade in the Eucosmini. There is no suggestion that the group is monophyletic, and at this stage it certainly does not deserve formal ranking. The group is characterised by a generalised morphology usually with a costal fold, a welldeveloped, often bifid uncus, long socii, a simple valva (cucullus shape modified within the genera), a sclerotised ring in the ductus bursae just posterior to the origin of the ductus seminalis, and two large horn-shaped signa. Fine strands connecting the anellus with the base of the socii could be a possible apomorphy for the group.
Distribution The Epinotia-group is most diverse in the Holarctic (Epinotia Hübner) and Neotropical (Crocidosema
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Zeller) regions, with only few representatives in the Oriental region, and only Crocidosema in Australia.
Biology Members of the Epinotia-group are mostly leaf rollers on numerous plant families, but several Crocidosema species bore in flowers and fruit of Malvaceae, and some Epinotia species mine leaves and conifer needles (Hannemann 1961).
Remarks Plesiomorphic Epinotia and Ancylis Hübner genitalia are strikingly similar due to many symplesiomorphies, but Epinotia has a ring-shaped sclerite in the ductus bursae at the origin of the ductus seminalis and always lacks the welldeveloped apodemes for muscle m4 present in the males of Ancylis.
Crocidosema Zeller (Figs 76, 627–640) Crocidosema Zeller, Isis Oken, Leipzig 1847: 721. Type species: Crocidosema plebejana Zeller, 1847, ibid. 1847: 721, by monotypy. [Sicily.] Crocidosoma Walker, 1863, List Specimens Lepid. Insects Colln Br. Mus. 27: 279. An incorrect subsequent spelling. Heligmocera Walsingham, 1892, Proc. Zool. Soc. Lond. 1891: 507. Type species: Heligmocera calvifrons Walsingham, 1892, ibid. 1891: 508, pl. 41 fig. 5, by original designation. [West Indies.] Parasuleima Clarke, 1965, Proc. US Natn. Mus. 117: 77. Type species: Crocidosema insulana Aurivillius, 1922, in Aurivillius, Prout & Meyrick, in Skottsberg, Nat. Hist. Juan Fernandez and Easter Island 3(2): 267, pl. 11, fig. 20, by original designation. [Juan Fernandez I.]
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the dorsal half. Males of the plebejana-group lack a costal fold but in most species have the cubital pecten modified to form a large, dense, erect scale tuft diagnostic for the genus if present. In males of the lantana-group the costal fold is present, the frons is concave with modified scaling and a scale pencil from the base of the hindwing margin is inserted into a longitudinal fold along the base of the abdomen. The labial palpi are long, sinuate and widened medially in all Australian species and the forewing has M-stem and chorda weakly developed and R4 and R5 separate or stalked. In the hindwing Rs and M1 are separate or stalked, M3 and CuA1 stalked or entirely fused, and 3A is well developed. The male genitalia are diagnostic for the genus with the hindmargin of S8 deeply divided into distally often pointed lobes, the uncus a dorsally hairy, down-curved process, the socii slender, drooping lobes with connecting strands from each side of the anellus to their base, the valva with a curved rib along outer surface, a triangularly projecting sacculus, a hump on the costal base and either an ovate or very large rhomboid cucullus, often with large bristles from its outer surface. The female genitalia also are diagnostic for the genus with narrowed ovipositor lobes and S7 either with two scale-filled large depressions or narrow and strongly reduced to two usually medially separated, sclerotised transverse bands with a serrate posterior margin and a projecting tooth or lobe on each side of the sterigma. The sterigma consists of a spinulose lamella postvaginalis and a smooth ring or large sclerotised funnel fused with the hindmargin of S7. The colliculum, if present, is of thickened membrane, the posterior portion of the ductus bursae is sclerotised, ending in an inverted Vshaped sclerite in the lantana-group, the anterior portion from origin of the ductus seminalis is membranous, with a small diverticulum in widened anterior end. The corpus bursae is lined with spinules and has two curved, sickle-shaped signa with a hollow base and often bluntly rounded tip.
Description Diagnosis Crocidosema species are small to medium-sized, typical eucosmines usually with well-developed costal strigulae ending in a large, V-shaped, white preapical pair followed by a dark apical spot, an ocelloid patch framed by two silver bands with at least traces of black bars, a dark, triangular, pretornal mark on dorsum and nearly in all species with a dark basal third of the wing, at least along
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Adults (Figs 627–630) Wingspan. 10.5–17.0 mm; small to mediumsized. Head (Fig. 637). Ocellus well developed. Frons either with a wide band of long, anteriorly projecting scales in upper part and remainder with small, appressed scales (plebejana-group), or strongly modified in male, concave with scales arranged in vertical, lateral ridges and medially in a feather-like pattern (C. lantana (Busck) and close
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Figs 627–630. Crocidosema, adults. 627, 628, C. plebejana. 627, , Wilton, NSW. 628, , nr Cocklebiddy, WA. 629, C. polyphaea, , Yeppoon, Q. 630, C. lantana, nr Yeppoon, Q. Scale = 2 mm.
relatives). Labial palpus moderately long to long (2.0–2.5× diameter of eye), usually porrect, strongly sinuate; second segment large, sinuate, teardropshaped, strongly widened in distal half by long, loose scales along ventral margin and distal half of dorsal margin; terminal segment long, slender, inclined, often concealed in long scales from second segment (in the New World Crocidosema longipalpana Möschler terminal segment modified, slender and upturned in male (Brown et al. 2002 as C. litchivora Baixeras)). Antennae to just beyond middle of forewing; scaled except for narrow band, with minute cilia in male of C. plebejana-group, moderately long in lantana-group. Wings (Fig. 76, 627–630). Forewing index 0.32–0.35. Forewing elongate, lightly widening towards termen, costa lightly curved near base and apex, apex projecting, termen oblique and lightly sinuate; male with (lantana-group) or without (plebejana-group) costal fold; chorda from halfway between R1 and R2 to between R5 and M1, M-stem to M3, both only weakly developed; all veins
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separate in plebejana-group, R4 and R5 connate or stalked in lantana-group, M2, M3 and CuA1 converging towards margin; CuP present at margin; in C. plebejana anal loop 0.30× length of 1A+2A; wing pattern diverse, sexually dimorphic in C. plebejana; except for one unnamed Australian species that is whitish speckled with few greyish scales, Crocidosema has well-developed costal strigulae ending in a large, V-shaped, white preapical pair followed by a dark apical spot, an ocelloid patch framed by two silver bands with at least traces of black bars, a dark, triangular, pretornal mark on dorsum and in nearly all species basal third of wing darker, at least dorsal half. Hindwing with long, scattered hairscales across anal region in male, modified and curved near loop of anal vein, with cubital pecten in males of plebejana-group usually a very large, dense, erect tuft of long, distally spatulate scales; in males of lantana-group with a long scale pencil from base of anal margin; Rs and M1 parallel and closely approximated towards base or stalked, base of M2
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curved close to base of M3, M3 and CuA1 stalked or entirely fused; CuP present at margin; 1A+2A and 3A well developed; whitish to brownish grey. Pregenital abdomen. Males of lantana-group with T1–3 strongly modified (Fig. 632), with a
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longitudinal fold along each side filled with sex scales, accommodating the scale pencils from the hindwing base. Hindmargin of S8 in male deeply divided, with long, distally pointed lobes in plebejana-group (Fig. 635), T8 with a weak, inverted Y-shaped sclerotisation.
Figs 631–637. Crocidosema, male genitalia, abdomen and head. 631, 632, C. lantana, Rous, NSW, slide ANIC 3143. 631, genitalia. 632, dorsal abdomen base. 633–635, C. polyphaea, Yeppoon, Q, slide T2399. 633, genitalia. 634, aedeagus. 635, abdomen, 8th segment. 636, 637, C. plebejana. 636, genitalia, Black Mt, ACT, slide ANIC 12995. 637, head.
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Male genitalia (Figs 631, 633, 634, 636). Tegumen weakly sclerotised, wide and low. Uncus a downcurved medial process, finger- to lobe-shaped, dorsally hairy. Socii long, narrow, densely hairy, drooping lobes, with two weak, parallel connections from each side of anellus to base of socii. No trace of gnathos visible. Valva highly modified with a narrow short ‘neck’, a curved rib on outer surface from base to apex, with sacculus triangularly projecting, a subtriangular hump on base of costa beyond costal process, and a very large cucullus, subtriangular to rhomboid in the plebejana-group, ovate to band-shaped in lantana-group, hairy on inner surface and with a series of large, ventrally directed bristles from outer surface in plebejana-group. Juxta subtriangular to crescentic, caulis short, with narrow lateral keels towards anellus, anellus ring-shaped, laterally connected by weak strands to base of socii. Aedeagus curved, slightly narrowed towards apex; vesica with a sheaf of deciduous, lanceolate cornuti.
Female genitalia (Figs 638–640). Ovipositor lobes strongly narrowed, flat or erect and parallel. Sternum 7 in the lantana-group with two round, scale-filled depressions with a serrate ridge along their hindmargin; S7 in the plebejana-group reduced to narrow remnant, very weakly sclerotised except for two medially usually separated, strongly sclerotised narrow transverse bands with a serrate posterior margin and with paired, sclerotised processes varying from two small teeth to two flat, ear-shaped lobes (C. plebejana, Hawaiian species), one on each side of the sterigma. Sterigma a spinulose, longitudinal band in intersegmental membrane behind entrance to ostium, often with bristles, and a smooth, sclerotised ring or large funnel fused with hindmargin of S7. Ductus bursae with a colliculum of thickened membrane in plebejana-group, followed by a smooth, sclerotised tube to origin of ductus seminalis, in lantana-group colliculum not apparent and sclerotised posterior section ending in an inverted V-shaped sclerite near middle; membranous anterior portion widening
Figs 638–640. Crocidosema, female genitalia. 638, C. lantana, Mt Bartle Frere, Q, slide ANIC 13751. 639, C. polyphaea, Yeppoon, Q, slide ANIC 13749. 640, C. plebejana, Black Mt, ACT, slide ANIC 12991.
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towards corpus bursae with a small diverticulum at ‘neck’ of corpus bursae; corpus bursae inverted pear-shaped, partially spinulose, with two large, curved, sickle-shaped signa with hollow base and often bluntly rounded tip.
Distribution Apart from the cosmopolitan C. plebejana, Crocidosema is known from the southern US, central and South America, Hawaii and Australia where it occurs in all states except Tasmania. Two endemic Australian species have been collected, one near Geraldton in Western Australia and one (C. apicinota (Turner)) from Cairns to Yeppoon in Queensland.
Biology The larva of C. plebejana, the cotton tip worm, is a pest on cotton (Gossypium hirsutum) in Australia and is generally reported to feed in seeds, fruits and flowers of numerous Malvaceae (Malva, Hibiscus, Sida). However, in Australia it has repeatedly been reared from heads of wheat (Poaceae) and once from Cassia occidentalis (Caesalpiniaceae) in the Northern Territory. There is a large literature on C. plebejana with regard to cotton (Bishop and Blood 1978; Hamilton and Zalucki 1993, etc.), and it has been reported as injurious to okra (Abelmoschus esculentus) from Japan (Nasu and Yasuda 1993). Zimmerman (1978) reported several species of Abutilon and Sida (both Malvaceae) as hosts of C. plebejana (as C. blackburnii (Butler)) and Abutilon sandwicense and Hibiscus arnottianus as hosts of Crocidosema marcidella (Walsingham). Whilst the species close to C. plebejana have a strong affinity with Malvaceae, the New World species recently transferred to Crocidosema in the tortricid part of the Checklist of the Atlas of Neotropical Lepidoptera (Powell et al. 1995) have a much wider host spectrum, illustrated by Crocidosema lantana, introduced from Mexico to Hawaii as a biocontrol agent for Lantana (Verbenaceae), and by Crocidosema aporema (Walsingham) feeding on soybean (Glycine max, Fabaceae) (Liljesthrom et al. 2001).
Remarks Given the wide distribution of C. plebejana, numerous descriptions and illustrations are available, including those by Heinrich (1923) and Razowski (1989). Parts of the description of
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C. plebejana and the figures given for the male genitalia and wing venation by Hannemann (1961) are incorrect and must refer to another genus, probably Acroclita. Zimmerman (1978) gave a detailed and well-illustrated discussion of the Hawaiian Crocidosema species (some synonyms of C. plebejana), including information on immatures. Clarke (1965) figured head, venation and female genitalia of C. insulana (as Parasuleima) from Juan Fernandez Is with M3 and CuA1 usually entirely fused. The concept of Crocidosema has recently been expanded to include a number of New World species, including C. lantana (Powell et al. 1995). Both in its restricted and extended scope the genus could well be merely a derived group within Epinotia Hübner, characterised by numerous apomorphies but leaving Epinotia a paraphyletic assemblage. However, as Epinotia is not part of the Australian fauna, it seems best to leave the status quo until the entire group is subject to a cladistic analysis. Brown et al. (2002) provided a comprehensive description, including immatures, of C. longipalpana, a member of the lantana-group. The plebejana- and lantana-groups are each characterised by a number of derived features, in particular different secondary sexual characters in the male. However, the transformation series of the female S7 with the paired scale-filled depressions in C. lantana (Fig. 638) and C. longipalpana, each with a serrate margin and a proximal tooth in C. longipalpana, to the remnant of S7 in Crocidosema polyphaea (Turner) (Fig. 639) and C. plebejana (Fig. 640), consisting largely of the enlarged serrate ridges and teeth, is a strong indication that the group is monophyletic. Other derived features of Crocidosema include the valva with a hump at the base of the costa, a projecting sacculus, a rib along the outer surface, and the downcurved, hairy uncus. On the other hand, Epinotia and Crocidosema share overall very similar wing venation and genitalia in both sexes, in particular a similar tegumen structure including connecting strands between the anellus and the base of the socii, and a partially spinulose bursa with similar signa. Different species of the C. plebejana-group may look very similar superficially and the male genitalia generally provide few diagnostic characters. Judging from the Australian species and the figures given by Zimmerman (1978), the female genitalia, in particular the shape of S7, provide the best diagnostic characters for species separation. Presence of two quite distinct, localised, endemic
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Australian species of the plebejana-group, one on the Queensland coast and one in Western Australia, and the occurrence of C. plebejana throughout the continent, suggest that Crocidosema has been present in Australia for a long time. The fact that several malvaceous species are salt tolerant and a frequent feature of beach vegetation may well be relevant to the cosmopolitan distribution of C. plebejana. Eucosma polyphaea Turner in Horak et al. (1996) was erroneously treated as a senior synonym of Crocidosema apicinota (Turner) even though Common (1957) had previously synonymised C. polyphaea with Epinotia lantana.
Constituent species Twenty four valid species of Crocidosema are listed in Brown (2005). Two named and one unnamed species are known from Australia, in addition to the deliberately introduced C. lantana. plebejana Zeller, 1847, Isis Oken, Leipzig 1847: 721 (Crocidosema). [Sicily.] altheana (Mann, 1855), Verh. Zool.-bot. Ges. Wien 5: 555 (Penthina). [Germany.] lavaterana (Millière, 1863), Ann. Soc. Linn. Lyon (N. S.) 9: 34 (Paedisca). [France.] peregrinana (Möschler, 1866), Berl. Entomol. Z. 10: 139 (Grapholitha). [France.] obscura (Wollaston, 1879), Ann. Mag. Nat. Hist. (5)3: 341 (Steganoptycha). [St Helena.] blackburnii (Butler, 1881), Ann. Mag. Nat. Hist. (5)7: 393 (Proteopteryx). [Hawaiian Is.] plebeiana Meyrick, 1881, Proc. Linn. Soc. NSW 6: 659 (Crocidosema). [unjustified emendation] excitana (Möschler, 1891), Abh. Senckenb. Naturforsch. Ges. 16(1889): 333 (Grapholitha). [Puerto Rico.] ptiladelpha Meyrick, 1917, Trans. Entomol. Soc. Lond. 1917: 18 (Crocidosema). [Peru.] insulana Aurivillius, 1922, in Skottsberg, Nat. Hist. Juan Fernandez and Easter Island 3: 267 (Crocidosema). [Chile, Juan Fernandez Is.] synneurota Meyrick, 1926, Trans. Entomol. Soc. Lond. 74: 276 (Crocidosema). [Galapagos Is.] charmera (Turner, 1946), Trans. R. Soc. S. Aust. 70: 202 (Eucosma). [Qld: Toowoomba; NSW: Ebor.]
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tornocycla (Turner, 1946), Trans. R. Soc. S. Aust. 70: 210 (Eucosma). [Qld: Eungella, Toowoomba.] iris Diakonoff, 1984a, Entomol. Basil. 9: 389 (Crocidosema). [Indonesia, West Sumba.] bostrychodes Diakonoff, 1992, Ann. Soc. Entomol. Fr. (N. S.) 28: 46 (Crocidosema). [Madagascar.] apicinota (Turner, 1946), Trans. R. Soc. S. Aust. 70: 206 (Eucosma). [Qld: Mackay.] Sp. rev. eridela (Turner, 1946), Trans. R. Soc. S. Aust. 70: 202 (Eucosma). [Qld: Mackay.] perversa (Turner, 1946), Trans. R. Soc. S. Aust. 70: 207 (Eucosma). [Qld: Yeppoon.] lantana Busck, 1910, Proc. Entomol. Soc. Wash. 12: 132 (Crocidosema), intro. delib. [Hawaiian Is; introduced from Mexico.] polyphaea (Turner, 1926), Trans. R. Soc. S. Aust. 50: 138 (Eucosma). [Qld: Yeppoon.] tornocosma (Turner, 1946), Trans. R. Soc. S. Aust. 70: 205 (Eucosma). [Qld: Noosa, Brisbane.] phaedropa (Turner, 1946), Trans. R. Soc. S. Aust. 70: 209 (Eucosma). [Qld: Toowoomba.] corynetes (Diakonoff, 1982), Zool. Verh. (Leiden) 193: 59 (Epinotia). [Sri Lanka.]
Epiblema-group Status remarks This is an informal genus-group, corresponding to one of Razowski’s (1987) tentative eucosmine lineages in which he included Epiblema Hübner, Notocelia Hübner, Blastesthia Obraztsov (as Coccyx Treitschke), Retinia Guenée, Barbara Heinrich, Gravitarmata Obraztsov and Rhyacionia Hübner.
Evidence for monophyly It is not suggested that this grouping is monophyletic, and at this stage it certainly does not deserve formal ranking. The Epiblema-group of genera, including Coenobiodes Kuznetzov and the absconditana-group in Australia and several other mainly Palaearctic and/or Nearctic genera, is characterised by drooping, elongate socii, a valva with a single or bifid process (harpe, clasper) near the ventral margin of the basal excavation, and usually a very short caulis in the male. The less derived taxa have a well-developed costal fold, the sterigma separate from S7 in the intersegmental
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membrane and some have retained a bifid uncus. A process from or near the ventral margin of the basal excavation is also found in some other Eucosmini (Gypsonoma Meyrick, Rhopobota Lederer), and only a phylogenetic analysis will show whether this structure has developed independently more than once. However, the three Australian genera including Epiblema may well be part of a monophyletic group.
Distribution The Epiblema-group is most diverse in the Holarctic region, reaching Australia only with a few rather derived taxa.
Biology The larvae of the Epiblema-group are either leaf rollers or borers, with those of Epiblema boring in roots and stems of Asteraceae and those of several other genera in twigs or cones of conifers. Epinotia absconditana (Walker) was reared from a larva joining leaves on Rapanea howittiana (Myrsinaceae) at Wingham, NSW, by J. Stockard.
Remarks Epiblema did not naturally occur in Australia. The North American E. strenuana (Walker) has been deliberately introduced for biological control of the weed Parthenium hysterophorus (Asteraceae).
Epiblema Hübner (Figs 77, 641–646) Epiblema Hübner, [1825], Verz. Bekannter Schmett: 375. Type species: Phalaena foenella Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 536, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 6, 54. [Europe.] Cacochroea Lederer, 1859, Wien. Entomol. Monatschr. 3: 331. Type species: Paedisca grandaevana Lienig, 1846, in Lienig & Zeller, Isis Oken, Leipzig 1846: 238, by monotypy. [Europe.] Monosphragis Clemens, 1860, Proc. Acad. Nat. Sci. Philad. 12: 354. Type species: Monosphragis otiosana Clemens, 1860, ibid. 12: 354, by monotypy. [USA.] Euryptychia Clemens, 1865, Proc. Entomol. Soc. Philad. 5: 140. Type species: Euryptychia
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saligneana Clemens, 1865, ibid. 5: 141, fig. 8, by monotypy. [North America.]
Diagnosis The medium-sized Epiblema strenuana (Walker), introduced as a biocontrol agent and the only species of this genus in Australia, is superficially easily recognisable by its minutely speckled forewing covered with dark grey scales with pale ochreous tips and with a sparse wing pattern consisting of white, silvery grey and brownish ochreous costal strigulae towards the apex, a large, roughly triangular, uniformly grey mark on the dorsum with a pale basal margin and reaching nearly to the costa, and a whitish ocelloid patch with silvery grey marks and a curved silvery band along its inner margin edged with black scales. A long, slender costal fold is present in the male and the labial palpi are porrect, sinuate, rather short and with a short terminal segment. All veins are separate in the forewing and in the hindwing only M3 and CuA1 are stalked to about 56M. A conspicuous, rod-like sclerotisation runs laterally along the entire length of the second tergite on both sides. The male genitalia of E. strenuana are characterised by a tegumen with broad ‘shoulders’, a lobe-shaped uncus and large, long drooping socii narrowed beyond the middle, a valva with two processes on the margin of the basal excavation consisting of a flattened hump ventrally and a large, erect, triangular semi-cone distally. The female genitalia have modified ovipositor lobes with their inner sides appressed to each other, a large sterigma formed by a longitudinal, spinulose band behind the ostium with upturned lateral margins carrying a few bristles on each side, and large, blunt, hornshaped cornuti with flattened, round apices.
Description Adults (based on strenuana, Figs 641, 642) Wingspan. (10)12–19 mm; medium-sized. Head (Fig. 645). Ocellus well developed. Upper part of frons with anteriorly projecting scales, remainder with small, appressed scales. Labial palpus porrect, sinuate, rather short (1.6–1.8× diameter of eye), second segment sinuate, gradually widening to just before broadly rounded apex, with loosely projecting scales ventrally and distally, terminal segment short. Antenna to about middle of wing, entirely scaled in male, without visible cilia. Thorax. Smooth. Legs unmodified.
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Figs 641–646. Epiblema strenuana, adults, head and genitalia. 641, 642, adults. 641, , Roma, Q. 642, , Parkhurst, Q. 643, 644, male genitalia and aedeagus, nr Taroom, Q, slide ANIC 12970. 645, head, male. 646, female genitalia, nr Taroom, Q, slide ANIC 12971. Scale = 2 mm.
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Wings (Figs 77, 641, 642). Forewing index 0.38. Forewing elongate, subrectangular (subtriangular in many Palaearctic species), widest before tornus, with costa weakly curved in basal half, apex lightly protruding, termen sinuate; with well-developed, slender costal fold to beyond 56: costa, enclosing scale pencil; discal cell with chorda rudimentary, from between R1 and R2 to between R4 and R5, M-stem a weak trace only; all veins separate with R3, R4 and R5 roughly equidistant at base and with M2, M3 and CuA1 equidistant; CuP present at margin; anal loop 0.33 length of 1A+2A; E. strenuana with ground colour of dark grey scales with pale ochreous tips, with costal strigulae of increasing length beyond costal fold consisting of alternate silvery grey and brownish ochreous lines separated by white lines converging below costa before apex; with a large, irregular, roughly triangular mark of entirely grey scales on middle of dorsum reaching nearly to costa and edged basally by an indistinct, sinuate paler band; ocelloid patch whitish containing some silver grey marks and a black longitudinal streak at its apex, preceded by a curved silvery band basally edged with some black scales. Hindwing with well-developed cubital pecten and axillary tuft; Rs and M1 closely approximated at base and sometimes anastomosing, base of M2 moderately close to stalk of M3 and CuA1 that are stalked to about 56:; CuP present at margin, 1A+2A and 3A well developed; grey. Pregenital abdomen. Conspicuous, rod-shaped sclerotisations laterally along entire length of T2. Hindmargin of S8 bilobed in male with shallow medial emargination; T8 with weak, inverted T-shaped sclerotisation at base and convex hindmargin. Male genitalia (Figs 643, 644). Tegumen with very prominent ‘shoulders’ and wide pedunculi. Uncus a rounded lobe with some hairs dorsally. Socii large, drooping, elongate lobes. Gnathos laterally narrow bands medially fused to sclerotised ventral surface of anal tube, forming a projecting, groove-shaped process. Valva medially strongly narrowed with ventral margin angled in basal half and a large, kidney-shaped cucullus projecting on ventral margin as a rounded lobe, set with increasingly stout bristles towards ventrodistal margin; basal excavation with a conspicuous, flattened, hump-shaped process on ventral margin and a very large, erect, triangular process bent to form a semi-cone near distal margin (harpe); costal process inconspicuous, down-curved. Juxta triangular, caulis very short. Aedeagus tapering towards apex; vesica with a bundle of sinuate, flattened cornuti.
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Female genitalia (Figs 646). Ovipositor lobes in E. strenuana modified, narrow, their inner surfaces pressed together. Hindmargin of S7 with deep, U-shaped emargination. Sterigma a narrow ventral rim around entrance to ostium and a large, longitudinal, spinulose band in intersegmental membrane behind ostium with lateral margins raised to form projecting flanges, with a few bristles on each side. Ductus bursae very short (less than 56O length of corpus bursae), somewhat twisted and widening towards corpus bursae, membranous except for sclerotised central portion; colliculum two small, indistinct plates of thickened membrane just below ostium; ductus seminalis from anterior third; corpus bursae large, teardrop-shaped, with two large, short and wide, horn-shaped signa with flattened, round apex.
Distribution Epiblema strenuana was introduced into Australia from North America (Mexico) where the species has a very wide distribution from southern Canada to Mexico. Material in the ANIC has been collected from much of Queensland, from Toowoomba in the south-east to Moreton on Cape York Peninsula, inland at Carnarvon Gorge and at Musselbrook in the north near the border with the Northern Territory. The genus Epiblema is distributed throughout the Holarctic and large parts of the Oriental region.
Biology In its native range E. strenuana, the ragweed borer, bores in stems and branches of Ambrosia, Xanthium and Chenopodium (Heinrich 1923; Miller 1987). It has been deliberately introduced into Australia in 1982–1984 to control exotic weeds in the Asteraceae and has widely established on Ambrosia artemisiifolia, Parthenium hysterophorus and Xanthium strumarium, the last a cosmopolitan weed and the first two introduced from North America and Mexico respectively (Julien 1992; Dhileepan 2001). Larvae of Epiblema generally are borers in stems and roots of Asteraceae.
Remarks Epiblema strenuana was diagnosed and figured by Heinrich (1923) and Miller (1987), and the genus Epiblema was discussed in detail and illustrated by Heinrich (1923), Obraztsov (1965) and Kuznetzov (1978). Obraztsov (1965) divided the genus into the
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three subgenera: Epiblema s. str.; Cacochroea Lederer (characterised by a second process from the ventral margin of the basal excavation in addition to the more distal tooth), and Notocelia Hübner with two fixed cornuti. Notocelia is now treated as a distinct genus (Kuznetzov 1978; Brown 1979; Razowski 1989; Karsholt and Razowski 1996) and Cacochroea as a synonym of Epiblema (Kuznetzov 1978; Razowski 1989). The rounded, more basal process used by Obraztsov (1965) to define Cacochroea is well developed in E. strenuana, and probably belongs to the groundplan of the Epiblema-group of genera. There is no single, unique apomorphy for Epiblema, but the combination of a costal fold, conspicuous ‘shoulders’ to the tegumen, elongate, drooping socii and a medially constricted valva with a sclerotised, usually triangular process (harpe) from near the distal margin of the basal excavation (in E. strenuana in addition to a humplike more basal process) characterises the genus. Epiblema shares a similar valva with a process from near the distal margin of the basal excavation with the absconditana-group and Coenobiodes Kuznetzov, both present in Australia, and Notocelia and Blastesthia Obraztsov. Notocelia, as stated above, is characterised by two fixed cornuti and the absconditana-group by a bifid uncus, strongly reduced socii and a different configuration of M2 and M3 in the forewing. Blastesthia differs from Epiblema only in derived features such as absence of costal fold and its specialised feeding mode in conifer buds.
Constituent species Epiblema is a large genus with nearly 90 species in the Holarctic and Oriental regions (Brown 2005). There is only one deliberately introduced species in Australia: strenuana (Walker, 1863), List Specimens Lepid. Insects Colln Br. Mus. 28: 383 (Grapholita). [North America.] exvagana (Walker, 1863), List Specimens Lepid. Insects Colln Br. Mus. 28: 383 (Grapholita). [North America.] flavocellana (Clemens, 1865), Proc. Entomol. Soc. Philad. 5: 138 (Steganoptycha). [USA.] subversana (Zeller, 1875), Verh. Zool-bot. Ges. Wien 25: 318 (Grapholita). [USA.] minutana (Kearfott, 1905), Proc. US Natn. Mus. 28: 356 (Eucosma). [USA.]
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antaxia (Meyrick, 1920), Exotic Microlepid. 2: 344 (Eucosma). [unecessary replacement name for E. minutana.]
The absconditana-group (Figs 78, 647–657)
Diagnosis The two species of the absconditana-group are small to medium-sized, diversely marked greyish brownish moths with variably developed, outwardly oblique, darker transverse lines or bands beyond base and beyond middle, with a series of dark spots (between strigulae that may or may not be developed) from middle of costa to apical spot, and hardly any trace of an ocelloid patch. The labial palpi are long, porrect and hardly sinuate, the forewing has costa and M-stem well developed, the bases of M2 and M3 closely approximated and lacks a costal fold in the male, the hindwing has Rs and M1 closely approximated and parallel at base and M3 and CuA1 stalked to about 56:. There are no external diagnostic features for the group. However, the male genitalia are diagnostic with a bifid uncus, the socii hairy, drooping, membranous lobes, weak gnathos arms connected with two sclerotised rods laterally along the anal tube, and, most importantly, valvae with a bipartite process along the ventral margin of the basal excavation consisting of a series of bristles along the basal excavation overlaid by a keel-shaped projection with a more basal rounded lobe and a more distal point, if well developed. The female genitalia are very characteristic with a complex sterigma in the intersegmental membrane (connected to S7 in one of the two species), a sclerotised ring connected to a large, ear-shaped, membranous and spinulose lobe on each side, recalling the sterigma of the Palaearctic Notocelia Hübner, a variable but always posteriorly sclerotised ductus bursae and two thorn- to horn-shaped signa with a hollow base.
Description Adults (Figs 647, 648, 651) Wingspan. 10–18 mm. Small to medium-sized. Head (Figs 649, 650). Ocellus well developed. Frons with long, anteriorly projecting scales in upper part, remainder of frons with minute, appressed scales. Proboscis probably slightly longer
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Figs 647–651. The absconditana-group, adults, head and antenna. 647, 648, Epinotia absconditana, adults. 647, , nr Casino, NSW [image reversed]. 648, , Toowoomba, Q. 649, 650, E. absconditana, male. 649, head. 650, antenna, ventral view. 651, Eucosma symploca, paratype, , Cape York, Q. Scale = 2 mm.
than labial palpus. Labial palpus long (2.2–2.3× diameter of eye), porrect, hardly sinuate; second segment strongly widened near or beyond middle by long, loose scales along ventral margin and distal half of dorsal margin, ovate to subtriangular; terminal segment long, slender, porrect. Antennae reaching to middle of wing, entirely scaled in male (Fig. 650), no cilia visible. Thorax. Without crest. Legs unmodified. Wings (Figs 78, 647, 648, 651). Forewing index 0.36–0.40. Forewing moderately wide, costa and dorsum roughly parallel in distal half, costa lightly curved throughout but more so at base, apex slightly to moderately projecting, termen weakly oblique and lightly sinuate; without costal fold in male; chorda from below R2 to between R5 and M1, M-stem to base of M2, both well developed; all
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veins separate beyond discal cell, M2 and M3 very close at base with M3 closer to M2 than to CuA1; CuP present near margin; anal loop 0.28× length of 1A+2A; wing pattern quite different in the two Australian species but both brownish with outwardly oblique, transverse fasciae either indistinct and only partially developed on paler brownish background (Epinotia absconditana (Walker)) or more or less complete though irregular, wide and partially edged with blackish brown on whitish background (Eucosma symploca Turner), always with well-developed dorsal portion of subbasal fascia and with a series of dark spots (between strigulae that may or may not be developed) from middle of costa ending in a dark apical spot. Hindwing with cubital pecten and axillary tuft well developed; Rs and M1 parallel and
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Figs 652–657. The absconditana-group, genitalia. 652, 653, Epinotia absconditana, male genitalia and aedeagus, Wollongong, NSW, slide T2152. 654, Eucosma symploca, paratype male, Cape York, Q, slide ANIC 12984. 655, 656, E. absconditana. 655, female, Mt Keira, NSW, slide T2146. 656, female, signa, slide ANIC 4992. 657, E. symploca, female, Thursday Is, Q, slide ANIC 12981.
closely approximated at base, base of M2 bent close to base of stalked M3 and CuA1; CuP present at wing margin, closer to CuA2; 1A+2A well developed, no trace of 3A; brownish grey. Pregenital abdomen. Unmodified. Hindmargin of S8 in male strongly bilobed, T8 with indistinct, inverted Y-shaped sclerotisation. Male genitalia (Figs 652–654). Tegumen high, moderately sclerotised; pedunculi hardly tapering towards base. Uncus distally bifid. Socii long, drooping, hairy lobes. Gnathos weak lateral arms connected with two long, parallel, sclerotised rods laterally along anal tube. Valva of typical Epiblemashape with stout costal process, base of costa convex to angled and ventral margin of basal excavation modified, in E. absconditana with a row of bristles
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along distal portion of margin overlaid by a keelshaped projection from inner valva surface forming a round lobe ending distally in a triangular point, reduced to a bristled lobe on margin of basal excavation in E. symploca; valva slightly narrowed beyond sacculus, cucullus ovate to reniform with a row of spines along ventral margin. Juxta large, pentagonal, caulis very short, anellus a wide ring. Aedeagus short to moderately long, narrowing towards apex; vesica with few deciduous cornuti. Female (Figs 655–657). Ovipositor lobes subreniform, anteriorly narrower. Sternum 7 wide, hindmargin lightly concave. Sterigma complex, in intersegmental membrane, but fused with hindmargin of S7 in E. absconditana; a sclerotised ring leading to ostium and laterally connected with
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a large, ear-shaped, membranous, spinulose lobe on each side; sterigma partly invaginated behind S7 in E. symploca, in E. absconditana fused with modified hindmargin of S7. Ductus bursae variable, very long and complex in E. absconditana but shorter and apparently reduced in E. symploca; posteriormost part of ductus bursae a smooth, at least partially sclerotised tube to ductus seminalis that arises from closer to ostium than to corpus bursae, in E. absconditana with a membranous ring below ostium; no trace of colliculum; remainder of ductus bursae straight and membranous in E. symploca, gradually widening towards corpus bursae, but very long and tightly coiled in E. absconditana with the first half-coil sclerotised, the rest membranous. Corpus bursae ovate with two signa with hollow base, long, nearly straight, distally pointed, bladeto horn-shaped in E. absconditana, and short, subtriangular to hook-shaped in E. symploca.
Distribution The two species occur at Thursday I. and Cape York and from the Bunya Mts in Queensland to Kiama in New South Wales.
Biology Epinotia absconditana was reared from a larva joining leaves on Rapanea howittiana (Myrsinaceae) at Wingham, NSW, by J. Stockard, and has only been collected in rainforest localities. Eucosma symploca is known from near the tip of Cape York only where eucalypt as well as monsoon forest is present.
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and differ in the configuration of M2 and M3 in the forewing. The absconditana-group is left as an informal species-group until the closely related Plutographa Diakonoff from Madagascar and related Oriental Eucosmini have been studied. Coenobiodes is the closest genus to the absconditana-group in the Australian fauna, sharing a very similar valva with a stout costal process, a process from the ventral margin of basal excavation and a very short caulis in the male. The female genitalia in both groups have a posteriorly sclerotised and anteriorly twisted ductus bursae, but the Australian species of Coenobiodes all have a conspicuous constriction somewhat below the ostium.
Constituent species The following two named Australian species form the absconditana-group: absconditana (Walker, 1863), List. Specimens Lepid. Insects Colln. Br. Mus. 28: 351 (Sciaphila) [NSW: Sydney.] perplexa Turner, 1916, Trans. R. Soc. S. Aust. 40: 526 (Eucosma). [Qld: Brisbane, Killarney.] aulacota Turner, 1946, Trans. R. Soc. S. Aust. 70: 212 (Eucosma). [NSW: Acacia Plateau.] symploca Turner, 1946, Trans. R. Soc. S. Aust. 70: 203 (Eucosma) [Qld: Cape York.] euprepes Turner, 1946, Trans. R. Soc. S. Aust. 70: 204 (Eucosma). [Qld: Cape York.]
Coenobiodes Kuznetzov Remarks The two species of the absconditana-group with a well-developed, bipartite process from the ventral margin of the basal emargination clearly belong to the Epiblema-group of genera. Early on during this revision they were treated as species of Nuntiella Kuznetzov on the basis of the bifid uncus and a forewing venation with M2 and M3 closely approximated at base. This assignment proved to be incorrect once the female of Nuntiella became known (Zhang and Li 2004) and the genus turned out to belong to the Acroclita-group. The sterigma in the two species of the absconditana-group is highly derived with large, spinulose, membranous lobes on each side, recalling the sterigma of Notocelia Hübner. However, the Australian species lack the two fixed cornuti diagnostic for Notocelia
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(Figs 79, 658–666) Coenobiodes Kuznetzov, 1973, Entomol. Obozr. 52: 687. Type species: Coenobiodes acceptana Kuznetzov, 1973, ibid. 52: 689, by original designation. [Japan.]
Diagnosis Species of Coenobiodes are medium-sized, fairly broad-winged and have a dark brownish, typically eucosmine wing pattern on whitish ground with a lilac-grey, greyish blue or ochreous tinge, with basal third of wing darkened though sometimes interrupted by transverse pale bands, a large triangular or more rarely a small hourglass-shaped mark on dorsum next to ocelloid patch often giving
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rise from its tip to a dark band curving around ocelloid patch to middle of termen, and a blackish spot in apex usually fused with a dark triangle over costal half of termen. Coenobiodes has moderately long, sinuate labial palpi, white-tipped and white on inner surface, with the second segment medially widened and with a slender terminal segment. All veins are separate in the forewing, with R3, R4 and R5 close at base and M2, M3 and CuA1 roughly equidistant, in the hindwing Rs and M1 are stalked to 56M in all Australian species but separate in the Japanese C. acceptana, and M3 and CuA1 are stalked to about 56M. The male genitalia are characterised by a tegumen with distinct ‘shoulders’ and very long and slender socii (except C. melanocosma (Turner) with wider sclerotised socii toothed on median margin). The valva is medially strongly constricted and has a triangular to hump-shaped process on ventral margin of basal excavation and a tuft of persistent scales from outer surface of sacculus, and the aedeagus is either short and wide and/or has lateral flanges. The female genitalia are characterised by ovipositor lobes that are either slender throughout or narrow anteriorly with much wider posterior lobes, by a very variable sterigma ranging from a sclerotised ring in an emargination of S7, often with small dorsal or ventral lobes, to a projecting triangular structure fused with S7, by a conspicuous constriction in the ductus bursae below ostium formed by two membranous, lens-shaped structures, and by two long, slender, horn-shaped signa in the Australian species (much smaller in C. acceptana).
slightly oblique termen; discal cell with chorda and usually also M-stem well developed; all veins separate with R3, R4 and R5 rather close at base, likewise M2, M3 and CuA1, which are all about equidistant; CuP present at margin; anal loop 0.30× length of 1A+2A; ground colour whitish transversely strigulated and partially suffused with pale grey, lilac-grey or greyish blue, with typical eucosmine, brownish and blackish brown pattern elements: costa strigulated throughout and with five conspicuous pairs of long white strigulae in distal half and with a blackish spot in apex; basal
Description Adults (Figs 658, 659) Wingspan. 11–18 mm; medium-sized. Head (Fig. 662). Ocellus moderate. Vertex scales anteriorly projecting between antennae, merging with downcurved scales of upper frons, remainder of frons with appressed, small scales. Labial palpus porrect, sinuate, moderately long (1.8–2.3× diameter of eye), second segment sinuate, teardrop-shaped, gradually widening to beyond middle with rounded apex of long, loose scales, terminal segment slender, moderately long. Antennae to just beyond middle of forewing, entirely scaled in male, without visible cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 79, 658, 659). Forewing index 0.38–0.44. Forewing elongate subtriangular to subrectangular, widest just before tornus with weakly curved costa, lightly protruding apex and
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Figs 658, 659. Coenobiodes, adult males. 658, C. sp., Mt Keira, NSW. 659, C. melanocosma, Lamington Nat. Park, Q. Scale = 2 mm.
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third of wing with several, variable, transverse, parallel, blackish brown lines fused distally to form an angulate or outwardly oblique dark band, particularly in dorsal half; an elongate triangular to hourglass-shaped dark mark on dorsum before tornus reaching to beyond middle of wing and anterior to ocelloid patch usually connected with a curved dark line skirting ocelloid patch and running to middle of termen, usually confluent with a dark triangle across anterior half of termen including apical spot; ocelloid patch a pale ovate patch with a vertical dark zig-zag line in a darker distal area roughly from beyond 36: dorsum to apex. Hindwing with well-developed cubital pecten and anal tuft; Rs and M1 closely approximated at base (C. acceptana) or stalked to 56M (all Australian species), base of M2 close to base of stalk of M3 and CuA1, which are stalked to about 56M; CuP present at margin, 1A+2A well developed, no trace of 3A in male of C. melanocosma; grey to greyish coppery. Pregenital abdomen. Short, rod-like sclerotisations laterally on T2 from base to middle. Hindmargin of S8 strongly bilobed in male with a deep V-shaped emargination; T8 with transverse triangular sclerotisation at base and convex hindmargin. Male genitalia (Figs 660, 661, 663, 666). Tegumen with prominent rounded ‘shoulders’ lateral to socii and wide pedunculi. Uncus varying from a short bifid process (C. acceptana) to a rounded hump (ending in a ventral, vertical, blade-like process in C. melanocosma) to a small, rounded cone. Socii very long and slender, drooping, hairy throughout, in C. melanocosma wider, sclerotised and with a strongly toothed median margin. Gnathos two weak lateral bands, medially fused or rising. Valva long and fairly slender, medially narrowed, with large, triangular to hump-shaped projection from ventral margin of basal excavation (harpe) and stout costal process; basal half of valva narrow, with a large tuft of long, persistent scales or bristles from outer surface; cucullus delineated by conspicuous ‘neck’, ovate to kidney-shaped, sometimes with dense sheaf of bristles from outer surface. Juxta semicircular to triangular, caulis very short. Aedeagus nearly always rather short and wide, often with conspicuous lateral flanges; vesica with a large bundle of flattened, distally sinuate cornuti. Female genitalia (Figs 664, 665). Ovipositor lobes either very slender throughout or slender anteriorly with a much larger posterior lobe. Ostium usually in emargination of S7, in C. melanocosma integrated within S7; S7 next to
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sterigma with scales in large, modified sockets. Sterigma very variable, a variously modified rim around entrance to ostium, sometimes with short ventral and/or dorsal lobes, in C. melanocosma a smooth, triangular projection fused with S7 with entrance to ostium in its base and with a posteriorly widening, longitudinal strip of sclerotised spinules to base of S8. Ductus bursae with a conspicuous constriction (?colliculum) shortly below ostium formed by thickened, lensshaped, membranous structures on each side (in one undescribed Australian species this is far below ostium and the intervening ductus bursae forms a rigid tube to ostium), usually followed by a lightly sclerotised portion to origin of ductus seminalis, anteriormost part curved and widening towards ovate corpus bursae; two horn-shaped signa, long and slender in Australian species, but very small in C. acceptana.
Distribution Coenobiodes is known from Japan, Vanuatu and Australia, where it occurs along the eastern coast from McIlwraith Range in Queensland to Kiama in New South Wales.
Biology No host is known for Coenobiodes, but all the Australian material is from rainforest localities.
Remarks Unlike the Australian species of Coenobiodes the type species C. acceptana from Japan has a bifid uncus with two small lateral tips and Rs and M1 in the hindwing are not stalked. Otherwise, however, the Australian species have very similar genitalia in both sexes and share the same characteristic forewing pattern. Coenobiodes acceptana is well documented in the original description by Kuznetzov (1973) with figures of male and female genitalia and Kawabe (1982) gave a colour illustration of the adult. Bradley (1962) provided illustrations of adults and genitalia of the species from Vanuatu. There is no single, unique apomorphy for Coenobiodes, but a combination of the following derived features defines the genus: pronounced ‘shoulders’ to the tegumen, extremely long and slender socii (except in C. melanocosma where they are highly derived), a hump-shaped process (harpe) on the lower margin of the basal excavation, a tuft
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Figs 660–666. Coenobiodes, genitalia and head. 660, 661, C. sp., male genitalia and aedeagus, Mt Keira, NSW, slide T2396. 662, head, C. melanocosma, male. 663, C. melanocosma, aedeagus, slide T2226. 664, C. sp., female genitalia, Mt Keira, NSW, slide T2406. 665, C. melanocosma, female genitalia, Lamington Nat. Park, Q, slide T2376. 666, C. melanocosma, male genitalia, Lamington Nat. Park, Q, slide T2226.
of persistent scales from the outer surface of the valva, aedeagus often with lateral flanges, narrow or modified ovipositor lobes, a ring-shaped sterigma unless sterigma fused with S7, scales with modified sockets on S7 next to sterigma, ductus bursae with conspicuous constriction below ostium formed by two membranous lens-shaped
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structures, long and slender horn-shaped signa, a characteristic forewing pattern, and labial palpi white-tipped and white on inner surface. The character combination in the various species suggests that the sclerotised, serrate socii and the sterigma fused with S7 in C. melanocosma as well as the long slender aedeagus with large lateral flanges
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and the posteriorly strongly sclerotised ductus bursae in an unnamed Australian species are infrageneric apomorphies. The only close relatives of Coenobiodes in the Australian fauna are the absconditana-group with a very similar valva and shared traits in the female genitalia, and the introduced Epiblema Hübner. Coenobiodes shares with Epiblema, Notocelia Hübner, Hendecaneura Walsingham and other principally northern hemisphere genera a medially constricted valva with a hump-shaped process (harpe) from the ventral margin of the basal excavation and elongate, drooping socii. The enigmatic Plutographa Diakonoff from Madagascar (Diakonoff 1970, 1989a) with its bifid uncus, long slender socii and usually with a process from the lower margin of the basal excavation also looks closely related, though the strongly spinulose sterigma and especially the presence of coremata in some species are unusual features in eucosmines. The Japanese Coenobiodes abietella (Matsumura) (Kuznetzov 1976; Kawabe 1982) has less slender socii and a differently shaped valva lacking the hump on the margin of the basal excavation. This, together with very different sterigma, ductus bursae and wing pattern could well exclude the species from Coenobiodes. In addition to the Japanese type species C. acceptana, a species from Vanuatu is here referred to Coenobiodes (Bradley 1962; Diakonoff 1959b, 1989): euryochra (Bradley, 1962), Bull. Br. Mus. (Nat. Hist.), Entomol. 12: 252 (Eucosma). [Vanuatu.] Comb. nov.
Constituent species There are four named species of Coenobiodes, one in Australia, in addition to two unnamed Australian species. melanocosma (Turner, 1916), Trans. R. Soc. S. Aust. 40: 526 (Eucosma). [Qld: Brisbane.] Comb. nov.
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Evidence for monophyly Several apomorphies are each shared between variably overlapping subsets of the Acroclita-group, together forming a possibly monophyletic grouping. At least some species of most genera have modified, usually flattened bristles from the outer surface of the valva, the anellus has prominent laterobasal angles, there is a strong tendency to sclerotised socii, the ductus bursae anteriorly often has a characteristic, bi-pronged sclerite, and several genera have modified scales near the anterior margin of the discal cell or the costa of the hindwing. The group shares several other traits that, however, are also found elsewhere in the Eucosmini: raised scales on the fore wing dorsum, R3 closely adjacent at base to the usually stalked R4 and R5, very long labial palpi, and a paired or bifid uncus if present. Acroclita and Tritopterna both often have sexually dimorphic hindwing venation.
Distribution The Acroclita-group is centred on the Oriental region and well represented in Australia. Rhopobota extends across the Holarctic region, Acroclita reaches to the Western Palaearctic, Megaherpystis Diakonoff is known from the Seychelles, Madagascar and the African region (Diakonoff 1989b), and Proteoteras Riley from the Nearctic region.
Biology Species of Rhopobota and Acroclita are leaf rollers, Melanodaedala feeds in flowers that are spun together, Proteoteras and Fibuloides larvae are borers, those of Tritopterna and Noduliferola are fruit borers, and Heleanna larvae are reported as both borers and leaf rollers.
Remarks Several features refer Peridaedala Meyrick and Assulella Kuznetzov to the Acroclita-group, probably close to Melanodaedala, gen. nov.
Acroclita-group Status remarks This is a new, informal genus-group including, in Australia, Melanodaedala, gen. nov., Rhopobota Lederer, Fibuloides Kuznetzov, Acroclita Lederer, Tritopterna Meyrick, Noduliferola Kuznetzov and Heleanna Clarke.
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Melanodaedala, gen. nov. (Figs 80, 667–672) Type species: Bathrotoma (?) scopulosana Meyrick, 1881, Proc. Linn. Soc. NSW 6: 677, hereby designated. [Australia.]
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Diagnosis Species of Melanodaedala are small to mediumsized, rather narrow-winged eucosmines with subrectangular greyish ochreous or greyish brown forewings with a simple but usually characteristic wing pattern of long costal strigulae, a brown apical spot, an indistinct ocelloid patch and a white basal line to the fringe, and on the dorsum usually with a dark suffusion along basal third ending in a narrow, erect, dark triangle edged with white scales on outer face and a low, dark semicircle edged with white scales before tornus and with scattered white scales between these two dark marks. The hindwings are dark grey and conspicuously thinly scaled and translucent between the densely scaled veins in the male, a diagnostic feature for males of Melanodaedala together with the black scaling on the forewing underside. The labial palpi are rather short and porrect with the second segment moderately widened and a short third segment with a diagnostic black sclerotisation at its tip in the male. All veins are separate in the forewing, though R4 and R5 and M3 and CuA1 are closely approximated, and M2 is bent close to the stalk of M3 and CuA1 in the hindwing. The male genitalia of Melanodaedala are diagnostic with their large, flat, strongly sclerotised uncus and socii, the prominent but separate gnathos arms with strongly sclerotised, band-shaped distal portion, and a valva with a narrow ‘neck’ carrying a group of blunt pegs on its inner surface and with a fan-shaped cucullus. The female genitalia (M. scopulosana only) are characterised by modified, narrow but swollen ovipositor lobes, the sterigma an hourglass-shaped, medially narrowed funnel with a prominent ventral lip and dorsally connected with a semicircular, spinulose lamella postvaginalis with two lateral scale patches, a short, lightly sclerotised ductus bursae and a corpus bursae with fine spinules, including a conspicuous band around its entrance, and with two shortish, slender, horn-shaped signa.
Description Adults (Figs 667, 668) Wingspan. 10–15 mm; small to medium-sized. Head (Fig. 671). Ocellus well developed. Labial palpus porrect, sinuate, rather short (1.5–1.7× diameter of eye), second segment slightly sinuate, teardrop-shaped, gradually widening towards rounded apex, with long, loose scales ventrally and distally, terminal segment short, in male with a conspicuous black sclerotisation dorsally on apex. Antenna to beyond middle of wing, scaled in male
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except for a narrow, naked band with scattered, very short cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 80, 667, 668). Forewing index 0.33–0.36. Forewing elongate, subrectangular distally but narrowed towards base, widest before tornus, with costa lightly curved throughout but especially at base, apex slightly projecting, termen sinuate, little oblique; without costal fold; with chorda from below R2 forming a very narrow accessory cell and M-stem to between M2 and M3, distally very close to posterior margin of cell; all veins separate with basal portion of R4 and R5 close and parallel from angle of cell and M2, M3 and CuA1 all three close and about equidistant at base with M3 and CuA1 close and parallel for some distance; CuP present at margin; anal loop 0.33× length of 1A+2A; greyish ochreous to greyish brown finely speckled with pale-tipped scales throughout, with long, very oblique pale grey, brown, leaden and blackish brown costal strigulae, a round, brown, apical spot and an indistinct ocelloid patch containing some fine dark brown lines and divided from tornus by a band containing scattered leaden scales; fringe conspicuously whitish at base along entire termen but distally blackish brown in costal half; dorsum usually with characteristic markings: a dark brown, low semicircle just before tornus and an irregular dark mark along basal third ending in a more distinct, erect, outwardly oblique brown triangle with its outer face edged with white scales continuing along dorsum and around dark semicircle before tornus; sometimes with a conspicuous patch of whitish suffusion between two dark marks on dorsum; lower surface with black scales in male. Hindwing with moderate cubital pecten and axillary tuft, wing especially in male very thinly scaled and partially translucent between veins, with scattered long hairscales along anal margin in male; Rs and M1 closely approximated at base, base of M2 bent close to base of M3 and CuA1 that are stalked to nearly 56O; CuP present at margin; 1A+2A well developed distally but part of basal loop missing (base of 1A), 3A only a faint indication near margin; dark brown to blackish brown, thinly scaled and translucent between densely scaled, conspicuous veins. Pregenital abdomen. Unmodified. Hindmargin of S8 in male with a deep, semicircular emargination; T8 with rounded hindmargin. Male genitalia (Figs 669, 670). Tegumen high and robust, strongly sclerotised. Uncus a strongly sclerotised, flat, downcurved, more or less deeply
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bilobed process. Socii strongly sclerotised and rigidly projecting, elongate, slightly sinuate, bandshaped, lateral, flat lobes with very few bristles on outer surface. Gnathos well developed, with strongly sclerotised, band-shaped, lightly twisted distal portion, porrect or rising. Valva very long
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and slender except for greatly widened, fan-shaped cucullus; basal part of valva with a dense tuft of bristles from outer surface near ventral margin and with a large, sinuate costal process; sacculus with a row of single bristles between extremely large basal excavation and ventral margin and a large,
Figs 667–672. Melanodaedala scopulosana, adults, genitalia and head. 667, 668, adults. 667, , Upper Allyn R., NSW. 668, , Dorrigo, NSW. 669, 670, male genitalia and aedeagus, nr Hope Vale Mission, slide T2222. 671, head, male. 672, female genitalia, Mt Keira, NSW, slide ANIC 13750. Scale = 2 mm.
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triangular tooth from its angle; ‘neck’ of valva long and slender with a row or patch of conspicuous, blunt pegs on inner surface; cucullus bristled to spined on inner surface and with a few long, flattened bristles ventrally on outer surface. Juxta small, triangular; caulis moderately long; anellus a small cup with small ventral angles. Aedeagus rather slender, curved in M. scopulosana; vesica with a moderate bundle of flattened cornuti. Female genitalia (Figs 672) (M. scopulosana). Ovipositor lobes modified, very narrow but swollen, nearly round in cross section. Hindmargin of S7 concave and fused with sterigma, laterally produced into a low crest. Sterigma a large, medially constricted, hourglass-shaped funnel leading to ostium, with curved and projecting ventral lip and dorsally fused with a spinulose, lightly sclerotised, inverted semicircular lamella postvaginalis with two patches of scale sockets on each side; anteriormost portion of hourglassshaped funnel membranous, followed by narrow, weakly sclerotised colliculum and wider, sclerotised and lightly twisted ductus bursae with ductus seminalis originating somewhat below colliculum. Corpus bursae ovate with fine spinules except near the two slender, shortish, horn-shaped signa, with a ring of larger spinules near entrance to corpus bursae.
Distribution Melanodaedala is known from India, Myanmar, Thailand (Kawabe 1989), Vietnam (Kuznetzov 1988a), Japan (Kawabe 1974), the Solomon Is, Papua New Guinea (ANIC material) and Australia where it occurs along the eastern coast from Wollongong in New South Wales to Heathlands on Cape York Peninsula in northern Queensland.
Biology Fletcher ([1921]) described the biology of Melanodaedala melanoneura (Meyrick) in India in detail: spinning together the flowers of Rhus semialata (Anacardiaceae), and feeding within a silken tube. Most Australian specimens have been collected in rainforest localities and a few in riverine vegetation in drier regions.
Remarks The following two non-Australian species are here included in Melanodaedala:
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diffusa (Bradley, 1957), Nat. Hist. Rennell Island, Br. Solomon Islands 2(19): 89 (Allohermenias). [Rennell I.] Comb. nov. melanoneura (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 866 (Eucosma). [India.] Comb. nov. The three species upon which Melanodaedala is based are very characteristic and also remarkably similar in genitalia as well as in external appearance, and their generic identity has clearly puzzled earlier authors. Meyrick described M. scopulosana as ?Bathrotoma, obviously unconvinced but relying on similarities of certain wing pattern elements. Bradley (1957) referred M. diffusa to Allohermenias Diakonoff presumably on the strength of a similarly shaped valva that he discussed in detail, though M. diffusa lacks a costal fold and has R4 and R5 separate unlike Allohermenias. Details of the genitalia in both sexes of Melanodaedala preclude a close relationship with the Hermenias-group. Melanodaedala melanoneura was initially described as an Eucosma species and the Indian lectotype was figured by Clarke (1958). Kuznetzov (1988a), figuring a specimen from Hanoi, referred the species to Megaherpystis Diakonoff from the Seychelles (Diakonoff 1969). Megaherpystis belongs to the Acroclita-group and has similar valva and sclerotised socii but very different wing and labial palpus shape, and lacks the enlarged gnathos arms and several other Melanodaedala apomorphies. The monophyly of Melanodaedala is based on several conspicuous apomorphies: males with black sclerotisation dorsally on apex of third labial palpus segment, hindwings thinly scaled and translucent between densely scaled veins and forewing underside with black scales; socii rigid, flat and sclerotised, gnathos arms medially not connected but with strongly sclerotised, porrect or rising, band-like distal portion, basal part of valva with a tuft of bristles ventrally from outer surface and a row of bristles below basal excavation, ‘neck’ of valva long and slender and with a row or patch of blunt pegs, and cucullus fan-shaped with several flattened bristles from outer surface. Based only on M. scopulosana no generic apomorphies can be identified for the female. The wing pattern is similar in all known species of Melanodaedala and may well be apomorphic to some extent. Peridaedala Meyrick and Assulella Kuznetzov are closely related, both genera sharing apomorphies with Melanodaedala. All three genera have a robust, strongly sclerotised tegumen with a wide, flat uncus and modified, distally strongly sclerotised, porrect or rising gnathos arms. Assulella also shares a
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similar valva and absence of a costal fold with Melanodaedala but it has small, hairy socii, the gnathos arms medially connected, no tuft of bristles from the outer surface of the valva near its base and no pegs on the ‘neck’ of the valva, and M3 and CuA1 in the hindwing connate only (Kuznetzov 1973; Diakonoff 1983a). Assulella females have a modified S7, a projecting sterigma and a very long and thin ductus bursae, totally unlike M. scopulosana. Peridaedala shares flat, sclerotised socii with Melanodaedala but it is distinguished by greatly enlarged and medially fused gnathos arms, a costal fold, stalked R4 and R5 and a notched antenna in the male (Diakonoff 1953), and has M3 and CuA1 in the hindwing connate only.
Constituent species Two species from outside Australia are above referred to Melanodaedala, and the South African Eucosma tumulata Meyrick could well be congeneric but needs further study. M. melanoneura has been reported from India to Japan, but given the similar habitus of the Melanodaedala species known to date the identity of the species throughout this range will have to be confirmed. One named species is known from Australia: scopulosana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 677 (?Bathrotoma). [NSW: Sydney, Parramatta.] Comb. nov. pachyneura (Turner, 1916), Trans. R. Soc. S. Aust. 40: 525 (Eucosma). [Qld: Sandgate.]
Rhopobota Lederer (Figs 81, 673–683) Rhopobota Lederer, 1859 April, Wien. Entomol. Monatschr. 3: 124 (key). Nomenclaturally available but without included nominal species until Lederer, 1859 December, ibid. 3: 366. Type species: Tortrix naevana Hübner, [1817], Samml. Eur. Schmett. 7: pl. 41, fig. 261, by subsequent monotypy. [Europe.] Erinaea Meyrick, 1907c, J. Bombay Nat. Hist. Soc. 18: 141. Type species: Erinaea chlorantha Meyrick, 1907c, ibid. 18: 141, by monotypy. [Sri Lanka.] Norma Heinrich, 1923, US Nat. Mus. Bull. 123: 12 (key), 191. Type species: Epinotia dietziana
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Kearfott, 1907, Trans. Am. Entomol. Soc. 33: 92, by original designation. [USA.] Kundrya Heinrich, 1923, US Nat. Mus. Bull. 123: 11 (key), 192. Type species: Kundrya finitimana Heinrich, 1923, ibid. 123: 192, figs 8, 34, 415, by original designation. [USA.] Eumarissa Clarke, 1976, Insects Micronesia 9: 10 (key), 32. Type species: Eumarissa leucognoma Clarke, 1976, ibid. 9: 32, text-fig. 10, pl. 2, fig. g, by original designation. [Guam.]
Diagnosis Rhopobota species are small to large moths with rather narrow, subovate forewings, nearly always without costal fold (absent in all Australian species), with a projecting apex containing a blackish spot and contrasting and well-developed costal strigulae. The wing pattern is usually characteristic, with paler dorsal and terminal area and an oblique dark band from apex to near middle of dorsum where it is angled and runs to the base of the wing, often extended to the basal part of the costa. This band may be reduced to the two well-defined, blackish, lightly curved streaks that form its distal half. The hindwing in the male usually has smaller, modified scales in its central part and below a band of modified, blackish scales along the anterior margin of the cell, a combination that, together with the characteristic wing pattern, is diagnostic for the genus. Rhopobota has sinuate, porrect, moderately widened labial palpi. The wing venation is diverse, usually without an accessory cell but with the Mstem present in the forewing, with M4 and M5 usually stalked (all Australian species) but rarely only approximated or fused into a single vein, with M1, M2 and M3 usually roughly equidistant at base but M3 distally converging with M2, with M3 fairly close to CuA1 at base. In the hindwing Rs and M1 are parallel and approximated at base, M3 and CuA1 are stalked to 56O or more and CuA2 originates from not far beyond the middle of the cell. In the male genitalia the widely spaced, lightly sclerotised uncus processes, the very long band-like socii with densely hairy apical lobes that are either medially paired or bent upward on long stalks and closely associated with the gnathos, a slender valva with a widened cucullus without any ‘neck’ but nearly always with a conspicuous finger- to bandshaped process distally from the ventral margin of the basal excavation, and with a modified, upwardly pointing costal process are diagnostic for Rhopobota. In the female genitalia the sterigma is well-
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sclerotised and diverse, either fused with S7 or fitted into a deep emargination in its hindmargin. The ductus bursae with a folded sclerite below the ostium and a bipartite sclerite at the entrance to the corpus bursae consisting of two longitudinal sclerotised bands on either side reaching deep into the corpus bursa and in the bursa ‘neck’ connected by a transverse, arch-shaped sclerite are diagnostic for Rhopobota, either in this typical configuration or in variably reduced form, together with two curved, flattened, horn-shaped signa of usually unequal size, sometimes reduced to one signum or two small tubercles (R. hortaria (Meyrick)) only.
Description Adults (Figs 673–676) Wingspan. 10–20 mm (Australian species); small to large. Head (Fig. 679). Ocellus well developed. Scales of upper part of frons long, anteriorly projecting, remainder of frons with minute, appressed scales. Proboscis about as long as labial palpus. Labial palpus porrect, sinuate, moderately long (1.8–2.0× diameter of eye), strongly widened beyond middle, second segment subtriangular with long, loose scales along ventral and anterior third of dorsal margin, terminal segment moderately long. Antenna to just beyond middle of wing, entirely scaled, no cilia visible. Thorax. Smooth. Legs unmodified. Wings (Figs 81, 673–676). Forewing index 0.37–0.40. Forewing elongate-subovate except for projecting apex, widest in middle or before tornus, costa evenly curved, apex acute, projecting, termen strongly sinuate, vertical to little oblique; rarely with costal fold (absent in Australian species); discal cell with chorda usually absent and M-stem usually to base of M3; R4 and R5 usually stalked (all Australian species), seldom only approximated (Norma) and very rarely fused into one vein (Kundrya, Erinaea); M1, M2 and M3 usually roughly equidistant at base but M3 distally converging with M2, M3 fairly close to CuA1 at base and the two veins either parallel or distally diverging; CuP well developed near margin; anal loop 0.31× length of 1A+2A; with diverse wing pattern with paler dorsal and terminal areas and nearly always with a variably expressed oblique dark band from apex to above middle of dorsum or to dorsum proper just before middle and then angled and running to wing base, often widened at base and including basal part of costa; with welldeveloped, contrasting strigulae, a blackish spot in
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apex and often with two well-defined, blackish, lightly curved streaks forming distal half of dark band; ocelloid patch indicated by two parallel, vertical bands of glossy scales. Hindwing with cubital pecten present and axillary tuft well developed, males often with modified, small, widely spaced scales in central part of wing and underside with a band of dense, modified, blackish scales along anterior margin of cell; bases of Sc+R1 and Rs converging and touching near base; Rs and M1 parallel and approximated at base; base of M2 curved to close to stalk of M3 and CuA1 that are stalked to about 56O or more; CuA2 from not far beyond middle of cell; CuP a very weak trace at wing margin only, 1A+2A and 3A well developed; usually shades of greyish brown, scaling sexually dimorphic. Pregenital abdomen. Usually unmodified, but Rhopobota clivosa (Meyrick) with a tuft of large, modified scales at base of abdomen (Clarke 1958). In male hindmargin of S8 with deep medial emargination, medially strongly projecting lobe in T8. Male genitalia (Figs 677, 678, 680, 683). Tegumen robust, well sclerotised, ovate to subrectangular, usually with a weakly bilobed apex. Uncus arms widely separate, weakly sclerotised, one on each apical lobe, varying from short, finger-like to large, inflated, irregular structures. Socii long membranous bands laterally from below apex with densely hairy rounded tips paired medioventrally in less derived condition (R. hortaria), with gnathos two lateral bands medially fused, parallel and apparently connected with socii; in more derived state socii arms increasingly slender and elongate and apparently fused with gnathos arms to form sclerotised rods carrying paddle-shaped, densely hairy tips, bent upwards in slide preparation. Valva slender except for strongly widened apical third but without a ‘neck’, nearly always with a long digit- to band-shaped process from basal excavation or just beyond (all Australian species), and with a conspicuously modified, upwardly pointing costal process; cucullus varying from a slightly enlarged, rounded apex to a transverse, often crescentic structure with projecting ventral and/or dorsal lobe, sometimes with prominent spines from dorsal angle; in some species with flattened bristles from outer surface of valva. Juxta large, caulis moderately long, anellus large. Aedeagus small, usually tapering towards apex; vesica with a few or a sheaf of lanceolate cornuti. Female genitalia (Figs 681, 682). Ovipositor lobes unmodified. Sterigma fitted into deep
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emargination in hindmargin of S7 or entirely fused with S7, very diverse but always well sclerotised; in Australian species either a complex invertedsemicircular or rectangular sclerite with lamella antevaginalis protuberant and with two projecting angles at base (R. hortaria) or a flat, subtriangular, partially spinulose lamella postvaginalis joined to a projecting, ring-shaped lamella antevaginalis. Ductus bursae of variable length, usually gradually widening towards corpus bursae; membranous except for short, crumpled, sclerotised region between weakly developed ostium of thickened membrane and origin of ductus seminalis, and for a variably developed prong-shaped sclerite at entrance to corpus bursae, ranging from two longitudinal bands of sclerotisation along either side of bursa neck reaching to middle of corpus bursae and connected
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in bursa neck by a transverse, arch-shaped sclerite in typical configuration, to variably reduced condition with only two sclerotised bands on either side (R. hortaria). Corpus bursae ovate, with two curved, flattened, horn-shaped signa of usually unequal size, sometimes reduced to two small tubercles (R. hortaria).
Distribution Rhopobota is most diverse in the Oriental and Palaearctic regions but the genus extends into the Nearctic and even Neotropic regions and is present in New Guinea and Australia where it occurs along the eastern coast from Cooktown in northern Queensland to near Dungog north of Newcastle in New South Wales. A single specimen of an
Figs 673–676. Rhopobota, adults. 673, 674, R. hortaria. 673, , Bunya Mts, Q [image reversed]. 674, , Elizabeth Beach, NWS. 675, 676, R. sp. 675, , Mission Beach, Q. 676, , Russel R. Palm Swamp, Q [image reversed]. Scale = 2 mm.
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unnamed species is known from the Northern Territory.
Biology Rhopobota larvae are leafrollers on a wide range of plants, with several species known to be polyphagous. Brown (1983) provided a checklist with known host plants including several genera of the Ericaceae and Ilex spp. in the Aquifoliaceae for two species in North America and one on the Kuril Is, but also Berberidaceae, Cornaceae, Fagaceae, Rosaceae, Rhamnaceae and Dipsacaceae. Rhopobota hortaria has been reared by I. F. B. Common and J. S. Dugdale from a larva joining leaves of Rhodosphaera rhodanthema (Anacardiaceae) in Cooyar, Queensland, and from joined leaves of Polyscias elegans (Araliaceae) by J. Stockard at Wingham, New South Wales. All Australian species have been collected at rainforest or monsoon forest localities.
Remarks Rhopobota is a large, widely distributed and diverse genus, reflected in the four generic synonyms. There are numerous illustrated treatments mainly of subsets of the genus (Heinrich 1923; Hannemann 1961; Bentinck and Diakonoff 1968; Clarke 1976; Miller 1987; Razowski 1989) and many species are figured by Clarke (1958) as species of Acroclita Lederer, but also as Erinaea. Brown (1979, 1983) studied the group in a world-wide context and on the basis of convincing genitalic apomorphies synonymised four genera that had been largely based on venational differences. The North American Norma and Kundrya, both with larvae feeding on Ilex spp., lack the digit-shaped process from the inner valva surface according to Heinrich (1923), and whilst R4 and R5 are approximated in Norma they are fused into one vein in Kundrya. Erinaea has peculiarly modified forewing venation, figured by Clarke (1958), with either the fused R4 and R5 stalked with M1 or one vein missing. In his description of Erinaea chlorantha Meyrick (1907c: 141) says ‘Posterior tibia in with brush of dense long projecting scales on basal half beneath’. According to Clarke (1958) all the male syntypes are missing and Brown’s (1979) synonymy based on the female genitalia is for the time being accepted. Only males are available of the type species of Clarke’s (1976) Eumarissa and only females of his second species, thus further material will be needed to confirm that the sexes are correctly assigned and
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the characteristic sclerotisation at the bursa entrance is indeed absent in these species. The monophyly of Rhopobota is supported by several unique apomorphies, in particular the shape of the bipartite sclerite at the entrance to the corpus bursae in the female, and in the male the widely separate and weakly sclerotised uncus arms, the very long socii apparently associated with the gnathos and either ventrally paired or with narrow arms and upturned in slide preparations, the characteristic valva with a large finger- or bandshaped process from the ventral margin of the basal excavation (absent in two North American species) and a modified, dorsally projecting costal process. Rhopobota is highly derived, but several potential apomorphies link it to the Acroclita-group of genera, with the prong-shaped sclerite in the bursa neck the most conspicuous one. The two groups also share a keel- or blade-like process from beyond the basal excavation, flattened bristles from the outer valva surface (present only in some Rhopobota species) and a similar sterigma derived from a dorsal spinulose plate and a ventral raised rim. There is some uncertainty whether the dorsal teguminal processes of Rhopobota are homologous with the bifid uncus in Epinotia (Brown 1983), but until convincing evidence is produced I follow Heinrich (1923) and treat them as uncus arms in contrast to Razowski’s (1989) interpretation of ‘no trace of uncus but with a pair of sublateral processes’. The label on the holotype of Acroclita hortaria says ‘Victoria TPL.90’, and the description states ‘Vic.: Locality not recorded; one specimen’. Lucas lived in Melbourne but before 1890 had moved to Brisbane. There is no doubt about the identity of the female holotype, which has been dissected. The ANIC has no material of R. hortaria from south of Newcastle and as the species is quite common where it occurs and sufficiently attractive to be readily collected it is assumed that the holotype has been mislabelled.
Constituent species Brown (2005) records 42 species of Rhopobota worldwide, including two names for the one named Australian species. At least three unnamed species are known from Australia. hortaria (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 241 (Acroclita). [Vic.?] philobrya (Meyrick, 1920), Exotic Microlepid. 2: 344 (Acroclita). [Qld: Toowong.] Syn. nov.
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Figs 677–683. Rhopobota, genitalia and head. 677, 678, R. hortaria, male genitalia and aedeagus, Bunya Mts, Q, slide T2136. 679, head, R. hortaria, male. 680, R. sp., aedeagus, slide T2295. 681, R. hortaria, female genitalia, Bunya Mts, Q, slide T2137. 682, R. sp., female genitalia, Mt Bartle Frere, Q, slide ANIC 12967. 683, R. sp., male genitalia, Mission Beach, Q, slide T2295.
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Fibuloides Kuznetzov (Figs 82, 684–696) Fibuloides Kuznetzov, 1997b, Entomol. Obozr. 76(4): 810. Type species: Fibuloides modificana Kuznetzov, 1997b, ibid. 76(4): 810, fig. 17. [Vietnam.] Eucoenogenes auctt.
Diagnosis Fibuloides species are rather narrow-winged, brownish eucosmines with diverse wing pattern but always with costal strigulae ending in a prominent, V-shaped white preapical pair and with a line of blackish scales along the middle of the termen. Most Australian species have the costal half of the forewing dark brown in contrast to the paler dorsal half that is either ochreous or greenish whilst the males in several Oriental and Papuan species have the basal half of the wing covered with modified black scales with an iridescent blue sheen. Males usually have a notch at the base of the flagellum and in nearly all species modified, long and dense fringe along the anal margin of the hindwing or a pencil of long hairscales from its base together with transverse bands of modified scaling dorsally on the abdomen, a combination that is diagnostic for the genus if present. The labial palpi are long, sinuate and porrect with a widened second segment except in the male of few species (F. phycitipalpia, sp. nov.) with modified, slender, rising palpi closely appressed to frons. The forewing rarely has a costal fold and has either all veins separate or R4 and R5 stalked, even in very closely related species, and in the hindwing M3 and CuA1 are always and Rs and M1 are sometimes stalked. The male genitalia are characterised by a usually rather weakly sclerotised, subtriangular tegumen, by a well-developed, often bifid uncus, by drooping, membranous, densely bristled socii, by a diagnostic gnathos with weak, lateral arms usually from below middle of tegumen and medially angled to form two conspicuous, parallel, rising, long, sclerotised bands, by a medially constricted valva with a large basal excavation, large, flattened bristles from outer surface and a transversely ovate to bilobed cucullus sometimes ending in a ventral spine, by an often strongly modified anellus with projecting ventrolateral corners and a dorsal, tongue-shaped process parallel to the aedeagus, and by an often distally upcurved aedeagus with sinuate, lanceolate
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cornuti. The female genitalia are characterised by a rather simple sterigma, a membranous cup, either in a deep key-hole-shaped emargination of S7 or more anterior and entirely fused with S7 near its anterior margin, with the spinulose sclerotised lamella postvaginalis usually fused with S7 and S7 sometimes with teeth or a transverse ridge or a scaled medial hump anterior to sterigma. The very characteristic ductus bursae in the female is diagnostic for the genus, short and membranous except for two small, paired, lens-shaped sclerites (colliculum) just below the ostium and a large, bipartite sclerite in its anterior half with a prong on each side projecting towards the corpus bursae that has two large, horn-shaped signa.
Description Adults (Figs 684–687) Wingspan. 9–21 mm; small to large. Head (Figs 688–690). Ocellus well developed. Chaetosema often with very long bristles. Vertex with long, loosely appressed scales, anteriorly projecting between antennae and forming with scales of upper part of frons a large, obliquely flattened, forward projecting tuft (except in F. phycitipalpia, sp. nov., where the long scales on upper frons form a tuft rising parallel to tip of labial palpus), remainder of frons with minute, appressed scales. Proboscis about as long as labial palpus. Labial palpus except in F. phycitipalpia, sp. nov. moderately long to long (1.9–2.5× diameter of eye), porrect, strongly sinuate, with second segment sinuate and distally greatly widened by long, loose scales along entire ventral margin and distally along dorsal margin; terminal segment moderate to very long and slender; in male of F. phycitipalpia, sp. nov. labial palpus strongly modified, appressed to frons, long, slender and short-scaled, with second segment long, slender, nearly spindle-shaped, dorsoventrally compressed, rising vertically to level of vertex, with terminal segment minute, angled forwardly; in female of F. phycitipalpia, sp. nov. labial palpus porrect and unmodified as in rest of genus. Antenna to beyond middle of wing, flagellum in male with often inconspicuous dorsal notch at base, scaled except for a narrow ventral band with minute cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 82, 684–687). Forewing index 0.33–0.38. Forewing elongate subovate (F. phycitipalpia, sp. nov.) to elongate subrectangular, widest in middle or before tornus, with costa weakly to moderately curved
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throughout, apex weakly projecting, termen lightly sinuate, slightly oblique; discal cell with chorda or remnants close to anterior margin, M-stem a trace only; rarely with narrow costal fold to beyond
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middle (F. aestuosa (Meyrick), F. deltostoma (Diakonoff)), but usually with three tufts of raised scales on dorsal half of wing; rarely all veins separate with R4 and R5 closely approximated
Figs 684–690. Fibuloides, adults and heads. 684, 685, F. phycitipalpia, Cooloola, Q. 684, , holotype [image reversed]. 685, , paratype [image reversed]. 686, F. minuta, , paratype, Mt Bellenden-Ker, Q [image reversed]. 687, F. sp., , Moses Ck nr Mt Finnigan, Q. 688–690, heads. 688, F. phycitipalpia, male. 689, F. phycitipalpia, female. 690, F. minuta, male. Scale = 2 mm.
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Figs 691–696. Fibuloides, genitalia and abdomen. 691, F. phycitipalpia, paratype, Cooloola, Q, female genitalia, slide ANIC 4916. 692, 693, F. minuta, holotype, male genitalia and aedeagus, Mt Bellenden-Ker, Q, slide T2349. 694, F. phycitipalpia, abdomen, male (III–VI, segments 3–6). 695, F. phycitipalpia, holotype, male genitalia, Cooloola, Q, slide ANIC 4919. 696, F. minuta, paratype, female genitalia, nr Gympie, Q, slide T2405.
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(F. cyanopis (Meyrick)) to connate but usually R4 and R5 stalked (F. phycitipalpia, sp. nov.), often to beyond 56O, with base of R3 often close to stalk of R4 and R5; M2 much closer to M3 than to M1 with M2 and especially M3 and CuA1 rather close together from posterior angle of cell; CuP present near margin; anal loop 0.27× length of 1A+2A; wing pattern diverse with few obvious generic elements but always with costal strigulae ending in a prominent, V-shaped pair of white strigulae before apex and with a line of blackish scales along middle of termen; F. phycitipalpia, sp. nov. mottled grey and ochreous brown, dorsum somewhat darker in basal third, followed by a paler, outwardly oblique wide band over middle third and an indistinct, dark-edged triangular mark before tornus, the two other Australian species with costal half of wing dark brown in contrast to the paler dorsal half that is either ochreous or greenish. Hindwing with cubital pecten and anal tuft well developed, in male modified either with extremely long fringe along anal margin, particularly dense and erectile in F. phycitipalpia, sp. nov. or combined with a narrowed anal area in one unnamed Australian species, or with a large scale pencil from base of anal area in some species close to F. cyanopis; Rs and M1 closely approximated at base or stalked (F. phycitipalpia, sp. nov.), base of M2 curved towards base of stalk of M3 and CuA1; CuP present at margin, 1A+2A well developed, 3A present or absent; dark brownish grey, partly translucent due to minute scales in both sexes of several species. Pregenital abdomen. Male dorsally nearly always with transverse bands of modified scales across anterior part of two to five tergites, on T4–T6 in F. phycitipalpia, sp. nov. (Fig. 694), T4–T7 in an unidentified species from Papua New Guinea and T5–T8 in an unnamed Australian species with two kinds of modified scales: long and slender in a narrow anterior band and short and swollen on rest of tergite except for a narrow posterior band. Anterior margin of S2 deeply V-shaped between short apodemes and with a conspicuously wide rim, particularly at base of apodemes. In male hindmargin of S8 deeply bilobed with rounded medial excavation. Male genitalia (Figs 692, 693, 695). Tegumen usually moderately to weakly sclerotised and subtriangular, only in one Australian species strongly sclerotised and with pronounced ‘shoulders’. Uncus variable, a subrectangular flat lobe with two variably produced lateral points, or two long, diverging, paddle-shaped lobes fused at base, or a
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single, ventrally curved hook with a ventral keel to the widened apex. Socii moderately large, membranous, densely bristled, drooping lobes. Gnathos arms usually from below middle of tegumen, laterally weak but medial portion usually two parallel, rising, long, sclerotised bands. Valva very variable in detail, but always medially constricted, with a large basal excavation and with a line of very large, flattened bristles from outer surface of basal half of valva; cucullus from transversely ovate to irregularly bent to bilobed with ventral lobe sometimes ending in a large spine. Juxta triangular, caulis of variable length, anellus progressively modified with projecting ventrolateral corners and a dorsal, tongue-shaped process parallel to aedeagus, resulting in large, triangular structure several times the size of the aedeagus in some species (often ventrally displaced in dissections). Aedeagus from short to very long, with distal half weakly to strongly and abruptly upcurved (up to a right angle); vesica with a bundle of sinuate, lanceolate cornuti. Female genitalia (Figs 691, 696). Ovipositor lobes flat, with large, triangular posterior lobes. Eighth segment in some species dorsally strongly sclerotised. Sterigma in deep key-hole shaped emargination of S7 or entirely fused with S7 and situated near its anterior margin; sterigma a membranous cup fused with S7, lamella postvaginalis a sclerotised transverse band carrying spinules and some scales and increasingly fused with S7 as ostium moves anteriorly, sometimes with a transverse ridge or a curved line of teeth across S7 anterior to sterigma and/or with a medial hump covered with modified scales or a huge, three-lobed spinulose process just anterior to entrance to ostium. Ductus bursae distinctly shorter than corpus bursae, gradually widening towards corpus bursae, membranous except for two well-defined areas of sclerotisation, the first two small, paired, lens-shaped sclerites just below ostium, the second a large, bipartite sclerite in anterior half, with a prong on each side projecting to entrance of corpus bursae, with ductus seminalis originating from base of prongs. Corpus bursae ovate with two large, horn-shaped signa.
Distribution Fibuloides has been reported from Sri Lanka, India, Thailand, China, Japan, Indonesia, New Guinea and Australia, where it has been collected along the eastern coast from Elizabeth Beach just north of Newcastle in New South Wales to the McIlwraith Range in northern Queensland.
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Biology No host records are known for any Australian species, but all the localities are in at least marginal rainforest. Oku (1967) reported that larvae of F. aestuosa are of economical importance feeding in seeds of Castanea crenata (Fagaceae) in Hokkaido, Japan. Fibuloides japonica (Kawabe) has been reared from galls of Schlechtendahlia chinensis (Bell) (Aphididae) on Rhus javanica (Anacardiaceae) (Yoshiyasa 1986). An unidentified species of Eucoenogenes reported feeding on carambola in Taiwan (Ho 1985) is probably a species of Fibuloides.
Remarks A large number of non-Australian Fibuloides species have been described mostly as either Acroclita, Eucosma or Eucoenogenes Meyrick, with illustrations associated with the original description or in Clarke’s (1958) catalogue of Meyrick’s types in the BMNH. The following species are here transferred to Fibuloides, but the synonymies will need confirmation: aestuosa (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 854 (Spilonota). [India.] Comb. nov. ligyropis (Meyrick, 1937), in Caradja and Meyrick, Dt. Entomol. Z. Iris 51: 176 (Acroclita). [China.] biuncana (Kuznetzov, 1997b), Entomol. Obozr. 76(4): 805, figs 12, 13 (Peridaedala). [Vietnam.] Comb. nov. corinthia (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 858 (Acroclita). [Sri Lanka, India] Comb. nov. crocoptila (Diakonoff, [1968] 1967), US Nat. Mus. Bull. 257: 79, figs 110, 549 (Peridaedala). [Philippines.] Comb. nov. cyanopis (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 866 (Eucosma). [India.] Comb. nov. melanochlaena (Meyrick, 1936), Exot. Microlepid. 4: 611 (Eucosma). [Indonesia.] deltostoma Diakonoff, [1968], US Natn. Mus. Bull. 257: 82, figs 68, 102, 105, 551 (Eucoenogenes). [Philippines.] Comb. nov. euphlebia (Kawabe, 1989), Microlepid. Thailand 2: 53, figs 79, 117, 146, 147 (Eucoenogenes). [Thailand.] Comb. nov. japonica (Kawabe, 1978), Tinea 10: 185, figs 11, 25, 37 (Eucoenogenes). [Japan.] Comb. nov. levatana (Kuznetzov, 1997a), Entomol. Obozr. 76(1): 197, fig. 11 (Eucoenogenes). [Vietnam.] Comb. nov. metaspra (Diakonoff, 1983a), Zool. Verh. (Leiden) 204: 31, fig. 20, pl. 6 fig. 19 (Hermenias). [Sumatra.] Comb. nov.
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macrosaris (Meyrick, 1938), in Caradja and Meyrick, Deuts. Entomol. Z. Iris 52: 84 (Eucosma). [Java.] Comb. nov. munda (Diakonoff,1983a), Zool. Verh. (Leiden) 204: 40, fig. 23a, pl. 5 fig. 12 (Epinotia (Asthenia)). [Sumatra.] Comb. nov. neaera (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 859 (Acroclita). [Sri Lanka.] Comb. nov. nigrovenana (Kuznetzov, 1988a), Trudy Zool. Inst. Leningr. 176: 88 (Acroclita). [Vietnam.] Comb. nov. pythonias (Meyrick, 1910), Trans. Entomol. Soc. Lond. 1910: 434 (Rhopobota). [Indonesia.] Comb. nov. segregana (Kuznetzov, 1997b), Ent Obozr. 76(4): 804, fig. 10 (Eucoenogenes). [Vietnam.] Comb. nov. thysanota (Meyrick, 1912b), J. Bombay Nat. Hist. Soc. 21: 860 (Acroclita). [India.] Comb. nov. Diakonoff ([1968]) associated his Philippine F. deltostoma with Eucoenogenes on the basis of labial palpus shape though he was well aware of the tenuous nature of this assignment given that he had only a single male of F. deltostoma to compare with the single female holotype of the Indian Eucoenogenes melanancalis (Meyrick). Subsequently, several Oriental and Japanese species have been treated as Eucoenogenes (Kuznetzov 1976; Kawabe 1978; Kawabe 1982; Razowski 1989), some of them obviously closely related to F. deltostoma. However, the female genitalia of E. melanancalis differ too much from the very consistent and derived genitalia of all these species to be treated as congeneric. The genus Fibuloides (Kuznetzov 1997b) has recently been established for a Vietnamese species of this group. The anterior margin of S7 in E. melanancalis ends in a conspicuous rim and the ductus bursae, albeit weakly sclerotised, lacks the two-pronged sclerite diagnostic for Fibuloides species. Furthermore, E. melanancalis has been ‘bred from larvae mining leaves of Eugenia jambolana’ (Clarke 1958) whilst known Fibuloides larvae are borers in Castanea and other fruit (see Biology). The monophyly of Fibuloides is supported by a number of conspicuous apomorphies including a notch at the base of the male antennae, the angled, modified gnathos arms with their distal portion forming paired, sclerotised, rising bands, a valva with a very large basal excavation and flattened bristles from its outer surface, the characteristic spinulose sterigma and the two-pronged anterior sclerite in the ductus bursae. In the majority of species the anal margin of the male hindwing
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carries a long, modified fringe associated with transverse bands of modified scales dorsally on the abdomen. The presence of flattened bristles from the outer surface of the valva that Fibuloides shares with Tritopterna and Acroclita is a synapomorphy linking the three genera. A group of species including ‘Epiblema’ ancyrota Meyrick, 1907a, referred to Eucoenogenes by Kawabe (1982) but lacking the crucial Fibuloides apomorphies, nevertheless shares not only these flattened bristles but also a similar venation with Fibuloides and could possibly be the sister group. Rhopalovalva Kuznetzov also is a close relative, but its uncus is not bifurcate and it lacks the bi-pronged sclerite anteriorly in the ductus bursae. The male genitalia of Rhopaltriplasia Diakonoff, described as a genus of the Olethreutini (Diakonoff 1973), show considerable similarities to those of Fibuloides, but no females were available for the original description. Two species subsequently included in the genus, Rhopaltriplasia macrorhis Diakonoff from Sumatra (Diakonoff 1983a) with a notched antenna and Rhopaltriplasia insignata Kuznetzov from Vietnam (Kuznetzov 1997b) with Acroclitagroup female genitalia, undoubtedly are close relatives of Fibuloides. However, given the single uncus and coremata of the two earlier described species, it is not a foregone conclusion that they all are congeneric. Fibuloides has a surprising number of secondary sexual modifications in the male ranging from the notched antenna to a costal fold in some few species to the appressed palpi in F. phycitipalpia, sp. nov. and to the modified fringe on the anal margin of the hindwing associated with modified scaling dorsally on the abdomen. Interestingly, F. aestuosa and F. deltostoma, the two species with a costal fold, seem to lack the modified anal fringe and abdominal scales. The notched antenna is shared with Peridaedala Meyrick within the Acroclitagroup but is otherwise characteristic for the Spilonota-group. The description of Acroclita catharoptis Lower appears to be based on a mixed series including a species of Fibuloides closely related to F. phycitipalpia, sp. nov. and a Tritopterna species with the conspicuously snow-white inner surface to its labial palpus. The original description refers to five specimens from Kuranda and Cooktown but only one possible syntype has been found in the South Australian Museum, a greatly damaged male without the tip of the abdomen, labelled ‘Kuranda’ but without a label indicating that it originated from the Lower collection. This specimen is closely
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related to F. phycitipalpia with similar wing pattern and exactly the same modified labial palpi, but clearly not conspecific given the quite different wing shape, a more ochreous colouring and paler anal fringe. Given that it is not even certain that this specimen did indeed belong to the type series it serves taxonomic stability best to treat F. catharoptis as a junior synonym of the widely distributed Tritopterna capyra (Meyrick) on the basis of the unmistakable labial palpi mentioned in the description. This means that the Kuranda specimen of F. catharoptis is being treated as a possible paralectotype and not as the lectotype.
Constituent species Seventeen non-Australian species are assigned above to Fibuloides. The synonymies established by Clarke (1958) and the identity of some of the more widely distributed taxa will require confirmation. Eucoenogenes telifera Chrétien lacks several of the generic apomorphies and is not included in Fibuloides. Four species are known from Australia, two described below, a third represented by the damaged paralectotype of F. catharoptis and a fourth remains unnamed. phycitipalpia, sp. nov. minuta, sp. nov.
Fibuloides phycitipalpia, sp. nov. (Figs 684, 685, 688, 689, 691, 694, 695)
Material examined Holotype. , ‘26.01S 153.05E, Broutha Scrub, Cooloola, Qld [Queensland], 31 Aug 1978, I.F.B. Common’, ‘ANIC genitalia slide No. 4919 ’; (ANIC). Paratypes. Queensland: 4, same label data as holotype, ANIC genitalia slide No. 4916 (ANIC, QM).
Diagnosis The modified male labial palpi of F. phycitipalpia, slender, vertically rising and appressed to the frons, are unique and diagnostic for the species. Externally, F. phycitipalpia is characterised by R3 closely approximated to the stalk of R4 and R5 in the forewing, by Rs and M1 stalked and M2 at base bent close to the stalk of M3 and CuA1 in the hindwing, and by its greyish ochreous brown colouring with typically eucosmine pattern, with the pale rhomboid patch over the middle third of the dorsum, edged by a dark line proximally and
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distally, the most prominent feature. The male has a dorsal notch right at the base of the antenna, long hairscales from the hindwing anal margin and transverse bands of modified scales on T4–T6. The male genitalia of F. phycitipalpia are characterised by a subrectangular, flat uncus with triangularly produced corners, slender and widely distant rising gnathos arms, a sacculus with a distal, triangular, bristled projection with a few short spines, a transversely triangular cucullus with both lobes rounded and ventral one with short spines along its margin, with flattened scales from outer valva surface in a more basal clump followed by an evenly spaced series, and by a long aedeagus with scobinations on ventral surface. The diagnostic features of the female genitalia are a strongly sclerotised T8, S7 with a transversely sinuate furrow partially edged with keel-shaped crenulations and with a key-hole-shaped emargination in hindmargin fused with the spinulose lamella postvaginalis, a well-defined prong-shaped sclerite with arms tapering to points, and short, slender horns as signa.
Description Male (Fig. 684) Wingspan 15 mm. Head modified, long scales on upper part as two vertically rising appressed tufts in front of antennal base, brownish ochreous touched with dark grey on tip of labial palpus and across vertex; ocellus well developed, chaetosema normal; labial palpi modified and short-scaled, angled at base of second segment, vertically rising to beyond top of head, appressed to frons, long second segment slender, narrowing towards tip, terminal segment minute, angled forward (Fig. 688); antennae with small deep notch dorsally close to flagellum base, scaled except for narrow anterior band, cilia minute. Thorax brownish ochreous except for blackish brown metathorax with scattered ochreous scales. Forewing with R3 at base close to stalk of R4 and R5 (stalked to 56:); greyish ochreous brown, with intricate typically eucosmine pattern of white, dark brown and brownish ochreous pale-tipped scales: with long red-brown, white and grey strigulae along costa, red-brown in basal third followed by two pairs of white ones, and from beyond middle to apex with a series of increasingly shorter and less oblique red-brown and silvery grey lines, interspersed with five pairs of short white strigulae, with the more apical grey lines curved and confluent, running parallel to termen to above tornus; with a narrow blackish brown line along anterior 36: of termen interrupted by a white dot below apex; basal third of dorsum
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with a band of white-tipped grey scales followed on central third by a large outwardly oblique rhomboid patch of similar but more whitish scales bordered proximally and distally by an oblique dark brown line, with the proximal one variably darkened to a blotch and the distal one recurved and running to tornus, forming an inverted V-shaped mark; central part of wing from base to ocelloid patch of mixed grey and brownish ochreous pale-tipped scales with a few short brown lines; ocelloid patch indicated by short, obliquely transverse dash and a dot below. Fringe concolorous with adjacent wing. Hindwing with Rs and M1 stalked, M2 at base close to stalk of M3 and CuA1 (stalked to middle); with dense fringe of long ochreous hairscales from anal margin; dark brown with coppery gloss, fringe paler. Underside of wings with blackish brown scales variably covered by ochreous scales. Abdomen blackish brown overlaid with ochreous scales; with transverse band of modified scales anteriorly on T4–T6 (Fig. 694). Male genitalia (Fig. 695). Tegumen subtriangular, weakly sclerotised. Uncus a subrectangular flat lobe with triangular lateral points. Socii densely bristled, teardrop-shaped lobes. Rising gnathos arms narrow, weakly sclerotised. Valva very long and slender with distal portion of sacculus a bristled triangular projection with a group of short thorns, cucullus ‘neck’ slender and curved, cucullus bilobed, transversely triangular with short spines along outer margin near ventral angle; with a recurved blade from lower margin of basal excavation and a series of flattened bristles from outer surface of sacculus, a dense tuft just beyond basal angle followed by five larger and evenly spaced ones. Juxta triangular, caulis rather short, anellus small with basal angles slightly projecting. Aedeagus large and long, upwardly angled, slightly narrowing towards apex, with few scobinations on ventral surface; no trace of cornuti. Female (Fig. 685) Wingspan 15.5–16.5 mm. As for male except for secondary sexual modifications: labial palpus long, sinuate, porrect, second segment triangular, distally greatly widened, third segment long and slender (Fig. 689); metathorax and abdomen as rest of thorax; hindwing brownish grey with concolorous fringe. Female genitalia (Fig. 691). Tergum 8 simple but strongly sclerotised. Sternum 7 strongly sculptured, with a strongly sinuate deep transverse furrow below hindmargin and ostium, edged by some keel-shaped crenulations on both sides; hindmargin with a deep, key-hole shaped
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emargination with ostium in centre of S7. Sterigma a spinulose, hour-glass shaped plate behind ostium, fused into emargination of S7, with a crenulate hindmargin. Ductus bursae widening towards corpus bursae, less than 56O its length; colliculum two semicircular lateral sclerites; prong-shaped sclerite well-defined and strongly sclerotised with arms tapering and tip reaching beyond bursa neck. Corpus bursae ovate, weakly spinulose and less so at anterior end and around signa; signa two short, slender, smooth, straight horns with hollow base.
Distribution and biology The species is only known from the Broutha Scrub, Cooloola, Queensland.
Remarks According to male genitalic structure, F. segregana (Kuznetzov) from Vietnam is the most closely related species of apparently very similar appearance but with a costal fold and without modified labial palpi. The species name refers to the unusually modified male labial palpi.
Fibuloides minuta, sp. nov. (Figs 686, 690, 692, 693, 696)
Material examined Holotype. , ‘17.16S 145.52E, Cableway Tower 3, Mt Bellenden-Ker, Q[ueensland], 1020 m, 27 Oct 1981, E.D. Edwards’, ‘Genitalia slide No. T2349’ (ANIC). Paratypes. Queensland: 1, 13°44´S 143°20´E, Golden Nugget Ck camp site, McIlwraith Range, 520 m, 7. vii.1989, Nielsen, Edwards & Horak; 1 , Moses Ck, 4 km N by E of Mt Finnigan, 360 m, 14.x.1980, E. D. E.; 1 , 17°16´S 145°53´E, lower slopes Mt Bellenden-Ker, 500 m, 21.x.1981, E. D. E., ANIC genitalia slide No. 13762 ; 1 (Genitalia slide T2349), 1, same label data as holotype; 1 , 17°16´S 145°51´E, 0.5 km WSW of Mt Bellenden-Ker Centre Peak, 1500 m, 2.x.1981, E. D. E.; 1, 18 mls S of Gympie, 23.iv.1964, I. F. B. C. & M. S. U., Genitalia slide T2405. New South Wales: 2 , 3, 32°17´S 152°31´E, Elizabeth Beach, 27.iv.1979, I. F. B. C., ANIC genitalia slide No. 13760 (ANIC, AM, QM, BMNH).
Diagnosis The combination of the small size, the greenish forewings with R3 close to the stalk of R4 and R5 and the thinly scaled, partially translucent hindwings with M2 at base close to the stalk of M3 and CuA1 should identify F. minuta in the
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Australian fauna, especially the male with a shallow antennal notch and long hairscales from hindwing anal margin. The male genitalia of F. minuta are characterised by a strongly sclerotised tegumen with pronounced ‘shoulders’, a deeply bifid uncus with large diverging rabbit-ear-shaped lobes, a gnathos with medially parallel, slender rods, a valva with large basal excavation, a very long, slender, downcurved cucullus ‘neck’ with a small ovate cucullus ending in a drawn out apex with a single blunt spine, a row of flattened bristles from outer sacculus surface and a larger one from neck of cucullus, a moderately long, upwardly angled aedeagus with a bundle of cornuti. The diagnostic features of the female genitalia are S7 with a transverse ridge below the ostium and an U-shaped emargination in its hindmargin, a sterigma formed by a shallow cup and a band-shaped, spinulose lamella antevaginalis fused with S7, a prong-shaped sclerite with an ill-defined top and band-shaped arms, spinulose corpus bursae, and two long, slender, nearly straight horn-shaped signa.
Description Male (Fig. 686) Wingspan 9.5–11.0 mm. Head (Fig. 690) greenish brown; with long loose scaling; labial palpi long, sinuate, porrect, second segment distally strongly widened, terminal segment long, slender; antennae with small, rather shallow notch dorsally right at base. Thorax mixed greenish brown, legs cream with scattered blackish brown scales. Forewing with R3 close to stalk of R4 and R5 (stalked to 56:), with raised scales in dorsal half and distally along veins; greyish green with typical eucosmine pattern, white costal strigulae and darkened with blackish brown in costal half; ground colour in costal half dark brown sprinkled with brownish green, with two indistinct pairs of long whitish green strigulae from costa at 56: and four pairs of short white strigulae along distal half of costa, the last before blackish apex forming a conspicuous v-shaped white mark, their tips linked below costa by an irregular narrow band of grey and ochreous green; ground colour in dorsal half ochreous green with a dark brown line along middle of termen, a dark brown outline of an irregular triangle over end of dorsum. Hindwing with Rs and M1 adjacent, M2 at base close to stalk of M3 and CuA1 (stalked to 56:); hindwing scales modified except around margin and white scales along costa, dark brown, minute and widely spaced with translucent wing membrane visible, fringe concolourous with paler tips, along anal margin
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longer but not denser than elsewhere. Underside of wings ochreous overlaid with blackish scales, very densely on hindwing. Abdomen dark grey above and ochreous beneath, overlaid with blackish scales. Male genitalia (Fig. 692, 693). Tegumen subrectangular, with pronounced ‘shoulders’, strongly sclerotised. Uncus bifid nearly from base, two large, diverging rabbit-ear-shaped lobes with a notch in outer margin near apex. Socii densely bristled, teardrop-shaped lobes. Lateral part of gnathos arms broad, weakly sclerotised bands, rising portion two closely parallel, slender rods, outwardly bent near socii. Valva widest in basal part, with very large basal excavation with a recurved, large, projecting keel from its basal margin, with a row of very large flattened bristles from outer surface of sacculus and a single, larger one from neck of cucullus; cucullus ‘neck’ long, slender, downcurved; cucullus small, ovate, bristled on inner surface and ventrally ending in a drawn out apex with one (rarely two) short, blunt spine. Juxta smallish, caulis slender and moderately long, anellus small with basal angles slightly projecting. Aedeagus moderately long, upwardly angled, apical part narrower; with a bundle of lanceolate cornuti. Female Wingspan 10–12 mm. As for male except for more slender antennae without basal notch, paler hindwing scales and without blackish scales on abdomen; triangle on end of hindwing dorsum often not only an outline but entirely dark brown. Female genitalia (Fig. 696). T8 simple, moderately sclerotised. Sternum 7 with nearly straight transverse ridge below ostium and deep Ushaped emargination in hindmargin. Sterigma a wide shallow cup ventrally fused into emargination of S7, lamella postvaginalis a spinulose, sclerotised, broadly transverse band fused to S7 on both sides. Ductus bursae widening towards corpus bursae, only about 56O its length; colliculum two small lateral sclerites followed first by a short membranous portion and then a widening sclerotised region ending in two narrow, band-like prongs with their points in bursa neck. Corpus bursae ovate, spinulose but less so at anterior end and around signa; signa two long, slender, smooth, flattened, nearly straight horns with hollow base.
Distribution and biology The species is known from several rainforest locations along the eastern coast from the
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McIlwraith Range on Cape York Peninsula in Queensland to Elizabeth Beach on the central coast of New South Wales.
Remarks According to the genitalia of both sexes, the Vietnamese F. biuncana (Kuznetzov) is very closely related. Fibuloides biuncana was described as a Peridaedala species and differs by the presence of a costal fold and brownish grey rather than greenish colouration. The species name refers to its small size.
Acroclita Lederer (Figs 83, 697–702) Acroclita Lederer, 1859 April: Wien. Entomol. Monatschr. 3: 123 (key). Nomenclaturally available but without included nominal species until Lederer, 1859 November, ibid. 3: 329. Type species: Paedisca arctana Staudinger, 1859, Entomol. Ztg, Stettin 20: 232, by subsequent monotypy. [Spain.] Acrocita Turner, 1946, Trans. R. Soc. S. Aust. 70: 197. An incorrect subsequent spelling of Acroclita Lederer, 1859. Hylotropha Turner, 1946, Trans. R. Soc. S. Aust. 70: 197. Type species: Hylotropha leptotypa Turner, 1946, ibid. 70: 197, by monotypy. [Australia.] Syn. nov.
Diagnosis Acroclita bryopa Meyrick is a smallish, narrowwinged olethreutine with forewings mottled with bluish and olive greenish and blackish on whitish background, without costal fold but with two raised scale tufts on the dorsal half near middle, and with dark grey hindwings transparent between the scales. Acroclita has long, porrect labial palpi with the second segment strongly widened beyond middle and the third segment long and slender. R4 and R5 are stalked in the forewing and the base of R3 is closely approximated to their stalk. The hindwing has Rs and M1 short-stalked, M2 usually not close to M3 at base, with the position of M3 and CuA1 sexually dimorphic with M3 and CuA1 connate to stalked in females and M3 separate from CuA1, which arises from just below the angle of the cell in males. In A. bryopa, the base of M2 is much
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AUSTRALIAN OLETHREUTINE GENERA
closer to M3 in the female than in the male. The male hindwing of A. bryopa is modified with the costa deeply sinuate in its basal third and carrying a long, projecting hairpencil, with a row of long scales with modified sockets on the upper side within the discal cell parallel to the anterior margin, and with an elliptic depression containing distally curled, modified scales on the underside of the anal area. The male genitalia are characterised by a stout tegumen with lateral, usually deeply bifid, strongly sclerotised socii, a slender and medially constricted valva with a triangular to tooth-like process from ventral margin of basal excavation, flattened bristles from outer surface of valva and by a wide juxta, a short caulis and a small aedeagus. The female genitalia are characterised by a spinulose swelling in the lamella postvaginalis, a short, wide ductus bursae containing folded sclerites in its entire length, and two horn-shaped signa. In A. bryopa T8 is strongly sclerotised and rigid.
Description Adults (Figs 697, 698) Wingspan. 10.0–13.5 mm (Australian species); small to medium-sized. Head (Fig. 700). Ocellus moderate. Vertex with very long, loosely appressed scales, anteriorly projecting between antennae and forming with scales of upper part of frons a large, obliquely flattened, anteriorly projecting tuft, remainder of frons with small, appressed scales. Proboscis about as long as labial palpus. Labial palpus porrect, sinuate, long (2.1–2.2× diameter of eye), second segment sinuate, strongly widened beyond middle by long, loose scales forming broadly rounded apex, terminal segment very long and slender. Antenna reaching to beyond middle of wing, stout in male, scaled except for very narrow band, cilia not visible. Thorax. Smooth. Legs unmodified. Wings (Figs 83, 697, 698). Forewing index 0.34–0.39. Forewing elongate subrectangular, widest before middle, with costa lightly curved in basal half then nearly straight to pointed, protruding apex, termen oblique and sinuate, dorsum weakly curved throughout; discal cell in A. bryopa Meyrick gradually widening towards lightly oblique closing vein, with M-stem very faint, distally close to hindmargin of cell to M3, chorda close to anterior margin of cell, originating from below R2; R3 from just before angle of cell, at base parallel and close to long stalk of R4 and R5; M1, M2 and M3 about equidistant, M3 close to CuA1,
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with CuA1 from just below angle of cell; CuP well developed near margin; anal loop 0.30× length of 1A+2A in A. bryopa; A. bryopa with glossy fore wing scaling and two patches of raised scales on dorsal half near middle, with complex, mottled pattern of olive green, pale bluish green, creamy white and blackish brown. Hindwing in male of A. bryopa modified with deep, rounded indentation in basal half of costa giving rise to a conspicuous scale pencil, upper side with a row of long scales from modified sockets in discal cell parallel to anterior margin, and under side of anal area with an elliptic depression anteriorly deflecting the course of 1A+2A and filled with long, distally curved, pale ochreous scales and surrounded by modified, melanic scales, this depression often hidden beneath downwardly folded anal margin; venation in A. bryopa with Rs and M1 shortstalked, arrangement of M2, M3 and CuA1 sexually dimorphic with base of M2 bent close to short stalk of M3 and CuA1 in female, but in male with base of M2 distant from M3, and CuA1 from before angle of cell, separate from M3; CuP halfway between CuA2 and 1A+2A, 3A not clearly visible in male of A. bryopa; with small, blackish scales, membrane translucent between scales. Pregenital abdomen. Unremarkable. Hindmargin of S8 bilobed in male with rounded lobes and shallow, V-shaped medial emargination; T8 with weak, inverted Y-shaped sclerotisation and lightly convex hindmargin. Male genitalia (Figs 699, 701). Tegumen squat, broadly rounded. Acroclita subsequana HerrichSchäffer (=arctana) (Diakonoff and Wolff 1976) and Japanese species (Oku 1979) with top of tegumen two paired, hairy humps associated with a ventral, bilobed bristled process, A. bryopa with tegumen top flattened, lightly hairy, with a pair of small teeth ventrally below apex. Socii large, wellsclerotised, usually distally bifurcate structures, in A. subsequana arms naked and tapering, in A. bryopa more finger-shaped and apically hairy, especially ventral arm. Gnathos in A. subsequana only a membranous, transverse band, in A. bryopa much more developed with two dorsal, spinulose lobes. Valva long, slender, medially narrowed, with triangular or tooth-shaped projection on ventral margin of basal excavation and very small costal process; in A. subsequana cucullus ovate, more heavily bristled towards base, in A. bryopa cucullus ventrally angulate and with very large, spike-like spines, one on pointed lobe in centre of cucullus and two from margin below cucullus angle; ventral margin angled near base, with a tuft of flattened
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bristles from outer surface beyond base, and in A. bryopa with a small, triangular keel on ventral margin below ‘neck’, close to a very large, curved spike from outer surface of valva. Juxta wide, caulis very short, anellus in A. bryopa with pronounced ventrolateral angles. Aedeagus short, with bulbous base in A. bryopa; vesica with sheaf of distally sinuate, deciduous cornuti. Female genitalia (Fig. 702). Ovipositor lobes subreniform. In A. bryopa T8 rigid, strongly sclerotised, as posterior margin of T7. Hindmargin of S7 concave. Sterigma a spinulose, raised rim in A. subsequana with a small spinulose tumescence in lamella postvaginalis, in A. bryopa sterigma more complex with lamella antevaginalis a short spinulose tongue and lamella postvaginalis a round, spinulose hump with two smaller, lateral knobs anterior to a vertical, spinulose ridge on each side. Ductus bursae short and wide, narrowest below ostium, entire length with an irregular, longitudinally folded sclerite with ventral margins partially scalloped, ductus seminalis arising from near middle; corpus bursae ovate, with two horn-shaped signa.
Distribution Acroclita s. str. has only been confirmed from southern Europe, Madeira, Japan, Vietnam and Australia, where A. bryopa occurs on the eastern coast from near Dorrigo in northern New South Wales to the Cairns region in Queensland.
Biology Larvae of the European A. subsequana feed on leaves and seeds of Euphorbia spp. (Euphorbiaceae) (Bradley et al. 1979) and two Japanese species tightly spin young leaves of Elaeagnus umbellata (Elaeagnaceae) (Oku 1979); no host is known for the Australian A. bryopa but the species is restricted to rainforest localities.
Remarks Paedisca arctana Staudinger, a junior synonym of Acroclita subsequana (Herrich-Schäffer), is the type species of Acroclita by subsequent monotypy (Nye and Fletcher 1991). Descriptions and illustrations of the species can be found in Diakonoff and Wolff (1976), Kuznetzov (1978) and Razowski (1989, 2003a). Based on stalked R4 and R5 a large number of eucosmines have been referred to Acroclita by Meyrick and subsequent authors, but genitalic characters show that the genus has to be restricted to a few species only. It was a surprise to find an
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Acroclita s. str. in Australia, but the strikingly similar bifurcate processes laterally on the tegumen, together with the entirely congruent female genitalia provide convincing apomorphies to link A. bryopa with A. subsequana. Turner (1946) erected a new genus for Hylotropha leptotypa Turner, a junior synonym of A. bryopa, based on a single female. He misinterpreted the closely approximated R3 as stalked with R4 and R5, but considered Hylotropha to be a development of Acroclita. Acroclita bryopa shares conspicuous apomorphies with A. subsequana but has more derived male genitalia and remarkably specialised hindwings in the male with three different scale tufts and a modified venation with M2 widely separate from M3 at base. However, to treat Hylotropha as a separate genus would leave Acroclita as a paraphyletic group. The homology of the complex tegumen structures in Acroclita is not obvious and has been variously interpreted. The lateral, bifurcate processes have often been treated as gnathos arms (Diakonoff and Wolff 1976), but the gnathos is clearly present as a well-developed transverse band in A. bryopa and also in the Japanese species as remarked on by Oku (1979). Whilst Oku (1979) treats the medial hairy processes as socii and the lateral arms as additional projections from the tegumen the latter are here tentatively considered to be derived socii based on comparison between tegumen structures in Acroclita, Tritopterna Meyrick and Proteoteras Riley (Heinrich 1923). Acroclita has always been isolated within the Palaearctic fauna but there are links to the Oriental Tritopterna Meyrick and the New World Proteoteras Riley. Tritopterna shares the raised scale tufts on the fore wing, the squat tegumen, sclerotised socii, the short caulis, the close approximation of R3 to the stalk of R4 and R5 (connate in some Tritopterna species), and with A. bryopa the sexually dimorphic hindwing with modified scales along the anterior margin of the cell and the flattened bristles from the ventral margin of the outer surface of the valva. Acroclita subsequana and Tritopterna capyra (Meyrick) both use Euphorbiaceae as hosts. Proteoteras shares with A. bryopa the presence of flattened bristles from outer valva surface, shape of anellus with ventrolaterally produced angles and modified scales along hindwing costa in male. The holotype of A. leptotypa is labelled in Turners hand ‘TYPE Hylotropha leptotypa Turner’ and ‘Ebor, NSW, 14–12–39’, but in the description Mittagong is given as the type locality. However, Mittagong is far outside the known range of
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AUSTRALIAN OLETHREUTINE GENERA
A. bryopa whereas a second specimen is available from Ebor, close to other rainforest localities where A. bryopa is found. The type locality has been altogether omitted for two species in this very late paper by Turner (1946), so the contradictory type localities for A. leptotypa are probably due to an error in manuscript preparation.
337
Constituent species Very few species occurring in southern Europe, Madeira and Tenerife have currently been confirmed as Acroclita s. str. (Diakonoff and Wolff 1976), together with two Japanese species (Oku 1979), and one named species from Australia. Acroclita trachynota Meyrick from Borneo appears
Figs 697–702. Acroclita bryopa, adults, head and genitalia. 697, 698, adults, Mt Bartle Frere, Q. 697, [image reversed]. 698, . 699, male genitalia, Mt Bellenden-Ker, Q, slide T2211. 700, head, male. 701, aedeagus, slide T2211. 702, female genitalia, Mt Bellenden-Ker, Q, slide T2212. Scale = 2 mm.
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OLETHREUTINE MOTHS OF AUSTRALIA
to be an Acroclita s. str. according to external features and the genitalia of the single female holotype, but examination of a male is required to confirm this. bryopa Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 242 (Acroclita). [Qld: Mt Tamborine.] Comb. rev. leptotypa (Turner, 1946), Trans. R. Soc. S. Aust. 70: 197 (Hylotropha). [NSW: Ebor.]
Tritopterna Meyrick (Figs 86, 703–708) Tritopterna Meyrick, 1921, Zool. Meded. (Leiden) 6: 151. Type species: Tritopterna chionostoma Meyrick, 1921, ibid. 6: 152, by monotypy. [Java.]
Diagnosis Tritopterna species are small to medium-sized, narrow-winged olethreutines with variably patterned or mottled greyish to dark brownish forewings with several tufts of raised scales on dorsal half, with 4–6 conspicuous, paired, whitish costal strigulae, with blackish streaks along parts of M1 and M2 and often with a dark erect triangle on the dorsum before tornus, wings sometimes variably suffused with whitish from dorsum. The male has a projecting tuft of recurved scales from the base of the forewing dorsum. Tritopterna has large, sinuate, porrect labial palpi with the second segment strongly widened near the middle and triangular, the terminal segment long and slender with white inner surface. R4 and R5 are stalked in the forewing and the base of R3 is closely approximated to or connate to short-stalked with the stalk of R4 and R5. The hindwing has Rs and M1 closely approximated and M2 bent at base more or less close to M3, and it is sexually dimorphic with regard to M3 and CuA1 with the two veins connate to short-stalked in the female but clearly separate with CuA1 from before the angle of the cell in the male. Males of Tritopterna have a band of modified scales forming a pencil along the base of the anterior margin of the hind wing cell. The male genitalia are characterised by a squat, subrectangular tegumen with two curved, strong bristles on each side of the flattened apex, by sclerotised naked and hook-like socii, by long, slender and medially narrowed valvae with a series or large tuft of flattened bristles from outer surface near ventral margin in basal half, with a large,
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triangular, keel-shaped process from inner surface below end of basal excavation and usually with the base of the cucullus with a patch of short spines on ventral margin, often as a projecting angle, by a short caulis and a smallish aedeagus without cornuti. The female genitalia are characterised by a ventrally more or less raised, sclerotised ring around the entrance to the ostium, surrounded by spinulose membrane and fused into the V- or Ushaped excavation in the hindmargin of S7 and by a short, sclerotised ductus bursae that is longitudinally folded and twisted. All Tritopterna females examined have a modified ovipositor with very small, slender lobes and a narrow, barrelshaped 8th segment with an anteromedially infolded T8, and lack signa.
Description Adults (Figs 703, 704) Wingspan. 10–16 mm; small to medium-sized. Head (Fig. 706). Ocellus well developed. Vertex with very long, loosely appressed scales, anteriorly projecting between antennae and forming with scales of upper part of frons a large, obliquely flattened, forwardly projecting tuft, remainder of frons with small, appressed scales. Proboscis about as long as labial palpus. Labial palpus porrect, sinuate, long (2.2–2.5× diameter of eye), strongly widened in middle, second segment large, triangular (T. chionostoma, T. capyra (Meyrick)) to teardrop-shaped, widest near middle with long scales from ventral and dorsal margin in distal 36:, terminal segment very long and slender; inner surface of second segment snow-white in T. capyra and T. chionostoma. Antenna reaching to beyond middle of wing, stout in male, scaled except for narrow band, cilia not visible. Thorax. Smooth. Legs unmodified. Wings (Figs 86, 703, 704). Forewing index 0.35–4.0. Forewing elongate, subrectangular, widest in middle, with costa weakly curved in basal third, apex moderately pointed, termen sinuate and oblique; base of dorsum in male with a projecting tuft of recurved scales from margin; discal cell at most with weak traces of M-stem and chorda; base of R3 closely approximated and parallel to stalk of R4 and R5 (T. capyra) or R3 connate to shortstalked with stalk of R4 and R5 (T. chionostoma), R4 and R5 stalked to beyond 36: with R5 to termen; M1, M2 and M3 about equidistant; base of M3 close to base of CuA1 with the latter from before angle of cell; CuP present at margin; anal loop 0.22–0.31× length of 1A+2A; with 2–3 patches of raised scales in dorsal half of wing, wing pattern variable and
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AUSTRALIAN OLETHREUTINE GENERA
complex, grey-brown to red-brown (T. capyra) with blackish marks, variously suffused with whitish from dorsum, with 4–6 conspicuous, paired, whitish costal strigulae, an irregular, partly interrupted band or J-shaped leaden mark parallel to termen and usually with a dark, high triangle on dorsum before tornus and dark streaks along M1 and, closer to cell, along M2, sometimes confluent.
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Hindwing with cubital pecten present (T. capyra) or absent, with sexually dimorphic venation and in males on upper side with modified scales along anterior margin of cell, conspicuous, long, bandlike scales in T. capyra and T. chionostoma; Rs and M1 closely approximated and parallel at base (T. capyra); M2 bent at base toward M3 but variable from quite distant (Clarke 1971) to closely
Figs 703–708. Tritopterna capyra, adults, head and genitalia. 703, 704, adults. 703, , nr Bungwahl, NSW. 704, , nr Port Macquarie, NSW. 705, male genitalia, Port Macquarie, NSW, slide T2216. 706, head, male. 707, aedeagus, slide T2216. 708, female genitalia, Port Macquarie, NSW, slide T2207.
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approximated to M3 (T. capyra); M3 and CuA1 connate to short-stalked in female but clearly separate in male with CuA1 from below angle of cell; CuP well developed and halfway between CuA1 and 1A+2A, 3A not visible; brownish grey. Pregenital abdomen. Unremarkable. Tip of abdomen in male with conspicuous, large, tripartite, expandable scale tuft. Hindmargin of S8 deeply bilobed; T8 with convex hindmargin and inverted T-shaped sclerite. Male genitalia (Figs 705, 707). Tegumen squat, subrectangular with a flat top with isolated bristles, often with two particularly large, curved bristles on each side. Uncus variable, downcurved, from a large, tapering, beak-shaped process to a short, apically T-shaped digit. Socii wellsclerotised, naked, hook-shaped to medially widened bars. Gnathos a weakly sclerotised, medially narrowed, transverse band usually in basal third of tegumen. Valva long, slender, medially narrowed; cucullus semicircular, partially bristled and with some short, scattered spines along ventral margin, often with projecting ventral corner densely set with spines; ventral half of valva with a series or large tuft of flattened, often deciduous spines from outer surface near ventral margin, and with a large, triangular, keel-shaped process from inner surface below end of basal excavation; costal process long, straight, slender; with projecting angle in costa near base. Juxta triangular, caulis very short. Aedeagus lightly curved and tapering towards apex in lateral view; vesica without trace of cornuti. Female genitalia (Fig. 708). Ovipositor lobes modified, small and slender and apophyses posteriores stout. Eighth segment modified, barrelshaped with anteromedian portion of T8 infolded to form a complex, diamond-shaped structure. Hindmargin of T7 medially concave, bilobed; S7 with V-to U-shaped hindmargin. Sterigma fused with hindmargin of S7, a ventrally more or less raised, sclerotised, transversely ovate ring around entrance to ostium, surrounded by a spinulose area. Ductus bursae short, sclerotised, longitudinally folded and twisted, resembling a plait, ductus seminalis originating from close to corpus bursae; corpus bursae ovate, without signa.
Distribution Tritopterna occurs from Sri Lanka (Meyrick 1927b), Sumatra, Java and Australia to the western Pacific in Samoa, Rapa, the Marquesas Is, the Carolines (Clarke 1971, 1976, 1986) and the southern Cook Is
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(Dugdale personal communication 2005). In Australia it is found in the Cairns district in northern Queensland to Kiama in New South Wales.
Biology Tritopterna anachastopa (Meyrick) has been reared by L. G. E. Kalshoven from fruits of Glochidion and flowers of Mallotus (both Euphorbiaceae) in Telawa, Java (Clarke 1958), and in Australia T. capyra has been reared from Glochidion ferdinandi, from ‘larvae tunneling in fruit’ by I. F. B. Common at Church Point, NSW, 28.vi.1959 and from ‘seed capsules’ by K. M. Moore at Lisarow, NSW, 7-8.ii.1971.
Remarks As mentioned by Clarke (1971), Meyrick (1921) stated in his original description of Tritopterna that in the forewing R3 is stalked with R4 and R5, that M3 and CuA1 are connate in both wings, and that Rs and M1 in the hindwing are stalked. With only the single holotype of T. chionostoma available it is difficult to correctly interpret the venation without a wing slide. Except for M3 and CuA1, which are close but separate in both wings, from just before the angle of the cell, a re-examination of the type confirmed Meyrick’s description. Clarke (1971) discussed Meyrick’s description of Tritopterna, described and figured adults and genitalia of both sexes of Tritopterna galena Clarke from Rapa (later synonymised with Tritopterna eocnephaea (Meyrick) (Clarke 1986)), and included Tritopterna anastrepta (Meyrick) from Samoa in the genus. He did not mention any sexual dimorphism in wing venation. Clarke (1976) reported T. chionostoma from the Caroline Is and figured male and female genitalia. This identification, based on the snowwhite inner surface of the labial palpus, is questionable as the Caroline Is male figured has long, slender and sharply bent socii in contrast to the short, thick and nearly straight socii of the type of T. chionostoma from Java. The monophyly of Tritopterna is based on a large number of mostly unique apomorphies: an undivided uncus, large curved bristles from the top of the tegumen, strongly sclerotised but not bifurcate socii, a large keel from the inner surface of the valva below the end of the basal excavation, a cluster of spines at the base of the cucullus often on a ventrally projecting angle, modified ovipositor and T8, a ringshaped sterigma surrounded by spinulose membrane
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and fused into a deep excavation of S7, the sclerotised, longitudinally folded and twisted ductus bursae and absence of signa. Tritopterna shares numerous apomorphies with Acroclita bryopa Meyrick: large, porrect labial palpi, R3 closely approximated to stalk of R4 and R5, a hindwing with sexually dimorphic venation and a band of modified scales along anterior margin in discal cell in male, a squat tegumen, strongly sclerotised socii, flattened bristles from outer surface of valva near ventral margin in basal half, a keel from inner surface of valva near ‘neck’, and a short ductus bursae longitudinally folded and sclerotised throughout. Until further species of Acroclita Lederer linking the type of the genus and T. bryopa become known the synapomorphies for the two genera are unclear, but species of both genera have been reared from Euphorbiaceae. The description of Acroclita catharoptis Lower appears to be based on a mixed series including a Tritopterna species with the conspicuously snowwhite inner surface to its labial palpus and a species of Fibuloides closely related to T. phycitipalpia. The original description refers to five specimens from Kuranda and Cooktown but only one possible syntype has been found in the South Australian Museum, a greatly damaged male without the tip of the abdomen, labelled ‘Kuranda’ but without a label indicating that it originated from the Lower collection. This specimen is closely related to Fibuloides phycitipalpia but probably not conspecific, and there is no further material available. Given that it is not even certain that this specimen did indeed belong to the type series, though it is referred to as a lectotype in Brown (2005), it serves taxonomic stability best to focus on the description of the labial palpi that undoubtedly refer to a Tritopterna species and to treat T. catharoptis as a junior synonym of the widely distributed Tritopterna capyra.
Constituent species Four named non-Australian species of Tritopterna are known (Brown 2005), the type species T. chionostoma and Tritopterna anachastopa from Java, T. anastrepta from Samoa and T. eocnephaea (Meyrick) (Clarke 1986) from the Marquesas and Rapa Is, and one species is known from Australia. capyra (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 241 (Acroclita). [NSW: Sydney.]
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catharoptis (Lower, 1916), Trans. R. Soc. S. Aust. 40: 539 (Acroclita) [Qld: Cooktown, Kuranda.] Syn. nov.
Noduliferola Kuznetzov (Figs 84, 709–717) Noduliferola Kuznetzov, 1973. Entomol. Obozr. 52: 685. Type species: Noduliferola abstrusa Kuznetzov, 1973, ibid. 52: 685, by original designation. [China.] Duessa Clarke, 1976, Insects Micronesia 9(1): 10 (key), 26. Type species: Duessa pleurogramma Clarke, 1976, ibid. 9(1): 28, by original designation. [Micronesia.] Microclita Diakonoff, 1984a, Entomol. Basil. 9: 393. Type species: Microclita niphada Diakonoff, ibid. 9: 394, by original designation. [Sumba.]
Diagnosis Species of Noduliferola look like typical small to medium-sized eucosmines and can be recognised without dissection only if one is familiar with the species. They have large, porrect, sinuate labial palpi and narrow, subrectangular forewings with a produced apex and a narrow costal fold to the middle of the wing in the male. The wing pattern always includes a blackish spot in apex and often portions of a broad, dark, curved band from middle of costa to apex. In the forewing chorda and Mstem are absent, R4 and R5 are stalked and their stalk is closely approximated to R3 at base. The hindwing venation is variable with Rs and M1 stalked, the base of M2 bent towards M3, and M3 and CuA1 either stalked or fused into a single vein that is then connate with M2, and 3A is absent. The highly modified male T8 ending in a conspicuously enlarged, triangular, pointed lobe is diagnostic for Noduliferola. The male genitalia are unique, with the shield-shaped apex to the tegumen and the distally membranous and complex valva diagnostic for the genus. The large bundle of modified scales from a fold between tegumen and valva is unique in the Australian Eucosmini. The female genitalia with a strongly sclerotised, broadly ring-shaped sterigma, fitted into a deep, semicircular emargination in the hindmargin of S7 and with a spinulose hump in the lamella postvaginalis, are diagnostic for Noduliferola. The sclerotised ductus bursae in N. neothela with two prongs at the
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entrance to the corpus bursa is very similar to that of Fibuloides.
Description Adults (Figs 709–711) Wingspan. 9–14 mm (up to 20 mm in some non-Australian species); small to medium-sized. Head (Fig. 715). Ocellus moderate. Projecting tuft of long scales from upper part of frons, remainder of frons with minute, appressed scales. Proboscis short. Labial palpus porrect, long (1.9–2.3× diameter of eye) and strongly sinuate; second segment in distal half greatly widened by
loose scales on dorsal and ventral margin; third segment very long and thin. Antenna to beyond middle of wing, stout in male, entirely scaled, with minute cilia. Thorax. With a raised posterior crest. Legs unmodified. Wings (Figs 84, 709–711). Forewing index 0.30–0.33. Forewing elongate subrectangular, widest in middle, with costa evenly curved, apex produced and termen concave below apex; with a narrow costal fold to middle of wing in male, concealing modified yellowish scales; chorda and M-stem not developed; R3 at base close to long stalk of R4 and R5; M3 and CuA1 often closely
Figs 709–711. Noduliferola, adults. 709, 710, N. neothela, Lamington Nat. Park, Q. 709, . 710, . 711, N. sp., , Mt Bellenden-Ker, Q. Scale = 2 mm.
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approximated and parallel at base and sometimes distally converging again to the point of close approximation; CuP only a short trace near margin; anal loop 0.3× length of 1A+2A, base of 1A very faint; with tufts of raised scales and a complex wing pattern, pattern sexually dimorphic in N. neothela (Turner) (no other species with both sexes examined); ground colour white, grey, yellowish brown to brown or greyish green (N. neothela) with diverse brownish to black marks and always with a dark spot in apex, with the most frequent pattern element an often only partially expressed, strongly curved dark band from middle of costa to apex; costal strigulae well developed in distal half. Hindwing with dense cubital pecten and axillary tuft; Rs and M1 stalked, M2 at base bent close to stalk of M3 and CuA1 or M2 and fused M3 and CuA1 connate from angle; CuP present in distal half; 1A+2A well developed, 3A absent; uniformly greyish or brownish. Pregenital abdomen. Ventral apodemes large, conical. Sternum 8 in male with bilobed hindmargin, T8 with a transverse, roughly diamond-shaped sclerotisation at base and distally produced into a very large, triangular lobe of up to several times the length of S8 (Figs 716, 717). Male genitalia (Figs 712, 713). Vinculum a short, moderately wide band fused with tegumen. Tegumen strongly modified with distal portion elongated and shield-shaped with a pronounced medial ridge from pointed apex. A distinct, bifid uncus present in N. pleurogramma only. Socii two small knobs or well-defined patches of bristles laterally below ventral end of ridge from apex. Gnathos a curved transverse band in middle or basal third of tegumen. Angle between tegumen and valva with a membranous fold containing a large, well-defined bundle of very long, band-like scales. Valva very complex, narrower and more sclerotised in basal portion, distal part beyond ‘neck’ formed by excavation in ventral margin greatly swollen, of very thin membrane and separated into three lobes: tip of valva variably extended into a triangular point, with a second rounded to hump-shaped lobe with few to numerous strong bristles arising on inner surface below distal half of costa, and with a third small lobe ventrally from inner surface beyond ‘neck’, carrying heavy spines and often also one or two very large spikes; inner surface of distal portion often covered with modified, inflated scales; basal part of valva in some species with a small hump set with spines at base of costa or with a sharp tooth on ventral margin before neck; costal process long, straight and slender. Caulis very short, anellus a
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broad, nearly cup-shaped ring. Aedeagus wide and short; vesica with a large bundle of lanceolate cornuti. Female genitalia (Fig. 714). Ovipositor lobes elongate subtriangular. Hindmargin of S7 deeply excavated to accommodate large, strongly sclerotised, broadly ring-shaped sterigma, with lamella antevaginalis transversely wrinkled in N. neothela and lamella postvaginalis often much wider, with a hump and spinulose. Ductus bursae short and often gradually widening into corpus bursae, with a short colliculum followed by short membranous portion, remainder well sclerotised (to below inception of ductus bursae), in N. neothela ending in two long, sharp prongs projecting into corpus bursae. Corpus bursae ovate or pear-shaped, densely spinulose especially in apical part, with two large, hollow, horn-shaped signa, often dissimilar with one broader and blunt.
Distribution Noduliferola has been recorded from China, Vietnam, Indonesia, Micronesia, the Marquesas Is and Australia, where it occurs in rainforest and marginal rainforest localities from Brisbane to Cape York Peninsula, Queensland.
Biology A long series of N. neothela has been reared by the author in early December from fruit of Eugenia sp. (Myrtaceae) collected on the ground in Binna Burra, Lamington NP, Qld. Another series of the same species reared from fruit of Vitex lignum-vitae (Lamiaceae) in April by A. W. S. May from Yarraman, Qld, is in the ANIC. Robinson et al. (2001) reported shoots of Vitex pinnata (Lamiaceae) as host for N. spiladorma (Meyrick).
Remarks Kuznetzov (1973) and Diakonoff (1984a) provided drawings of head, venation and male genitalia of N. abstrusa and N. niphada respectively, and Clarke (1976, 1986) figured adults, head, wing venation, and male and/or female genitalia for four species of Duessa. Kuznetzov (1997a, 1997b, 2003) synonymised Microclita and Duessa and figured the male genitalia of N. insuetana Kuznetzov. The monophyly of Noduliferola is based on a narrow costal fold covering characteristically modified scales, a highly derived T8 with a long, triangular posterior lobe, a scale pencil on a fold
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Figs 712–717. Noduliferola, head, abdomen and genitalia. 712, N. neothela, male genitalia [holotype N. nimbata], slide ANIC 8633. 713, N. sp., male genitalia, nr Mt Tip Tree, Q, slide T2334. 714, N. neothela, female genitalia, Yarraman, Q, slide T2217. 715, head, N. neothela, male. 716, 717, abdomen, 8th segment, male. 716, N. neothela. 717, N. sp.
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between the tegumen and the valva, the uniquely modified apex of the tegumen and the position and shape of the strongly sclerotised, ring-shaped sterigma with a spinulose hump in the lamella postvaginalis. The complex valva has a uniquely derived distal portion with additional lobes bearing bristles or spines, though secondary modifications have led to two quite distinct shapes within the genus. The synonymy of Duessa and Microclita is obvious with the type species of both genera having M3 and CuA1 in the hindwing fused into a single vein and with very similar male genitalia. The type species of Noduliferola has M3 and CuA1 stalked and a differently shaped valva. An unidentified Australian species combines a valva of the Duessa/ Microclita type with a hindwing with both M3 and CuA1 present and stalked. There may well be some indication for two species-groups in Noduliferola based on genitalic structure, but the numerous apomorphies shared by the entire group define a tightly monophyletic entity. Furthermore, males and females are associated with certainty only for N. neothela, so the following observations are tentative. The species-group including N. abstrusa and N. neothela is characterised by extremely elongated tegumen and valva, the latter distally triangular, and in the female probably by a twopronged sclerite in the ductus bursae. The second group seems to be characterised by a distally more rounded valva that is ventrally sclerotised in the basal half and often has a projecting tooth, and the sclerotisation in the ductus bursae has no anteriorly projecting points. In most species of the second group M3 and CuA1 are fused into a single vein. Noduliferola is included in the Acroclita-group on the strength of the bipronged sclerite in the ductus bursae, the wing venation and the raised scales on the dorsum, an assignment supported by the cladistic analysis. Whereas Peridaedala crocoptila Diakonoff, [1968], from the Philippines could either be a species of Noduliferola or Fibuloides Kuznetzov, there is no doubt that the following species (Clarke 1958) belongs to Noduliferola: spiladorma (Meyrick, 1932), Exotic Microlepid. 4: 221 (Acroclita). [Java.] Comb. nov.
Constituent species Brown (2005) lists eight named species from outside Australia, in addition to the species referred to Noduliferola above and one named and one unnamed species from Australia:
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neothela (Turner, 1916), Trans. R. Soc. S. Aust. 40: 524 (Acroclita). [Qld: Brisbane.] nimbata (Turner, 1946), Trans. R. Soc. S. Aust. 70: 199 (Acroclita). [Qld: Toowoomba.]
Heleanna Clarke (Figs 85, 718–723) Heleanna Clarke, 1976, Insects Micronesia 9: 10 (key), 11. Type species: Rhopobota physalodes Meyrick, 1910, Trans. Entomol. Soc. Lond. 1910: 368, by original designation. [Sri Lanka.]
Diagnosis Heleanna species are small moths with subrectangular forewings, widest in basal third and with two small tufts of raised scales on dorsum, and in the male always with a longitudinal, ovate depression above at the base of the cell filled with whitish, modified scales and covered by a dome of long scales, and sometimes with a costal fold covered by a dense fringe of curved, large scales. The forewings are of pale ochreous or greyish green ground colour with ill-defined costal strigulae and a variable and complex pattern of dark brown speckling and marks, with some marks brownish olive and edged with black in the Australian H. chloreis (Turner). In the forewing R3 is close to the long stalk of R4 and R5 or stalked with it, and M3 is close to CuA1 at base but distally approaching M2, in the hindwing Rs and M1 are close but separate at base, M2 is bent close to the stalk of M3 and CuA1, and CuA2 originates from 6M of cell or from beyond. The head of Heleanna is characteristic in both sexes with scaling of sculpted appearance on the frons forming a large, flat, narrowly triangular area of tightly appressed scales anteriorly projecting over the labial palpi, with a medial, nose-shaped ridge in the male, and with long, porrect, sinuate labial palpi of triangular outline, neatly fitting the frontal scale projection. In the male genitalia (strongly asymmetrical in H. chloreis) a robust, medially notched tegumen without any trace of an uncus (except possibly in the Japanese H. fukugi Nasu), arm-like, strongly sclerotised, drooping socii, naked in basal half and distally usually variably bristled and often with terminal thorn, and usually a long and extremely slender valva medially narrowed to a band-like ‘neck’ with a narrowly elliptical cucullus with 1–2 spines along the ventral margin and one on the
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apex are diagnostic for Heleanna. The female genitalia of Heleanna are characterised by a ring- or funnel-shaped sterigma posterior to S7 or in an excavation of its hindmargin, a twisted and longitudinally grooved or folded ductus bursae, strongly sclerotised except for short portions at both ends, and an inverted pear-shaped corpus bursae with narrowed anterior end, with two nearly straight, horn-shaped cornuti.
Description Adults (Figs 718, 719) Wingspan. 8–15 mm (Australian species 8–10 mm); small. Head (Fig. 721). Ocellus well developed. Head scaling with a sculpted appearance; vertex with long, loosely appressed scales, projecting between antenna and, together with scales of upper frons, forming a large, flat, narrowly triangular area of tightly appressed scales anteriorly projecting over labial palpi, with a medial, nose-shaped ridge in male; remainder of frons minutely scaled. Maxillary palpus minute, scaled. Proboscis about as long as labial palpus. Labial palpus porrect, sinuate, long (1.9–2.3× diameter of eye), of triangular outline and with dense, loosely appressed scaling, second segment in distal half greatly enlarged by long, dense scales along ventral and distal portion of dorsal margin and by an additional crest of loose scales dorsally from outer surface at widest point, terminal segment long but partially hidden, forward or downward pointing. Antenna to beyond middle of wing, entirely scaled, thicker in male, no cilia visible. Thorax. Smooth. Legs unmodified. Wings (Figs 85, 718, 719). Forewing index 0.29–0.40 (0.29–0.32 in Australian species). Forewing elongate-subrectangular, very slender in Australian species, widest in basal third, with costa curved in basal third then weakly concave, apex rectangular to slightly projecting, termen weakly to conspicuously sinuate, vertical to slightly outwardly oblique; with two rounded tufts of raised scales, one on each side of fold, at about 369 dorsum, in male small costal fold covered by a dense fringe of curved, large scales present (H. physalodes) or absent (H. chloreis) and an ovate deep depression above (visible also from below) at base of cell always present, filled with whitish, modified scales and covered by a dome of large scales; discal cell with chorda and M-stem (to base of M3) reduced or absent; R3 close to or stalked with long stalk of R4 and R5; M1, M2 and M3 about equidistant; M3
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very close to CuA1 at base but the two veins distally strongly diverging with M3 bent towards apex; CuA2 from beyond middle of cell; CuP present at margin; pale ochreous ground colour often with a greenish tinge with ill-defined costal strigulae and a variable and complex pattern of dark brown speckling and marks, in particular 2–3 small, usually squarish marks on costa at base, at 56M and at 56O, an often indistinct, outwardly oblique streak from 56M dorsum to middle of wing (to end of scalefilled depression in male) or beyond, a small black spot in apex and an oblique dark streak or mark in distal fourth from centre of wing to near apex; in H. chloreis the ground colour is partly iridescent greyish green and some of the dark marks are brownish olive edged with black, especially in the female. Hindwing with well-developed cubital pecten and axillary tuft, sometimes with small, widely spaced scales resulting in a semitranslucent wing (H. chloreis) and distal half of costa with persistent scales; adjacent bases of Sc+R1 and Rs connected by crossvein R1, Rs and M1 closely approximated and parallel at base, M2 bent close to stalk of M3 and CuA1; CuA2 from 6M of cell or beyond; CuP a weak trace at wing margin, 1A+2A well developed, 3A not visible; greyish ochreous to brownish grey, in H. chloreis greyish ochreous around margin and along veins, with isolated, minute, black-tipped scales between veins. Pregenital abdomen. Unmodified. In male hindmargin of S8 with deep medial emargination, T8 with weak, inverted-T shaped sclerotisation at base. Male genitalia (Figs 720, 722). Tegumen robust, usually rounded-ovate or squarish, but strongly asymmetrical in chloreis with lateral portion nearly 156O times as long on right side. Top of tegumen flat or with a medial notch, without any trace of uncus. Socii modified, two long, slender, strongly sclerotised, drooping arms, naked in basal half and distally variably bristled except in Heleanna fukugi, usually with a distinctive row of short bristles often ending in a terminal, long, outwardly-pointing thorn; in H. chloreis socii arms asymmetrical, much longer on the right side, sinuate, distally hairy and with a flattened, crescentic apical lobe ending in a bundle of bristles. Gnathos a weak, membranous remnant only. Valva long and extremely slender throughout and medially narrowed to a band-like ‘neck’ except in H. fukugi where it is parallel-sided, with large, flattened bristles ventrally from outer surface of valva; usually with a clump of small bristles on tubercle dorsally near distal end of basal excavation
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Figs 718–723. Heleanna chloreis, adults, head and genitalia. 718, 719, adults, Moses Ck nr Mt Finnigan, Q. 718, . 719, . 720, male genitalia, nr Kuranda, Q, slide ANIC 1289. 721, head, male. 722, aedeagus, slide ANIC 1289. 723, female genitalia, nr Kuranda, Q, slide ANIC 12987. Scale = 2 mm.
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and with a longitudinal row of bristles parallel to ventral margin at base; cucullus narrowly elliptical, densely bristled on inner surface, with 1–2 spines along ventral margin and one on apex; costal process moderately large. Juxta very small, caulis short, anellus ovate, cup-shaped. Aedeagus small, slender; vesica with a small sheaf of slender cornuti. Female genitalia (Fig. 723). Ovipositor lobes elongate ovate. Hindmargin of S7 straight, weakly concave or with a U-shaped medial excavation. Sterigma variable, from a simple, medial, variously sclerotised ring or collar to a more or less complex short funnel, but genitalia in H. chloreis strongly asymmetrical and entrance to ostium on the left body side posterior to straight hindmargin of S7, entirely membranous without any sclerotisation. Ductus bursae moderately long to long, strongly sclerotised except for short portions at both ends, irregularly twisted and longitudinally grooved or folded, with colliculum of thickened membrane or absent, with ductus seminalis from sclerotised part near its anterior end; sclerotised portion of ductus bursae particularly long and sinuate in H. chloreis; corpus bursae inverted pear-shaped with narrowed anterior end, with two nearly straight, horn-shaped cornuti.
Distribution Heleanna is known from the Chagos Archipelago (south of the Maldives), the Seychelles, Sri Lanka, India, Taiwan, Caroline Is, Mariana Is, Solomon Is, Fiji, Austral. Is (Clarke 1976), the Ryukyus (Japan) (Nasu 1999), and Australia, where it occurs only in the Cairns region in northern Queensland, north nearly to Cooktown.
Biology Fletcher (1932) reports that H. physalodes has been bred in Fiji from larvae in flowerbuds of Barringtonia (Lecythidaceae). Clarke (1976) gives label data for H. physalodes ssp. physalodes from Guam citing Barringtonia and fruit of Cordia (Boraginaceae) as hosts and for H. physalodes ssp. tricia Clarke from Caroline Is atolls citing Calophyllum inophyllum (Clusiaceae) without any further details. Nasu (1995, 1999) gives detailed information about the biology and immatures of two species. Heleanna melanomochla (Meyrick) rolls young mango leaves (Anacardiaceae) in Taiwan (Nasu 1995) and H. fukugi bores into buds and rolls young leaves of Garcinia subelliptica (Clusiaceae) in the Ryukyus (Nasu 1999). An unidentified species of Heleanna has been reared at
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Madang, Papua New Guinea, from Pimelodendron amboinicum (Euphorbiaceae) (Miller personal communication 2003). No host is known for the Australian H. chloreis, but all the specimens have been collected in rainforest.
Remarks Meyrick’s (1910) original description of H. physalodes was based on two specimens from Ile du Coin, Chagos Archipelago, and one female specimen from Galle, Sri Lanka. Diakonoff (1950) selected one of the Chagos Is specimens as the lectotype that was figured by Clarke (1958). Diakonoff (1969) subsequently transferred H. physalodes to Herpystis Meyrick, figuring the genitalia of both sexes from material from the Seychelles though, judging from the figure, there is some doubt whether the female really is congeneric. In his description of Heleanna, Clarke (1976) summarised the available literature and extensively figured H. physalodes based on material from the Caroline Is, including three new subspecies. Nasu (1995) redescribed and illustrated H. melanomochla from Taiwan, including immatures, and more recently provided an equally comprehensive treatment of H. fukugi from the Ryukyus (Nasu 1999). The monophyly of Heleanna is based on the modified, sculpted head scaling, the ovate depression above on the basal third of the forewing filled with whitish, modified scales and covered with large, domed scales, stalked R4 and R5, a dorsally notched, robust tegumen without uncus, arm-like, drooping, strongly sclerotised socii bristled at most near the apex, and a sclerotised, twisted and longitudinally grooved or folded ductus bursae. There are three genera closely related with Heleanna, all insufficiently known and based on type material from islands either in the Indian Ocean or the Pacific, and the entire group will have to be reevalued once more material becomes available. Pseudoclita Bradley, originally based on a single female from the Solomon Is (Bradley 1957) and present on Norfolk I. off Australia, shares with Heleanna very similar female genitalia and a somewhat similar valva with a row of bristles below the basal excavation but has a paired, welldeveloped uncus set with large spines and small, lateral, membranous socii lobes. Pseudoclita also has modified, sculpted head scaling, though less pronounced than Heleanna, but it has shorter labial palpi, R4 and R5 separate, lacks the scale-filled depression on the forewing and has the tegulae scaling produced into conspicuous long pencils in
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the male. The following species from Norfolk I. is here referred to Pseudoclita: macroma (Turner, 1918), Trans. R. Soc. S. Aust. 42: 287 (Acroclita). [Norfolk Is.] Comb. nov. Cosmetra Diakonoff, described from Réunion (Diakonoff 1977), shares a very similar tegumen with exactly the same socii as Heleanna but lacks the modified head scaling, the secondary sexual structures on the forewing and has R4 and R5 separate as well as a much more plesiomorphic valva and different female genitalia. Psegmatica Meyrick from Fiji, known only from the single undissected male holotype (Meyrick 1930) and a female without abdomen in the BMNH, looks superficially rather similar to Heleanna and Pseudoclita but lacks the secondary sexual structures of the forewing, has unmodified tegulae in the male and, most importantly, M3 and CuA1 are separate in the hindwing whilst R4 and R5 are stalked in the forewing. The highly asymmetrical genitalia in H. chloreis are most unusual but a consistent feature of the species. The conflicting messages of an allegedly very wide distribution of H. physalodes, from the Seychelles to the Solomon Is, and the fact that Clarke (1976) recognised four different subspecies for the Caroline Is alone suggest that the type species of Heleanna deserves closer scrutiny. Heleanna is included in the Acroclita-group on the strength of the sclerotised socii, the flattened bristles from the outer surface of the valva in H. fukugi, the wing venation and the raised scales on the dorsum, an assignment supported by the cladistic analysis.
Constituent species Three named species of Heleanna are known from outside Australia (Brown 2005), the type species H. physalodes with four subspecies (Clarke 1976), H. melanomochla (Meyrick) from Taiwan (Clarke 1958, Nasu 1995), and H. fukugi from the Japanese Ryukyus (Nasu 1999). One named species is known from Australia. chloreis (Turner, 1916), Trans. R. Soc. S. Aust. 40: 523 (Acroclita). [Qld: Kuranda.]
Spilonota-group Status remark This is a new, informal genus-group including, in Australia, Spilonota Stephens, Strepsicrates Meyrick,
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Holocola Meyrick, Hermenias Meyrick and Eccoptocera Walsingham.
Evidence for monophyly The monophyly of this group is supported by several apomorphies, even though they are absent in some taxa: antennal notch in male, extended anal region in male hindwing with upturned anal margin folded over long scale pencil from base, funnel-shaped sterigma usually fused with S7, tendency for signa to be reduced and lost and bursa to become strongly spinulose.
Distribution The Spilonota-group is centered on the Australian region, with Strepsicrates occurring in all faunal regions and Spilonota extending into the western Palaearctic and introduced into North America.
Biology Members of the Spilonota-group are generally leaf rollers, with a strong emphasis on Myrtaceae as host plants. However, the northern hemisphere apple pest Spilonota ocellana ([Denis & Schiffermüller]) is polyphagous on numerous plant families and Spilonota laricana Heinemann feeds on shoots of Larix decidua (Pinaceae). The larvae of the Australian Spilonota constrictana-group spin one or several of the hollowed-out myrtaceous flower buds or seed capsules together to make a portable case. Strepsicrates Meyrick usually is a leaf roller on Myrtaceae, but the larva of the Australian S. macropetana (Meyrick) frequently tunnels in eucalypt galls. Though the majority of host records for Holocola Meyrick from Australia involve leaf rolling on Myrtaceae, H. triangulana Meyrick and two other species of this group have been reared from Acacia species (Mimosaceae).
Remarks Three highly derived genera from New Zealand, Protithona Meyrick, Parienia Berg and Hendecasticha Meyrick, also belong to the Spilonotagroup, as does the minute Lepidunca Diakonoff from Sumba (Diakonoff 1984a) and Xenosocia Diakonoff from Madagascar (Diakonoff 1989b). Presence of an antennal notch, costal fold and modified anal hindwing region, together with a bifid uncus, large membranous socii, a medially constricted slender valva and two horn-shaped signa
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are here ascribed to the groundplan of the Spilonotagroup. Within the group, the uncus has repeatedly developed into large and widely separate arms or become lost, the socii often are conspicuously modified, swollen and/or sclerotised, and various large spines have developed on the cucullus. Reduction of the signa and the parallel development of strong spinules in the bursae also have occurred parallel several times within the genus-group.
Spilonota Stephens (Figs 87, 724–738) Spilonota Stephens, 1829 [June], Nom. Br. Insects: 46. Type species: Tortrix comitana [Denis & Schiffermüller] sensu Hübner, [1799], [ = Tortrix ocellana [Denis & Schiffermüller], 1775, Ankündung syst. Werkes Schmett. Wienergegend: 130], by subsequent designation by Curtis, 1835, Br. Entomol. 12: folio 551 (as ‘Tortrix comitana’ Hüb.). [Austria.] Tmetocera Lederer, 1859 April, Wien. Entomol. Monatsschr. 3: 124 (key). Type-species: Tortrix ocellana [Denis & Schiffermüller], 1775, Ankündung syst. Werkes Schmett. Wienergegend: 130, by subsequent monotypy. [Austria.] Bathrotoma Meyrick, 1881, Proc. Linn. Soc. NSW 6: 635 (key), 675. Type species: Bathrotoma constrictana Meyrick, 1881, ibid. 6: 675, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 44, 61. [Australia.] Syn. nov. Batrotoma Turner, 1946, Trans. R. Soc. S. Aust. 70: 197 (incorrect subsequent spelling).
Diagnosis Spilonota species are small to medium-sized, very diversely marked eucosmine moths with a dark apical spot (except white species) and with a notch at the base of the antenna and a costal fold in all males except those of S. ocellana and close relatives and S. quietana (Meyrick). The latter have their forewing divided by a somewhat sinuate oblique line from near base to apex into a darker grey costal portion and a very pale dorsal half. S. constrictana (Meyrick) and related species can be recognised by a dark, narrowly subtriangular mark along the termen and a dark row of basal scales in the terminal fringe present also in S. ocellana. Spilonota
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has moderately long, porrect, sinuate labial palpi, a forewing with R4 and R5 nearly always separate and a hindwing with Rs and M1 either separate or stalked, M3 and CuA1 nearly always fused into a single vein (rarely stalked only) and in the male often with a scale pencil concealed by the upturned anal margin. The male genitalia are diagnostic for the genus, with a well-developed uncus (absent in S. ocellana and close relatives), nearly always dorsoventrally flattened and often deeply cleft medially, with naked and sclerotised socii originating from the inner side of the teguminal ‘shoulders’, either swollen and triangular, band-like or slender and elbowed, and with a long, slender valva always with at least one prominent thorn or spike ventrally or subdorsally on the cucullus. Some species of Holocola Meyrick also have rigid socii but they are positioned widely separate, much more laterally on the tegumen, and their cucullus does not have a prominent spine. The female genitalia have a sclerotised, funnel-shaped sterigma fused with the concave hindmargin of S7 in the Australian species and two large, thorn-shaped signa with a sclerotised, round or triangular base. The ductus bursae usually is posteriorly sclerotised up to the small but conspicuous colliculum of thickened membrane, followed by a short membranous portion then by an often irregularly sclerotised anterior half, with the exception of S. quietana where the ductus bursae is narrow and entirely membranous except for the colliculum and a small sclerite near the origin of the ductus seminalis.
Description Adults [based predominantly on Australian species] (Figs 724–727) Wingspan. 8–19 mm; small to medium-sized. Head (Figs 728, 729). Ocellus moderate. Frons with moderate to long, projecting scales in upper part, remainder of frons with minute, appressed scales. Maxillary palpus minute, scaled, 3segmented. Proboscis about as long as labial palpus. Labial palpus moderately long to long (2–2.4× diameter of eye), more compact in S. constrictana and related species, sinuate, porrect; second segment large, sinuate, strongly widened in distal half by long, loose scales along ventral margin and distal half of dorsal margin; terminal segment from stout and rather short (S. constrictana) to very long and slender, sometimes concealed in scales from second segment. Antennae reaching to beyond middle of forewing; flagellum with basal notch in
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male (Fig. 729) except in S. quietana and related species: second and third flagellomeres narrow and fused, third and sometimes fourth with a dorsal spur; in male either entire antenna scaled (S. constrictana) or with a ventral band with minute cilia. Thorax. Without crest. Legs unmodified. Wings (Figs 87, 724–727). Forewing index 0.29–0.41. Forewing narrowly subrectangular, sometimes slightly widening distally, costa from weakly and evenly curved to faintly sinuate, apex slightly projecting, termen sinuate and slightly oblique; male with moderately wide costal fold to near middle or slightly beyond, absent only in S. ocellana and close relatives and in S. quietana and
351
related species; discal cell long, accessory cell narrow and often very short with chorda variable, from halfway between R1 and R2 to between R5 and M1 in S. ocellana but much weaker, from just below R2 to just beyond R3 in other species, Mstem usually only a weak trace in membrane to M3; usually all veins separate beyond discal cell, rarely R4 and R5 connate to stalked (connate or shortstalked in S. ruficomana (Meyrick), stalked in S. quietana and related species), with CuA1 strongly curved and distally approximated to M3 in more broad-winged species; CuP present near margin; anal loop 0.25× length of 1A+2A; wing pattern diverse, with well-developed costal strigulae and nearly always with a dark spot in apex (absent only
Figs 724–727. Spilonota, adult males. 724, S. constrictana, Cook, ACT. 725, S. sp. D, Jim Jim Ck nr Mt Cahill, NT. 726, S. sp. B, nr Millstream, WA. 727, S. sp. nr quietana, Swansea, NSW. Scale = 2 mm.
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in whitish species) and with at least some parts of the ocelloid patch present, usually the two silvery bands but often also remnants of one or more black dashes; frequently with dark transverse bands or remnants thereof on dorsum just below middle and at 6M, and in S. constrictana and related species always with a dark, narrowly subtriangular mark along termen and a dark basal line along terminal fringe. Hindwing with cubital pecten and axillary tuft developed, males with anal margin usually folded over scale pencil from base of wing (not in S. ocellana) and anal region expanded to form prominent angle in S. constrictana and S. quietana, males of S. constrictana with modified, diverging scales on upperside of discal cell, arranged in fishbone-pattern; Rs and M1 usually closely approximated at base, rarely stalked (S. ruficomana), base of M2 strongly bent towards M3, M3 and CuA1 rarely stalked (S. ocellana and close relatives, S. acrosema), nearly always fused into one vein; CuP halfway between CuA2 and 1A+2A, the latter distant from anal margin, no trace of 3A visible in Australian species examined; pale to dark grey or greyish brown. Pregenital abdomen. Small, lateral, membranous pockets in female of some species (S. ocellana, S. constrictana) between S6 and S7. In male hindmargin of S8 strongly bilobed, T8 with an inverted T-shaped sclerotisation. Male genitalia (Figs 731–734). Tegumen high, robust, with strong ‘shoulders’. Vinculum generally small, but forming a prominent ventral triangle in S. constrictana and related species. Uncus nearly always a broad, dorsoventrally flattened process, undivided and of subtrapezoid shape in
S. constrictana and deeply cleft with two band- or paddle-shaped processes elsewhere except for the single, rod-shaped uncus of S. quietana; uncus absent in S. ocellana and close relatives. Socii naked, sclerotised processes from inner side of tegumen ‘shoulders’, pointed and three-dimensional in S. ocellana, flat and band-like in S. constrictana, and long, slender and elbowed in other Australian species. Gnathos two weak, lateral bands, not always apparent. Valva long and slender, of very variable shape but always with at least one large ventral spine, with costal process often small; valva usually narrowed beyond basal excavation into long, slender ‘neck’ then expanding into rounded or subtriangular cucullus with a spine or large spike from ventral margin or adjoining outer surface; cucullus in S. ocellana and S. constrictana and related species small, with rounded and sometimes swollen dorsal lobe and curved, fingerlike ventral process ending in thorn or spine; in S. acrosema and related species cucullus with a large subdorsal or dorsal spine, and rarely sacculus ending in a triangular process and rest of valva very slender, distally rounded with one or few small ventral thorns. Juxta triangular; caulis short and wide in S. ocellana and S. constrictana, very long in S. quietana; anellus a broad ring. Aedeagus lightly curved, short and wide in S. ocellana and S. constrictana, very long, slender and curved in S. quietana; with a sheaf of lanceolate, deciduous cornuti except in S. quietana and related species. Female genitalia (Figs 735–738). Ovipositor lobes subreniform, anteriorly narrower. Sterigma fused with medially concave hindmargin of S7,
Figs 728–730. Spilonota constrictana, head, antenna and larval case. 728, head, male. 729, base of male antenna with notch (arrow), dorsal side. 730, portable case of S. constrictana larva.
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forming a wide, sclerotised funnel leading to ostium, with ventral rim straight or medially projecting as rounded lobe or truncate point; in S. ocellana sterigma very wide, invaginated and lamella antevaginalis a medially deeply notched, broadly triangular lobe; S. quietana and related species with a curved, transverse fold across S7, ending in a medial, keel-shaped process. Posteriormost portion of ductus bursae sclerotised to conspicuously narrow, split colliculum of thickened membrane, followed by short membranous portion then by sclerotised anterior half, gradually widening, somewhat twisted and irregularly sclerotised in Australian species; ductus seminalis originating from anterior 56: of ductus bursae; in S. quietana ductus bursae narrow and entirely membranous except for two minute colliculum sclerites below ostium and a small, round sclerite at origin of ductus seminalis, with asymmetrical lateral dilation near corpus bursae; in S. ocellana with weakly sclerotised area in ‘neck’ of corpus bursae. Corpus bursae roundish, teardropshaped, minutely spinulose or dimpled; signa two strong thorns with a sclerotised, roundish or triangular base.
Distribution Spilonota occurs throughout the Palaearctic, Oriental and Australian regions, and S. ocellana is an introduced pest in North America. In Australia, Spilonota occurs throughout the continent, except for very dry areas, and is present also in Tasmania.
Biology The northern hemisphere S. ocellana is a polyphagous leafroller on shoots and flowers of numerous plant families, and a pest on apple, and Spilonota laricana Heinemann feeds on shoots of Larix decidua (Pinaceae). Spilonota constrictana and related species have a remarkable biology with their larvae feeding in flowers of several species of Melaleuca and Callistemon (Myrtaceae) and using one or several of the hollowed-out flower buds or seed capsules, spun together, as a portable case (Fig. 730) that is then fixed to a twig for pupation. A species fairly close to S. acrosema (Turner) has been reared from larvae binding together leaves of Melaleuca uncinata near Barellan, NSW, and the larvae of one species near S. quietana have been collected ‘ex Leptospermum’, without any further details given.
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Remarks Spilonota ocellana and Bathrotoma constrictana share not only a similar antennal notch, sclerotised socii originating from the inner side of the tegumen ‘shoulders’, a spike from the ventral margin of the cucullus and very similar female genitalia with membranous lateral pockets between S6 and S7, but also the wing pattern of S. ocellana agrees well with S. constrictana and related Australian species. Whereas Spilonota s. str. can be characterised by several apomorphies, mostly reductions such as loss of costal fold, anal pencil and uncus in the male, and by a more highly derived sterigma with a projecting and medially deeply notched lamella antevaginalis, no convincing apomorphy has been found to define the Australian species associated with S. constrictana, and Bathrotoma is consequently treated as a synonym of Spilonota. A cladistic analysis including all the undescribed species may reveal that a subgroup including S. acrosema that has the large spike on the cucullus in a subdorsal or dorsal position, the colliculum far below the ostium and the signa with a triangular base, deserves generic rank. Parienia Berg (replacement name for Exoria Meyrick) from New Zealand shares characteristically modified and sclerotised socii with Spilonota as well as a ductus bursae that is sclerotised in its anterior half. The slender, elbowed socii of Parienia could be considered a synapomorphy shared with an Australian subgroup within Spilonota, but on the other hand Parienia is characterised by numerous apomorphies (ventral process on aedeagus, highly modified cucullus and anellus, modified ovipositor lobes, ostium within S7) and, more importantly, all known species of Spilonota have a large thorn or spike on their valva that is absent in Parienia. Until relationships are resolved in much more detail it is in the interest of taxonomic stability to treat Spilonota and the New Zealand Parienia as separate genera and assume they are probable sister groups. Xenosocia Diakonoff from Madagascar (Diakonoff 1989b) also is very close to both Parienia and Spilonota and will have to be included in any detailed cladistic analysis of this group. The type species Xenosocia panegyrica Diakonoff (Diakonoff 1989b: fig. 14, not fig. 13 as stated) is characterised by sclerotised, slender and elbowed socii, a long, slender, single uncus, and a ventrally triangularly expanded cucullus without any thorns. According to the description, the costal fold and the notch at the base of the antenna are present in some males of the genus and absent in others, as in the
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Figs 731–736. Spilonota, male (731–734) and female (735, 736) genitalia. 731, S. ocellana, UK, slide BM 24889. 732, S. constrictana, Wilton, NSW, slide ANIC 13822. 733, S. sp. C., nr Borden, WA, slide ANIC 13805. 734, S. sp. D, nr Mudginbarry Homestead, NT, slide ANIC 13820. 735, S. ocellana, UK, slide ANIC 13818. 736, S. constrictana, Mt Keira, NSW, slide ANIC 13823.
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355
Spilonota squarely falls into a genus-group including Strepsicrates Meyrick, Holocola etc. centred on myrtaceous host plants in the Australian region and characterised in the groundplan by a notched antenna, costal fold and anal fold with a scale pencil.
Constituent species
Figs 737, 738. Spilonota, female genitalia. 737, S. sp. D, nr Borroloola, NT, slide ANIC 13821. 738, S. sp. F, Yeppoon, Q, slide T2251.
Australian Spilonota, and there is a similar tendency for M3 and CuA1 in the hindwing to become entirely fused. Description and illustrations of Spilonota ocellana are found in numerous publications (Heinrich 1923; Bentink and Diakonoff 1968; Kuznetzov 1978; Miller 1987; Razowski 1989, etc.). Of the species described by Meyrick and figured by Clarke (1958) only a minority appear to be true Spilonota (Spilonota calceata (Meyrick), Spilonota lechriaspis Meyrick), a few others are either Holocola or Strepsicrates species, while the rest are quite unrelated. The monophyly of Spilonota is based on sclerotised, naked socii originating from the inner side of the teguminal ‘shoulders’ (in contrast to the rigid socii of some Holocola species) and a cucullus with at least one prominent ventral thorn or spike. Both S. ocellana and S. constrictana have lateral membranous pockets between S6 and S7 in the female. In Australia the group is further characterised by a tendency of M3 and CuA1 to fuse into a single vein in the hindwing. Spilonota constrictana and related species not only possess numerous structural apomorphies, they are also distinctly set apart by a larva with a portable case.
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Razowski (1989) reported about 15 described true Spilonota species from beyond Australia and Brown (2005) includes 29 species in Spilonota and three in Bathrotoma. The Oriental taxa currently classified as Spilonota will have to be re-examined, they could include species of Fibuloides Kuznetzov, Strepsicrates and Holocola. The male genitalia illustration of Spilonota penechra Diakonoff from Madagascar (Diakonoff 1989b) suggests a species of Strepsicrates rather than of Spilonota. Only four species of Spilonota have been named from Australia, with at least 16 unnamed species present in the ANIC. Bathrotoma angelica Diakonoff, described from New Guinea (Diakonoff 1953), with the antennal notch distant from base and with large, hairy, rather flimsy socii, is probably a species of Holocola. constrictana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 675 (Bathrotoma). [NSW: Sydney, Parramatta.] Comb. nov. panolbia (Turner, 1946), Trans. R. Soc. S. Aust. 70: 200 (Ancylis). [Qld: Springbrook.] ruficomana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 676 (Bathrotoma). [NSW: Sydney.] Comb. nov. phaeoloma (Turner, 1946), Trans. R. Soc. S. Aust. 70: 203 (Eucosma). [Qld: Lake Barrine, Atherton Tableland.] acrosema (Turner, 1946), Trans. R. Soc. S. Aust. 70: 208 (Eucosma). [NSW: Brunswick Heads.] Comb. nov. quietana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 673 (Holocola). [Qld: Brisbane.] Comb. nov. ochronota (Turner, 1925), Trans. R. Soc. S. Aust. 49: 56 (Acroclita). [Qld: Townsville.]
Strepsicrates Meyrick (Figs 88, 739–759) Monilia Walker, 1866, List Specimens Lepid. Insects Colln Br. Mus. 35: 1741. Type species: Monilia semicanella Walker, 1866, ibid. 35: 1741, by monotypy. [Australia]
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Strepsiceros Meyrick, 1881, Proc. Linn. Soc. NSW 6: 678. Type species: Sciaphila ejectana Walker, 1863, List Specimens Lepid. Insects Colln. Br. Mus. 28: 350, by subsequent designation. [Australia.] A junior homonym of Strepsiceros Smith, 1827, in Griffith, Animal Kingdom … by the Baron Cuvier, with additional descriptions 5: 365 (Mammalia). Strepsicrates Meyrick, 1888, Trans. Proc. NZ Inst. 20: 73. Objective replacement name for Strepsiceros. Phthinolophus Dyar, 1903, Proc. Entomol. Soc. Wash. 5: 307. Type species: Phthinolophus indentatus Dyar, 1903, ibid. 5: 307, by monotypy. [USA.]
Diagnosis Strepsicrates species are inconspicuous, mediumsized, rather narrow-winged greyish or brownish eucosmine moths with a dorsal notch at some distance from the base of the antenna or close to its base (infensa-group) and with a costal fold as well as a hindwing anal fold containing a scale pencil in the male. Strepsicrates has moderately long (very long in the infensa-group), porrect, sinuate labial palpi, all veins separate in the forewing and M3 and CuA1 stalked in the hindwing. Species of the infensa-group can usually be superficially identified by their characteristic wing pattern and the conspicuously long labial palpi combined with separate R4 and R5 and a notch at the base of the antenna in the male. Unless recognised at the species level by wing pattern, the remaining Strepsicrates species can superficially be separated only tentatively on the basis of the forewing structure, namely a distally widened cubital cell and a tuft of raised scales on the dorsum in the male, from those Holocola species with separate R4 and R5 and an antennal notch distant from the base. However, the male genitalia are diagnostic for the genus with a triangular to digit-shaped ventral process on the cucullus closely associated with a single spine, a bipartite uncus often developed into strongly modified and widely separate arms (entirely reduced in a few species), reduced, small, membranous socii, and a large, barrel-shaped anellus. The female genitalia are characterised by a signum fused with the deeply concave hindmargin of S7, a short ductus bursae narrowed below the ostium by a split colliculum of thickened membrane, then widened, sclerotised and somewhat twisted to the origin of the ductus seminalis, followed by a short, membranous anterior portion usually with a pronounced bulge
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next to the sclerotised section. The corpus bursae is partially lined with sharp spinules and lacks signa.
Description Adults (Figs 739–742) Wingspan. 11–18 mm; medium-sized. Head (Figs 743–745). Ocellus well developed. Frons with a band- or horseshoe-shaped tuft of strongly projecting scales in upper part; scales on lower part small, appressed and upwardly directed. Maxillary palpus minute, scaled, 3-segmented. Proboscis c. 1.2× length of labial palpus. Labial palpus moderate to long (1.7–2.5× diameter of eye), generally longer in females and particularly large in infensa-group, sinuate, porrect, segmental ratio 1:2.5:0.8 (S. semicanella (Walker), ) and 1:2.8:1.5 (S. ejectana (Walker), ); second segment sinuate, subtriangular to teardrop-shaped, strongly widened by long, loose scales along ventral and distally along dorsal margin, terminal segment moderate to long, with apical vom Raths organ reaching to middle or below. Antenna to middle of forewing, dorsally scaled, ventrally with short cilia in male; flagellum with basal (infensa-group) or sub-basal notch in male (rest of genus): first six flagellar segments widened, laterally compressed and dorsally fused, with deep dorsal incision over segments 7–10, followed by a large dorsal spur on segments 10 and/ or 11 (Fig. 745). Thorax. Without crest. Legs unmodified. Wings (Figs 88, 739–742). Forewing index 0.32–0.37. Forewing long and relatively narrow, with costa and dorsum sub-parallel or only little divergent towards strongly oblique termen; male with well-developed costal fold often to beyond middle of costa containing conspicuously modified scent scales and, except for the infensa-group, with large patch of raised scales on dorsum near anal vein; cell c. 0.65× length of wing, M-stem and chorda visible only as trace in stained membrane; with all veins present and separate beyond cell; anal loop 0.35× length of 1A+2A; wing pattern diverse, often sexually dimorphic, variously greyish or brownish, usually with a triangular dark mark on dorsum before tornus and often paler along dorsum, with ocelloid patch recognisable at most as a series of parallel short black lines. Hindwing with cubital pecten and anal tuft well developed, in male with anal margin narrowly folded over pencil of long scales on upper surface and often (S. infensa (Meyrick), S. semicanella etc.) with modified, very long and slender scales on upperside of discal cell; with Rs and M1closely approximated at base; M2
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bent at base to close or moderately so to stalked M3 and CuA1; CuP present at margin, 1A+2A halfway between CuP and anal margin; 3A not visible in stained wing; greyish. Pregenital abdomen. Unmodified. In male hindmargin of S8 deeply bilobed; T8 with a transversely H-shaped sclerotisation. Male genitalia (Figs 746–753, 757–758). Vinculum narrow. Tegumen robust, pedunculi strong. Uncus, if present, always bipartite, but very variably developed: a median bifid uncus of two slender, curved points (S. macropetana (Meyrick)) or the two uncus arms developed into large pointed and hairy processes, arising widely separate (infensa-group, S. semicanella, S. ejectana), or no trace of lateral arms left. Socii small or vestigial, long and finger-like (S. macropetana) or round, hairy knobs (S. ejectana, S. semicanella) or merely lateral tufts of bristles (infensa-group). Gnathos arms usually two weakly sclerotised curved bands apically fused with anal tube, but with distal parts strongly developed in infensa-group. Valva very variable in shape, straight and medially little constricted (S. ejectana) to very long and thin with downcurved cucullus (infensa-group), but cucullus always with a triangular or digitate process from ventral margin followed by a prominent spine; costal process well developed. Caulis short and often wide, anellus large and barrel-shaped. Aedeagus hardly curved and rather short; vesica with numerous, spindle-shaped deciduous cornuti. Female genitalia (Figs 754–756, 759). Ovipositor and apophyses unremarkable. Sternum 7 with posterior margin strongly emarginate and fused with sterigma, sometimes with two large sharp thorns close to ventral angles of sterigma (S. infensa). Sterigma ranging from a small cup to a large, rectangular depression, rarely a triangular process from lamella antevaginalis; lamella antevaginalis usually a strongly sclerotised rim, lamella postvaginalis a sclerotised transverse band. Ductus bursae shorter than length of corpus bursae, narrowed below ostium by split colliculum of crescent-shaped, thickened membrane, sometimes asymmetrically sclerotised, then widened and sclerotised to inception of ductus seminalis, often somewhat twisted or with longitudinal folds and anteriorly often with some spinules, anteriormost portion near corpus bursae usually membranous, often with a bulge at end of sclerotised section. Corpus bursae round to teardrop-shaped; without signa but bursa lined with numerous, conspicuous, long and sharp spinules, sometimes reduced in certain areas.
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Distribution Strepsicrates is extremely widely distributed and has been reported from southern USA, British Guiana, Hawaii, Polynesia, Melanesia, Australia, New Zealand, New Guinea, Philippines, Indonesia, Malaysia, Thailand, China, Siberia, Japan, India, Sri Lanka, Saudi Arabia, Mauritius, Madagascar, and the Canary Is; in Australia the genus occurs throughout the mainland and Tasmania.
Biology Nearly all the hosts reported for Strepsicrates belong to the Myrtaceae: Eugenia and Psidium in the New World; Eucalyptus, Leptospermum and numerous other myrtaceous genera in the Australian/Oriental region. Heinrich (1923) reports Myrica cerifera (Myricaceae) as one of the hosts for the North American S. indentata (Dyar). The larvae of the Australian species are usually leaf-tiers (S. semicanella reported from Eucalyptus and Lophostemon, S. ejectana from Leptospermum and S. macropetana and S. infensa from Eucalyptus), but S. macropetana larvae have repeatedly been found tunnelling in eucalypt galls. Psidium guajava is mentioned as food plant of S. rhothia (Meyrick) in Mauritius, Sri Lanka and India. Both S. semicanella and S. macropetana cause problems in eucalypt nurseries, with several studies on S. macropetana (as the eucalyptus leafroller) from New Zealand (Mauchline et al. 2001).
Remarks Meyrick (1911b) synonymised Monilia and Strepsicrates with Spilonota Stephens, and later authors who considered Strepsicrates to be a valid separate genus did not realise that Monilia was the senior synonym. Given the economic relevance of Strepsicrates in eucalypt plantations and the large literature now linked with this name it would be unreasonable to resurrect Monilia, a name that has not been used since it was synonymised in 1911. The name Strepsicrates is therefore maintained in the interest of stability and universality. Strepsicrates has been repeatedly studied and numerous species have been illustrated, among others in contributions by Heinrich (1923), Clarke (1958, 1971, 1986), Diakonoff ([1968]), Zimmerman (1978), Razowski (1989) and Robinson et al. (1994). The monophyly of Strepsicrates is demonstrated by the unique combination of a triangular to digitshaped process ventrally on the cucullus next to a single prominent spine, and by the absence of signa,
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Figs 739–745. Strepsicrates, adults, heads and base of antenna. 739, 740, S. semicanella. 739, , Menangle Park, NSW. 740, , Maclean, NSW [image reversed]. 741, 742, S. ejectana. 741, , nr Braidwood, NSW. 742, , Grampians, Vic. [image reversed]. 743–745, S. ejectana, male, head and detail of antennal base with antennal notch (arrow), detail in ventral view. Scale = 2 mm.
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Figs 746–753. Strepsicrates, male genitalia and aedeagus. 746, 747, S. macropetana, Black Mt, ACT, slide T2132. 748, 749, S. semicanella, Brisbane, Q, slide T2156. 750, 751, S. ejectana, nr Perth, WA, slide ANIC 13736. 752, 753, S. limnephilana, nr Gympie, Q, slide T2241.
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a loss shared by Hermenias Meyrick and parts of Holocola Meyrick. Strepsicrates is also characterised by a bifid uncus either lost or increasingly developed into two widely separate arms, and by small, membranous, gradually reduced socii, in contrast to several related genera with their highly derived swollen and fuzzy (Holocola, Hermenias) or rigid, sclerotised socii (Spilonota Stephens, some groups of Holocola). No obvious sister group has been identified (apart possibly from the New Zealand Hendecasticha Meyrick, see below), but Strepsicrates and Spilonota share the presence of at least one large spine on the cucullus and modified scaling on the upper surface of the forewing discal cell in the males of some species. The strongly derived Hendecasticha from New Zealand, with a vein lacking in each wing and a notch at the base of the antenna, has only a single pair of tegumen processes, distantly spaced, strongly sclerotised, parallel rising arms tapering to a point that strongly recall the uncus arms in Strepsicrates. The cucullus in Hendecasticha ends in a ventral point but lacks the spine next to this process present in Strepsicrates. Some species in the triangulana-group
of Holocola also have an uncus somewhat recalling that of Hendecasticha, but it is always associated with well-developed socii and a differently shaped cucullus. But probably most importantly, the female genitalia of Hendecasticha, though they are not very informative and largely membranous, have the ductus seminalis originating from near the middle of the ductus bursae as in Strepsicrates and not from the neck of the corpus bursae as in the triangulana-group. Assuming that the tegumen processes do represent the uncus and based on the cucullus shape, Hendecasticha is tentatively treated as very close to Strepsicrates, but given the very derived structure of this minute insect molecular studies will probably be required to assess the relationship of the two genera. Strepsicrates is a large, widely distributed genus badly in need of revision. Reliable identification will require the study of ample material because of the sexual dimorphism in wing pattern and often subtle differences in the male genitalia of related species. In the Australian fauna, the shape of the sterigma is the most useful character for separating species. The scope of S. semicanella, the species
Figs 754–756. Strepsicrates, female genitalia. 754, S. macropetana, Black Mt, ACT, slide T2133. 755, S. semicanella, Rockhampton, Q, slide T2363. 756, S. ejectana, Wilton, NSW, slide ANIC 13739.
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most frequently associated with damage to eucalypts, is unclear even within Australia. A thorough study is needed to establish the range of this species that has possibly already been enlarged by eucalypt plantings throughout south-east Asia, and the presence in the region of other, related species attacking eucalypts.
Constituent species Brown (2005) lists 29 species of Strepsicrates, with Strepsicrates cryptosema Diakonoff from Saudi Arabia (Diakonoff 1983b) not convincingly assigned to this genus. Nine named and at least four unnamed species are known from Australia. macropetana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 683 (Strepsiceros). [Qld: Duaringa; NSW: Blackheath, Sydney, Parramatta, Bulli, Newcastle; Vic.: Melbourne.] brachytycha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 190 (Spilonota). [WA: Mt Dale.] liphaema (Turner, 1946), Trans. R. Soc. S. Aust. 70: 191 (Spilonota). [Qld: Toowoomba.] tornosema (Turner, 1946), Trans. R. Soc. S. Aust. 70: 196 (Spilonota). [WA: Mt Dale.] semicanella (Walker, 1866), List Specimens Lepid. Insects Colln. Br. Mus. 35: 1741 (Monilia). [Qld: Moreton Bay.]
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lasiophora (Lower, 1908), Trans. R. Soc. S. Aust. 32: 323 (Strepsiceros). [Qld: Townsville.] euploca (Turner, 1946), Trans. R. Soc. S. Aust. 70: 190 (Spilonota). [Qld: Brisbane.] dyselia (Turner, 1946), Trans. R. Soc. S. Aust. 70: 191 (Spilonota). [Qld: Cairns, Atherton, Townsville, Brisbane.] transfixa (Turner, 1946), Trans. R. Soc. S. Aust. 70: 206 (Eucosma). [Qld: Kuranda.] ebenocosma (Turner, 1946), Trans. R. Soc. S. Aust. 70: 203 (Eucosma). [Qld: Kuranda.] Comb. nov. ejectana (Walker, 1863), List Specimens Lepid. Insects Colln. Br. Mus. 28: 350 (Sciaphila). [NSW: Sydney.] servilisana (Walker, 1863), List Specimens Lepid. Insects Colln. Br. Mus. 28: 356 (Sciaphila). [New Zealand.] infimana (Walker, 1863), List Specimens Lepid. Insects Colln. Br. Mus. 28: 357 (Sciaphila). [New Zealand.] saxana (Walker, 1863), List Specimens Lepid. Insects Colln. Br. Mus. 28: 357 (Sciaphila). [New Zealand.] ligniferana (Walker, 1863), List Specimens Lepid. Insects Colln. Br. Mus. 28: 363 (Conchylis). [NSW: Sydney.] infimama (Walker, 1864), List Specimens Lepid. Insects Colln. Br. Mus. 29: 986 (Sciaphila). [New Zealand.]
Figs 757–759. Strepsicrates infensa, genitalia, Black Mt, ACT. 757, 758, aedeagus and male genitalia, slide T2124. 759, female genitalia, slide T2125.
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metabola (Turner, 1946), Trans. R. Soc. S. Aust. 70: 191 (Spilonota). [WA: Denmark.] zophotypa (Turner, 1946), Trans. R. Soc. S. Aust. 70: 192 (Spilonota). [WA: Denmark.] mesosticha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 194 (Spilonota). [WA: Albany.] subpallida (Turner, 1946), Trans. R. Soc. S. Aust. 70: 195 (Spilonota). [Qld: Stradbroke I.] igniferana; McQuillan, 1992, Pap. Proc. R. Soc. Tasmania 126: 86 (Strepsicrates). [Misspel.] sphenophora (Turner, 1946), Trans. R. Soc. S. Aust. 70: 190 (Spilonota). [NSW: Brunswick Heads.] Sp. rev. limnephilana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 680 (Strepsiceros). [NSW: Waratah on the Hunter R.] infensa (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 228 (Spilonota). [Qld: Brisbane.]
Holocola Meyrick (Figs 89–91, 760–788) Holocola Meyrick, 1881, Proc. Linn. Soc. NSW 6: 635 (key), 669. Type species: Holocola thalassinana Meyrick, 1881, ibid. 6: 670 (key), 672, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 43, 61. [Australia.] Allohermenias Diakonoff, 1953, Verh. K. Ned. Akad. Wet. (2) 49 (3): 88 (key), 146. Type species: Allohermenias tenuitexta Diakonoff, 1953, ibid. (2) 49 (3): 146, figs 356, 359, by original designation. [New Guinea.] Syn. nov. Neohermenias Diakonoff, 1966, Zool. Verh. (Leiden) 85: 73. Type species: Hermenias scoliomelas Diakonoff, 1953, Verh. K. Ned. Akad. Wet. (2) 49 (3): 122 (key), 125, figs 337, 338, 342, by original designation. [New Guinea.] Syn. nov.
Diagnosis Species of Holocola are rather narrow-winged, very diversely marked and sexually dimorphic eucosmines with a triangularly expanded apical region and a scale pencil beneath the upturned anal margin of the hindwing in the male, with a costal fold present or absent and an antennal notch either present near the base, at some distance from the base or absent. Superficially, only individual species or some species-groups may be recognised by their
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wing pattern as there are no generically diagnostic external characters separating Holocola from related genera. Holocola species have moderately long to long, porrect and sinuate labial palpi, a forewing with a narrow discal cell and R4 and R5 separate or stalked but without raised scales on the dorsum in the male in contrast to Strepsicrates Meyrick, and a hindwing with Rs and M1 separate and M3 and CuA1 stalked or fused into a single vein. Some subgroups of Holocola can be recognised by externally visible apomorphies such as a very long anal pencil protruding beyond the hindwing, usually associated with long scales from the anal margin, or modified scales on the upper surface of the costal fold or the base of the costa. The male genitalia are diagnostic for the genus, with a robust tegumen with very slender pedunculi at base, a variable bifid or single uncus often entirely reduced in the plinthinana-group, with modified socii ranging from large inflated structures with dense fuzz on their upper surface to sclerotised, scarcely bristled, flat lobes (plinthinana-group), a slender valva strongly narrowed between sacculus and cucullus with at most one to few thorns on the cucullus but never with a long, prominent spine and with the angle of the sacculus sometimes produced into a triangular to tongue-shaped process, the cucullus downcurved, from trapezoidal to subtriangular to broadly recurved or narrowly paddle-shaped, the caulis with wide lateral flanges and, most importantly, the anellus with a prominent, often hood-shaped dorsal lobe. Development of the dorsal region of the anellus to a lobe longer than the lateral parts is the best diagnostic feature for Holocola. Separation of Spilonota Stephens and the plinthinana-group, both with flat, sclerotised socii and often with M3 and CuA1 entirely fused in the hindwing, is facilitated by the position of the socii, from the inner side of the tegumen ‘shoulders’ in Spilonota but more laterally in the plinthinana-group, and by the presence of a series of small thorns along the margin of the broad, subtriangular cucullus in the plinthinana-group rather than a single, prominent spike or thorn on a finger-like process in Spilonota. The female genitalia have the sterigma fused with the hindmargin of S7, a sclerotised funnel with its ventral margin often formed by a projecting lobe from the hindmargin of S7. Sternum 7 sometimes has crests, humps or long thorns anterolaterally to the ostium. The ductus bursae is constricted below the ostium and variably sclerotised (entirely membranous in thalassinana-group) to the origin of the ductus seminalis, and the membranous
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section between the ductus seminalis and the corpus bursae is usually very short with two scobinate depressions, more rarely (thalassinanagroup) long and with a single sclerotised patch at the entrance to the corpus bursae or the ductus seminalis originates from the corpus bursae together with the ductus bursae (triangulanagroup). The corpus bursae is partially spinulose and usually lacks signa, but two horn-shaped signa are present in the plinthinana-group and a few other taxa.
Description Adults (Figs 760–769) Wingspan. 8–27 mm; small to large. Head (Fig. 788). Ocellus well developed. Vertex with two moderate, lateral tufts of scales with their tips medially convergent. Frons with transverse tuft of long, anteriorly projecting scales in upper part, remaining scales minute, appressed. Maxillary palpus minute, scaled. Proboscis c. 1.5× length of labial palpus. Labial palpus moderate to long (1.9– 2.9× diameter of eye), sinuate, porrect; second segment sinuate, teardrop-shaped, strongly widened in distal half by long, loose scales along ventral margin and sometimes also along distal half of dorsal margin, terminal segment moderate to long and slender, sometimes concealed in scales from second segment. Antenna to beyond middle of forewing, dorsally scaled, ventrally with minute to fairly long (close to diameter of flagellum) cilia in male; flagellum often with notch, either close to base or at some distance; very rarely base of flagellum dorsoventrally compressed rather than notched. Thorax. Without crest. Legs unmodified. Wings (Figs 89–91, 760–769). Forewing index 0.27–0.37. Retinaculum in male a hook from subcosta, in female a row of large scales behind cubitus. Forewing usually long and slender with costa little curved and roughly parallel to dorsum, termen usually lightly sinuate and more or less oblique with apex slightly projecting, rarely costa strongly curved in distal half and apex nearly falcate (H. chlidana (Turner)); male with or without costal fold varying from very small to reaching to middle of costa, often with modified scales beneath costal fold and sometimes also upperside of costal fold or base of costa with modified scaling; discal cell long and slender, accessory cell narrow, M-stem and chorda both present but sometimes only weak traces in membrane; all veins separate beyond cell or R4 and R5 stalked, M3 and CuA1 converging
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towards margin; CuP present near margin; basal loop 0.25–0.31× length of 1A+2A; wing pattern very diverse and often strongly sexually dimorphic, nearly always with costal strigulae visible at least in distal half and with a dark apical spot, often with a dark triangular mark on dorsum before tornus, ocelloid patch often indicated by some parallel black lines. Hindwing with cubital pecten and axillary tuft developed; males with anal region expanded triangularly usually forming a projecting angle, anal margin narrowly folded upward over scale pencil from base of wing, rarely with a fringe of long scales from anal margin, or hindwing except for apex with modified scales, either dense and whitish or very narrow and distantly spaced; Rs and M1 parallel and closely approximated at base, M2 curved towards base of M3, M3 and CuA1 stalked or entirely fused (many species in the plinthinanagroup); CuP present at margin halfway between CuA2 and 1A+2A, the latter distant from anal margin, weak trace of 3A sometimes visible in anal fold; whitish to dark grey, anal pencil often conspicuously dark. Pregenital abdomen. Unmodified. In male hindmargin of S8 bilobed, T8 with a transverse sclerotised bar at base and sometimes with a weak, medial, V-shaped sclerotised area. Male genitalia (Figs 770–781). Upper half of tegumen robust, pedunculi tapering to slender base. Uncus variable, from two slender, short, curved points joined at base to broadly spatulate and distally bifid, or a single median process varying from broadly paddle-shaped or capitate to a reduced stump, or uncus entirely lost in some species of the plinthinana-group; uncus more rarely two widely separated processes (triangulana-group) or two parallel, curved, flat blades (H. pellopis (Turner)). Socii always present and modified, mostly large, inflated, three-dimensional structures with dense fuzz on upper surface, sometimes with a sclerotised ventral rim or produced apex or entirely sclerotised and with only a few bristles, either threedimensional or flat lobes (plinthinana-group). Gnathos usually two weak bands, rarely base of each arm conspicuously sclerotised. Valva long and slender, of variable shape but always strongly narrowed between sacculus and cucullus, sometimes with one to few thorns on cucullus margin but never with a long, prominent spine, with large basal excavation and slender costal process; angle of sacculus from obtuse to produced into triangular to tongue-shaped process, cucullus separated by conspicuous and often long ‘neck’, downcurved, from trapezoidal to subtriangular to
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Figs 760–765. Holocola, adults. 760, 761, H. deloschema, Clyde Mt, NSW. 760, . 761, . 762, 763, H. spodostola, Stirling Range, WA. 762, . 763, . 764, 765, H. sp. nr pachnitis, Mt Bellenden-Ker, Q. 764, [image reversed]. 765, . Scale = 2 mm.
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broadly recurved or narrowly paddle-shaped. Juxta small, subtriangular, caulis moderately long with wide lateral flanges, anellus with prominent dorsal lobe, tongue- to hood-shaped. Aedeagus from nearly straight to apex strongly downwardly bent; usually with a sheaf of lanceolate, deciduous cornuti. Female genitalia (Figs 782–787). Ovipositor lobes usually subreniform, ventrally narrower, more rarely broadly ovate or narrowly crescentic. Sterigma fused with hindmargin of S7, forming a variably shaped funnel leading to ostium, its ventral margin often formed by a projecting lobe from hindmargin of S7. Sternum 7 sometimes with crests, humps or long thorns anterolaterally to ostium. Ductus bursae narrowed below ostium by
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colliculum of two small crescentic lateral thickenings of wall, usually not sclerotised, followed by variably sclerotised central portion to origin of ductus seminalis, often with more strongly sclerotised longitudinal folds and sometimes divided by a more membranous area into an often only weakly sclerotised posterior section and a short, strongly sclerotised anterior portion; in the triangulana-group a lateral sclerite only in anteriormost portion; membranous section between origin of ductus seminalis and corpus bursae variable, from very short (rarely with a small projecting pouch) to long and gradually widening, usually with two scobinate depressions or with a single sclerotised patch (thalassinana-
Figs 766–769. Holocola, adults, males. 766, H. plinthinana, Niagara Park, NSW. 767, H. sp. triangulana-group, nr Braidwood, NSW. 768, H. nitida, Tinderry Mts, NSW. 769, H. sp. nr hypomolybda, nr Hotel Kosciusko, NSW [image reversed]. Scale = 2 mm.
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Figs 770–777. Holocola, male genitalia and aedeagus. 770, 771, H. nitida, nr Nerriga, NSW, slide T2401. 772, 773, H. nr thalassinana, Como, NSW, slide ANIC 12966. 774, 775, H. deloschema, Mt Bartle Frere, Q, slide T2395. 776, 777, H. triangulana, Black Mt, ACT, slide T2144.
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group) at entrance to corpus bursae; more rarely (triangulana-group) ductus seminalis originating directly from corpus bursae next to ductus bursae. Corpus bursae round to ovate, with a broad band of spinules from entrance to apex and back to entrance, leaving two smooth or at most dimpled areas occupied each by a horn-shaped signum, if signa present (plinthinana-group and some other species); signa absent in most species.
Distribution India, Indonesia, New Guinea, Samoa and Australia, where the genus occurs throughout the continent with strong emphasis on the southern half and Tasmania, in particular at higher altitude locations. In India, Indonesia and New Guinea the genus is generally found at higher altitudes, up to 3300 m in New Guinea (Clarke 1958: as Spilonota; Diakonoff 1954: as Hermenias).
Biology The majority of host records for Holocola from Australia involve binding leaves on Myrtaceae, with
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several species reared from Leptospermum (H. thalassinana (Meyrick), H. fluidana (Meyrick), H. honesta (Meyrick), H. zopherana (Meyrick)), several undescribed species reared by J. Balciunas from Melaleuca and one species near H. hypomolybda (Turner) from Eucalyptus pauciflora. Species of the thalassinana-group have also been reared from a seed capsule of Eucalyptus maculosa on Black Mt, ACT, and from flowers of Melaleuca quinquenervia in Ballina, NSW. However, a species near H. obeliscana (Meyrick) (plinthinanagroup) has been reared from a larva binding shoots of Melichrus urceolatus (Epacridaceae) by I. F. B. Common in Canberra, and H. triangulana and two other species of this group have repeatedly been reared from Acacia species (Mimosaceae) (ANIC).
Remarks Holocola has several conspicuously homoplasious external characters that are traditionally accorded considerable taxonomic importance such as presence or absence of costal fold and antennal notch, and a venation with R4 and R5 either separate or stalked and M3 and CuA1 in the
Figs 778–781. Holocola, male genitalia with aedeagus. 778, H. spodostola, Margaret R., WA, slide ANIC 13018. 779, H. pericyphana, Killarney, Q, slide T2397. 780, H. plinthinana, nr Braidwood, NSW, slide ANIC 13016. 781, H. pellopis, Buringbah, NSW, slide ANIC 13029.
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Figs 782–787. Holocola, female genitalia. 782, H. plinthinana, nr Braidwood, NSW, slide T2233. 783, H. sp. nr thalassinana, Richmond R., NSW, slide T2390. 784, H. spodostola, Stirling Range, WA, slide ANIC 13011. 785, H. sp. nr pachnitis, Bellenden-Ker, Q, slide T2404. 786, H. deloschema, Bellenden-Ker, Q, slide ANIC 13007. 787, H. sp. nr triangulana, nr Broome, WA, slide ANIC 13009.
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Fig. 788.
Holocola triangulana, head, male.
hindwing stalked or entirely fused. Outgroup comparison leaves little doubt that presence of both costal fold and antennal notch pertains to the groundplan of Holocola, together with R4 and R5 separate and M3 and CuA1 both present and stalked. Several obviously monophyletic groups defined by both male and female genitalia demonstrate that all these characters have repeatedly changed within the genus to a derived configuration, i.e. secondary absence of the relevant structures through loss, or stalking and fusion of veins. Only a cladistic analysis far beyond the scope of this study will convincingly resolve the relationships of the numerous Holocola species. Based on present evidence the type species of Holocola, Allohermenias and Neohermenias all belong to groups subordinate within a larger entity that defies subdivision into monophyletic groups and is here treated as a single genus, with Holocola the oldest available name. The identity of H. thalassinana, the type species of Holocola, has not yet been conclusively established with several closely related and strongly sexually dimorphic species as possible candidates. Meyrick named several Holocola species and some of the types have been figured by Clarke (1958; as Spilonota). Diakonoff (1953) described and figured 19 species of Holocola from the mountains of New Guinea as Hermenias, all later transferred to his new genus Neohermenias (Diakonoff 1966), and two species as Allohermenias and one as Bathrotoma. More recently, Diakonoff (1983a) described or redescribed and figured two new and one named Holocola species from Atjeh, Sumatra, also as Hermenias. The monophyly of Holocola is based on the dorsally elongated, tongue- to hood-shaped anellus
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and the presence of two granulate depressions in the ‘neck’ of the corpus bursae. The latter are absent in the triangulana-group where the anteriormost portion of the ductus bursae is reduced and the ductus seminalis originates directly from the corpus bursae, and in the thalassinana-group, where this portion has become greatly lengthened and contains a single sclerotised patch rather than two granulate depressions. As these absences coincide in both cases with a generally derived ductus bursae they are considered to be secondary derivations. Based on genitalia morphology (most importantly shape of uncus, socii and cucullus) and some external features such as wing pattern or an elongated anal pencil, several quite distinct speciesgroups can be identified within Holocola, but they do not at present allow subdivision of the genus as their removal would leave a paraphyletic assemblage. The present arrangement in the checklist below tries to group closely related species but is no doubt partially artificial as for example quite diverse taxa all characterised by the plesiomorphic presence of signa in the female are heading the list. Whilst presence or absence of an antennal notch is uninformative with the loss having occurred repeatedly, the position of the notch (near the base or at some distance from it, without any intermediate states) seems taxonomically much more relevant. The antennal notch is near the base in Spilonota, Hermenias, part of Strepsicrates and part of Holocola, and distant from the base in most Strepsicrates and part of Holocola. Presence of signa and a notch at the base are generally correlated and both these states are here assumed to be plesiomorphies, with the notch having independently moved to a more distal position within both Holocola and Strepsicrates. The plinthinana-group, with longitudinal pattern elements forming paler costal and dorsal regions separated by a dark streak from base to apex, is well characterised by rigid, flat and often only sparsely bristled socii, a rather small, single uncus that may be entirely reduced and a tendency for M3 and CuA1 in the hindwing to be fused into a single vein. These derived features are combined with presence of signa in the female, a notch at the base of the antenna in the male and separate R4 and R5. The Papuan species described in Allohermenias by Diakonoff (1953) could on the basis of their male genitalia belong to the plinthinana-group though they lack the antennal notch and have R4 and R5 stalked. A small group of Spilonota constrictana look-alikes without any named species also has the
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signa present, the antennal notch at the base and R4 and R5 separate, but is characterised by a Y-shaped uncus and a modified S7. The Acacia-feeding triangulana-group has two usually pointed and widely separated uncus arms (Fig. 776), and in the female the ductus bursae is strongly modified with an anterior lateral sclerotisation only and with the ductus seminalis arising directly from the corpus bursae. The type of Neohermenias belongs to a strongly sexually dimorphic group of species centred on tropical upland localities in New Guinea and Australia with large, inflated, flimsy socii, a distally bifid uncus, no signa, a large costal fold, the antennal notch at some distance from the base and with stalked R4 and R5. The type species of Holocola, also with R4 and R5 stalked but without antennal notch, has rather similar male genitalia though with a variably produced sacculus angle, while the female genitalia are derived with a largely membranous ductus bursae and the portion between ductus seminalis and corpus bursae lengthened and with a sclerotised patch instead of two scobinate depressions. The synonymies in the checklist below are preliminary only and will have to be reassessed in a future in-depth revision.
Constituent species About 30 species of Holocola are described from outside Australia, mainly from the mountains of New Guinea, and 42 named and many more unnamed species are known from Australia. chlidana (Turner, 1927), Pap. Proc. R. Soc. Tasmania 1926: 126 (Epiphyas). [Tas.: Rosebery.] pellopis (Turner, 1946), Trans. R. Soc. S. Aust. 70: 196 (Spilonota). [NSW: Brunswick Heads.] tapina (Turner, 1946), Trans. R. Soc. S. Aust. 70: 211 (Eucosma). [NSW: Brunswick Heads.] sollicitana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 687 (Strepsiceros). [NSW: Sydney, Parramatta.] callimita (Turner, 1946), Trans. R. Soc. S. Aust. 70: 189 (Hermenias). [Qld: Toowoomba.] obeliscana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 690 (Strepsiceros). [NSW: Sydney, Parramatta.] perfixa (Turner, 1916) Trans. R. Soc. S. Aust. 40: 525 (Eucosma). [NSW: Sydney.] ochronephes (Turner, 1946) Trans. R. Soc. S. Aust. 70: 192 (Spilonota). [Qld: Brisbane, Stradbroke I., Toowoomba, Stanthorpe, Coolangatta; NSW: Brunswick Heads.]
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euryptycha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 192 (Spilonota). [NSW: Brunswick Heads.] leucopleura (Turner, 1946), Trans. R. Soc. S. Aust. 70: 208 (Eucosma). [Qld: Stanthorpe, Brunswick Heads.] sicariana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 691 (Strepsiceros). [NSW: Sydney.] peraea (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 247 (Eucosma). [Vic.: Sale.] plinthinana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 689 (Strepsiceros). [NSW: Parramatta.] vitiosa (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 230 (Spilonota). [SA: Mt Lofty.] zopherana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 688 (Strepsiceros). [NSW: Sydney; lectotype designated by Dugdale (1988: 117).] clastomochla (Turner, 1946), Trans. R. Soc. S. Aust. 70: 193 (Spilonota). [WA: Albany, Nornalup.] euthytoma (Turner, 1946), Trans. R. Soc. S. Aust. 70: 194 (Spilonota). [Tas.: Strahan.] stenophylla (Turner, 1946), Trans. R. Soc. S. Aust. 70: 196 (Spilonota). [WA: Albany, Denmark.] poliophylla (Turner, 1946), Trans. R. Soc. S. Aust. 70: 197 (Spilonota). [WA: Denmark.] niphosticha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 195 (Spilonota). [WA: Perth.] tranquilla (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 233 (Spilonota). [Tas.: Deloraine.] mediocunea (Turner, 1946), Trans. R. Soc. S. Aust. 70: 193 (Spilonota). [Tas.: Waratah.] triangulana Meyrick, 1881, Proc. Linn. Soc. NSW 6: 670 (Holocola). [Vic.: Melbourne.] opsia (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 247 (Eucosma). [SA: Port Lincoln.] phaeoscia (Turner, 1916), Trans. R. Soc. S. Aust. 40: 524 (Eucosma). [Qld: Nambour.] leuconota (Turner, 1946), Trans. R. Soc. S. Aust. 70: 208 (Eucosma). [NSW: Brunswick Heads.] chalcitis (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 231 (Spilonota). [Tas.: Blue Tier.] periptycha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 201 (Eucosma). [WA: Albany.] aethalostola (Turner, 1946), Trans. R. Soc. S. Aust. 70: 190 (Bathrotoma). [WA: Tammin.] honesta (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 233 (Spilonota). [Vic.: Geelong, Macedon, Melbourne; Tas.: Deloraine.] leuconephela (Turner, 1925), Trans. R. Soc. S. Aust. 49: 56 (Eucosma). [NSW: Barrington
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Tops.] Nec (Turner, 1946), Trans. R. Soc. S. Aust. 70: 205 (Eucosma). nitida (Turner, 1946), Trans. R. Soc. S. Aust. 70: 201 (Eucosma). [NSW: Mittagong.] spanistis (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 232 (Spilonota). [NSW: Mt Kosciusko.] argyrotypa (Turner, 1927), Pap. Proc. R. Soc. Tasmania 1926: 135 (Spilonota). [Tas.: Cradle Mt] ebenostigma (Turner, 1946), Trans. R. Soc. S. Aust. 70: 191 (Spilonota). [Vic.: Mt Buffalo.] leucopyga (Turner, 1946), Trans. R. Soc. S. Aust. 70: 194 (Spilonota). [NSW: Mt Kosciusko.] fluidana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 686 (Strepsiceros). [NSW: Sydney, Bulli, Blackheath.] striphromita (Turner, 1946), Trans. R. Soc. S. Aust. 70: 202 (Eucosma). [NSW: Brunswick Heads.] polymita (Turner, 1946), Trans. R. Soc. S. Aust. 70: 206 (Euosma [sic]). [NSW: Brunswick Heads.] polyplega (Turner, 1946), Trans. R. Soc. S. Aust. 70: 208 (Eucosma). [Qld: Caloundra.] ammostigma (Turner, 1946), Trans. R. Soc. S. Aust. 70: 193 (Spilonota). [Qld: Stanthorpe.] baeodes (Turner, 1946), Trans. R. Soc. S. Aust. 70: 195 (Spilonota). [Qld: Brisbane.] hypomolybda (Turner, 1927), Pap. Proc. R. Soc. Tasmania 1926: 135 (Spilonota). [Tas.: Launceston.] notosphena (Turner, 1946), Trans. R. Soc. S. Aust. 70: 192 (Spilonota). [Qld: Springbrook.] diplostigma (Turner, 1946), Trans. R. Soc. S. Aust. 70: 193 (Spilonota). [Qld: Brisbane.] tarachodes (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 231 (Spilonota). [Tas.: Mt Wellington.] pericyphana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 685 (Strepsiceros). [NSW: Sydney.] deloschema (Turner, 1914), Proc. Linn. Soc. NSW 39: 554 (Spilonota). [NSW: Ebor.] catapasta (Turner, 1915), Proc. Linn. Soc. NSW 40: 192 (Bathrotoma). [NSW: Ebor.] liturata (Turner, 1925), Trans. R. Soc. S. Aust. 49: 56 (Acroclita). [Qld: Charleville.] melanographa (Turner, 1916), Trans. R. Soc. S. Aust. 40: 523 (Bathrotoma). [NSW: Hornsby, near Sydney.] atmophanes (Turner, 1946), Trans. R. Soc. S. Aust. 70: 194 (Spilonota). (Qld: Toowoomba, Bunya Mts.] seditiosana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 684 (Strepsiceros). [NSW: Sydney, Bulli Pass.]
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morosa (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 232 (Spilonota). [NSW: Bathurst; Tas.: Deloraine.] perspectana (Walker, 1863), List Specimens Lepid. Insects Colln. Br. Mus. 28: 393 (Grapholita). [Qld: Moreton Bay.] thalassinana Meyrick, 1881, Proc. Linn. Soc. NSW 6: 672 (Holocola). [NSW: Sydney; Newcastle.] ischalea (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 240 (Acroclita). [NSW: Sydney, Newcastle.] spodostola (Turner, 1946), Trans. R. Soc. S. Aust. 70: 189 (Bathrotoma). [WA: Margaret R.] commatica (Turner, 1946), Trans. R. Soc. S. Aust. 70: 199 (Acroclita). [WA: Collie.] lucifera (Turner, 1946), Trans. R. Soc. S. Aust. 70: 195 (Spilonota). [Tas.: Cradle Mt.] imbrifera (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 226 (Hermenias). [Vic.: Gisborne.]
Hermenias Meyrick (Figs 92, 789–800) Hermenias Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 225. Type species: Hermenias epidola Meyrick, ibid. 36: 225, by original designation. [Australia.] Herminias Lower, 1896, Victorian Naturalist 12: 149. Nomen nudum.
Diagnosis Hermenias species are medium-sized to large, narrow-winged grey olethreutines of somewhat crambine-like appearance with inconspicuous markings formed by small dark longitudinal striae often confluent into an oblique dark streak from base to apex. Hermenias has long, narrow and porrect labial palpi and the males are characterised by usually conspicuously ciliate, basally notched antennae and a dark hair pencil in the anal fold of the hindwing. The long costal fold reaching to 36: of the wing, the very long discal cell distally narrowed beyond the short and steeply angled R2, and the veins R3 with the stalked R4 and R5 and M1 all arising closely together from the anterior angle of the discal cell are, as a combination, diagnostic for Hermenias. The socii in the male genitalia are rigid and dorsocaudally covered with dense, fine, fur-like hair as in Holocola Meyrick, they may also be similarly large and inflated, but the uncus in Hermenias is always single, never
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bipartite or apically cleft. The female genitalia with the bursa wall covered with long sharp spinules are typically those of a member of the Spilonota-group. Absence of signa and a characteristic sterigma formed by a projecting, rounded median lobe from the hindmargin of S7 are shared with Holocola, but Hermenias is characterised by its ductus bursae with a split colliculum below the ostium and a sclerotised area with some spinules between the origin of the ductus seminalis and the bursa neck. Hermenias lacks the paired, slightly depressed patches of conspicuous granulation at the neck of the corpus bursae typical for Holocola.
Description Adults (Figs 789–792) Wingspan. 14–23 mm; medium-sized to large. Head (Fig. 793–795). Ocellus moderate. Frons with dense long scales in upper part, projecting as a broad triangular wedge with labial palpi appressed to its ventral surface. Maxillary palpus minute, scaled, appressed, 3-segmented. Proboscis as long as labial palpus. Labial palpus very long (1.9–3.2× diameter of eye), slender, weakly sinuate, porrect or drooping, segmental ratio 1:4:1.8; second segment large, sinuate, widest in middle; terminal segment long, thin, with apical vom Raths organ to below middle. Antenna reaching to beyond middle of forewing; flagellum with deep basal notch in male: dorsal excision in second and third segment followed by dorsal spur on third and fourth segments (Figs 794, 795); with whorls of conspicuous cilia in male on ventral surface at least at base of flagellum, 0.5–1.0× flagellar diameter or longer. Thorax. Without crest. Legs unmodified. Wings (Figs 92, 789–792). Forewing index 0.25–0.28. Forewing very long and narrow, costa and dorsum parallel or wing distally slightly widened, with costa evenly curved, apex obtusepointed, termen strongly oblique; male with extremely long costal fold to 36:; discal cell very long (0.73 length of wing in a typical species), distally narrowed beyond R3, chorda and M-stem visible only as trace in stained wing; R2, R3 and distal part of R1 parallel and running steeply to costa; R4 and R5 stalked, closely approximated to R3 and M1 at base; CuP present near margin; anal loop 0.27× length of 1A+2A; wing pattern of small, grey, longitudinal striae, often partly confluent, especially along an oblique line from wing base to apex, rarely with prominent white subapical strigulae along costa. Hindwing with cubital
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pecten, with axillary scale tuft and with anal margin in male narrowly folded over large pencil of long, dark grey scales from small process at base; with Rs and M1 parallel and closely approximated in basal half, M2 at base close to stalk of M3 and CuA1; CuP halfway between CuA2 and 1A+2A, the latter distant from anal margin; 3A not present (unless within anal fold and concealed by scale pencil); pale grey. Pregenital abdomen. Unmodified. Sternum 8 in male medially with a U- to V-shaped excision in hindmargin. Male genitalia (Figs 796–798). Vinculum a moderate band, articulated with tegumen. Tegumen high, with slender pedunculi. Uncus a single, small median process, parallel-sided or apically slightly expanded. Socii smallish to moderate, rigid and three-dimensional, with the lateral edge produced blade-like and the dorsocaudal surface covered with a mat of very dense, long, fur-like hair. Gnathos hardly discernible. Valva with large basal excavation, long and narrow, strongly constricted into more or less curved neck beyond middle then abruptly expanded into subtriangular cucullus with a row of short, strong spines along distal margin; rarely one of the apical spines greatly enlarged (H. epidola). Caulis moderate, anellus a broad ring. Aedeagus slender and little curved, with small coecum; vesica with a cluster of deciduous, lanceolate cornuti. Female genitalia (Figs 799–800). Ovipositor lobes large, semicircular or ovate, hardly narrowed ventrally; apophyses short. Sternum 7 sometimes with traces of two warts anterolaterally of sterigma, with strongly projecting rounded median lobe from hindmargin forming lamella antevaginalis of sterigma; entrance to ostium a flat, funnel-shaped pocket covered by posteromedial lobe of S7. Ductus bursae membranous except for a strongly sclerotised and often very long colliculum just below ostium and a slightly widened, sclerotised, irregularly folded and partially spinulose anteriormost part between corpus bursae and origin of ductus seminalis. Corpus bursae ovate to round, inner surface covered with long sharp spinules except towards bursa neck; signa absent.
Distribution Hermenias is only known from the southern half of Australia, from New South Wales to Western Australia, including Tasmania, mainly at higher altitudes up to 1700 m.
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Figs 789–795. Hermenias, adults, head and antennal notch. 789, 790, H. rivulifera. 789, , nr Bateman’s Bay, NSW. 790, , Gibraltar Range Nat. Park, NSW. 791, 792, H. sp. E, Black Mt, ACT. 791, . 792, . 793–795, H. sp., male. 793, head. 794, antennal notch, dorsal side. 795, antennal notch, ventral side. Scale = 2 mm.
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Biology A single rearing record ‘larva binding terminal leaves of Eucalyptus coccifera’ (Myrtaceae) and one pupa found ‘beneath loose bark of Eucalyptus’ and one ‘among snow grass’ are the only food plant indications (ANIC).
Remarks Adult and genitalia of H. epidola have been figured by Clarke (1958), but the many Papuan species whose genitalia were illustrated by Diakonoff (1953) as Hermenias species were subsequently transferred to Neohermenias Diakonoff (Diakonoff 1966), a junior synonym of Holocola. Hermenias is closely related to Protithona Meyrick from New Zealand, with both genera sharing extremely narrow forewings with a very long discal cell and similar male genitalia with inflated socii and an undivided uncus. Though Protithona looks superficially quite distinct, much smaller, with very different wing pattern and one
vein lacking in each wing, much of the difference between the two genera is due to the extremely derived nature of Protithona and probably partly correlated with its small size. The two genera doubtlessly are sister groups and it cannot be entirely ruled out that Protithona arose from within Hermenias. The evidence is inconclusive at present and in the interest of taxonomic stability the two genera are here treated as distinct. The extremely long costal fold of Hermenias and the crumpled sclerite anteriorly in its ductus bursae are considered apomorphies for the genus that separate it from Protithona. The whorls of long cilia ventrally on the male antenna of Hermenias are a further apomorphy of the genus, present in all species but reduced to short remnants in epidola with its strongly derived cucullus. As indicated by the many Papuan species of Holocola initially described as Hermenias by Diakonoff (1953), the two genera share several apomorphies such as an antennal notch, swollen socii covered with fuzzy fur-like hair, similar
Figs 796–799. Hermenias, genitalia. 796, 797, H. rivulifera, male genitalia and aedeagus, Black Mt, ACT, slide T2252. 798, H. epidola, male genitalia, Swansea, T, slide ANIC 12993. 799, H. rivulifera, female genitalia, Black Mt, ACT, slide T2253.
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cucullus shape, a lobe-like projecting lamella antevaginalis and a spinulose corpus bursae without signa. However, Holocola is characterised by two granulose dimples at the neck of the corpus bursae (absent in few species), a tongue-shaped dorsal extension of the anellus, a forewing of average width with a much shorter (underived) costal fold than Hermenias and, at least in plesiomorphic species, a bifid uncus. Meyrick (1911b) described H. epidola from three syntypes of which two are still present in the BMNH, a male from Deloraine (Tasmania) and a female from Gisborne (Victoria). No lectotype has been previously designated. The two remaining syntypes are apparently not conspecific, and given the paucity of characters in the female genitalia to distinguish species in contrast to the distinctive male genitalia it has been decided to designate the Deloraine male as a lectotype. This male with the label data ‘Deloraine, Tasmania, 26/11/82’ (in Meyrick’s handwriting) is here designated as the lectotype. The Gisborne female had previously been labelled as an informal lectotype. The Deloraine
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male lacks the code numbering usually present on Meyrick’s BMNH material. Tuck (personal communication 2005) suggested that ‘The absence of a specimen code number implies that Diakonoff had probably borrowed it at the time when the other material of that group in Meyrick’s collection was labelled up, so it was missed in the allocation of code numbering’. Clarke (1958: pl. 210) figured this specimen apparently without realising that it was a syntype. Lower’s nomen nudum, epidolana (Lower, 1896): 149 (Herminias), strictly represents the now nameless population from Gisborne (Victoria).
Constituent species None of the Hermenias species described from outside Australia appear to be congeneric with H. epidola, nor are the more recently described species from Sumatra (Diakonoff 1983a), Thailand (Kawabe 1995) and Taiwan (Razowski 2000). Some 8–10 species from Australia are represented in the ANIC, only two of which are named: epidola Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 225 (Hermenias). Lectotype ‘Deloraine, Tasmania, 26/11/82’ [in Meyrick’s handwriting], ‘Hermenias epidola Meyr., E. Meyrick det. in Meyrick Coll.’, ‘ genitalia on slide 21.vi.1949 JFGC 9563’ is hereby designated. [Tas.: Deloraine.] rivulifera Turner, 1946, Trans. R. Soc. S. Aust. 70: 189 (Hermenias). [NSW: Maryland, near Stanthorpe.]
Eccoptocera Walsingham (Figs 93, 801–805) Eccoptocera Walsingham, 1907, Fauna Hawaii 1(Part V): 673. Type species: Steganoptycha foetorivorans Butler, 1881, Ann. Mag. Nat. Hist. (5. series): 394, by monotypy and original designation. [Hawaii.] Alcina Clarke, 1976 Insects Micronesia 9(1): 21. Type species: Alcina stenotes Clarke, 1976, ibid.: 9(1): 21, by monotypy and original designation. [Guam.] Syn. nov.
Diagnosis Fig. 800. Hermenias sp. D, female genitalia, nr Braidwood, NSW, slide T2316.
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Eccoptocera are small to medium-sized, narrowwinged and variably patterned eucosmines with a large costal fold and an inconspicuous antennal notch in the male. A highly derived wing venation
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with loss of one vein in both fore- and hindwings characterises Eccoptocera. The monotypic New Zealand genus Hendecasticha Meyrick, also belonging to the Spilonota-group of genera, shares loss of a vein in both wings. The genitalia of both sexes are typical and very consistent within the genus. The short broad uncus with its somewhat swollen appearance together with the vestigial socii and the strongly sinuate valva ending in a long ventrally directed spine are diagnostic for Eccoptocera. The valvae of Spilonota Stephens and Eccoptocera have exactly the same shape, possibly a measure of the close relationship between the two genera though Spilonota has a complete set of wing veins and has sclerotised, derived socii. The superficially close similarity of the valva of Strepsicrates infensa (Meyrick), however, is probably due to convergence as its spine is subapical and sits on a separate process from the pointed apex proper, as in all Strepsicrates. The female of Eccoptocera are less diagnostic and share presence of signa with Spilonota and some groups in Holocola Meyrick and, like Spilonota ocellana ([Denis & Schiffermüller]), have a sclerotised patch in the bursa neck.
Description Adults (Figs 801, 802) Wingspan. 6–17 mm; very small to mediumsized. Head (Fig. 803). Ocellus well developed. Frons in upper half with long, strongly forward and downward projecting scales. Proboscis slightly longer than labial palpus. Labial palpus moderately long to long (1.8–2.4× diameter of eye), porrect and strongly sinuate; second segment large and teardrop-shaped, strongly widened in apical half by loose long scales along dorsal and ventral margin that nearly conceal the long thin terminal segment. Antenna to beyond middle of forewing; flagellum in male with shallow dorsal notch just beyond base, scaled dorsally and with short cilia ventrally. Thorax. Without crest. Legs with particularly long, loose and rigid scales on dorsal surface of mid tibia. Wings (Figs 93, 801, 802). Forewing index 0.25–0.33. Forewing long and narrow, subrectangular; male with rounded costal fold to beyond middle (0.5–0.65× length of wing), containing modified scales; discal cell 0.66–0.68× length of wing, chorda and M-stem absent except for a faint remnant of the latter proximal to trace of closing vein; one vein absent and the vein arising
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from anterior angle of discal cell stalked: presumably R4 and R5 fused and stalked with M1, M2 straight and far distant from stalk of R4, R5 and M1 at base, M3 and CuA1 distally both strongly bent towards apex; CuP present near margin; anal loop 0.3× length of 1A+2A; without any raised scales, greyish or brownish with a typically Eucosmini-type pattern with well-developed ocelloid patch, dorsal half of wing often conspicuously paler than costal half. Hindwing with well-developed cubital pecten and axillary tuft, in male anal region expanded with pronounced anal angle and anal margin thickened and narrowly folded over small pencil of hair-scales from base of wing; closing vein only as a trace in membrane, Rs and M1 connate or short-stalked, one vein absent, presumably M3 and CuA1 entirely fused and closely approximated at base to M2; CuP present towards margin; 1A+2A about halfway between CuP and anal margin; 3A not visible; greyish to brownish. Pregenital abdomen. Unmodified. Sternum 8 in male medially with a shallow V-shaped excision in hindmargin, T8 with a thin, medially bent transverse ridge at base of segment. Male genitalia (Fig. 804). Vinculum relatively wide and little sclerotised. Tegumen high and slender, weakly sclerotised. Uncus a single, short, swollen, ventrally curved median process, usually with a widened and dorsoventrally somewhat compressed apex and a ridge-like dorsal expansion to its neck. Socii small, ill-defined membranous lateral lobes with few bristles that extend to tegumen and dorsal neck of uncus. Gnathos short lateral remnants only. Valva long, very slender and strongly sinuate, tapering to narrow and strongly curved ‘neck’ beyond middle then abruptly expanded into small cucullus; with very large basal excavation and reduced costal process; cucullus with round dorsodistal lobe and drawn-out ventral apex ending in long spine. Juxta triangular, caulis moderately long and wide, fused with anellus to a broad ring. Aedeagus thick and short, often curved, narrowed towards bevelled or drawn-out curved apex, often ending in a sharp ventral point; vesica with a sheaf of sinuate, lanceolate, deciduous cornuti. Female genitalia (Fig. 805). Ovipositor lobes narrowly crescentic. Sterigma a shallow sclerotised funnel with irregular margin, entirely fused with S7, with long lamella postvaginalis with scalloped posterior margin seemingly continuous with hindmargin of S7; lamella antevaginalis short and concave laterally, rising to triangular median process. Ductus bursae shorter than corpus bursae,
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Figs 801–805. Eccoptocera australis, paratype adults, head and genitalia, nr Rounded Hill, Cooktown, Q. 801, 802, adults. 801, . 802, . 803, head, male. 804, male genitalia, slide T2354. 805, female genitalia, slide T2371. Scale = 2 mm.
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gradually widening from two small crescentic lateral sclerites below ostium (colliculum), lightly sclerotised from there to inception of ductus seminalis at about midlength and with a variably developed sclerite or lightly sclerotised area laterally in the transitional region between ductus and corpus bursae. Corpus bursa ovate, with two blunt spines with granulose surface as signa; inner surface of bursa with an area of granulation around each signum, otherwise covered with minute, short spinules.
Distribution Eccoptocera was described from Hawaii where it seems represented by a number of closely related species (Zimmerman 1978). The genus has also been reported from Guam in the southern Mariana Is (Clarke 1976; as Alcina), from Sri Lanka (Meyrick 1912b as species of Hermenias Meyrick) and now from Australia. In Australia, the genus has been collected from Cape York Peninsula in Queensland as far south as Cooktown.
Biology Eccoptocera species are leaf rollers and Zimmerman (1978) gives Metrosideros (Myrtaceae) as the preferred host of E. foetorivorans Butler in Hawaii. The genus has also been reared from other Myrtaceae (Psidium guajava and Syzygium sandwicensis) and from Cheirodendron (Araliaceae) in Hawaii (Zimmerman 1978), and from Styphelia sp. (Epacridaceae) in Guam (Clarke, 1976). No hosts are known for the Australian species.
Remarks Zimmerman (1978) provided a detailed discussion of Hawaiian Eccoptocera, figuring adults and genitalia of several presumably distinct and new species in addition to E. foetorivorans and E. osteomelesana (Swezey). The comprehensive description and illustrations of Alcina stenotes Clarke (Clarke 1976) further add to the documentation of Eccoptocera. Clarke (1958) figured two species from Sri Lanka as Hermenias, that are discussed below. The monophyly of Eccoptocera is based on the characteristic uncus shape and the reduced venation in both fore- and hindwing. Retention of signa in the female and reduced but otherwise unmodified socii refer Eccoptocera to the less derived genera within the Spilonota-group, despite
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its highly modified venation. The similarities in the shape of valva and sterigma and the shared presence of a sclerotised area at the transition from ductus to corpus bursae in Eccoptocera and Spilonota may well indicate a close relationship between the two genera. The identity of the missing vein in each wing can only be guessed at. In the closely related Spilonota M3 and CuA1 in the hindwing are either stalked or, more often, fused into a single vein, and the same condition is assumed here for Eccoptocera. The forewing condition is less clear, with the relative position of the veins in the narrow-winged Hermenias suggesting that the missing vein in Eccoptocera may be due to fusion of R4 and R5, that in turn is stalked with M1. The following species, two from Sri Lanka described as Hermenias (Clarke 1958, Diakonoff 1982) and the type species of Alcina from Guam, are here referred to Eccoptocera: implexa (Meyrick, 1912b): 852 (Hermenias). [Sri Lanka.] Comb. nov. palmicola (Meyrick, 1912b): 853 (Hermenias). [Sri Lanka.] Comb. nov. stenotes (Clarke, 1976): 21 (Alcina). [Guam.] Comb. nov.
Constituent species Five species of Eccoptocera are known from outside Australia, two from Hawaii and the three referred to the genus above. Zimmerman (1978) recognises nine distinct ‘new’ species of Eccoptocera from Hawaii in addition to the two described ones, but he refrains from naming them. One species named below is known from Australia: australis, sp. nov. sp. A; Horak et al. 1996, Monogr. Aust. Lepid. 4: 134 (Eccoptocera).
Eccoptocera australis, sp. nov. (Figs 93, 801–805) sp. A; Horak et al. 1996, Monogr. Aust. Lepid. 4: 134 (Eccoptocera).
Material examined Holotype. , ‘15.17S 145.13E, 1 km N of Rounded Hill, Cooktown, Qld [Queensland], 5 Oct 1980, E.D. Edwards’, ‘Genitalia slide T2354’ (ANIC).
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Paratypes. Queensland: 1, Red Is Point, Cape York, 29.iii.1964, I. F. B. C. & M.S.U.; 2, 11°41´S 142°42´E, 15 km NE by E Heathlands, 22.iii.1992, E. D. E.; 2, 11°43´S 142°42´E, 13 km E by N Heathlands, 12.iii.1992, E. D. E.; 1, 11°45´S 142°35´E, Heathlands, 10.iii.1992, E. D. E.; 1, 11°46´S 142°41´E, 11 km E of Heathlands, 11.iii.1991, E. D. E.; 1, 11°48´S 142°31´E, 9 km SW Heathlands, 20. iii.1992, E. D. E.; 1 , 13°43´S 143°19´E, Edge Rainforest McIlwraith Ra., 500 m, 10.vii.1989, Nielsen, Edwards & Horak; 1, 13°44´S 143°19´E, 2 km W Golden Nugget Ck, McIlwraith Ra., 520 m, 2.vii.1989, Nielsen, Edwards & Horak; 3 , 1, 13°44´S 143°20´E, Golden Nugget Ck Camp Site, McIlwraith Ra., 520 m, 26.vi–9.vii.1989, Nielsen, Edwards & Horak; 1, 13°44´S 143°21´E, 1 km E Golden Nugget Ck, McIlwraith Ra., 520 m, 30.vi.1989; Nielsen, Edwards & Horak; 1, 13°45´S 143°22´E, Upper Leo Ck, McIlwraith Ra., 500 m, 1.vii.1989, Nielsen, Edwards & Horak; 2 , 2, same data as holotype, but 1 6.x.1980; (ANIC, QM, BMNH).
Diagnosis The wing pattern of Eccoptocera species is so similar that comparative material is needed for reliable separation. However, E. osteomelesana is clearly much paler (Zimmerman 1978) than E. australis. In Australia, absence of a forewing vein will separate E. australis, the only known species of Eccoptocera, from some very similar-looking Australian Holocola and Spilonota species of the same size. The male genitalia of E. australis are characterised by their uncus, valva, caulis and aedeagus shape. The uncus of E. implexa has much longer lateral projections and that of E. stenotes is much smaller and triangular, the valva of E. stenotes is wider and that of E. implexa and E. palmicola narrows much more abruptly beyond the sacculus, the latter two species also have a shorter caulis than E. australis, and the aedeagus of E. foetorivorans has a drawnout, conspicuously curved apex. In the female genitalia the shallow excavation in the hindmargin of S7, the broadly triangular lamella antevaginalis, the sclerite laterally in the bursa neck and the short, blunt signa are diagnostic for E. australis.
Description Male (Figs 801, 803) Wingspan 9–10 mm. Head and thorax brownish grey with pale-tipped scales and paler mottling, especially on outer surface of labial palpi (Fig. 803), with their inner surface whitish. Legs with paletipped dark grey scaling except for paler grey hind tibia, mid tibia with particularly dark, spiky scaling, tips of tarsal segments pale.
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Wings. Forewings with large costal fold to middle; a distally narrowing dark grey band along costa, reaching to middle in basal 369 with a triangular extension towards dorsum below end of costal fold, thence narrowing to apex, with 3–4 blackish strigulae outlined by a few pale scales along distal half of costa and a white spot on costa just before black apex; ocelloid patch an erect band of mixed white and dark grey scales with 3–4 longitudinal parallel black bars, delineated on both sides by a wide, erect silvery band, the inner one slightly oblique and the outer one interrupted near tornus; with a sinuate black mark along termen, widening and curved into fringe at apex; dorsal region of mixed white, pale grey and dark greybrown scales, reaching 36: towards costa beyond middle of wing and ending in an ill-defined darker grey-brown triangle on dorsum adjacent to inner silvery band of ocelloid patch. Fringe blackish at apex, remainder with a white basal row and mottled pale grey distal row. Hindwings rather thinly scaled, dark grey with coppery gloss, with similarly coloured fringe. Abdomen as hindwings above, paler grey below. Male genitalia (Figs 804). Of typical Eccoptocera shape; uncus with a short, relatively broad ‘neck’ with a dorsal ridge and a laterally slightly expanded flattened apex; valva gradually tapering from widest point, cucullus ‘neck’ moderately bent only, base of ventral spine nearly at right angle to cucullus axis; caulis rather long; aedeagus somewhat tapering to bevelled apex; with large sheaf of lanceolate cornuti. Female (Fig. 802). Wingspan 9.0–12.5 mm. As for male, but without costal fold. Female genitalia (Fig. 805). Of typical Eccoptocera shape; emargination in hindmargin of S7 much wider than high, not reaching to centre of S7, lamella antevaginalis a well-defined broad triangle; ductus bursae 69 length of corpus bursae, lightly sclerotised between colliculum and inception of ductus seminalis; corpus bursae with an elongate, slightly longitudinally folded sclerite on one side of bursa ‘neck’, and with two short, blunt, thorn-shaped signa.
Distribution Eccoptocera australis is only known from the Cape York Peninsula, Queensland, as far south as Cooktown.
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UNASSIGNED TO GENUS-GROUP Herpystis Meyrick (Figs 94, 806–813) Herpystis Meyrick, 1911a July, Trans. Linn. Soc. Lond. (2) Zool. 14: 268. Type-species: Herpystis rusticula Meyrick, 1911a, ibid. (2) Zool. 14: 268, by monotypy. [Seychelles.] Herpystis Meyrick, 1911b November, Proc. Linn. Soc. NSW 36: 225 (key), 244. Type-species: Herpystis avida Meyrick, 1911b, ibid. 36: 245, by monotypy. A junior homonym of Herpystis Meyrick, 1911b July. [Australia.] Diplonearcha Meyrick, 1914, Exot. Microlepid. 1: 274. Type-species: Diplonearcha insinuans Meyrick, 1914, ibid. 1: 274, by monotypy. [Sri Lanka.] Syn. nov.
Diagnosis Species of Herpystis are small, very narrow-winged, brownish moths with variable wing pattern but always with dark costal strigulae in basal half, a dark blotch or oblique band near middle of the costa, a series of paired white strigulae in apical half, a prominent, subapical dark strigula on the costa usually linked with a series of parallel dark, longitudinal dashes in the ocelloid patch that are preceded and followed by transverse, silver-grey marks, always with a white line along the termen and usually with a short, longitudinal, blackish
streak in the middle of the wing beyond the cell. Herpystis has sinuate, moderately long, slender to medially strongly widened labial palpi and an unusual wing venation with all veins separate in the forewing (except in the non-Australian H. insinuans with R5 and M1 stalked) and with CuA1 in the hindwing from well before the angle of the cell, distant from the connate M2 and M3. The female frenulum has two bristles only, probably sometimes reduced to one. The male genitalia are characterised by a robust, simple tegumen with elongate, very flimsy socii, a wide vinculum with pronounced internal angles, usually slender valvae without a costal process but with a large spine on the ventral margin in apical half and with the ventral margin often with a cluster of spines near its middle and with a variably developed emargination distally and sometimes also proximally to the spine cluster. The female genitalia are unique with the ostium at the base of the medially divided S7, rod-like sclerotisations along the anterior margin of S7 and the hindmargin of S6, the sterigma a spinulose band or two spinulose humps anterior to the ostium and a long funnel, a short colliculum of thickened membrane, the bulla seminalis directly joined to the entrance of the corpus bursae with a short, wide connection, and the two horn-shaped signa that may be reduced to small remnants.
Description Adults (Figs 806, 807) Wingspan. 6–10 mm; small to very small.
Figs 806, 807. Herpystis, adults, males. 806, H. sp. nr jejuna, Elizabeth Beach, NSW. 807, H. sp. nr avida, Rous, Richmond R., NSW. Scale = 2 mm.
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Figs 808–813. Herpystis, genitalia and head. 808, H. sp. nr avida, male genitalia, Richmond R., NSW, slide ANIC 13789. 809, head, H. sp. nr avida, male. 810, 811, H. sp. A, male genitalia and aedeagus, nr Port Macquarie, NSW, slide T2337. 812, H. sp. C, female genitalia, Lake Eacham, Q, slide ANIC 13794. 813, H. sp. A, female genitalia, nr Port Macquarie, Q, slide ANIC 12975.
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Head (Fig. 809). Ocellus moderate. Vertex scales anteriorly merging with downcurved scales of upper frons, remainder of frons with appressed, small scales. Labial palpus porrect, sinuate, moderately long (1.9–2.2× diameter of eye), slender to moderately wide, second segment sinuate and lightly to strongly widened by long scales along ventral margin and distal half of dorsal margin, terminal segment long, slender, downward pointing. Proboscis as long as labial palpus. Antenna to beyond middle of wing, entirely scaled in male, without visible cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 94, 806, 807). Forewing index 0. 27–0.35. Forewing long and slender, hardly widening towards apex, with costa weakly curved to nearly straight except near base and apex, apex roundedpointed, termen oblique and faintly concave below apex; discal cell without trace of chorda or M-stem; all veins separate except in H. insinuans where R5 and M1 are stalked, bases of M3 and CuA1 closely approximated; CuP present at margin; anal loop 0.24× length of 1A+2A, with anterior branch weak and only a trace in membrane; typically eucosmine pattern in greyish or reddish brown, with cream, silver grey and some blackish brown, with numerous, parallel, outwardly oblique and angled, transverse lines and bands either well developed (rarely forming a broad dark band across basal half beyond base) or suffused except for dark costal strigulae in basal half, a dark blotch or oblique band near middle of costa, a series of paired white strigulae in apical half, a prominent, subapical dark strigula on costa usually linked with parallel series of dark, longitudinal dashes in ocelloid patch that are preceded and followed by transverse, silver-grey marks, always with a white line along termen and usually with a short, longitudinal, blackish streak in middle of wing beyond cell. Hindwing narrow, with costa only weakly sinuate, with moderate cubital pecten and axillary tuft, without modified anal margin in male, frenulum in female with two bristles only; Rs and M1 only approximated for a short distance at base, M2 and M3 connate from angle of cell, CuA1 from far below angle of cell, but closer to M3 than to CuA2; CuP present at margin halfway between CuA2 and 1A+2A, the latter halfway between CuP and anal margin, without any trace of 3A in slide of Australian species; light to dark brownish grey, sometimes with scales minute and wing transparent in between. Pregenital abdomen. Unmodified. Hindmargin of S8 bilobed in male with a V-shaped medial emargination; T8 with an inverted-Y-shaped
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sclerotisation near base, hindmargin roundedconvex. Male genitalia (Figs 808, 810, 811). Tegumen robust, ovate, apex usually truncate, rarely weakly triangular, but without trace of uncus; vinculum a wide band with projecting inner angles. Socii membranous, very flimsy, moderately large to small, elongate, bristled lobes. Gnathos arms slender, weakly sclerotised, strongly upcurved and fused to conically projecting anal tube. Valva without costal process, with a large basal excavation, a large spine from ventral margin at some distance from apex and often with a cluster of spines on a projection near middle of ventral margin, sometimes including one particularly prominent one; valva either very slender (Micronesian species) or once (H. tinctoria Meyrick) or twice (H. jejuna Meyrick) narrowed by ventral emarginations, with the more distal one between spiny projection on middle of ventral margin and apex of valva usually at least indicated but a more basal emargination separating an angled sacculus from a prominent, spiny projection near middle of valva only present in some species. Juxta triangular, caulis a long wide band. Aedeagus long, widest at base, gradually narrowing towards apex; vesica with a sheaf of lanceolate cornuti. Female genitalia (Figs 812, 813). Ovipositor lobes reniform. Ostium at base of S7, anterior margin of S7 and posterior margin of S6 ventrally with rod-shaped sclerotisations; S7 medially divided into two posteriorly rounded, lateral sclerites connected only at base just below entrance to ostium; the wide, longitudinal, depressed, smooth, medial band dividing S7 and leading to ostium is spinulose posteriorly and the anterior margin of S7 is narrowly sclerotised and carries a band of spinules or two spinulose humps below entrance to ostium. Sterigma (posterior part of ductus bursae) a long, smooth, sclerotised funnel ending in a short colliculum of thickened membrane followed by a sclerotised, short, wide tube or ring opening to bifurcation of roughly equal-sized corpus bursae and bulla seminalis; corpus bursae with two signa, ranging from long and horn-shaped, distally pointed and flattened and hollow at base to only small, spine-shaped remnants; bulla seminalis with ductus seminalis originating from close to broad connection with corpus bursae.
Distribution Herpystis has been recorded from the Seychelles, Sri Lanka, India, Pakistan, Micronesia, the Solomon Is,
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Vanuatu and Australia, where it has been collected along the eastern coast from Cooktown in Queensland to Batemans Bay in New South Wales.
Biology Herpystis cuscutae Bradley was reared at Rawalpindi, Pakistan, from larvae feeding in the stem of Cuscuta reflexa (Cuscutaceae), an unnamed Australian species has been reared by A. W. S. May in Innisfail from fruit of Breynia stipitata (Euphorbiaceae), and the holotype of H. insinuans has been ‘bred from a psyllid gall on Ficus [Moraceae] (Rutherford)’ in Peradenyia, Sri Lanka. All the Australian material is either from rainforest or localities with some rainforest plants.
Remarks Meyrick described Herpystis twice in the same year, first based on Herpystis rusticula from the Seychelles (Meyrick 1911a) in July 1911, then based on the Australian Herpystis avida (Meyrick 1911b) in November 1911. There is no doubt that the two species are congeneric. Clarke (1958) figured the types and their genitalia for H. rusticula, H. jejuna Meyrick and H. tinctoria Meyrick, Bradley (1968) illustrated male and female genitalia of H. cuscutae Bradley, Clarke (1976) provided illustrations of adults and genitalia of three new species from Micronesia and Kuznetzov (1988b) a drawing of the male genitalia of a new species from Vietnam. The male genitalia figured by Diakonoff (1982) for H. jejuna from Sri Lanka appear not to be conspecific with those of the type from India. According to Meyrick (1914) Diplonearcha has Rs and M1 stalked, but examination of the female holotype (not male as stated in the description) shows these two veins to be closely approximated as in other Herpystis species. The female genitalia of H. insinuans are exactly like those of Herpystis, and apart from stalked R5 and M1 the wing venation is the same. Treating Diplonearcha as a separate genus would render Herpystis a paraphyletic group, and the two genera are here synonymised. The monophyly of Herpystis is supported by several unusual apomorphies, including the position of CuA1 in the hindwing from below the angle of the cell well distant from M3, a frenulum of at most two bristles in the female, a valva without a costal process but with a spine distally on the ventral margin, and especially the unique female genitalia with a characteristic ductus bursae and
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the large bulla seminalis directly joined to the neck of the corpus bursae. No sister group has been identified and the affinities of the genus are unclear. Clarke (1976) included Herpystis in the Olethreutini, as suggested by the hindwing venation (CuA1 separate from M3) and the valva structure with a discrete spine tuft near the middle of the ventral margin, but Diakonoff (1973) did not discuss the genus in his monograph on the southeast Asian Olethreutini and later treated the genus as an eucosmine (Diakonoff 1982). The genus is here also tentatively included in the Eucosmini based on the absence of 3A in the hindwing, the absence of any trace of teguminal muscle attachments, the short colliculum of thickened membrane and the wing pattern with a recognisable ocelloid patch. Herpystis avida was described from a single male from Cairns, northern Queensland, and the holotype has lost its abdomen. Several specimens from southern Queensland (Toowong) and northern New South Wales (Rous, Richmond R.) look rather similar, but their identity can only be confirmed once their genitalia can be compared with newly collected material from Cairns. In the figure legends they are here referred to as ‘sp. near H. avida’.
Constituent species Nine named species of Herpystis are confirmed from outside Australia and one named and probably four unnamed species from Australia. Herpystis iodryas Meyrick, 1937 (Clarke 1958: pl. 213, figs 1–1b) and Herpystis pallidula Meyrick, 1912b (Clarke 1970: pl. 59, figs 1, 1a) are both not Herpystis species, with the latter probably belonging to the genus Parabactra Meyrick. The male genitalia of Herpystis assimulatana Kuznetzov (Kuznetzov 1997a) also suggest it is not congeneric with H. rusticula. avida Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 245 (Herpystis). [Qld: Cairns.]
Icelita Bradley (Figs 95, 814–825) Icelita Bradley, 1957, Nat. Hist. Rennell Is. 2: 90. Type species: Icelita tatarana Bradley, 1957, ibid. 2: 91, pl. l, fig. 3; pl. 3, figs 35–38, by original designation. [Solomon Is: Rennell Is.]
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Diagnosis Icelita species are smallish greyish or brownish moths with elongate, distally characteristically narrowed forewings with a bluntly rounded apex, a short, hardly oblique termen with a notch near its middle and a pattern of parallel, oblique, broad bands across wing. The males can be distinguished from those of Eucosmophyes Diakonoff and some other similar-looking eucosmines by a modified anal region in their hindwing with a scale pencil from its base either covered by the upfolded anal margin or associated with additional scale tufts further distally on anal region, and by modified head scaling with upwardly oriented scales even in upper part of frons, often resulting in a dorsally projecting tuft. Icelita has short, slightly upcurved labial palpi with the second segment slightly
widening towards tip and the round apex of the third segment only just visible. Radius 4 and R5 in the forewing are separate or stalked, in the hindwing Rs and M1 are closely approximated or stalked, M2 is bent close to stalk of M3 and CuA1 that are either stalked to 6M or entirely fused. The male genitalia are characterised by a high and weakly sclerotised tegumen with the apex projecting far beyond the socii but without differentiated uncus, by socii set with slender scales and by complex valvae with two large, modified scale tufts, one from its outer surface and one from the base of the costa, and with a conspicuous band- to mushroom-shaped process from the sacculus. The female genitalia are characterised by a sterigma of either two membranous pockets posterior to S7 or paired,
Figs 814–817. Icelita, adults. 814, 815, I. monela, Clump Point, Q. 814, [image reversed]. 815, . 816, 817, I. indentata, Clump Point, Q. 816, . 817, [image reversed]. Scale = 2 mm.
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spinulose plates ending in a small, round invagination, a complex ductus bursae with a long colliculum of thickened membrane below ostium, often reduced, and with a sclerotised ring or funnel in its anterior half, and by two long, hollow, hornshaped signa.
Description Adults (Figs 814–817) Wingspan. 9.0–13.5 mm; small to mediumsized.
385
Head (Fig. 823). Ocellus moderate. Vertex with long, loose scales often separated into a prominent tuft on each side; scaling on frons modified in male, upwardly directed even in upper part of frons, either very short, appressed and surrounded by an U-shaped band of longer, erect scales along eyes and across base of frons (I. indentata Bradley), or of long, slender scales ending in a medial tuft dorsally projecting between antennae (I. monela Clarke); in female scaling on frons unremarkable, downwardly directed in upper part. Maxillary palpus minute, scaled. Labial palpus short
Figs 818–823. Icelita, male genitalia, head and abdomen. 818, 819, I. monela, male genitalia and aedeagus, Gap Ck, Q, slide ANIC 12974. 820, 821, 8th segment, male. 820, I. monela. 821, I. indentata. 822, I. indentata, male genitalia, Gap Ck, Q, slide ANIC 13775. 823, head, I. indentata, male.
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(1.3–1.6× diameter of eye), weakly upcurved, with second segment lightly curved, widening towards apex, terminal segment very short with only rounded tip visible. Antennae to just beyond middle of forewing, in male scaled dorsally only, with short cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 95, 814–817). Forewing index 0.34–0.38. Forewing slender, elongate-subovate to subrectangular, widest in middle, narrowed towards base and apex, with costa weakly curved throughout, apex slightly protruding, bluntly rounded, termen hardly oblique, with notch below apex and portion of termen behind notch very short, tornus oblique, weakly rounded, transition to
dorsum indistinct; discal cell with M-stem a trace in membrane only and chorda present (I. tatarana, I. monela) or a trace only (I. indentata); R4 and R5 from separate to stalked, M1, M2, M3 and usually also CuA1 converging towards margin and ending at or near terminal notch, bases of M3 and CuA1 separate; CuP present at margin; anal loop 0.42× length of 1A+2A; greyish or brownish with pronounced costal strigulae and a characteristic wing pattern of variably developed, broad, parallel, outwardly oblique transverse bands with usually two more prominent lines often separating a dark proximal area from a paler area distally, first line weakly curved or angled from beyond 56: costa and second from about 36: costa to before tornus with a
Figs 824, 825. Icelita, female genitalia. 824, I. indentata, Clump Point, Q, slide ANIC 1299. 825, I. monela, Gap Ck, Q, slide ANIC 13777.
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projecting ‘step’ near middle; dark apical scales extending halfway along termen and for a similar distance along costa, there interrupted by a pronounced pair of white strigulae; ocelloid patch indicated and sometimes with a few small, longitudinal black dashes. Hindwing with strongly sinuate costa and weakly to moderately developed cubital pecten and axillary tuft, in male with modified anal region with a variable hair pencil from base of upper side of anal region, large and covered by the upfolded anal margin in I. indentata, smaller and exposed but with additional scale tufts more distally on upper surface in some other species (I. monela); Rs and M1 separate at base but closely approximated (I. indentata) to stalked (I. tatarana), base of M2 very close to stalk of M3 and CuA1 that are stalked to about 6M or entirely fused (I. monela); CuP present at margin, 1A+2A far from anal margin, no trace of 3A visible; grey to grey-brown, scales in basal half often small and wing translucent between scales. Pregenital abdomen. Ventral apodemes of S2 widened by a membranous keel. In male hindmargin of S8 strongly bilobed with a deep U-shaped medial emargination; T8 very long and slender, projecting tongue-like, little sclerotised except for transverse basal sclerite (Figs 820, 821). Male genitalia (Figs 818, 819, 822). Tegumen very high, distally triangular, rounded or bifid, very weakly sclerotised, its apex far beyond origin of socii but no differentiated uncus apparent. Socii membranous, covered with long, slender scales, variable from large reniform lobes to narrow, pendant rods. Gnathos two slender, medially joined arms, with sublateral portion sometimes conspicuously sclerotised. Valva long and slender, strongly narrowed beyond basal third; with two large tufts of modified scaling, one from an invagination at base of costa, concealing costal process, the second from outer surface from below middle; with a conspicuous band- to mushroomshaped process with spinulose apex from lower margin of basal excavation; cucullus variable, with projecting ventral angle and broadly rounded apex in I. indentata, narrowly crescentic in I. monela and widest distally in I. tatarana. Juxta small, pentagonal; caulis a long band. Aedeagus long, curved, tapering towards apex; vesica with a bundle of deciduous, lanceolate cornuti and sometimes with a large, fixed cornutus protruding from aedeagus tip. Female genitalia (Figs 824, 825). Ovipositor modified, telescopic, with very small ovipositor lobes and greatly lengthened apophyses except for
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I. tatarana. Hindmargin of S7 weakly concave, ostium in intersegmental membrane. Sterigma invaginated between S7 and ductus bursae, either two membranous, shallow pockets as in I. monela or paired, spinulose, sclerotised lobes each anteriorly ending in a small round invagination as in I. indentata. Ductus bursae with very long colliculum of thickened membrane in distal third or posterior portion simply thick-walled; with a ring- or funnel-shaped sclerotisation in anterior half near middle with ductus seminalis originating ventrally from its base; corpus bursae ovate, partly dimpled or spinulose, with a spinulose patch ventrally on transition from corpus bursae to widened end part of ductus bursae, and with two long, hollow, curved, horn-shaped signa.
Distribution Icelita is known from Vietnam, Micronesia (Marshall Is, Mariana Is, Palau Is), the Solomon Is and Australia where it has been collected in northeastern Queensland near Cooktown and near Innisfail and in the Northern Territory on the Wessel Is off Arnhem Land.
Biology Clarke (1976) reports Afzelia bijuga (as Intsia bijuga) (Caesalpiniaceae) as food plant of Icelita tatarana coppelia Clarke on Guam. All Australian material has been collected in rainforest or monsoon forest habitats. The strongly telescopic ovipositor with very small, soft ovipositor lobes suggests that eggs of both Australian species are laid into crevices.
Remarks In his original description of Icelita Bradley (1957) figured I. tatarana as well as I. indentata, the latter as a Spilonota species. Clarke (1976) redescribed and comprehensively illustrated the genus based on all known species of Icelita, including two new ones from Micronesia. Icelita is unique and characterised by several conspicuous apomorphies: distally narrowed forewings with a short termen and in the male modified scaling on the frons, a large but weakly sclerotised tegumen, valvae with two large tufts of modified scales and with an apically spinulose sacculus process, a complex ductus bursae with a very long colliculum of thickened membrane and with a ring- or funnel-shaped sclerotisation in
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anterior half. The genus is highly derived and no sister group has been identified, though there are some shared characters with members of the Acroclita-group. The following species described from Vietnam as a Rhopobota (Kuznetzov 1988b), is here transferred to Icelita: antecellana (Kuznetzov, 1988b), Entomol. Obozr. 67: 627 (Rhopobota). [Vietnam.] Comb. nov.
Constituent species Including I. antecellana, five named species of Icelita are known, with I. tatarana and I. indentata originally described from the Solomon Is and later reported from Micronesia (Clarke 1976), together with the Micronesian species I. monela and I. cirrholepida Clarke. I. indentata and I. monela both occur in Australia, but the identity of the Micronesian specimens of I. indentata will have to be confirmed as their valva seems to lack the deep ventral emargination present in the Solomon Is and Australian material. indentata (Bradley, 1957), Nat. Hist. Rennell Is. 2: 89 (Spilonota). [Solomon Is: Rennell Is.] sp. A; Horak et al. 1996, Monogr. Aust. Lepid. 4: 135 (Icelita). monela Clarke, 1976, Insects Micronesia 9: 39 (Icelita). [Micronesia.]
Eucosmophyes Diakonoff (Figs 96, 826–832) Eucosmophyes Diakonoff, 1982, Zool. Verh. (Leiden) 193: 45. Type-species: Eucosmophyes icelitodes Diakonoff, 1982, ibid. 193: 45, figs 29, 29A; pl. 11, fig. 22, by original designation. [Sri Lanka.]
Diagnosis Eucosmophyes species are small grey moths with moderately slender forewings with a roundedpointed apex, an oblique, slightly concave but not conspicuously notched termen and with a characteristic pattern of pale and dark grey, parallel, outwardly oblique and lightly angled bands and lines across entire wing. All veins are separate in the forewing, with M-stem and chorda present, whereas in the hindwing Rs and M1 are stalked or so closely approximated as to appear stalked, and M3 and CuA1 are stalked. Superficially Eucosmophyes looks very similar to Icelita Bradley
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and several undescribed eucosmine species of unknown generic affinity with stalked R4 and R5 in the forewing. From those Icelita species with separate R4 and R5, Eucosmophyes may be separated by a forewing not narrowed distally, with a termen not conspicuously notched near middle and with M2, M3 and CuA1 not strongly converging towards margin. Eucosmophyes has white-tipped, moderately long, sinuate labial palpi with a rather slender terminal segment. The male genitalia are very similar in all species, characterised by a simple tegumen with small, roundish socii, simple valvae with a deep V-shaped ventral emargination, a partially bristled sacculus with a large basal excavation, a small costal process, and a semicircular to subtriangular cucullus, densely bristled on inner surface. The female genitalia are characterised by a sclerotised, funnel-shaped sterigma fitted into a deep, V-shaped emargination in the hindmargin of S7, by a short colliculum of thickened membrane, a ductus bursae nearly entirely sclerotised, of two parallel strands coiled around each other, and by a corpus bursae with a patch of spinules ventrally at its entrance and two unequal, horn-to blade-shaped signa.
Description Adults (Figs 826, 827) Wingspan. 6.5–10.0 mm; small to very small. Head (Fig. 830). Ocellus moderate. Scaling on upper part of frons anteriorly projecting, remainder of frons with minute, appressed scales. Labial palpus moderately long (1.7–1.9× diameter of eye), sinuate, widened in middle, basal 36: dark grey, apex of second and entire third segment white, second segment curved and gradually widening towards apex, third segment rather long and slender, downwards pointing. Antennae to just beyond middle of forewing, in male scaled also ventrally, without visible cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 96, 826, 827). Forewing index 0.34–0.37. Forewing slender, subtriangular, without costal fold, costa weakly curved throughout, apex rounded-pointed, termen oblique, weakly concave below apex; chorda present, M-stem to base of M2; all veins separate, R3, R4 and R5 equidistant at base, also M2, M3 and CuA1 that are distally at most slightly converging; CuP present at margin; anal loop 0.35× length of 1A+2A; with characteristic pattern of pale and dark grey, parallel, outwardly oblique and lightly angled bands and lines across entire wing, forming a dark basal area to nearly 56:
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and including a narrow, angled, dark, transverse line at about middle, widening towards dorsum and connected with a distally projecting, longitudinal dark blotch near middle of wing; with costal strigulae well developed and a prominent, curved, blackish line or band from well before apex to middle of termen, linked by a curved line from near its middle to blackish spot in apex. Hindwing narrow, evenly scaled, with moderately developed cubital pecten and in male with projecting anal area and anal margin very narrowly rolled over upperside covering small hairpencil (obvious only in slide preparation); bases of Rs and M1 very closely approximated in Australian E. commoni, sp. nov., stalked according to Diakonoff (1982) in E. icelitodes, base of M2 moderately close to stalk of M3 and CuA1 that are stalked to 56: or to middle; CuP present at margin, 1A+2A far from anal margin, no trace of 3A; evenly light to dark grey. Pregenital abdomen. Unmodified. In male hindmargin of S8 bilobed with a shallow, V-shaped medial emargination; hindmargin of T8 convex. Male genitalia (Figs 829, 831). Tegumen subovate, apex sometimes truncate, a narrow band except across apex. Uncus not present. Socii two small, roundish, sublateral, bristled lobes. Gnathos two weakly sclerotised lateral arms supporting a membranous, ventrally projecting cone (?anal tube). Valva simple, long, slender, with deep, V-shaped ventral emargination near middle resulting in narrow ‘neck’ to cucullus; ventral half with row of fine bristles below basal excavation and dense tuft of strong bristles in distal part of sacculus and with very small costal process; cucullus semicircular to subtriangular, densely bristled on inner surface. Juxta triangular, caulis a wide, long band. Aedeagus long, slender, slightly curved, widest at base; vesica with a sheaf of lanceolate cornuti.
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Female genitalia (Figs 828, 832). Ovipositor lobes unremarkable. Sternum 7 small, posteriorly narrowed and subtriangular, more strongly sclerotised along anterior margin and with a deep V-shaped emargination in hindmargin into which wide, sclerotised, funnel-shaped sterigma is fitted. Ductus bursae with short colliculum of thickened membrane, followed by a short membranous section and then widened into complex, sclerotised main part consisting of two parallel strands coiled around each other in one full turn, with ductus seminalis originating laterally near middle; corpus bursae ovate with patch of long dense spinules ventrally at entrance and with two unequal, hornto blade-shaped signa, hollow at base and flattened towards apex, with the ventral one distally wider.
Distribution Eucosmophyes is so far known from Brisbane to Yeppoon and in the coastal northern tip of the Northern Territory.
Biology An unidentified species of Eucosmophyes has been reared from the fruit of Psydrax lamprophylla (Rubiaceae) by C. Wilson near Darwin, and all Australian species have been collected in either rainforest or monsoon forest.
Remarks Diakonoff (1982) figured head, wing venation and genitalia of both sexes of E. icelitodes, the type species of the genus from Sri Lanka. All known species of Eucosmophyes have very similar appearance and genitalia in both sexes. The
Figs 826, 827. Eucosmophyes commoni, adults, Toowoomba, Q. 826, , holotype [image reversed]. 827, [image reversed]. Scale = 2 mm.
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monophyly of the genus is based on the small, rounded, sublateral socii, the simple valva with a deeply V-shaped ventral excavation and a very small costal process, and, foremost, on the female genitalia with the funnel-shaped sterigma, the sclerotised, twisted ductus bursae, the spinulose patch ventrally in the neck of the corpus bursae and the two unequal-sized signa. Even in the Australian fauna, there are several unnamed, eucosmine species superficially very similar to Eucosmophyes, mostly with R4 and R5 stalked, and one with very similar valvae but with a small bristled process near the base of the costa. Further material will be required before the relationships of these small eucosmines can be resolved. The spinulose patch in the bursa neck could indicate a close relationship with Icelita, as suggested by the wing pattern, even though the much more reduced male genitalia of Eucosmophyes are very different from the highly derived ones of Icelita. The twisted ductus bursae recalls that of some taxa in the Acroclita-group.
Constituent species The type of the genus, E. icelitodes from Sri Lanka, is the only species of Eucosmophyes named so far. Of the at least three species occurring in Australia, one is described below. commoni, sp. nov.
Eucosmophyes commoni, sp. nov. (Figs 826, 827, 830–832)
Material examined Holotype. , ‘27.36S 151.59E, Middle Ridge, Toowoomba, Q[ueensland], 660 m, 29 Oct. 1982, I.F.B. Common’, ‘ANIC genitalia slide No. 13792 ’ (ANIC). Paratypes. Queensland: 2 , 3, 27°33´S 151°59´E, Prince Henry Heights, 620 m, 16.i.1983, 22.xii.2003, 18. & 28.i.2004, 8.ii.2004, I. F. B. C. (ANIC, QM).
Diagnosis Eucosmophyes commoni has a wing pattern typical for the genus but characterised by white, grey and black colouring rather than the more greyish brown of other Australian species, and it has conspicuously dark grey hindwings. The male genitalia of E. commoni are characterised by a
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distally truncate and dorsally spinulose tegumen apex, a strongly angled sacculus, a wide cucullus and a submedially narrowed aedeagus with a wide base and with wide, distally strongly narrowed cornuti. The female genitalia of E. commoni are characterised by the shape of the strongly sclerotised S7 with the lip of the funnel-shaped sterigma projecting, and a broad ventral signum. A second, undescribed species of Eucosmophyes has been collected at the same localities in Toowoomba and can be separated from E. commoni by paler hindwings, a more slender cucullus and an aedeagus narrowest at its apex in the male, and by a different sterigma in the female without a projecting ventral lip.
Description Male (Fig. 826) Wingspan 9.0–10.5 mm. Head (Fig. 830), thorax and tegulae grey except for tip (distal fringe of second segment and entire third segment) and inner surface of labial palpi, frons and anterior fringe of vertex scaling that are whitish; thorax and tegulae scales grey, pale-tipped. Forewing white with grey and blackish pattern elements: paired white costal strigulae separated by black wedgeshaped marks along entire costa, less distinct and greyish white in basal third; apex with large black spot; basal third of wing with two parallel, dark grey, transverse, outwardly curved bands, the first occupying wing base, the second with its outer blackish margin from 56: costa to 56: dorsum, sometimes obliterated in dorsal half of wing; black costal mark at 56O connected by slender, outwardly oblique black line in middle of wing with a sometimes medially interrupted, hourglass-shaped large blackish mark from dorsum before tornus; second last black costal mark before apex extended as an oblique black line to middle of termen, there widened and following termen towards dorsum, separated by silver grey scales from black apical spot; ocelloid patch not developed, silver grey with an indistinct, darker wavy line rising from tornus; fringe concolorous with wing. Hindwing very dark grey, nearly charcoal grey, with long dense hairscales from anal roll; distal half of fringe paler than wing. Male genitalia (Fig. 831). Hindmargin of S8 with two short and wide lobes. Top of tegumen subtriangular, dorsally with parallel rows of fine spinules (visible in compound microscope only). Socii small, round, with few long bristles. Valva with sacculus strongly angled, cucullus nearly as
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wide as long. Aedeagus with wide base and narrower beyond middle than at apex, with a large sheaf of wide, bottle-shaped cornuti with narrow, drawn-out apex.
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Female (Fig. 827) Wingspan 10–11 mm. As for male. Female genitalia (Fig. 832). S7 strongly sclerotised, with anterior margin slightly convex in
Figs 828–832. Eucosmophyes, genitalia and head. 828, E. sp. A, female genitalia, Coburg Peninsula, NT, slide ANIC 13782. 829, E. sp. A, male genitalia, Coburg Peninsula, NT, slide ANIC 13783. 830, head, E. commoni, male. 831, E. commoni, paratype, male genitalia, Toowoomba, Q, slide ANIC 17064. 832, E. commoni, paratype, female genitalia, Toowoomba, Q, slide ANIC 12977.
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middle third and with hindmargin a shallow, evenly curved emargination; ventral lip of wide, funnel-shaped sterigma projecting beyond hindmargin of S7. Colliculum rather long. Ductus bursae bipartite and strongly sclerotised, with second strand forming a complete coil around straight central strand. Patch of spinules at bursa entrance large and dense; ventral signum very wide, broadest beyond middle, dorsal one much more slender.
Eucosmophyes commoni has only been collected in Toowoomba, in gardens at the top of the range adjoining rainforest.
large basal excavation and an ovate cucullus densely bristled along its projecting ventral margin, a very short caulis, and an aedeagus with an extremely long ejaculatory duct, some fixed thorns distally on its outer surface and numerous deciduous cornuti. The female genitalia are characterised by a distally narrowed S7 with its lateral margins developed into keel-shaped ridges and meeting medially to form a short triangular process covering the ostium, by an extremely long, coiled ductus bursae, membranous except for posteriormost sclerotised portion connected with S7 and giving rise to ductus seminalis, and by a small, roundish corpus bursae with two flat, dagger-shaped signa with round, hollow bases sitting in externally protruding membranous bulges of the bursa wall.
Remarks
Description
The species is named for Ian Common who 20 years after his retirement is still collecting exciting new species such as this small eucosmine for the ANIC.
Adults (Figs 833, 834) Wingspan. 8–13 mm; small. Head (Fig. 839). Ocellus moderate. Vertex with very long, dense scales, anteriorly strongly projecting between antennae, upper part of frons with long, anteriorly projecting scales, remainder of frons with short scales. Maxillary palpus relatively large, scaled. Proboscis about as long as labial palpus. Labial palpus strongly sinuate, porrect, rather short (1.6–1.9× diameter of eye), strongly widened beyond middle, with second segment sinuate, greatly expanded by long, loose scales along ventral and apical half of dorsal margin, terminal segment long but in female sometimes largely concealed by apical scales from second segment. Antenna to middle of wing, entirely scaled. Thorax. Smooth, with moderately long scale tuft from ventral process of tegulae. Legs unmodified. Wings (Figs 97, 833, 834). Forewing index 0.28–0.31. Forewing slender, subrectangular, costa and dorsum roughly parallel except near base, costa very weakly curved, apex pointed and strongly projecting, nearly falcate, termen sinuate; with two tufts of raised scales in dorsal half, one at about 56M and one just beyond cell; M-stem straight and very close to hindmargin of cell, to between M3 and CuA1, no trace of chorda; R3 from roughly halfway between R2 and angle of cell, R4 and R5 stalked to just beyond middle, M2 much closer to M3 than to M1, M3 and CuA1 moderately close and parallel near base, then diverging to converge again, together with M1 and M2, towards margin; CuP present near margin; anal loop 0.27× length of 1A+2A; wing pattern sexually dimorphic, ground
Distribution and biology
Whittenella, gen. nov. (Figs 97, 833–839) Type species: Palaeobia peltosema Lower, 1908, Trans. R. Soc. S. Aust. 32: 323, hereby designated. [Australia.]
Diagnosis Whittenella peltosema is a narrow-winged, small moth with pale greyish forewings with a greenish tinge when fresh and with a dark grey-brown to blackish streak (male) or large triangle (female) along distal 36: of the costa, with the terminal fringe white just behind the nearly falcate apex and a black basal line along its central part. The labial palpi are rather short, porrect, sinuate, with the second segment greatly widened by long, loose scales, and the terminal segment long. The M-stem is distally very close to the hindmargin of the cell and R4 and R5 are stalked in the forewing. The hindwing has persistent scales along distal half of costa and the base of M2 is curved towards the base of the stalked M3 and CuA1. The male genitalia are characterised by a unique tegumen with a cone-like process with two very long, slender, downcurved, distally widened and bristled arms from its apex, here interpreted as socii, a simple valva with a very large costal process, furcate scales just beyond the
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colour pale grey with a greenish tinge in fresh specimens, with a dark grey-brown to blackish streak along distal 36: of costa in male, expanded to a broad triangle with apex beyond middle of wing in female, and in male with an irregular dark suffusion sprinkled with blackish scales along basal third of dorsum; costal strigulae black dots in basal half and paired white oblique lines more distally in dark mark, with a vertical white mark near apex; termen with fringe conspicuously white just below apex and basal line of fringe black along central part. Hindwing with cubital pecten and axillary tuft well developed, with costa moderately sinuate and scales along distal half more persistent; Rs and M1 closely approximated in basal fourth, base of M2 curved towards base of stalked M3 and CuA1; CuP present at margin only; 1A+2A present but with 1A only a short stump, 3A weakly indicated near margin; light brownish grey with scales small and wing semitranslucent except near margin. Pregenital abdomen. Tip of ventral apodemes widened, lightly club-shaped. In male hindmargin of S8 with two projecting, rectangular lobes separated by U-shaped excavation, T8 shorter, with convex hindmargin. Male genitalia (Figs 836, 837). Tegumen robust, rectangular, with strongly projecting ‘shoulders’ and a cone-shaped median process with two very long, slender, downcurved arms separately from its apex, widened and loosely bristled in apical third, interpreted as socii. Gnathos two short, straight, lateral arms just below middle of tegumen, medially connected with indistinct sclerotisation along base of anal tube. Valva simple, rather narrow, only slightly narrowed
Figs 833, 834.
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beyond middle, with a very large costal process, a large basal excavation with a clump of furcate scales just beyond its distal edge, and with an ovate cucullus densely bristled along projecting ventral margin. Juxta triangular, caulis very short, anellus a broad band. Aedeagus short, straight, gradually tapering towards apex, with a few fixed thorns laterally in distal third; vesica with a very large bundle of deciduous cornuti; ejaculatory duct extremely long. Female genitalia (Figs 835, 838). Ovipositor lobes slender, elliptical. Distal portion of ductus bursae apparently fused with narrowed apex of S7, entrance to ostium behind apex of S7, simple, spinulose. Sternum 7 distally narrowed, subtriangular, with strongly sclerotised, keelshaped ridges along lateral margins meeting medially to form short triangular process ventrally to ostium. Ductus bursae extremely long and slender, with numerous coils, membranous except for posteriormost portion consisting of two lateral, elongate sclerites from each side of ostium (?colliculum) followed by short, laterally flattened, evenly sclerotised section with ductus seminalis originating from its middle; corpus bursae very small, roundish, conspicuously wrinkled, with two flat, dagger-shaped signa with round, hollow base, each sitting in small, externally protruding membranous bulge of bursa wall (Fig. 838).
Distribution Whittenella is so far only known from Australia where it has been collected in Queensland along the eastern coast from Cooktown to Lamington NP.
Whittenella peltosema, adults, Rockhampton, Q. 833, . 834, . Scale = 2 mm.
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Biology No host records are known for Whittenella, but all localities are in rainforest or monsoon forest habitats.
Remarks Whittenella is characterised by several apomorphies in the genitalia of both sexes, in particular by the highly derived tegumen without uncus but with long, finger-shaped, distally bristled socii, the patch of furcate scales on the valva, and the aedeagus with external, fixed thorns and an extremely long ejaculatory duct in the male, and the correspondingly long ductus bursae in the female, posteriorly sclerotised and fused with a modified S7.
Whittenella is highly derived and no sister group has been identified. The long, paired, distally hairy processes from the apex of the tegumen are here treated as socii, but it cannot be ruled out that they may represent the uncus. However, if they indeed are socii they may provide a link with Heleanna Clarke, an association supported also by the persistent scales distally on the hindwing costa. The wing venation and the raised scales on the forewing dorsum also would agree with a position in the Acroclita-group. This genus is named to honour Dr Max Whitten, a former Chief of CSIRO Entomology, who strongly supported taxonomic research at the ANIC and encouraged the phylogenetic study that has finally resulted in this book.
Figs 835–839. Whittenella peltosema, genitalia and head. 835, female genitalia, Yeppoon, Q, slide ANIC 13802. 836, 837, male genitalia and aedeagus, nr Mt Garnet, Q, slide T2228. 838, detail female genitalia with signa. 839, head, male.
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Constituent species peltosema (Lower, 1908), Trans. R. Soc. S. Aust. 32: 323 (Palaeobia). [Qld: Mackay.] Comb. nov. confusa (Turner, 1925), Trans. R. Soc. S. Aust. 49: 55 (Acroclita). [Qld: Brisbane.]
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UNASSIGNED TO GENUS ‘Acroclita’ tothastis Meyrick (Figs 840–844) is definitely not a species of Acroclita but can at present not be assigned with confidence to any existing eucosmine genus. In the forewing R4 and
Figs 840–844. ‘Acroclita’ tothastis, adults, head and genitalia. 840, 841, adults. 840, , nr Ravenshoe, Q [image reversed]. 841, , Cairns, Q. 842, head, male. 843, male genitalia, nr Ravenshoe, Q, slide ANIC 13781. 844, female genitalia, nr Cairns, Q, slide ANIC 13788. Scale = 2 mm.
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R5 are stalked and M3 and CuA1 entirely fused into a single vein, in the hindwing Rs and M1 and M3 and CuA1 are stalked. There are several apparently closely related named and unnamed Australian and Oriental species, all very small and narrow-winged with variably derived and often reduced venation, and with male genitalia with large, membranous socii and a rather simple valva (Fig. 843). The female genitalia of tothastis (Fig. 844) with a conspicuously triangular base to the signa and a weakly sclerotised prong in the bursa neck are more informative than those of the male. Protancylis Diakonoff from Saudi Arabia (Diakonoff 1983b) has very similar genitalia and wing pattern, though the latter could simply be a symplesiomorphy. However, tothastis has a much more derived wing venation, and only study of all the related Australian and Oriental taxa will resolve the identity and scope of the genera concerned.
Constituent species tothastis (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 243 (Acroclita). [Qld: Cairns.]
GRAPHOLITINI Guenée Furumi Komai and Marianne Horak Grapholithidi Guenée, 1845, Ann. Soc. Entomol. Fr. 2(3): 169. Ephippiphoridii Pierce & Metcalfe, 1922, Genitalia Group Tortricidae Lepid. Br. Is.: xxi, 74. Lipoptychidii Pierce & Metcalfe, 1922, Genitalia Group Tortricidae Lepid. Br. Is.: xxi, 93. Laspeyresiinae Heinrich, 1923, US Nat. Mus. Bull. 123: 10. Laspeyresiini: Obraztsov, 1946, Z. Wien. Entomol. Ges. 30(1945): 23.
Status remark The current concept of this group was established by Heinrich (1923) as a subfamily, Laspeyresiinae, but subsequent authors, e.g. Obraztsov (1946, 1958), Hannemann (1961), Danilevsky and Kuznetzov (1968), treated the same group as Laspeyresiini at tribal level. Razowski (1976) indicated that the family-group name, Laspeyresiinae or Laspeyresiini, is invalid, because
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its type genus Laspeyresia Hübner, 1825 is a homonym of Laspeyresia R. L., 1817 (an emendation of Laspeyria Germar, 1810, Noctuidae) and proposed the usage of Grapholitini stemming from the oldest family-group name, Grapholithidi Guenée, 1845. The tribe includes some 880 species in about 50 genera worldwide (Brown 2005).
Evidence for monophyly The Grapholitini traditionally have been defined by a hindwing venation with M2 distant and parallel to M3, and by reduced male genitalia including loss of uncus and socii (Heinrich 1923; Danilevsky and Kuznetzov 1968). However, there are some exceptions to this both in the Grapholitini and in non-grapholitine tribes. In Acanthoclita Diakonoff, Thaumatotibia Zacher, and some species of Cryptophlebia Walsingham (Grapholitini), the hindwing venation is the same as in the Eucosmini: M2 at its base curved towards M3, but uncus and socii are rarely present in species or genera undoubtedly belonging to the Grapholitini. Conversely, a similar hindwing venation as in Grapholitini is found in some species of the Olethreutini and the Enarmoniini. Horak and Brown (1991) regarded the tribe as a polyphyletic assemblage, i.e. representing not a clade but a grade characterised by a particular hindwing venation and atrophy of structures in the male genitalia. However, Komai (1999) demonstrated that most genera that have been assigned to the Grapholitini constitute a monophyletic group that is supported by the shared possession of a shortened S8 in the male with a straight posterior margin. In all other Olethreutinae, S8 in the male is as long as T8, and often has a bilobed posterior margin. Komai (1999) hypothesised that the grapholitine S8 has its distal half invaginated, forming a transverse fold. In some Grapholitini, a brush organ is formed between A8 and A9 and S8 is modified into an element of the brush organ (Grapholita-group). The brush organ consists of three elements: 1, a semicircular or crescent-shaped eighth sternite; 2, an intersegmental ventral sclerite between A8 and A9 that is folded into the abdomen except during mating, and 3, a pair of scale-tufts situated on the pleural area near the distal ends of the levers of the intersegmental ventral sclerite. This organ is regarded as a synapomorphy for the Grapholitagroup, and is referred to as ‘coremata’ (Komai 1999). The present study follows this usage.
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Diagnosis Very small to medium-sized moths with sometimes a rather simple forewing pattern consisting of a dorsal blotch or median fascia and a well-defined ocelloid patch on fuscous ground colour. These pattern elements are often reduced or lost. Forewing without costal fold except for Dichrorampha Guenée, often with well-developed chorda and M-stem. Hindwing with bases of Rs and M1 approximate or distant (Dichrorampha-group, Microsarotis Diakonoff, and Commoneria, gen. nov.), and usually with M2 distant and parallel to M3 (but in Acanthoclita, Thaumatotibia, and some species of Cryptophlebia, M2 at its base curved towards and approximated to the stalk of M3 and CuA1 as in the Eucosmini), with 3A usually present; sometimes with anal fold with modified fringe in male (Cydia-group). Female frenulum with two or three bristles. Sternum 2 usually of ‘tortricoid type’ consisting of a pair of anterolateral processes and a pair of apodemes; in Loranthacydia Horak, Common & Komai and Parapammene Obraztsov, anterolateral processes completely reduced (Fig. 977). Sternum 8 a short plate with an almost straight posterior margin (Fig. 881) (Dichrorampha and Cydia-groups) or S8 modified into an element of the coremata (Fig. 948) (Grapholita-group) (but in Loranthacydia-group S8 of non-grapholitine type). The Grapholitini, especially the Grapholita-group, are characterised by the possession of various types of abdominal scent organs. Beside the coremata, a transverse patch of scales or a pair of scale-tufts may be present on A4–A8, e.g. in Pammenopsis Kuznetzov with two large, deeply invaginated bundles of modified scales laterally between A6 and A7 and two bands of modified, broad scales anteriorly on T4 and T5 (Fig. 944); in Microsarotis Diakonoff two very large scale bundles deeply invaginated between T7 and T8 (Fig. 962); in many species of Parapammene a transverse patch of narrow, hairlike scales and modified black scales present on T4 and T5 respectively; in Matsumuraeses Issiki, two pairs of scale-tufts present on T7; in Cryptophlebia, Pseudopammene Komai, Strophedra HerrichSchäffer, and some species of Andrioplecta Obraztsov, a pair of scale-tufts arise from membranous, shallow pockets formed in the anterior corner of T8 (Fig. 925). The male genitalia of Grapholitini are best characterised by their simplification through degeneration or loss of the appendages of the tegumen: uncus usually absent, rarely rudimentary (some species of Cryptophlebia,
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Thaumatotibia, and Pammene Hübner); socius usually absent or only indicated by a patch of hairs, but in Leguminivora Obraztsov, Matsumuraeses and in some species of Fulcrifera Danilevsky & Kuznetzov, Notocydia, gen. nov., and Thaumatotibia, rudimentary socii present. Tegumen with pedunculi almost straight in the Loranthacydia-, Dichrorampha-, and Cydia-groups, but with pedunculi strongly curved and articulated with the anterior edge of vinculum in the Grapholita-group. Valva usually a well-sclerotised, flattened sac (but in Cryptophlebia greatly inflated, Fig. 919), usually with costal process indicated by short finger-like projection (absent in Loranthacydia, Pseudopammene and Dierlia Diakonoff). Cucullus usually well-defined and its inner surface with numerous bristles or hairs. Sacculus sometimes strongly concave on ventral margin near base (Cydia-group). Aedeagus well sclerotised, usually narrowed distally, sometimes curved. Bulbus ejaculatorius (distal part of the ductus ejaculatorius) very short in Grapholitini, while long and often coiled in Olethreutini and Eucosmini. Female genitalia with ostium bursae usually located between A7 and A8; ductus bursae usually sclerotised posteriorly to form a funnel, cup, or narrow ring, with a sclerotised ring posterior to the origin of the ductus seminalis; ductus seminalis arising from the middle to the anterior end of the ductus bursae; bulla seminalis always present, sometimes (e.g. in Selania Stephens, Andrioplecta, Ixonympha, gen. nov., and some species of Grapholita Treitschke) directly connected with corpus bursae or intervening ductus seminalis very short. Corpus bursae ovate, a diverticulum usually present in the Cydia-group, sporadically present in the Grapholita-group (Ecdytolopha Zeller, and some species of Thaumatotibia, Cryptophlebia, Selania, and Grapholita), always absent in the Loranthacydia and Dichrorampha-groups. Signa usually two horn- or blade-shaped structures; in Selania, Commoneria, gen. nov. and Ixonympha, gen. nov. signum reduced or absent. Sternum 7 usually a rectangular plate with an excavated posterior edge.
Distribution The tribe is distributed worldwide, adventive in New Zealand. The Loranthacydia-group is endemic to Australia. In the Dichrorampha-group, Dichrorampha has a Holarctic distribution with high diversity in the Palaearctic region (c. 100 species), Pammenemima is Indo-Australian, and
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‘Hemimene’, with many species referred to it by Meyrick, has its greatest diversity in the Neotropical region. The Cydia-group is concentrated in the Old World, with only Cydia present also in the New World. The Grapholita-group also is much more diverse in the Old World.
Biology The majority of Grapholitini larvae are borers in roots, stems, fruit, seeds, and buds of many plant families, but some are leaf tiers and leaf skeletonizers. The tribe includes many pest species, e.g. oriental fruit moth (Grapholita (Aspila) molesta (Busck)), codling moth (Cydia pomonella (Linnaeus)), false codling moth (Thaumatotibia leucotreta (Meyrick)) and macadamia nutborer (Cryptophlebia ombrodelta (Lower)). Female sex pheromones or sex attractants have been identified in 98 grapholitine species of 13 genera (Komai 1999; Witzgall et al. 2004). The pheromone components are 12 carbon acetate or alcohols with double bonding at various positions, which are roughly specific to each of three genusgroups: Dichrorampha-, Cydia-, and Grapholitagroups (Komai 1999).
Remarks Heinrich’s (1926) revision of the Nearctic fauna was the first comprehensive study of the Grapholitini, and his taxonomic concept was essentially followed by subsequent authors. Obraztsov (1958–1967) published a series of revisions of the Palaearctic olethreutine genera with checklists of species. Danilevsky and Kuznetzov (1968) produced a monumental monograph on the Palaearctic representatives, with a schematic phylogenetic tree illustrating generic relationships. Razowski (1989) reviewed the genera of the Palaearctic Olethreutinae. More recently, Komai (1999) revised the Palaearctic grapholitine fauna and provided a phylogenetical hypothesis for generic relationships. In addition to the comprehensive revisionary studies, there are many faunal treatments of specific regions or countries (Meyrick 1895, 1928; Kennel 1921; Benander 1950; Van Deurs 1956; Hannemann 1961; Bentink and Diakonoff 1968; Bradley 1972; Liu and Bai 1977; Kawabe 1982; Kuznetzov 1978; Liu and Li 2002; Razowski 2003a). Danilevsky and Kuznetzov (1968) divided the tribe into two subtribes: Dichroramphae and Laspeyresiae. Razowski (1989) proposed the use of
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Lipoptychina for the former and of Grapholitina for the latter subtribe, as the oldest family-group names for the relevant taxa. Komai (1999) recognised three genus-groups in the Palaearctic Grapholitini: Dichrorampha genus-group, Cydia genus-group, and Grapholita genus-group. These groupings roughly correspond to Danilevsky and Kuznetzov’s phylogeny scheme (1968). In the course of revisional studies of the Australian grapholitine genera, four genus-groups were recognised: Loranthacydia-group, Dichrorampha-group, Cydia-group, and Grapholitagroup. The Loranthacydia-group is endemic to Australia, based on the single genus Loranthacydia. The remaining three groups correspond to the groupings by Komai (1999). In Australia, the Dichrorampha-group includes only Pammenemima Diakonoff (=Titanotoca Diakonoff), the Cydiagroup is represented by five genera (Leguminivora Matsumura; Fulcrifera Danilevsky & Kuznetzov; Notocydia, gen. nov.; Apocydia, gen. nov.; Cydia Hübner), and the Grapholita-group by 12 genera (Acanthoclita Diakonoff; Centroxena Diakonoff; Commoneria, gen. nov.; Cryptophlebia Walsingham; Grapholita Treitschke; Gymnandrosoma Dyar; Ixonympha, gen. nov.; Microsarotis Diakonoff; Archiphlebia, gen. nov.; Pammenopsis Kuznetzov; Parapammene Obraztsov; Thaumatotibia Zacher).
Loranthacydia-group Status remarks This genus-group is here newly defined for the single genus Loranthacydia Horak, Common & Komai.
Evidence for monophyly The genus is very diverse, and three informal species-group are recognised (for detail, see Remarks for Loranthacydia). However, two significant synapomorphies link those three species-groups: 1, absence of the costal process of the valva, and 2, absence of the anterolateral processes on S2.
Distribution This group with 10+ species in a single genus, Loranthacydia, is endemic to Australia and has been recorded from Queensland, Northern Territory,
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New South Wales, South Australia, and Western Australia.
Biology Two species of the metallocosma-group have been reared from stems of mistletoes (Loranthaceae).
Remarks Komai (1999: 28) suggested that a possible apomorphy linking most genera previously placed in the Grapholitini is a short S8 in the male with its posterior margin straight or S8 modified into an element of the coremata. However, in Loranthacydia males S8 is not short and the posterior margin is bilobed as in Eucosmini or Olethreutini, and there are no coremata. The general appearance of Loranthacydia and the combination of a hindwing with M2 parallel to and widely separate from M3, and the genitalia configuration in both sexes refer the genus to the Grapholitini sensu auct. Therefore, we tentatively place this group in the Grapholitini.
Loranthacydia Horak, Common & Komai (Figs 98, 845–853) Loranthacydia Horak, Common & Komai, 1996, Monogr. Aust. Lepid. 4: 136. Objective replacement name for Leptarthra Lower. Leptarthra Lower, 1902, Trans. R. Soc. S. Aust. 26: 253. Type species: Leptarthra aulacodes Lower, 1902, ibid. 26: 253, by monotypy. [Australia.] Leptarthra Lower is a junior homonym of Leptarthra Baly, 1861, J. Entomol. 1: 202, – Insecta, Coleoptera.
Diagnosis Very small to medium-sized moths with an either typically grapholitine appearance or, in the majority of species, with semiovate, often quite narrow forewings with a long, strongly oblique, curved termen merging without pronounced tornus into dorsum, with an indistinct pattern of numerous, parallel, outwardly angled darker and interrupted silvery lines across wing, over a light greyish, yellowish or brownish ground colour, with a series of minute black dots along termen. The
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palpi are small to very small, slender and weakly sinuate to straight. Absence of anterolateral processes on S2 is a diagnostic character for Loranthacydia. The male genitalia of Loranthacydia are characterised by the absence of a costal process of the valva, and, in the majority of species, by a long, slender valva with a dense patch of spines ventrally to the distal part of the basal excavation, and by a few prominent bristles on the apex of the tegumen. The female genitalia are characterised by a sclerotised ring or rim as sterigma, by daggershaped signa (unless reduced to horn-shaped or absent), and usually a V-shaped, dorsal sclerotisation from T8 connecting the dorsal arms of the apophyses anteriores.
Description Adults (Figs 845, 846) Wingspan. 6.5–16.0 mm; very small to medium-sized. Head (Fig. 847). Ocellus well developed, chaetosema distinct. Proboscis about 1¼× to 1½× length of labial palpus. Labial palpus slender, from very small to small (0.8–1.6× diameter of eye), porrect, from lightly sinuate to straight (very short palpi); in larger palpi second segment distally slightly widened, apical segment fairly large, ovate, with appressed scales; very small palpi slender throughout, division between second and apical segment not discernible without dissection. Antenna reaching to middle of forewing or just beyond, cilia in male barely discernible. Thorax. Without crest. Legs unmodified. Wings (Fig. 98). Forewing index 0.37–0.50; forewing from subtriangular (pessota-group) to narrowly or broadly semiovate with termen curved, strongly oblique and merging into dorsum without well-defined tornus; cell 0.59–0.63× length of wing, with chorda absent and M-stem variably developed even within a species, often only a remnant to between M2 and M3 or entirely absent; all veins present and separate beyond cell, R1 from beyond middle of cell, M3 and CuA1 from closely approximated at base (pessota-group) to nearly as distant as M2 and M3 (metallocosma-group, group C), CuA2 from well beyond 36: cell; CuP present at margin; forewing in pessota-group pale grey and blackish, with typically grapholitine wing pattern with the dorsal blotch prominent and the ocelloid patch indicated, preceded by a wide silvery streak; remainder of genus with strongly modified forewing pattern, of often yellowish, reddish brown or greyish colour, with a series of oblique dark
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Figs 845–848. Loranthacydia, adult females, head and abdomen. 845, L. sp. nr metallocosma, Townsville, Q. 846, L. sp. pessota-group, Barren Grounds, NSW. 847, L. sp., head, male. 848, L. nr pessota, male, slide ANIC 8161, tip of abdomen with scale pencils (arrows). Scale = 2 mm.
strigulae along costa, often with some, especially those just beyond 56M and those at 56O, giving rise to outwardly angled dark lines across wing, and entire wing with numerous, subparallel, outwardly angled, interrupted silvery lines across wing, originating from between dark strigulae on costa, fringe along termen often with a series of very prominent black dots. Hindwing with only very ephemeral cubital pecten, or cubital pecten absent, with small axillary tuft and unmodified anal margin in male; frenulum of two bristles in female; with Rs and M1 from closely approximated at base to fairly distant (group C) and M3 and CuA1 from connate to distinctly separate (group C), M2 widely distant from M3, 3A well developed; from pale grey to blackish brown, yellow in several species of the metallocosma-group.
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Pregenital abdomen. Anterolateral processes on S2 absent; in males of some species of the pessota-group a dorsal pair of deeply invaginated bundles of long scales in the membrane between T6 and T7 (Fig. 848). Sternum 8 of male with posterior margin bilobed (see Remarks under Loranthacydia-group). Male genitalia (Figs 850, 851, 852). Vinculum a moderately narrow band. Tegumen high, subtriangular to ovate, a rather narrow band laterally, broader across apex, rarely with two lateral patches of short bristles below apex (C-group), usually with a few single, long bristles sublaterally and sometimes dorsally on apex. No trace of uncus present. Gnathos at most indicated by weakly sclerotised lateral triangles. Valva diverse, either long and slender, sometimes medially slightly
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constricted but otherwise of roughly equal width throughout, rarely upcurved but usually more or less straight, with a rounded apex and a patch of dense, strong spines below distal part of basal excavation, sometimes developed into a ventromedially projecting triangular lobe, or valva of more typically grapholitine shape with a deep emargination on the ventral margin at the base of the cucullus (L. metallocosma); distal half of valva densely hairy, with stronger bristles along distal margin; costal process of valva always absent.
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Caulis rather wide and short, fused with base of aedeagus. Aedeagus lightly curved or faintly sinuous, with bulbous base, tapering to bevelled apex; vesica with a dense bundle of long, slender cornuti in one species, but without traces of sockets in others. Female genitalia (Figs 849, 853). Ovipositor lobes from subreniform, flat and wide with particularly large rounded distal lobe (pessotagroup) to long, narrow and somewhat swollen (L. metallocosma), often with a triangular projection
Figs 849–853. Loranthacydia, genitalia. 849, L. nr pessota, female genitalia, nr Eucla, WA, slide ANIC 8161. 850, L. nr pessota, male genitalia, nr Yalata Mission, WA, slide ANIC 8610. 851, 852, L. sp. nr metallocosma, aedeagus and male genitalia, Townsville, Q, slide ANIC 4974. 853, L. sp. nr metallocosma, female genitalia, Townsville, Q, slide ANIC 4977.
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between the posterior lobes. Apophyses short, and in some species very broad, often with hooked or paddle-shaped apices. Dorsal arms of apophyses anteriores often connected with a V-shaped dorsal sclerotisation projecting anteriorly from T8. Sternum 7 with a curved, concave hindmargin, a rectangular to posteriorly narrowed plate or medially constricted with strongly excavate, concave lateral margins (metallocosma-group). Sterigma either a flat, sclerotised ring or a raised, round rim, in the metallocosma-group lamella postvaginalis with a tongue-shaped sclerotisation extending posteriorly from dorsal part of sclerotised ring. Ductus bursae usually shorter than corpus bursae, with a thick-walled colliculum just below sterigma and often with a small sclerite at inception of ductus seminalis (about halfway along ductus bursae); bulla seminalis very large in metallocosma-group; corpus bursae ovoid to teardrop-shaped, without diverticulum, with two signa except in group C, where signa are absent; signa dagger-shaped with a narrow, hollow base and a long, flattened blade, or variously reduced to small blades or often quite small, short, hollow horns
Distribution Endemic to Australia and recorded from all Australian states and territories except Tasmania, Victoria and the Australian Capital Territory, from arid areas such as Alice Springs to rainforest localities in northern Queensland (Babinda). Group C has only been collected in the northwestern corner of Australia.
Biology The labels of Dodd’s material in the ANIC of two species of the metallocosma-group from Townsville have the remark ‘bred’ on their label, and Meyrick (1911b), based on material from the same two series, reported that they both feed in ‘stems of Loranthus (Loranthaceae)’. The two species have a slightly different biology. One, an unnamed sibling species of L. metallocosma (Lower) and referred to as L. metallocosma by Meyrick (1911b: 287), ‘feeds in stems of Loranthus (Loranthaceae), a large gall forming round each, which splits open at the upper end in time for the moths to emerge; many on one stem, often only an inch or two apart (Dodd)’. The second Townsville species, identified as L. aulacodes by Meyrick, (though L. aulacodes has been described from Derby, WA, with the type not found) is said by Meyrick (1911b: 287) to feed ‘in stems of Loranthus, but without making a gall
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(Dodd)’. Since this time, these species of Australian mistletoes have been extensively revised and, while they are now placed in many genera, the genus Loranthus has been excluded from Australia. Almost certainly Dodd’s records pertain to Loranthaceae. If Loranthaceae are the hosts for all species of Loranthacydia it could possibly explain the extraordinarily wide distribution of the genus and its very high diversity suggested by the numerous species represented by single specimens in the ANIC.
Remarks Lower (1902) based Loranthacydia (as Leptarthra) on the single species L. aulacodes from Derby, Western Australia. None of the two syntypes of L. aulacodes could be found, and all material with the drawer label L. aulacodes in the SAMA is from Townsville and Mackay or has no locality label. Meyrick (1911b) identified material reared by Dodd from Townsville as L. aulacodes (part of the series in the ANIC), and Lower’s description agrees with the Townsville material. However, given that there is material of two other very similar-looking species from Queensland in the ANIC it is unlikely that the Townsville specimens are conspecific with material from Derby. There is, however, no doubt about the generic identity of Loranthacydia. Absence of the costal process of the valva and absence of the anterolateral processes on S2 are two very obvious apomorphies for Loranthacydia. The genus is very diverse and we recognise three informal species-groups, of which the first is probably merely a plesiomorphic assembly: 1, the pessota-group (containing L. pessota (Meyrick) and probably further unnamed species) with its typically grapholitine appearance, possibly characterised by a rim-shaped sterigma; 2, the metallocosma-group (containing L. aulacodes, L. metallocosma, L. multilinea (Turner), L. sinapichroa (Turner), and several unnamed species) with females with laterally concave margins of S7 and with a tongue-shaped lamella postvaginalis; 3, group C without any named species, with a strongly upcurved valva and no signa. The two species, L. aulacodes and L. pessota, were included in Loranthacydia in Horak et al. (1996) without the statement that this represented a new combination in both cases.
Constituent species Five named and many more unnamed species are known from Australia.
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aulacodes (Lower, 1902), Trans. R. Soc. S. Aust. 26: 253 (Leptarthra). [WA: Derby.] metallocosma (Lower, 1902), Proc. Linn. Soc. NSW 26: 670 (Tortrix). [Qld: Cooktown.] sinapichroa (Turner, 1926), Trans. R. Soc. S. Aust. 50: 139 (Laspeyresia). [Qld: Dalby.] multilinea (Turner, 1945), Trans. R. Soc. S. Aust. 69: 71 (Schoenotenes). [Qld: Brisbane.] pessota (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 290 (Laspeyresia). [SA: Port Lincoln.]
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Biology Larvae of Dichrorampha species are root-borers of Asteraceae, e.g. Achillea, Chrysanthemum, and Artemisia (Bradley et al. 1979). Pammenemima is associated with Desmodium (Fabaceae), with the larvae living in tied leaves. Known sex attractants for some Dichrorampha species are (E)–9-dodecen-1-ol acetate (E9–12Ac) and (E)–9,11-dodecadien–1-ol acetate (E9,11–12Ac) (Ulenberg et al. 1993).
Remarks
Dichrorampha-group Status remarks Danilevsky and Kuznetzov (1968) established the subtribe Dichroramphae for Dichrorampha Guenée. Razowski (1989), however, proposed the usage of Lipoptychina instead of Dichroramphae since the former name is older. Komai (1999) referred to the group as the Dichrorampha genus-group and added Pammenemima Diakonoff (=Titanotoca Diakonoff) to the group. This group contains about 140 species in three genera including ‘Hemimene’ sensu Meyrick worldwide, of which Pammenemima is present in Australia.
The Dichrorampha-group has a sister group relationship with the Cydia-group + the Grapholitagroup (Komai 1999).
Pammenemima Diakonoff (Figs 100, 854–858) Pammenemima Diakonoff, 1982, Zool. Verh. (Leiden) 193: 23. Type species: Lipoptycha ochropa Meyrick, 1905, J. Bombay Nat. Hist. Soc. 16: 587, by original designation. [Sri Lanka.] Titanotoca Diakonoff, 1984a, Entomol. Basil. 9: 380. Type species: Titanotoca pagerostoma Diakonoff, 1984a, ibid. 9: 381, by original designation. [Indonesia.] Syn. nov.
Evidence for monophyly Komai (1999) proposed the following three synapomorphies for the genera of the Dichrorampha genus-group: 1, Rs and M1 in hindwing separate and parallel or subparallel; 2, presence of a row of dots along termen on forewing; 3, sterigma, seventh sternite, and sclerotised part of ductus bursae completely fused in female genitalia.
Distribution Dichrorampha is Holarctic (112 spp.) and the majority of species occurs in the Palaearctic region (96 spp.), with concentrations in the Alps, the Caucasus, and Central Asia (Danilevsky and Kuznetzov 1968). Pammenemima (six spp.) is known from Sri Lanka, India, Indonesia, Philippines, Solomon Is and Australia (Northern Territory and Queensland). Many Neotropical species from Ecuador, Colombia, Guyana, Peru, Bolivia, and Brazil allocated to ‘Hemimene’ in ‘Exotic Microlepidoptera’ by Meyrick (1916, 1922, 1927c, 1932) (26 spp.) probably belong to the Dichrorampha-group.
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Diagnosis Species of Pammenemima are small to very small, rather broad-winged brownish grey moths with a faint coppery gloss, a bluntly rounded apex and a termen with a shallow notch between R5 and M2, with an indistinct grapholitine wing pattern including white costal strigulae, an indistinct dorsal blotch and two to three black dashes or dots close to the termen, but always with a row of blackish scales around apex and termen. The characteristically long white labial palpi of Pammenemima are strongly upcurved, slender, distally pointed and reaching to the upper margin of the eye. In the forewing chorda and M-stem are present and all veins are distant from each other at base, in the hindwing Sc+R1 and Rs are rather close to end of cell, Rs and M1 are at some distance from each other even at base, M2 is parallel and distant from M3, and M3 and CuA1 are either nearly connate or stalked. In both wings CuP is absent or a very weak trace only at margin. The male genitalia of Pammenemima are characterised by a weakly
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sclerotised, triangular tegumen with its apex produced into a short, tongue-shaped process, by a medially narrowed valva widest near base with a triangularly projecting sacculus, by a cucullus with spines along its inner ventral half, and by a large and strongly sclerotised aedeagus, diagnostic with its toothed or serrate apex with at least one large, additional spike. The female genitalia of Pammenemima are conspicuous and unique with a complex sterigma entirely fused with S7, with a lightly sclerotised, small, ovate accessory sac branching off below ostium, with the corpus bursae
broadly connected to the bulla seminalis and with two large, broadly hook-shaped signa.
Description Adults (Fig. 854) Wingspan. 7.0–9.5 mm; very small to small. Head (Fig. 855). Ocellus large, chaetosema with few long bristles. Labial palpus sharply upcurved, slender, pointed, rather long (1.9–2.1× diameter of eye), reaching to upper margin of eye, white, with second segment strongly curved at base,
Figs 854–858. Pammenemima tetramita, adult, head, genitalia and abdomen. 854, adult, female, nr Babinda, Q. 855, head, male. 856, male genitalia, syntype, Kuranda, Q, slide ANIC 8176. 857, male abdomen, 8th segment. 858, female genitalia, nr Mt Cahill, NT, slide ANIC 8175. Scale = 2 mm.
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slender and nearly parallel-sided with short, loose scales along ventral margin, terminal segment rather long, pointed. Proboscis about as long as palpus. Antenna to beyond middle of wing, stout in male, with short, semi-erect scales intermixed with short cilia. Thorax. Smooth. Legs unmodified. Wings (Figs 100, 854). Forewing index 0.41– 0.45. Forewing broadly subtriangular, slightly widening towards termen, with an evenly curved costa, a broadly rounded apex and a hardly oblique, slightly sinuate termen with a shallow notch between R5 and M2; with cell widening distally, chorda from halfway between R1 and R2 to between R4 and R5, M-stem to between M2 and M3, closing vein in P. tetramita (Turner) only trace in membrane; all veins separate and distant from each other at base; in P. tetramita at most a faint trace of CuP in membrane at margin; anal loop 0.26× length of 1A+2A; with a rather indistinct but typically grapholitine wing pattern, grey-brown ground colour with coppery gloss and with a series of whitish strigulae along costa, a row of blackish scales around apex and along notched termen, dorsal blotch only indicated as a series of indistinct, irregular paler bands, two or three black dashes or dots very close to termen. Hindwing usually without cubital pecten (present in P. ochropa) and with anal tuft of only a few hairs; frenulum with two bristles in female; venation not sexually dimorphic; course of Sc+R1 and Rs moderately close to end of cell, bases of Rs and M1 parallel but quite distant; M1, M2 and M3 subparallel and distant, M2 and M3 somewhat closer and slightly converging towards base; M3 and CuA1 almost connate (P. ochropa) or stalked (P. pagerostoma, P. tetramita); CuP a very faint trace at margin; 1A+2A and 3A well developed; grey-brown with a coppery gloss. Pregenital abdomen. Sternum 8 of male narrow, hindmargin nearly straight; T8 with a weak, inverted T- to Y-shaped sclerotisation in anterior margin, hindmargin projecting as a rounded lobe (Fig. 857). Coremata not developed. Male genitalia (Fig. 856). Vinculum a narrow band. Tegumen weakly sclerotised, triangular, apex rounded-triangular to produced into more strongly sclerotised, short tongue-shaped, sometimes haired lobe. No trace of socii. Gnathos two membranous lateral bands medially fused with anal tube. Valva widest near base, weakly to moderately narrowed below cucullus, with large basal excavation, sacculus projecting triangularly and sometimes produced into sharp point (P. tetramita) or with a cluster of spines (P. ochropa), inner surface of cucullus with
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spines along ventral half; costal process not apparent. Juxta directly fused with anellus, caulis not developed. Aedeagus large, wide and strongly sclerotised, in P. tetramita very short and not tapering except for oblique apex, sometimes surface covered with thorns of increasing size towards apex where they are very prominent (P. tetramita), always a somehow toothed or serrate apex with at least one large, additional spike; vesica with a sheaf of lanceolate, deciduous cornuti with a short basal process. Female genitalia (Fig. 858). Ovipositor lobes subreniform. Sternum 7 weakly sclerotised, with Vshaped, deep incision from hindmargin nearly to base of S7, with paired, concave, smooth fields posteriorly; anterior margin forming raised, ragged rim. Ostium at base of incision of S7. Sterigma entirely fused with S7, a complex, nose-shaped projection. A lightly sclerotised, small, ovate accessory sac branching off below ostium, ventrally to ductus bursae. Ductus bursae extremely short, sclerotised ventrally; teardrop-shaped corpus bursae with spinules in ‘neck’ and broadly connected with bulla seminalis, with two large, clavate to broadly hook-shaped signa with hollow base.
Distribution Pammenemima is known from Sri Lanka, India, Indonesia, Philippines, Solomon Is and Australia, where it has been collected in Darwin and Kakadu NP in the Northern Territory and the Cairns region in northern Queensland.
Biology The type material of P. exocentra (Meyrick) was reared from larvae on Desmodium ovalifolium (Fabaceae) in Bogor, Java. Fletcher (1932) described and figured the feeding mode of P. ochropa on a Desmodium sp. in Pusa (India), with larvae rolling together 4–5 leaves and sometimes boring into buds and stipules. F. Komai observed the adults of P. tetramita frequenting Desmodium sp. in Holmes Jungle Nature Park, Darwin, Northern Territory.
Remarks Diakonoff (1982) provided drawings of head, wing venation and genitalia of both sexes for P. ochropa, a drawing of the male genitalia of P. pagerostoma (Diakonoff 1984a), and a drawing of the female genitalia of P. ionia (Wileman & Stringer). The figure given in the same publication for the alleged male of P. ionia refers to a species of Grapholita.
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Fletcher (1932) gave drawings of larva, pupa and adult of P. ochropa and described its biology in detail. Clarke (1958) figured adult and genitalia of the types of P. exocentra (Meyrick), P. ochropa (Meyrick) and its junior synonym P. dissias (Meyrick). The monophyly of Pammenemima is based on several conspicuous apomorphies: a venation with CuP reduced or lost in both wing, a strongly sclerotised large aedeagus with a serrate apex with one or more additional spikes, S7 in the female with two paired, large, smooth, ovate patches posterior to sterigma, an ovate accessory sac from near ostium and broadly hook-shaped signa. Pammenemima pagerostoma, the type species of Titanotoca, differs from P. ochropa, the type species of Pammenemima, by stalked rather than separate M3 and CuA1 in the hindwing and by a medially less constricted valva with the sacculus ending in a sharp point but lacking the spine cluster present in P. ochropa. However, the many synapomorphies and the fact that both P. ochropa and P. exocentra, the latter with male genitalia very similar to those of P. pagerostoma, have been reared from Desmodium strongly support the synonymy of the two genera. Pammenemima shares the following characters with Dichrorampha, suggesting a close relationship between the two genera: forewing with two to three black dashes or dots close to the termen, the hindwing venation with Rs and M1 parallel and rather distant at base, the tegumen with short tongue-shaped lobe at the apex, and the sterigma entirely fused with S7. Superficially Pammenemima species recall some species of Dichrorampha. The following four species are here referred to Pammenemima: exocentra (Meyrick, 1939), Trans. R. Entomol. Soc. Lond. 89: 52 (Laspeyresia). [Java.] Comb. nov. ionia (Wileman & Stringer, 1929), Entomologist 62: 67 (Pammene). [Philippines.] Comb. nov. plumbosana (Bradley, 1961), Bull. Br. Mus. (Nat. Hist.), Entomol. 10: 127 (Laspeyresia). [Solomon Is.] Comb. nov. pagerostoma (Diakonoff, 1984a), Entomol. Basil. 9: 381 (Titanotoca). [Sumba.] Comb. nov.
Constituent species Five species of Pammenemima have been named from outside Australia, with P. plumbosana and P. pagerostoma probably both junior synonyms of P. tetramita, the only Australian species of the genus.
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tetramita (Turner, 1925), Trans. R. Soc. S. Aust. 49: 60 (Laspeyresia). [Qld: Kuranda.] Comb. nov.
Cydia-group Status remarks Komai (1999) proposed the Cydia genus-group for Leguminivora Obraztsov, Fulcrifera Danilevsky & Kuznetzov, Cydia Hübner, and Lathronympha Meyrick. This group, together with Komai’s (1999) Grapholita genus-group, roughly corresponds to the Laspeyresiae established by Danilevsky and Kuznetzov (1968). The Cydia-group contains about 240 species in six genera worldwide, of which two Australian genera, Apocydia, gen. nov. and Notocydia, gen. nov. are added here.
Evidence for monophyly The monophyly of the Cydia-group is supported by the following three apomorphies: 1, ventral margin of sacculus concave near base; 2, presence of anal fold on male hindwing, 3, vein 3A close to anal edge on male hindwing (Komai 1999).
Distribution Cydia is known from the Old and the New Worlds. Leguminivora and Fulcrifera have Palaearctic-Oriental-Australian distributions. Lathronympha, which Komai (1999) tentatively allocated to this group, is recorded from Europe, south China, and the Russian Far East. Apocydia, gen. nov. and Notocydia, gen. nov. are endemic to Australia.
Biology In the Palaearctic region, Cydia is associated with various plant groups, e.g. Coniferales, Fagales, Fabales (Danilevsky and Kuznetzov 1968). Larvae bore in stems, fruit, seeds, and cones or live under bark. In Australia, where this genus is represented by very few species including the accidentally introduced codling moth, Cydia pomonella (Linnaeus), one undescribed species has been reared from seed capsules of Dodonaea viscosa (Sapindaceae). Leguminivora, Fulcrifera, and Notocydia, gen. nov. have been reared from seed pods of Fabales.
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Sex pheromones or sex attractants have been identified for 37 species of Cydia. (E, E)–8,10dodecadien–1-ol acetate (E8E10–12Ac) is predominantly used as the main pheromone component (Komai 1999; Witzgall et al. 2004).
Remarks The Cydia-group is closely related with the Grapholita-group, as indicated by the common possession of the following two specialised traits: 1, crescent-shaped bulbus ejaculatorius; 2, presence of a diverticulum on the corpus bursae (Komai 1999). The latter character is prevalent in the Cydiagroup, but occurs sporadically in the Grapholitagroup.
Leguminivora Obraztsov (Figs 101, 859–863) Leguminivora Obraztsov, 1960, Tijdschr. Entomol. 103: 129. Type-species: Grapholita glycinivorella Matsumura, 1898, Dobutsugaku Zasshi 10: 127, by original designation. [Japan.]
Diagnosis Leguminivora species are small to medium-sized grapholitine moths with indistinct wing pattern, brown to dark brown densely irrorated with whitish, with whitish costal strigulae, obvious ocelloid patch and faint dorsal blotch. The labial palpus is moderately long and moderately wide, sinuate and slightly upcurved. The genus shares with other genera of the Cydia-group a hindwing with M3 and CuA1 stalked, in the male with 3A approximated to the thickened anal margin that is rolled upwardly to form an anal fold with modified fringe, enclosing a hairpencil from the base of the anal margin. Leguminivora is characterised by a male scent organ consisting of a pair of scale tufts from pockets in the pleuro-posterior region of an invaginated plate covered with numerous scales between A8 and A9. As in Fulcrifera Danilevsky & Kuznetzov, S2 has stout sternal apodemes in the female and usually at least remnants of sternal rods in the male. The male genitalia are characterised by a dorsolaterally widened, subrectangular, flattopped tegumen, the socii a pair of long hairpencils with converging tips and their bases from a concavity of the dorso-lateral expansion of the
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tegumen (shared by Notocydia, gen. nov.), narrow gnathos arms arising just below the top of the tegumen and running laterally along the subscaphium as in Fulcrifera, a short and wide valva, constricted near middle, a sacculus that is ventrally strongly concave near its base and forms an angular ridge, and a long, slender aedeagus, strongly curved in basal half and somewhat bulbous subbasally. The female genitalia are characterised by slender ovipositor lobes with a short posterior lobe (shared by Notocydia, gen. nov., Fulcrifera and Apocydia, gen. nov.), a long ductus bursae widened below inception of ductus seminalis with distal half transversely wrinkled and ending in a strongly sclerotised funnel- or tubeshaped sterigma, and a corpus bursae with a diverticulum and two horn-shaped signa.
Description Adults (Fig. 859) Wingspan. 9.0–15.5 mm (Australian species 10.5–14.5 mm); small to medium-sized. Head (Fig. 862). Ocellus well developed, chaetosema very obvious. Proboscis rather short, about 1.5× length of labial palpus. Labial palpus moderately long (1.7–2.1× diameter of eye), moderately wide, sinuate and lightly ascending, second segment rough-scaled beneath, nearly parallel-sided, third segment with short appressed scales. Antenna to beyond middle, anterior band without scales, cilia in male short. Thorax. Without crest. Legs unmodified. Wings (Figs 101, 859). Forewing index 0.43– 0.45. Forewing truncate-subovate, costa gently curved throughout, apex obtusely pointed, termen gently convex; all veins present and separate beyond cell, chorda from between R1 and R2 to between R4 and R5, M-stem to between M2 and M3, traceable only in distal part, R1 before 56O of cell, R2 nearer to R3 than to R1, R4 to costa, R5 to termen, M2 and M3 subparallel distally, the latter approximated to the former at base, CuA1 from lower angle of cell, CuA2 from beyond 36: of cell, CuP present at margin; brown to dark brown densely irrorated with whitish, with whitish costal strigulae, obvious ocelloid patch, and at most faint dorsal blotch. Hindwing subtrapezoidal or semi-oval, costa evenly curved, anal margin modified in male, thickened and rolled upwardly into a fold with modified fringe, containing a pencil of hair-like scales from base, with 3A closely parallel to anal margin; female frenulum three bristles; Rs and M1 approximated at base, M2 parallel to M3, M3 and CuA1 stalked,
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CuA2 from 36: of cell, CuP present at wing margin, 1A+2A with basal fork, 3A in male close and parallel to margin, 3A in female in normal position; greyish brown. Pregenital abdomen (Fig. 863). Sternum 2 with well-developed anterolateral processes and short and stout sternal apodemes, especially in female, with at least remnants of sternal rods in male. In male distal half of S8 invaginated, together with a pair of scale tufts in pleuro-posterior pockets, the invaginated plate densely covered with numerous, sometimes blackish scales (see Komai 1999: 15, 25). Male genitalia (Fig. 860). Vinculum a wide band with short saccus, V-shaped. Tegumen a narrow band, widening dorsolaterally, subrectangular with broad flat top. Socii a pair of long hair-pencils arising from concavity of dorso-lateral expansion of tegumen, with converging tips. Gnathos indicated by two narrow bands originating from just below top of tegumen and laterally running along subscaphium. Valva short and wide, somewhat narrowed in middle; basal excavation well-defined, costal process large; cucullus transversely ovate, densely bristled along outer margin; sacculus ventrally strongly concave near base, forming an angulate ridge. Aedeagus long, slender, wider in basal third, strongly curved in basal 56:–56O; caulis moderately long, fused with aedeagus near base (Fig. 860); vesica with some thin cornuti. Female genitalia (Fig. 861). Ovipositor lobes very slender, posterior lobes very short. Apophyses long and stout. Sternum 7 weakly sclerotised, shield shaped, ostium in deep excavation of its hindmargin. Sterigma a strongly sclerotised funnel or tube connected to a weakly sclerotised area with scale sockets (lamella postvaginalis). Ductus bursae long, very narrow and transversely wrinkled in posterior 36: to 6M, distally sclerotised to form sterigma; anteriorly much broader to inception of ductus seminalis where it is abruptly narrowed and partially sclerotised; bulla seminalis present; corpus bursae ovate, with a developed or reduced diverticulum (Fig. 861), with two horn-shaped signa.
Distribution Palaearctic, Oriental, Ethiopian and Australian regions. In Australia the genus has been found in Queensland from Brisbane to the Cairns region.
Biology The genus is associated with Fabaceae. Leguminivora glycinivorella is a well known pest of
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soybean, Glycine max, in north-east Asia. It also feeds on Maranta arundinacea (Marantaceae) and Sophora flavescens (Zhang 1994), and has been bred from larva webbing flowers of mango (Anacardiaceae) (Obraztsov 1967). Leguminivora ptychora (Meyrick) has been recorded as a sporadic pest of cowpea, Vigna unguiculata from Africa and Asia (Singh et al. 1990). The larvae bore into host seeds. No biological information is available on the Australian species.
Remarks Obraztsov (1960, 1967) and Danilevsky and Kuznetzov (1968) described and reviewed Leguminivora, providing illustrations of genitalia, wing venation, and pregenital abdomen. Clarke (1958) figured the types of L. ptychora and of two synonyms of L. glycinivorella, L. parastrepta Meyrick and L. anticipans Meyrick (as Laspeyresia). The monophyly of Leguminivora is supported by the configuration of the male scent organs, paired scale tufts from pockets in the pleuro-proximal area of the invaginated plate between A8 and A9. This configuration is not homologous to the ‘coremata’ shared among the Grapholita genusgroup (see Komai 1999: 25). The distally transversely wrinkled ductus bursae may also be an apomorphy for Leguminivora. Leguminivora is related to Fulcrifera and to the endemic Australian Notocydia, gen. nov. and Apocydia, gen. nov. Females of all four genera have slender ovipositor lobes with a short posterior portion, a probable synapomorphy. With Fulcrifera, Leguminivora shares overall similar male genitalia, especially the shape of the aedeagus, with Notocydia, gen. nov. similarly derived socii consisting of a pair of long hair-pencils from a concavity of the dorsolaterally expanded tegumen, with their tips converging.
Constituent species Three named species are known worldwide (Brown 2005). In addition, there are two species in Australia, one of which is described below. longigula, sp. nov.
Leguminivora longigula Komai & Horak, sp. nov. (Figs 859–861, 863)
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Figs 859–863. Leguminivora, adult, genitalia, head and abdomen. 859, L. longigula, paratype , Toowoomba, Q [image reversed]. 860, L. longigula, male genitalia, paratype, Brisbane, Q, slide ANIC 8140. 861. L. longigula, female genitalia, paratype, Brisbane, Q, slide ANIC 8138. 862, head, L. glycinivorella, Japan. 863, L. longigula, abdomen, 8th segment with bases of scale tufts (arrows). Scale = 2 mm.
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Material examined Holotype. : ‘27.33S 151.59E, Prince Henry Heights, 620 m, Toowoomba, Q, 25 Jan.1983, I.F.B. Common’, ‘ANIC genitalia slide No. 8087’ (ANIC). Paratypes. Queensland: 1, Mingela, 21.iv.1955, I. F. B. C.; 1, 12 mi SE of Bowen, 6.v.1955, I. F. B. C.; 1 , Brisbane, 31–03–[19]07, [Turner]; 1, Brisbane, Jan., [Turner]; 1, same data as holotype but 20.iii.1983 (ANIC).
Diagnosis Leguminivora longigula is of typically grapholitine appearance, with dark grey-brown forewings and equally dark hindwings with a coppery tinge, modified in male with scattered black scales in anal area and the anal margin upturned and fringed with dense, long, pale grey scales. Leguminivora species look superficially very alike, and there is a similar, probably slightly smaller unidentified species known from two specimens from northern Queensland, with distinct genitalia in both sexes. The male genitalia of L. longigula are characterised by socii of only a few bristles, a valva with both costa and ventral margin only weakly sinuate and a cucullus with a straight distal margin, and a sharply bent aedeagus with a weakly upcurved distal half. The female genitalia are characterised by the distal portion of the ductus bursae that is a narrow tube, and long, slender, horn-shaped cornuti with a ringshaped base.
Description Adults (single male rubbed and faded) (Fig. 859) Wingspan 11.5 mm (male), 11.5–14 mm (females). Head, thorax and tegulae with dark brownish grey scales with pale yellowish tips (denuded and faded to ochreous in male). Forewing ground colour dark brownish grey (faded in male) with scattered pale-tipped scales, pale tips long and yellowish white in basal half of wing, shorter and darker ochreous distally beyond an imaginary line from middle of costa to tornus; paired, rather stout, whitish strigulae along entire costa, separated by dark to blackish brown oblique streaks, with those from middle of costa and beyond edged by ochreous-tipped scales and distally converging, framing two oblique silvery lines originating from costal strigulae just beyond middle and at 6M costa respectively; ocelloid patch well developed, 3–4 black bars separated by ochreous-tipped grey scales, proximally delineated by an indistinct silver patch and distally by a broad silver band; dorsal blotch
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variably expressed as two outwardly curved paired lines of pale-tipped scales, sometimes indistinct; fringe partly pale-tipped, dark grey in apex, paler ochreous grey towards dorsum. Hindwing greybrown in male, anal area more densely scaled with scattered black scales and anal margin narrowly folded upwardly and with a long dense fringe of pale ochreous grey scales; hindwing dark greybrown with light coppery sheen in females; fringe paler, pale ochreous grey. Abdomen dark greybrown. In male S2 with a shield-shaped sclerotisation with remnants of sternal rods laterally, posteriorly followed by a smaller roundish sclerotised patch, both with scale sockets along midline; T8 long and narrow, with projecting, rounded hindmargin; S8 short, with lateral patches of scale sockets in a shallow membranous pocket posteriorly on each side; membrane behind S8 with a large sclerotised patch (Fig. 863). Male genitalia (Figs 860). Tegumen of rectangular outline. Socii very small patches of about nine long, close, parallel bristles forming a long narrow pencil each, slightly longer than tegumen width at top. Valva gradually widening towards nearly straight distal margin of cucullus, with both costa and ventral margin only very weakly sinuate. Aedeagus very long, sharply bent at about 56:; base elongate clavate, distal 36: slender, parallel-sided, only faintly upcurved; with a few very slender, bristle-shaped cornuti in tip. Female genitalia (Fig. 861). Ovipositor lobes very slender. Ostium in V-shaped emargination of hindmargin of S7. Ductus seminalis distally a slender, parallel-sided tube widening just at end and anteriorly followed by a slightly bulbous expansion and then a long, slender, transversely wrinkled, membranous portion to inception of ductus seminalis; anterior fourth much wider, with a small sclerite at inception of ductus seminalis and a scobinate dimple associated with a small dorsal diverticulum; corpus bursae ovate with two very long, slender, horn-shaped signa with a ring-shaped base.
Distribution and biology Leguminivora longigula has been collected in eastern Queensland from Brisbane to Bowen. No host records are known.
Remarks The single male is rubbed and strongly faded, and as the shape of the ductus bursae is the most
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obvious specific apomorphy, a female has been chosen as the holotype. The male and female paratypes from Brisbane both are from Turner’s collection, with his labels and the date in his handwriting.
Notocydia Komai & Horak, gen. nov. (Figs 99, 102, 864–874) Type species: Eucosma atripunctis Turner, 1946, Trans. R. Soc. S. Aust. 70(2): 207, hereby designated. [Australia.]
Diagnosis Small to medium-sized moths with light to dark grey or brownish grey forewings with typically eucosmine-grapholitine markings: five pairs of white strigulae from apex to middle of costa, dorsal blotch variable from absent to a large white area reaching well-developed ocelloid patch. Hindwing with M3 and CuA1 fused into a single vein. Male either with hindwing with anal fold, 3A approximated to margin and with hindtibia dorsally with erectile long scales, or male with an elongate tubular androconial fold filled with lanceolate scales above along the posterior edge of cell and CuA2 stalked with fused M3 and CuA1. The male genitalia are characterised by socii (if present) arising as long hair-pencils (sometimes only one or two hairs) from shallow concavities on dorso-lateral expansion of tegumen, by a very large semi-oval cucullus, and by a curved aedeagus with a dorsal process in distal half. The female genitalia are characterised by slender ovipositor lobes, a complex sterigma of a smooth plate connected anteriorly to a sclerotised pouch or cup and posteriorly with triangular scobination, and a short ductus bursae.
Description Adults (Figs 864, 870) Wingspan. 10–15 mm; small to medium-sized. Head (Figs 865, 871). Ocellus well developed, chaetosema obvious. Proboscis about as long as labial palpus. Labial palpus short (1.3–1.5× diameter of eye), porrect, sinuate, slender, second segment with short loose scales beneath, third segment short with short appressed scales. Antennae moderately long, reaching to beyond middle of forewing, band along anterior surface unscaled, cilia small.
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Thorax. Without crest. Legs: hindtibia in males of the N. atripunctis-group with erectile long scales dorsally. Wings (Figs 99, 102, 864, 870). Forewing index: 0.38–0.41; forewing truncate-suboval, costa gently curved throughout, apex pointed, termen gently convex; cell 0.57–0.65× length of forewing, chorda from between R1 and R2 (closer to R2) to R5, trace of M-stem to M3 or to between M2 and M3; all veins present and separate beyond cell; R1 from 56O of cell, R2 nearer to R3 than to R1, R4 to costa, R5 to termen; M2 and M3 subparallel distally, the latter approximated to the former at base; CuA1 from lower angle of cell; CuA2 from 36: of cell; CuP present at margin; light grey to dark brownish grey forewings with typical eucosmine-grapholitine markings: five pairs of white strigulae from apex to middle of costa, with dorsal blotch variable from absent to a large white area reaching to welldeveloped ocelloid patch. Hindwing subtrapezoidal or semiovate, costa evenly curved, with cubital pecten; anal margin in males of atripunctis-group thickened and rolled upwardly to form an anal fold with modified fringe (Fig. 102), in males of niveimacula-group anal margin unmodified but hindwing with an elongate androconial fold filled with lanceolate scales and a pencil of black scales above along posterior edge of cell (Fig. 99); frenulum of three bristles in female; Rs and M1 approximated at base, M3 and CuA1 fused into one vein, CuA2 from 36: of cell in male (in atripunctisgroup without androconial fold along posterior edge of cell) or CuA2 stalked with fused M3 and CuA1 in male (in niveimacula-group with androconial fold), in female CuA2 from 36: of cell, CuP present at wing margin, 1A+2A with basal loop, 3A in male close, curved and parallel to margin in atripunctis-group, in males of niveimacula-group 3A straight and unmodified as in all females. Pregenital abdomen (Fig. 873). Sternum 2 with well-developed anterolateral processes, with sternal apodemes often somewhat stouter in female, remnants of sternal rods often present. Sternum 8 in male unmodified, without coremata. Tergum 8 in male a long, distally protruding plate. Male genitalia (Figs 866, 867, 872). Vinculum becoming narrower dorsally. Tegumen with conspicuous dorso-lateral expansions and a flat top. Socii, when present, a pair of long hair-pencils or one or two prominent hairs arising from shallow concavities on dorso-lateral expansions of tegumen, their tips converging. Uncus absent. Gnathos a pair of narrow sclerotised bands near middle of tegumen,
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medially fused, often forming a pointed projection. Valva very large, spatulate with basal third narrow then ventrally abruptly widened at base of cucullus; basal excavation well-defined; costal process large; cucullus large, semi-oval, with a patch of strong spines near ventral angle; sacculus strongly concave ventrally near base, forming an angulate ridge (Fig. 866). Aedeagus short to moderately long, curved in middle, with a dorsal tooth or process at various positions in distal half; caulis moderately long; vesica with one to few very short to nail-shaped (?fixed) cornuti in tip of aedeagus. Female genitalia (Figs 869, 874). Ovipositor lobes slender. Sternum 7 triangular with ostium in deep to very deep excavation of posterior margin. Sterigma a smooth weakly sclerotised plate anteriorly ending in a small to large pouch or cup, often more strongly sclerotised near basal opening into ductus seminalis, and posteriorly connected with a scobinate triangular and often bipartite area (lamella postvaginalis). Ductus bursae short, especially posterior narrow membranous or partially sclerotised portion between sterigma and inception of ductus seminalis at middle to distal third of ductus bursae; anterior portion much wider, membranous; bulla seminalis present; corpus bursae ovate or pyriform, with a diverticulum (Fig. 874), and with two horn-shaped signa.
Distribution Notocydia is endemic to Australia, concentrated in the more drier regions. It is known from Western Australia, the Northern Territory, Queensland, and New South Wales.
Biology Larva of N. atripunctis (Turner) have been collected at Quilpie, Queensland, feeding on seed pods of Senna sp. (Caesalpiniaceae), and an undescribed species has been bred from dried flowers of Senna collected near Alice Springs, Northern Territory.
Remarks The monophyly of Notocydia is supported by: 1, M3 and CuA1 fused into one vein; 2, the very large, spatulate valva, and 3, an aedeagus with a dorsal process between middle and distal 56E. The genus is divided into two species-groups: 1, the atripunctis-group: hindwing with CuA2 from 36: of cell, in male with anal fold developed and 3A
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approximated and subparallel to margin; hindtibia in male with long erectile scales dorsally; socii only one or two prominent hairs from concavities on dorso-lateral expansion of tegumen or absent; 2, the niveimacula-group: hindwing in male without anal fold but with an elongate tubular androconial fold filled with lanceolate scales and a black hair pencil opening above along posterior edge of cell; CuA2 stalked with fused M3 and CuA1 in male; socii a pair of long hair-pencils arising from shallow concavities on dorso-lateral expansion of tegumen, with their tips converging. Leguminivora Obraztsov is considered to be the sister group of Notocydia, with the two genera sharing similar socii consisting of hair-pencils from a concavity of the dorso-laterally expanded tegumen.
Constituent species We are aware of at least 10 Australian species, of which six belong to the atripunctis-group, including the three named species, and four to the niveimacula-group, with niveimacula, sp. nov. described below. 1. The atripunctis-group atripunctis (Turner, 1946), Trans. R. Soc. S. Aust. 70: 207 (Eucosma). [Qld: Bunya Mtns.] Comb. nov. austera (Turner, 1946), Trans. R. Soc. S. Aust. 70: 209 (Eucosma). [Qld: Brisbane, Toowoomba.] lomacula (Lower, 1899), Proc. Linn. Soc. NSW 24: 95 (Laspeyresia). [NSW: Broken Hill.] Comb. nov. tephraea (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 287 (Laspeyresia). [SA: Peterborough as Petersburg.] Comb. nov. 2. The niveimacula-group niveimacula, sp. nov.
Notocydia niveimacula Komai & Horak, sp. nov. (Figs 870, 872–874)
Material examined Holotype. , ‘29.14S 116.00E, 4 km N of Morawa, WA, 9 Oct. 1992, E.D. Edwards’, ‘ANIC genitalia slide No. 18798 ’ (ANIC).
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Paratypes. Western Australia: 1 , 29°21´S 117°20´E, 15 km N by E Mt Singleton, 28.ix.1981, J. C. Cardale; same label data as holotype, 6 , 4; 1 , 27°43´S 114°10´E, Kalbarri, 6.x.1990, I. F. B. C. Northern Territory: 1 ,
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25°46´S 133°17´E, 8 km N Kulgera, 21.ix.1978, E. D. E.; 1 , 24°11´S 134°01´E, 56 km S by E of Alice Springs, 24. ix.1978, E. D. E.; 1 , 23°59´S 133°56´E, 32 km S by E of Alice Springs, 23.ix.1978, E. D. E. (ANIC).
Figs 864–869. Notocydia, gen. nov., atripunctis-group, adult, head and genitalia. 864, N. ? atripunctis, , 45 km NE of Wellbourn Hill, SA. 865, head, N. sp. nr atripunctis, male. 866, 867, N. atripunctis, male genitalia and aedeagus, Toowoomba, Q, slide ANIC 2238 [syntype of Eucosma austera]. 868, 869, N. atripunctis, Toowoomba, Q, slide ANIC 4967 [syntype of Eucosma austera]. 868, sterigma. 869, female genitalia. Scale = 2 mm.
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OLETHREUTINE MOTHS OF AUSTRALIA
Diagnosis Notocydia niveimacula is superficially recognisable by the combination of a typically grapholitine wing pattern, the modified hindwing venation and the broad white band across a dark forewing. In addition, the male has an invaginated fold with
modified scales and a black hairpencil along the hindmargin of the hindwing cell. In the male genitalia, N. niveimacula is characterised by a medially protruding hindmargin of T8, a rounded rather than triangular tegumen apex, narrow dorso-lateral expansions to the tegumen with socii
Figs 870–874. Notocydia, gen. nov., niveimacula-group, adult, head and genitalia. 870, N. niveimacula, paratype , 8 km N of Kulgera, NT. 871, head, N. sp. nr niveimacula, male. 872, 873, N. niveimacula, holotype, slide ANIC 18798. 872, male genitalia. 873, 8th abdomen segment. 874, N. niveimacula, paratype, nr Morawa, WA, female genitalia, slide ANIC 18801. Scale = 2 mm.
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long, slender hair pencils, a strongly projecting triangular keel from sacculus and usually two extremely short cornuti. In the female genitalia, N. niveimacula is characterised by a sclerotised, longitudinally wrinkled distal portion of the ductus bursae. All these characters separate N. niveimacula from a sympatric and closely related, unnamed Notocydia species with a narrower white band across the forewing, but also with a scale pencil along the hindmargin of the hindwing cell in the male.
Description Male (Fig. 870). Wingspan 11–13 mm. Head, thorax and tegulae grey-brown, lower part of head and labial palpi whitish ochreous, palpi light grey-brown on outer surface of second segment. Forewing subtriangular; base to beyond 56: mottled grey-brown with welldefined edge from middle of costa, inwardly angled below costa and running obliquely to 369 dorsum, with two pairs of sometimes very long white strigulae between 56: and middle of costa; with a broad, irregular, oblique band of snow white scales from distal half of costa to between 369 and 169 dorsum, incorporating dorsal blotch and costal strigulae that are delineated by four grey-brown spots along costa; ocelloid patch of 5–6 black dashes separated by whitish ochreous scales, the three dorsal ones parallel and aligned, the more costal ones partly closer to termen and interrupted, all framed by two subparallel silver bands proximally and distally and oblique and sometimes interrupted bands in tornus and apex; along termen with a band of pale-tipped scales, sometimes interrupted by white scales below apex and near dorsal end of ocelloid patch; some scattered silver scales in white area basal to ocelloid patch; fringe grey, darker towards tips. Hindwing with M3 and CuA1 fused into single vein and CuA2 apparently stalked with this fused vein due a fold on upper side along hindmargin of cell lined with modified scales and containing a slender pencil of jet black hairscales from near base; anal margin not modified, 3A straight; pale grey-brown, fringe pale grey-brown at base, whitish distally. Tergum 8 with hindmargin strongly convex, medially projecting (Fig. 873). Male genitalia (Fig. 872). Tegumen apex rounded, with narrow dorso-lateral expansions. Socii two small hair-pencils. Gnathos medially paired, upcurved, membranous arms. Valva with long, narrowly ovate cucullus with small,
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rounded-rectangular, projecting ventral angle, with a large, triangular, inwardly projecting keel from end of sacculus. Aedeagus large, strongly sinuate, widest near base and gradually tapering to tip, with a sharp keel-shaped point dorsally at about 169, two very short, stout and pointed cornuti in tip of aedeagus and a long slender rod-shaped structure within aedeagus from about middle to dorsal keel. Female. Wingspan 11.5–13.0 mm. Wing pattern as in male, but hindwing without fold along hindmargin of cell and CuA2 from 36: cell. Female genitalia (Fig. 874). Ovipositor lobes slender, closed against each other. Sternum 7 small, triangular, with rugose anterolateral corners, ostium in very deep emargination of hindmargin. Sterigma a narrow ring surrounding ostium and continued as a medial band in lamella postvaginalis, posteriorly widening, with bristles or at least their sockets. Ductus bursae moderately sclerotised, roughly parallel-sided but irregular and longitudinally wrinkled in posterior 69, beyond inception of ductus seminalis; anterior 369 membranous, wider but posteriorly narrowing, with a patch of sharp, sclerotised spinules near entrance to ductus seminalis. Corpus bursae pearshaped with a small, flap-shaped dorsal diverticulum and two slender, long, horn-shaped signa with a ring-shaped base.
Distribution and biology Notocydia niveimacula is known from dry areas of southern Western Australia (Kalbarri, Morawa, Mt Singleton) and the southern parts of the Northern Territory around Alice Springs. Nothing is known about its biology.
Remarks Three possibly conspecific specimens from southwestern Queensland have not been included, pending a revision of the entire genus.
Fulcrifera Danilevsky & Kuznetzov (Figs 103, 875–879) Fulcrifera Danilevsky & Kuznetzov, 1968, Fauna SSSR 98, Lepid. 5(1): 454. Type species:
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Laspeyresia luteiceps Kuznetzov, 1962, Entomol. Obozr. 41: 627, by original designation. [Siberia.]
Diagnosis Fulcrifera species are usually small grapholitine moths with indistinct wing pattern, light greyish brown to dark brown, densely irrorated with whitish, with whitish costal strigulae and distinct ocelloid patch, and in non-Australian species sometimes with dorsal blotch. The labial palpus is short and slender, porrect and sinuate to slightly upcurved. The genus shares with other genera of the Cydia-group a hindwing with M3 and CuA1 stalked, in the male with 3A approximated to the thickened anal margin that is rolled upwardly to form an anal fold with modified fringe and encloses a thin hairpencil from the base of the anal margin. S2 has stout sternal apodemes in the female and usually at least remnants of sternal rods in the male. 8th segment without coremata. The male genitalia are characterised by narrow gnathos arms arising just below the top of the tegumen and running laterally along the subscaphium, a sacculus that is ventrally strongly concave near its base and forms an angular ridge, and an aedeagus with an often parallel, strongly sclerotised process dorsally from anellus. The female genitalia are characterised by slender ovipositor lobes, a ring- to gutter-shaped sterigma in most Australian species with a heart-shaped spinulose plate in the lamella postvaginalis, a long ductus bursae widened below inception of ductus seminalis, and a corpus bursae with a diverticulum and two horn-shaped signa.
Description Adults (Fig. 875) Wingspan. 9–15 mm (Australian species 9.0– 14.5 mm; rarely to 18 mm); nearly always small. Head (Fig. 878). Ocellus well developed, chaetosema very obvious. Proboscis rather short, not much longer than labial palpus. Labial palpus short and slender (1.4×–1.7× diameter of eye), porrect to slightly upcurved, sinuate; second segment roughly parallel-sided, with long scales beneath; terminal segment short, with appressed scales. Antenna to beyond middle; anterior band unscaled, cilia short. Thorax. Without crest. Legs unmodified. Wings (Figs 103, 875). Forewing index 0.40– 0.45; forewing truncate-suboval, costa gently curved throughout, apex obtusely pointed, termen gently convex, chorda from between R1 and R2
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(closer to R2) to R5 or to between R4 and R5, Mstem to between M2 and M3, R1 from 56O of cell; all veins present and separate beyond cell, R2 nearer to R3 than to R1, R4 to costa, R5 to termen, M2 and M3 subparallel distally, the latter approximated to the former at base, CuA1 from lower angle of cell, CuA2 beyond 36: of cell, CuP present at margin; light greyish brown to dark brown, densely irrorated with whitish, with whitish costal strigulae and distinct ocelloid patch, and sometimes with distinct dorsal blotch (not in Australian species). Hindwing subtrapezoidal or semi-oval, costa evenly curved, anal margin in male thickened and often rolled upwardly, forming an anal fold with modified fringe, containing a very slender hairpencil from base of hindwing; frenulum of female consisting of three bristles; Rs and M1 approximated at base, M2 parallel to M3, M3 and CuA1 stalked, CuA2 from 36: of discal cell, CuP present at wing margin, 1A+2A with basal fork; 3A in male usually raised and close to margin, 3A in female in normal position (Fig. 103); light to dark grey-brown. Pregenital abdomen (Fig. 879). Sternum 2 with well-developed anterolateral processes, rather short and wide sternal apodemes, especially in female, and in male often with sternal rods. Sternum 8 in male unmodified, without coremata. Tergum 8 in male large expanded plate with a convex posterior margin. Male genitalia (Fig. 876). Vinculum a wide band with short saccus, U- or V-shaped. Tegumen a narrow band, with a rounded or pointed top. Socii a pair of hairy patches fused with diaphragma or absent. Uncus absent. Gnathos indicated by two narrow bands each originating from just below top of tegumen and running along each side of subscaphium. Valva heavily sclerotised, usually narrowed before or beyond middle but in Australian species basal two-thirds roughly parallel-sided and cucullus wider; basal excavation well-defined; costal process large; cucullus ovate or rounded, densely bristled along outer margin; sacculus ventrally clearly concave near base, forming an angulate ridge. Aedeagus of two types: one curved in basal 56:, the other bulbous at base and almost straight distally; caulis rather high, connected through anellus with base or basal 56E of aedeagus; anellus dorsally producing a short or long, often spiny, heavily sclerotised process often parallel to aedeagus (Fig. 876); vesica with some thin cornuti or without. Female genitalia (Fig. 877). Ovipositor lobes slender, posterior lobes very short. Sternum 7 often subtriangular, ostium in deep excavation of its
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AUSTRALIAN OLETHREUTINE GENERA
hindmargin. Sterigma variable, ring- to funnel- to gutter-shaped, posteriorly often expanding into a spinulose plate in lamella postvaginalis that is large, heart-shaped and lightly concave in most Australian species. Ductus bursae long, very narrow in posterior 36: to 6M and broad anteriorly, rarely with indication of bipartite colliculum below
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sterigma and in some species with irregular small sclerite close to inception of ductus seminalis; ductus seminalis arising from juncture of narrow and broad parts of ductus bursa, bulla seminalis present. Corpus bursae ovate, with a large or small diverticulum (Fig. 877), with two horn-shaped signa.
Figs 875–879. Fulcrifera persinuata, adult, head, abdomen and genitalia. 875, adult, , holotype. 876, male genitalia, holotype, slide ANIC 8067. 877, female genitalia, paratype, nr Borroloola, NT, slide ANIC 18779. 878, head, male. 879, male abdomen, 8th segment. Scale = 2 mm.
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OLETHREUTINE MOTHS OF AUSTRALIA
Distribution Palaearctic region; Oriental region; Ethiopian region; Australian region where Fulcrifera occurs north from Carnarvon in Western Australia, in the Northern Territory and south to Yeppoon in Queensland.
Biology Fletcher (1932) reported that the larvae of F. tricentra (Meyrick) feed on stems and branches of Tephrosia purpurea at Pusa, India. As hosts of F. tricentra, Zhang (1994) listed Crotalaria juncea (sunn hemp), Vigna mungo, Dolichos biflorus, and Sesbania bispinosa, and the species is known as a pest of cowpea Vigna unguiculata in southern India (Singh et al. 1990). According to Danilevsky and Kuznetzov (1968) the hosts of F. orientis Kuznetzov and F. luteiceps Kuznetzov are Sophora sp. and Caragana arborescens respectively. All the hostplants recorded belong to the Fabaceae and the larvae are borers. No biological information is available for the Australian species.
Remarks Danilevsky and Kuznetsov (1968) figured the genitalia of several species of Fulcrifera with their description of the genus. Clarke (1958) figured the types of F. tricentra and its alleged synonyms F. crocopa (Meyrick) and F. pseudonectis (Meyrick). The monophyly of Fulcrifera is supported by the short to very long, heavily sclerotised process that seemingly looks like a dorsal process of the aedeagus but originates from the dorsal portion of the anellus next to the aedeagus. The genus is divided into two species-groups: 1, the probably more plesiomorphic infirmana-group with a simple sterigma, a valva with elongate-ovate or rounded cucullus, a curved aedeagus and usually prominent socii; 2, the derived tricentra-group with dorsally expanded sterigma, straight aedeagus with a bulbous base, and a valva with small, transversely ovate cucullus, which includes the majority of the Australian species.
Constituent species Over 15 species are known worldwide, with at least seven unidentified species known from Australia, one of which is described below. persinuata, sp. nov.
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Fulcrifera persinuata Komai & Horak, sp. nov. (Figs 875–879)
Material examined Holotype. , ‘16.27S 136.05E, McArthur R., 48 km SW by S Borroloola, NT, 29 Oct. 1975, E.D. Edwards’, ‘ANIC genitalia slide No. 8067 ’ (ANIC). Paratypes. Northern Territory: 2 , 1, 16°08´S 136°06´E, 22 km WSW Borroloola, 2.xi.1975, E. D. E.; 1 , 16°19´S 136°05´E, 36 km SW Borroloola, 4.xi.1975, E. D. E.; 1 , 16°25´S 136°05´E, Surprise Ck, 45 km SW by S Borroloola, 5.xi.1975, E. D. E.; 1, 16°28´S 136°09´E, 46 km SSW Borroloola, 28.x.1975, E. D. E.; 1 , 16°34´S 135°41´E, Leila Ck, 14 km NW Cape Crawford, 6.xi.1975, E. D. E.; 1 , 16°47´S 135°45´E, McArthur R., 14 km S by W Cape Crawford, 25.x.1975, EDE (ANIC).
Diagnosis Fulcrifera persinuata is of inconspicuous, typically grapholitine appearance, grey-brown with an ochreous tinge due to the pale yellowish scale tips, with overall darker hindwings. The male is characterised by black glossy scales scattered across the abdomen and the anal region of the hindwing that also has a dense fringe of pale modified scales along its anal margin, and by modified hind tibiae with long scaling. A very similar-looking unidentified Fulcrifera species, also belonging to the tricentra-group, occurs in the same localities as F. persinuata, but its pale scale tips are whitish rather than yellowish as those of F. persinuata, its hindwing is hardly darker than the forewing and it lacks secondary sexual modifications in the male. The male genitalia of F. persinuata are characterised by a valva that is nearly parallel-sided in basal 36: with an only weakly convex ventral margin, a cucullus with a straight distal margin and an aedeagus with a doubly sinuate dorsal process as long as the aedeagus and with only a few bristleshaped cornuti. The female genitalia are characterised by a broadly triangular, deeply excavated S7, a large, heart-shaped sterigma with a long and narrow anteromedial process and broad, short, blunt, horn-shaped cornuti.
Description Male (Fig. 875) Wingspan 9.0–11.5 mm. Head (Fig. 878), thorax and tegulae with conspicuously pale-tipped greybrown scales, as forewing; lower frons and inner
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AUSTRALIAN OLETHREUTINE GENERA
side of labial palpi pale ochreous. Forewing with grey-brown scales with conspicuous, pale yellowish ochreous tips; with pairs of whitish strigulae nearly from base separated by leaden brown dashes with a purplish sheen, with one just beyond middle and one at 6M costa giving rise to two silver lines, initially strongly oblique then bent, widened and framing the ocelloid patch that consists of 3–5 parallel short black dashes; without any indication of a dorsal blotch. Hindwing dark grey-brown, anal area with scattered glossy black scales, from very few to many; 3A strongly developed, close and subparallel to anal margin that has a dense fringe of long, modified, distally widened pale grey to whitish scales mixed with few black scales; scale pencil from base of wing on upperside along base of anal fringe. Hind tibia modified with very long, loose scaling and scattered with black glossy scales. Abdomen dark grey-brown, variably scattered with glossy black scales. S2 with modified, sclerotised medial portion and a conspicuous medial band of scale sockets, sternal rods indicated; T8 nearly as long as wide, S8 much shorter and very wide (Fig. 879). Male genitalia (Fig. 876). Tegumen without uncus or socii, with weak lateral gnathos arms fused with a strongly projecting, lightly sclerotised anal tube. Valva in basal 36: nearly parallel-sided to basally only moderately narrowed, with ventral margin straight or only weakly convex; cucullus significantly wider, with straight distal edge. Caulis moderately long. Aedeagus very long, abruptly narrowed beyond strongly curved bulbous base, distal 169 straight, very narrow; with parallel, strongly sclerotised, band-like, apically spiny process originating from dorsal portion of anellus as long as aedeagus and doubly sinuate (unless twisted during preparation); vesica with a few slender, bristle-like cornuti in distal half. Female Wingspan 11.0–11.5 mm. As for male, but without the secondary sexual modifications of hindwing, hind tibia and abdomen. Sternal rods long, slender, well developed. Female genitalia (Fig. 877). Ovipositor lobes slender. Sternum 7 broadly triangular, with ostium at base of V-shaped emargination in hindmargin flanked by two longitudinal, parallel, strongly sclerotised bands, one on each side. Sterigma a large, heart-shaped, slightly concave, strongly sclerotised, spinulose plate with a long, slender anteromedian extension leading to ostium. Ductus
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bursae entirely membranous, very narrow in posterior 69, much wider anterior to inception of ductus seminalis, with a small diverticulum; corpus bursae ovate with two short, wide, blunt, hornshaped signa with hollow base.
Distribution and biology Pending a comprehensive revision of all Australian Fulcrifera species, F. persinuata is at present restricted to the type series from the vicinity of Borroloola in the Northern Territory. However, there is closely related or conspecific material in the ANIC ranging from Broome in Western Australia across the Northern Territory to near Townsville in Queensland. Nothing is known about the biology of F. persinuata, but the closely related F. pseudonectis (Meyrick) has been reared from larvae tunnelling in stems of Crotalaria juncea (Fabaceae) in India (Clarke 1958).
Remarks Fulcrifera persinuata belongs to the tricentra species-group, with similar secondary sexual modifications of hindwing, hind tibia and abdomen as the Indian F. pseudonectis (Meyrick). However, the valva of F. pseudonectis is much narrower at its base and has a more convex ventral margin, and the process above the aedeagus of F. persinuata is more strongly sinuate.
Apocydia Komai & Horak, gen. nov. (Figs 105, 880–885) Type species: Eucosma pervicax Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 250, hereby designated. [Australia.]
Diagnosis Apocydia are small to medium sized moths, with light greyish to dark red-brown forewings often paler in dorsal half, with long costal strigulae and several oblique silver lines from costa, and an elongate, in males partly whitish ocelloid patch. Their labial palpi are sinuate and porrect, with the second segment distally widened. The male antenna is conspicuously thickened by very dense scaling but otherwise Apocydia lacks any male secondary sexual characteristics. Chorda and M-stem are at best weak traces in the forewing, and hindwing M3 and CuA1
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are stalked. The male genitalia of Apocydia are characterised by a distally widened, hood-shaped tegumen with a pair of roundish, weakly sclerotised windows laterally below the apex, by long, slender valvae narrowest in middle, and the rather long aedeagus gradually narrowing towards its tip that contains 3–5 minute, bristle-like cornuti. The female genitalia have slender ovipositor lobes, a triangular S7 with deeply emarginate hindmargin, a strongly sclerotised, large, shallow, cup- to pocketshaped sterigma associated with a bristled bipartite patch of granulation, a moderately long ductus bursae twisted in middle, with a small sclerite just behind origin of ductus seminalis, and a corpus bursae with a diverticulum and two horn-shaped signa posteriorly.
Description Adults (Fig. 880) Wingspan. 11.5–16.0 mm; small to mediumsized. Head (Fig. 885). Ocellus well developed, chaetosema very obvious. Proboscis short. Labial palpus short to moderately long (1.4–1.8× diameter of eye), porrect, sinuate, second segment widening distally with rather long scales beneath, third segment short with short appressed scales. Antennae reaching beyond middle of forewing, in male thickened with dense scales and entirely scaled. Thorax. Without crest. Legs unmodified. Wings (Figs 105, 880). Forewing index: 0.37–0.38, forewing truncate-subovate, costa gently curved throughout, apex pointed, termen gently convex; cell 0.62–0.67× length of forewing; chorda absent or weak trace from between R1 and R2 (closer to R2) to R5, M-stem absent or a distal short remnant to between M2 and M3; all veins present and separate beyond cell; R1 from 56O of cell, R2 nearer to R3 than to R1, R4 to costa, R5 to termen, M2 and M3 subparallel distally but closer at base, CuA1 from lower angle of cell, CuA2 from 36: of cell, CuP present at margin; light greyish ochreous to grey-brown in males, orange ochreous to dark red-brown in females, often paler in dorsal half, with long costal strigulae and several oblique silver lines from costa, and an elongate ocelloid patch partly whitish in males. Hindwing subtrapezoidal or semiovate, costa evenly curved, anal margin without fold but thickened in male, normal in female; frenulum of three bristles in female; Rs and M1 parallel and approximated at base, M2 distant from and parallel to M3, M3 and CuA1 short-
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stalked, CuA2 from 36: of cell, CuP present at wing margin, 1A+2A with basal loop, 3A present in normal position; grey to dark greyish brown. Pregenital abdomen (Fig. 881). Sternum 2 with well-developed, anterolateral processes; remnants of sternal rods usually present in males. Sternum 8 in male unmodified, coremata not present. Tergum 8 in male large expanded plate with a convex posterior margin. Male genitalia (Figs 883, 884). Vinculum Vshaped, wide ventrally. Tegumen a narrow band laterally, strongly widened, hood-shaped distally with projecting hindmargin and usually with a pair of weakly sclerotised, roundish windows laterally below apex. No trace of socii and uncus. Gnathos two weak lateral arms merging with weakly sclerotised bands laterally along anal tube. Valva long, slender, medially narrower; basal excavation well-defined; costal process moderate; cucullus elongate-ovate, densely bristled on inner surface; sacculus ventrally weakly concave near base, with a ventral keel and a patch of dense, long spines distally. Caulis high and wide, anellus fused with base of aedeagus. Aedeagus rather long, curved, gradually narrowed towards apex; vesica with 3–5 minute, bristle-like cornuti in aedeagus tip (visible only at high magnification). Female genitalia (Fig. 882). Ovipositor lobes very slender. Sternum 7 small, triangular with hindmargin deeply excavated. Ostium in excavation of S7. Sterigma a large, shallow cup or pocket with strongly sclerotised ventral wall followed by paired patches of scobination with some bristles in lamella postvaginalis. Ductus bursae not longer than corpus bursae, narrowest and twisted in middle, with anterior 36: membranous; ductus seminalis with bulla seminalis arising from posterior third, immediately followed first by a variably sclerotised, sometimes bipartite portion (colliculum?) and then widening to sterigma; corpus bursae ovate, with a diverticulum, with two horn-shaped signa posteriorly.
Distribution Endemic to Australia, known from all mainland states except Victoria and South Australia, but with one specimen from Tasmania (Freycinet NP).
Biology Nothing is known about the biology of this monotypic genus, but it occurs through an
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extraordinary range of climates from Cape York to Tasmania’s eastern coast.
Remarks Apocydia is rather isolated but apparently belongs to the Cydia genus-group, with a concave ventral
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sacculus margin near its base as in the other genera of this genus-group and a modified hindwing anal margin, albeit only thickened in Apocydia and not folded as in Cydia Hübner. The thickened male antenna, the shape of the valva, the cup or pocket-shaped sterigma and the twisted ductus bursae are apomorphies for Apocydia.
Figs 880–884. Apocydia pervicax, adult, head, abdomen and genitalia. 880, adult, , nr Hope Vale Mission, Q. 881, male abdomen, 8th segment. 882, female genitalia, Florence Falls, NT, slide ANIC 4951. 883, 884, male genitalia and aedeagus, nr Cahill, NT, slide ANIC 8072. Scale = 2 mm.
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OLETHREUTINE MOTHS OF AUSTRALIA
Erminea Kirby & Spence, 1826, Introd. Entomol. 3: 123. Type species: Phalaena pomonella Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 538, by monotypy. [Europe.] A junior homonym of Erminea Haworth, 1811, Lepid. Br.: 512 (Lepidoptera, Yponomeutidae). Carpocapsa Treitschke, 1829, in Ochsenheimer, Schmett. Eur. 7: 231. Type species: Phalaena pomonella Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 538, by subsequent designation by Curtis 1831, Br. Entomol. 8: folio 352. [Europe.] Coccyx Treitschke, 1829, in Ochsenheimer, Schmett. Eur. 7: 230. Type species: Tortrix strobilana Hübner [1799], in Ochsenheimer, Schmett. Eur. 7: pl. 12, fig. 70, by subsequent designation by Desmarest, 1857, in Chenu, Encycl. Hist. Nat. (Papillons nocturnes): 224. [Europe.] Fig. 885.
Apocydia pervicax, head, male.
Constituent species The Australian A. pervicax is the only species known for this genus. pervicax (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 250 (Eucosma). [NT: Darwin.] Comb. nov. ferraria (Turner, 1916), Trans. R. Soc. S. Aust. 40: 535 (Laspeyresia). [Qld: Brisbane.] dolichosticha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 203 (Eucosma). [Qld: Cape York.] aspersa (Turner, 1946), Trans. R. Soc. S. Aust. 70: 207 (Eucosma). [Qld: Cape York.]
Cydia Hübner (Figs 104, 886–893) Cydia Hübner, [1825], Verz. Bekannter Schmett.: 375. Type species: Phalaena pomonella Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 538, by subsequent designation by Walsingham, 1897, Proc. Zool. Soc. London 1897: 130. [Europe.] Laspeyresia Hübner, [1825], Verz. Bekannter Schmett.: 381. Type species: Tortrix corollana Hübner, [1823], Samml. Eur. Schmett. 7: pl. 45, fig. 282, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 10, 57. [Europe.] A junior homonym of Laspeyresia R. L., 1817, Jenaische Allg. Lit.-Ztg 1: 288 (Lepidoptera, Noctuidae).
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Semasia Stephens, 1829, Nom. Br. Insects: 47. Type species: Phalaena pomonella Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 538, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 23, 58. [Europe.] Strobila Sodoffsky, 1837, Bull. Soc. Imp. Nat. Moscou 1837(6): 92. An unnecessary replacement name for Coccyx Treitschke, 1829. A junior homonym of Strobila Sars, 1829, Bidr. Söedyrenes naturh.: 17 (Coelenterata). Carpocampa Harris, 1841, Rep. Insects Mass. injurious to Vegn: 351. An unjusified emendation of Carpocapsa Treitschke, 1829. Cerata Stephens, 1852, List Specimens Br. Anim. Colln Br. Mus. 10: 77. Type species: Penthina servillana Duponchel, 1836, in Godart & Duponchel, Hist. Nat. Lépid. Papillons Fr. 9: 503, by monotypy. [France.] Melissopus Riley, 1882, Trans. Acad. Sci. St. Louis 4: 322. Type species: Carpocapsa latiferreana Walsingham, 1879, Illust. Typical Specimens Lepid. Heterocera Colln Br. Mus. 4: xi, 70, pl. 76, fig. 8, by monotypy. [USA.] Adenoneura Walsingham, 1907, in Sharp, Fauna Hawaii 1(5): 677. Type species: Adenoneura falsifalcellum Walsingham, 1907, ibid. 1(5): 677, pl. 10, fig. 17, by original designation. [Hawaii.] Crobylophora Kennel, 1908, Zool. Stuttg. 21(54): 50. Type species: Tortrix inquinatana Hübner, [1799], Samml. Eur. Schmett. 7: pl. 8, fig. 43, by original designation. [Europe.] A junior
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homonym of Crobylophora Meyrick, 1880, Proc. Linn. Soc. NSW 5: 177 (Lepidoptera, Lyonetiidae). Hedulia Heinrich, 1926, US Nat. Mus. Bull. 132: 6, 65. Type species: Hedulia injectiva Heinrich, 1926, ibid.: 132: 65, figs. 162, 334, by original designation. [USA.] Kenneliola Paclt, 1951, Revue fr. Lepid. 13: 127. Type species: Tortrix inquinatana Hübner, [1799], Samml. Eur. Schmett. 7: pl. 8, fig. 43, by original designation. [Europe.] An objective replacement name for Crobylophora Kennel. Pseudotomoides Obraztsov, 1959, Tijdschr. Entomol. 102: 200. Type species: Phalaena strobilella Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 539, by original designation. [Europe.] Collicularia Obraztsov, 1960, Tijdschr. Entomol. 103: 134. Type species: Catoptria microgrammana Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 188, by original designation. [France.] Phanetoprepa Obraztsov, 1968, Jl NY ent. Soc. 76: 236. Type species: Phanetoprepa agenjoi Obraztsov, 1968, ibid. 76: 237, by original designation. [Spain.] Danilevskia Kuznetzov, 1970, Entomol. Obozor. 49: 446. Type species: Danilevskia silvana Kuznetzov, 1970, ibid. 49: 446, by original designation. [Russia: Primorsky Krai.] Dicraniana Diakonoff, 1984b, Entomol. Gall. 1: 162. Type species: Semasia seriana Kennel, 1901, Dtsche Entomol. Z. 13: 270, by original designation. [Originally proposed as a subgenus of Cydia Hübner, [1825].] [Spain.]
Diagnosis Native Australian Cydia species are small to medium-sized moths, greyish brown, densely suffused with whitish, with whitish costal strigulae and a distinct ocelloid patch but no dorsal blotch, whilst C. pomonella has a characteristic wing pattern with a large coppery brown distal blotch (Fig. 886). Cydia has rather short and slender, porrect to slightly upcurved, slightly sinuate labial palpi. The genus is characterised by M3 and CuA1 stalked in the hindwing that is modified in the male, usually with a series of modified, glandular scales along either side of 3A and a thickened anal margin, often rolled upwardly, forming an anal fold with modified scales and holding a thin
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hairpencil from base of hindwing. The male abdomen lacks coremata, has S2 with variably developed sternal rods and T8 a large plate with a convex hind margin. The male genitalia are characterised by a sacculus that is subbasally usually concave on ventral margin and often has an angulate ridge at base, and a curved or sinuate aedeagus, often dorsoventrally flattened and/or with a lateral or dorsal process, or with a patch of thorns. The female genitalia are very diverse but Australian species have the corpus bursae with a diverticulum, a ring of granules around its posterior end and the ductus seminalis originating from the bursa ‘neck’.
Description Adults (Figs 886, 887) Wingspan. 8.5–22.0 mm (Australian species 10–20 mm); small to large. Head (Fig. 888). Ocellus well developed, chaetosema very obvious. Proboscis rather short, not much longer than labial palpus. Labial palpus smallish (1.4–1.9× diameter of eye, up to 2.1× in non-Australian species), porrect to slightly upcurved, sinuate and slender, second segment rather slender and nearly parallel-sided, with loose scales along ventral margin; terminal segment with short, appressed scales. Antenna to beyond middle of forewing in male, with unscaled band along anterior surface, cilia minute to small. Thorax. Without crest. Legs: Male hindtibia usually unmodified, rarely with a pencil of hair-like scales (not in Australian species). Wings (Figs 104, 886, 887). Forewing index 0.39–0.46 (Australian species); forewing truncatesuboval to subtriangular, costa gently curved throughout, apex sometimes obtusely pointed, termen gently convex, chorda from between R1 and R2 (closer to R2) to R5 or to between R5 and M1, M-stem to M2 or to between M2 and M3; all veins present and separate beyond cell, R1 from 56O of cell, R2 nearer to R3 than to R1, R4 to costa, R5 to termen, M2 and M3 subparallel distally, the latter approximated to the former at base, CuA1 from lower angle of cell, CuA2 beyond 56O of cell, CuP present at margin; colour and pattern quite variable within the genus, Australian species greyish brown, densely suffused with whitish, with whitish costal strigulae and distinct ocelloid patch, but without dorsal blotch; the introduced C. pomonella grey, with darker transverse lines and a large subtriangular brown mark with coppery sheen based on tornus. Hindwing subtrapezoidal, semi-oval, or
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subtriangular, sometimes sexually dimorphic in shape, costa evenly curved, anal margin in male thickened and often rolled upwardly, forming an anal fold with modified fringe and modified, glandular scales distributed along either side of 3A (Fig. 890); frenulum of female consisting of three
bristles; Rs and M1 approximated at base, M2 parallel to M3, M3 and CuA1 stalked, CuA2 from 36: of cell, CuP present at wing margin, 1A+2A with basal fork, 3A in male close to margin or in normal position, greyish brown. Pregenital abdomen. Sternum 2 with welldeveloped anterolateral processes, in males usually thin sternal rods from base of sternal apodemes. Sternum 8 in male unmodified, coremata not present. T8 in male large expanded plate with a convex posterior margin. Male genitalia (Figs 889, 893). Vinculum a wide band. Tegumen a moderately broad, entirely sclerotised band, with a rounded top. Socii usually absent, socii remnant of a few hairs sometimes present. Uncus absent. Gnathos two narrow sclerotised bars. Valva differentiated, generally spatulate in shape, narrowed in basal 56:– 56O; basal excavation well-defined; costal process large; cucullus usually ovate or semiovate, densely bristled; sacculus concave at base, sometimes with an angulate ridge. Aedeagus variable in shape and length, curved or sinuate, often flattened dorsoventrally, frequently with a lateral or dorsal process, or with a patch of thorns on lateral, dorsal, or ventral wall; caulis short to moderate, usually fused with base of aedeagus; vesica with some spiniform cornuti. Female genitalia (Figs 891, 892). Ovipositor lobes moderate to broad. Sternum 7 deeply incised posteriorly. Ostium bursae in posterior incision of S7. Sterigma a rectangular or circular plate in lamella postvaginalis, strongly elongate in Australian species, sometimes surrounded by a raised sclerite. Ductus bursae variable in length and width, often short and wide and transition into corpus bursae not clearly defined, variably sclerotised, without colliculum; inception of ductus seminalis variable in position but in Australian species from ‘neck’ of corpus bursae; bulla seminalis present; corpus bursae usually ovate, sometimes with a ring of granules posteriorly (Fig. 892), with an often well-defined diverticulum, with two horn-shaped signa.
Distribution
Figs 886–888. Cydia, adults and head, males. 886, C. pomonella. 887, C. sp. A, Mt Webb Nat. Park, Q. 888, head, C. sp. Scale = 2 mm.
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Afrotropical region; Holarctic region with rich fauna; Oriental region; Pacific including Hawaiian islands; Australian region with low species diversity. Some species of Cydia or Laspeyresia have been recorded from the Neotropical region, but confirmation is needed. In Australia the genus has been recorded from northern Queensland and the Australian Capital Territory, with the introduced
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C. pomonella present in all States except the Northern Territory and Western Australia.
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Australia. The only host record for a native Australian Cydia is for an unnamed species reared by I. F. B. Common from seed capsules of Dodonaea viscosa (Sapindaceae) in Canberra.
Biology Many plant families including Fagaceae, Fabaceae, and Rosaceae have been recorded as larval host plants for Cydia outside Australia, with C. pomonella a serious pest of pome fruit also in
Remarks The genus Laspeyresia Hübner, 1825, which was often used instead of Cydia until the late 1970s is
Figs 889–893. Cydia, genitalia and hindwing detail. 889, C. pomonella, male, slide ANIC 19281. 890, C. sp. A, male, hindwing anal area with modified 3A (arrow). 891, C. pomonella, female, slide ANIC 19282. 892, C. sp. A, female genitalia, Mt Webb Nat. Park, Q, slide ANIC 8133. 893, C. sp. A, male genitalia, Mt Webb Nat. Park, Q, slide ANIC 8070.
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held to be a junior homonym of Laspeyresia R. L., 1817, an emendation of Laspeyria Germar, 1810 (Noctuidae). Comprehensive reviews of Cydia have been given by Heinrich (1926), Obraztsov (1959), Danilevsky and Kuznetzov (1968), and Razowski (1989). Komai (1999) discussed the phylogenetic position within the tribe. This large genus is so diverse that no reliable autapomorphies for the genus have been found. A series of modified, glandular scales along either side of 3A in the male hindwing (Fig. 890) is the only feature present almost throughout the genus. Danilevsky and Kuznetzov (1968) divided the genus into the three subgenera, Endopisa, Laspeyresia s. str. and Kenneliola. This attempt appears unsuccessful since they are not all defined by synapomorphies but by a combination of some minor characters only. Cydia has relationships with Leguminivora Obraztsov, Notocydia, gen. nov., Fulcrifera Danilevsky & Kuznetzov, and Apocydia, gen. nov., with the five genera sharing the following characters: 1, ventral margin of sacculus concave near base; 2, presence of anal fold on male hindwing; 3, vein 3A close to anal edge on male hindwing (Komai 1999). The two unnamed Australian species are related to C. malesana (Meyrick) (Clarke 1958: 443), C. dadionopa (Diakonoff) (Diakonoff 1976), and C. pycnochra (Meyrick) (Clarke 1958: 452).
Constituent species There are at least 150 named species of Cydia worldwide (Brown 2005). Three species occur in Australia, the accidentally introduced pest species, C. pomonella, and two unnamed native species. pomonella (Linnaeus, 1758), Syst. Nat. (Edn 10) 1: 538 (Phalaena (Tortrix)). [Europe.] pomana (Fabricius, 1775), Syst. Entomol.: 653 (Pyralis), unjustified emendation of pomonella. pomonana ([Denis & Schiffermüller], 1775), Ankündung syst. Werkes Schmett. Wienergegend: 126 (Tortrix). [Austria.] aeneana (Villers, 1789), C. Linnaei Entomol. Faun. Suec. Descr. 2: 417 (Phalaena Tortrix). [Europe.] simpsonii Busck, 1903, Proc. entomol. Soc. Wash. 5: 235 (Cydia pomonella ssp.). [USA.] glaphyrana (Rebel, 1941), Mitt. Naturwiss. Inst. Sofia 14:1 (Carpocapsa splendana ab.), unavailable. [Macedonia.]
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Grapholita-group Status remarks Komai (1999) established the Grapholita genusgroup for 27 genera worldwide. This group contains about 400 species in 33 genera including Commoneria, gen. nov., Ixonympha, gen. nov., and Archiphlebia, gen. nov. added here.
Evidence for monophyly Komai (1999) found two synapomorphies linking the genera of the Grapholita-group: 1, tegumen strongly curved at base (pedunculus), and 2, presence of coremata consisting of three elements situated between A8 and A9 (Fig. 948).
Distribution The Grapholita genus-group is strongly concentrated in the Old World. Of 32 genera, only three genera, Grapholita Treitschke, Pammene Hübner, and Cryptophlebia Walsingham occur both in the Old and the New Worlds. One genus, Gymnandrosoma Dyar, has been recorded from the New World and Australia. Two genera are restricted to the New World: Pseudogalleria Ragonot (Nearctic) and Ecdytolopha Zeller (Nearctic and Neotropic). Pseudopammene Komai has been recorded in Japan. Four genera, Andrioplecta Obraztsov, Matsumuraeses Issiki, Selania Stephens, and Strophedra Herrich-Schäffer have a PalaearcticOriental distribution. Five genera, Agriophanes Meyrick, Dierlia Diakonoff, Karacaoglania Koçak, Spanistoneura Diakonoff, and Licigena Diakonoff are found only in the Oriental region. Parapammene has a Palaearctic-OrientalAustralian distribution and four genera, Acanthoclita Diakonoff, Centroxena Diakonoff, Microsarotis Diakonoff, and Pammenopsis Kuznetzov have Oriental-Australian distributions. Microsarotis has also been recorded from La Réunion and Madagascar. Three genera, Commoneria, gen. nov., Ixonympha, gen. nov., and Archiphlebia, gen. nov. are endemic to Australia. Thaumatotibia is present in the Ethiopian (leucotreta-group) and in the Palaearctic-OrientalAustralian regions (chaomorpha-group). Coniostola Diakonoff is known from south-east Asia and Madagascar. Six genera, Cryptoschesis Diakonoff, Dracontogena Diakonoff, Eucosmocydia Diakonoff, Hyposarotis Diakonoff, Pammenitis
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Diakonoff, and Thylacandra Diakonoff have been described from the Ethiopian region, especially from Madagascar.
Biology Komai (1999) reported that worldwide 29 dicotyledonous, three monocotyledonous, and three gymnospermous plant families are used as hosts by the Grapholita-group. Of these, two plant groups are the predominant hosts: Fabales (Mimosaceae, Caesalpiniaceae, and Fabaceae) and Fagaceae (Komai 1999). Boring into their hostplants is the prevailing feeding mode of the group, but leaf-rolling and leaf-skeletonising are sometimes observed. According to Komai (1999) and Witzgall et al. (2004), sex pheromones or sex attractants have been identified for 50 species of nine genera, namely Thaumatotibia, Cryptophlebia, Grapholita, Pseudogalleria, Ecdytolopha, Matsumuraeses, Acanthoclita, Pammene and Strophedra. E8–12Ac or Z8–12Ac are predominantly used.
Remarks The Grapholita-group has a sister group relationship with the Cydia-group, with the synapomorphies discussed in the Remarks for the Cydia-group. Centroxena is known from the Northern Territory and from near Mackay in Queensland, but there is not sufficient material available to describe any of the species.
Thaumatotibia Zacher (Figs 106, 107, 894–904) Thaumatotibia Zacher, 1915, Tropenpflanzer 18: 529. Type species: Thaumatotibia roerigii Zacher, 1915, ibid. 18: 529 [ = Argyroploce leucotreta Meyrick, 1913, Ann. Transv. Mus. 3: 276], by monotypy. [Africa.] Metriophlebia Diakonoff, 1969, Tijdschr. Entomol. 112: 89. Type species: Eucosma chaomorpha Meyrick, 1929, Trans. Entomol. Soc. Lond. 76: 495, by original designation. [Marquesas.]
Diagnosis Small to rarely large, greyish brown to blackish brown moths with broad forewings with a blackish
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triangular pretornal mark, without accessory cell in Australian species and with a hindwing with a short cell, especially in male. Thaumatotibia has long, sinuate labial palpi and a thoracic crest of often long, loose scales. T8 in the male is characteristic, a broadly sclerotised plate with a medially convex posterior margin laterally produced into curved points, with paired patches of long mane-like scales but without paired lateral scale tufts in shallow pockets as in Cryptophlebia. Coremata with or without deeply invaginated scale tufts, but intersegmental ventral sclerite never with levers. The male genitalia are characterised by a large, ovate valva with very long, curled scales on the outer surface, and with the sacculus often ending in distal ridge or teeth. In all Australian species the caulis base has the conspicuous, spinulose, paired lobes diagnostic for the chaomorpha-group. The female genitalia are characterised by a sterigma formed by an ovate or rectangular raised rim, posteriorly connected with a pair of ovate depressions with modified scales, by a ring of spinules at the juncture of the ductus and corpus bursae, and by large, horn-shaped signa.
Description Adults (Figs 894–897) Wingspan. 9–22 mm (Australian species 9–20 mm); small to rarely large. Head (Fig. 902). Ocellus well developed, chaetosema very obvious. Proboscis rather short, not much longer than labial palpus. Labial palpus long (2.0–2.4× diameter of eye), porrect and strongly sinuate; second segment widened distally, with scales appressed and rather short; terminal segment about one-third length of second, slender, with appressed scales, apex blunt. Antenna filiform, 36: length of forewing, entirely scaled; cilia small. Thorax. Posterior crest present, of loose, easily abraded scales. Hind tibia in male unmodified in the Australian species (chaomorpha-group), or with tufts of modified scales on inner side and the inner apical spur enlarged with a batch of scales in the Afrotropical leucotreta-group (Komai 1999). Wings (Figs 106, 107, 894–897). Forewing index: 0.42–0.45; forewing oblong subtriangular, costa little curved throughout, apex acutely rounded; termen almost straight, oblique; accessory cell small or absent, chorda if present from between R2 and R3 (closer to R3) or rarely from just below R2 to R4 or to between R4 and R5; M-stem to M2 or absent; all veins present and separate beyond cell; R1 from middle of cell
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margin; R2 arising nearer to R1 than to R3, R4 to costa, R5 to termen, in Australian species bases of R3, R4 and R5 approximated; M2 approximated to M3 at base; CuA1 from posterior angle of cell, CuA2 from 36: of cell, CuP present at wing margin; greyish brown to blackish brown, a blackish triangular pretornal mark usually present. Hindwing shape subtrapezoidal or sexually dimorphic in some species of Afrotropical leucotreta-group (e. g. T. leucotreta hindwing subtriangular in male), an ovate androconial patch sometimes present on distal 36: of CuA2 in the leucotreta-group; melanic androconial scales above along anterior and posterior margins of cell in one Australian species (Fig. 107); cubital pecten well developed; frenulum of female three bristles; cell short; Rs and M1 clearly separate at base, though approximated; M2 arising near stalk of M3 and
CuA1, M3 and CuA1 stalked, CuP present at wing margin; basal loop of 1A+2A only weakly indicated, 3A present; greyish brown. Pregenital abdomen (Figs 899, 900). Sternum 2 with well-developed anterolateral processes. Tergum 8 of male a broadly sclerotised plate with a medially convex posterior margin laterally produced into curved points, densely covered with easily removable, long mane-like scales; T6 and T7 of male sometimes with paired lateral patches of easily removable, long hair-scales. Coremata: eighth sternite a crescentic plate; intersegmental ventral sclerite lacking levers; with or without paired tufts of filiform scales deeply invaginated in membranous pockets on each side. Male genitalia (Figs 898, 901). Vinculum a moderate band. Tegumen ovate, a wide band, anterior margin sometimes sinuate, rounded at top.
Figs 894–897. Thaumatotibia, adults. 894, 895, T. zophophanes. 894, , Claudie R., Q. 895, , Quinn Park, Q. 896, 897, T. aclyta. 896, , Tooloom, NSW. 897, , Toowoomba, Q. Scale = 2 mm.
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Uncus usually absent, rarely a bipartite flat lobe with two sharp laterodistal points. Socii rarely present (not in Australian species). Gnathos weakly sclerotised, paired arms medially connected across base of anal tube, sometimes denticulate. Valva very large, little constricted before middle; basal excavation ill-defined; costal process narrow and weak; cucullus densely bristled on inner surface, with dense, very long, curled, easily removable scales on outer surface; sacculus often ending in distal ridge or teeth. Juxta triangular or trapezoidal; caulis very long, its base developed into a pair of conspicuous, spinulose lobes in the chaomorpha-group, without such lobes in the Afrotropical leucotreta-group. Aedeagus bulbous basally, narrowed at basal 56M to 56: and then almost straight; vesica with a series of fine spiniform cornuti. Female genitalia (Figs 903, 904). Ovipositor lobes moderate, subreniform. Sternum 7
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trapezoidal, ostium in shallow or deep, rounded or V-shaped posterior incision. Sterigma a protruding roundish to rectangular raised rim, posteriorly connected with a pair of ovate, sclerotised depressions with modified scales. Ductus bursae long and narrow, sometimes widening anteriorly, membranous except for narrow colliculum and an irregular small sclerite at posterior 56:–56:; ductus seminalis arising laterally, from posterior 56M–569 of ductus bursae; bulla seminalis present; corpus bursae ovate, with a ring of spinules around its entrance, sometimes with a small diverticulum; with two large, horn-shaped signa.
Distribution Ethiopian region (leucotreta-group), Oriental, Australian and eastern Palaearctic regions (chaomorpha-group). In Australia, Thaumatotibia occurs in the Darwin area of the Northern
Figs 898–902. Thaumatotibia, male genitalia, abdomen and head. 898, 899, T. zophophanes, Malanda, Q, slide ANIC 8185. 898, male genitalia. 899, 8th segment with coremata. 900, 901, T. aclyta, Rockhampton, Q, slide ANIC 8192. 900, 8th segment. 901, male genitalia. 902, head, T. zophophanes.
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Territory, Cape York Peninsula in Queensland to northern New South Wales as far south as the Allyn R.
Biology Thaumatotibia larvae are fruit borers, and a broad spectrum of plant families are known as larval host plants: Poaceae, Rutaceae, Lauraceae, Myrtaceae, Theaceae, Proteaceae, Sterculiaceae. Some species are polyphagous, which is unusual for Grapholitini, and known as pests, e.g. T. batrachopa (Meyrick) and T. leucotreta (Meyrick) (both in Africa). In Australia, T. zophophanes (Turner), which occurs from Cape York Peninsula to Eungella, is a pest on macadamia and avocado on the Atherton Tableland, Queensland, and has been reared from fruit of Salacia chinensis (Hippocrateaceae) by J. Lawson at Cape Tribulation. Sex pheromone components of Thaumatotibia have been reviewed by Komai (1999).
Remarks Komai (1999) has revised Thaumatotibia, providing illustrations of their genitalia and early stages. Adamski and Brown (2001) discussed the phylogenetic position of this genus in the context of a revision of the New World Ecdytolopha-group. Thaumatotibia is closely related to Cryptophlebia and the New World Pseudogalleria Ragonot, Gymnandrosoma Dyar, and Ecdytolopha Zeller (Adamski and Brown 2001), from which it can be separated by the following apomorphies: 1, tergum 8 in male with a broadly sclerotised plate with medially convex posterior margin laterally produced into curved points; 2, sterigma an ovate or rectangular raised rim posteriorly connected with a pair of ovate depressions with modified scales; 3, corpus bursae with a ring of spinules around its entrance (Komai 1999).
Constituent species The 20 described species worldwide are divided into two species-groups: the chaomorpha-group with an Indo-Australian distribution and the leucotretagroup that is restricted to the Ethiopian region. We are aware of three species from Australia of which the following two are named. zophophanes (Turner, 1946), Trans. R. Soc. S. Aust. 70: 217 (Argyroploce). [Qld: Kuranda.] Comb. nov. scioessa (Turner, 1946) Trans. R. Soc. S. Aust. 70: 218 (Articolla). [Qld: Lake Barrine.] Syn. nov. aclyta (Turner, 1916) Trans. R. Soc. S. Aust. 40: 529 (Alypeta). [Qld: Brisbane, Rosewood.] Comb. nov.
Gymnandrosoma Dyar (Figs 109, 905–908) Gymnandrosoma Dyar, 1904, Proc. Entomol. Soc. Wash. 6: 60. Type species: Gymnandrosoma punctidiscanum Dyar, 1904, ibid. 6: 60, by monotypy. [North America.]
Diagnosis Figs 903, 904. Thaumatotibia, female genitalia. 903, T. zophophanes, holotype, slide ANIC 16885. 904, T. aclyta, Buderim, Q, slide ANIC 16996.
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Australian Gymnandrosoma species are large, mottled dark grey olethreutines with triangular wings with an acute apex and nearly straight, oblique termen, with an outwardly oblique blackish
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band from beyond 56: dorsum often angled in middle of wing and running to costa at 56:, and with alternating dark and pale scales along termen suggesting a crenulated margin. Long, sinuate labial palpi, a conspicuous posterior crest on the thorax, hindwing venation with M2 distant from M3 at base and male hind tibia with long loose dorsal scales are typical for the genus in Australia. The male genitalia of Australian Gymnandrosoma species are characterised by a subtriangular dorsal tegumen process with groups of short bristles on posteroventral surface, a slender valva with a parallel-sided, narrower medial portion, a rounded cucullus and a semicircular depression on outer
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base with a well-defined, semicircular dorsal margin, and a long, slender aedeagus with a slightly bulbous base, bent nearly 90° and with a large bundle of lanceolate cornuti oblique to the aedeagus axis.
Description Adults (Fig. 905) (description based on Australian species) Wingspan. 19–23 mm (Australian species); large. Head. Ocellus well developed, chaetosema distinct. Frons in upper half with very dense, long,
Figs 905–908. Gymnandrosoma, adult, male genitalia and abdomen. 905, G. sp. nr gonomela, , Kalbarri, WA. 906, G. sp. nr gonomela, Kalbarri, WA, male genitalia, slide ANIC 16917. 907, 908, G. gonomela, syntype, Blackwood, SA [SAMA]. 907, 8th segment. 908, male genitalia. [908 at higher magnification than 906]. Scale = 2 mm.
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forwardly projecting, horseshoe-shaped scale tuft; remainder with short, upwardly oriented scales. Proboscis short, not longer than labial palpus. Labial palpus long (2.2–2.4× diameter of eye), sinuate, porrect; second segment strongly widened dorsomedially, triangular; terminal segment long, slender, downward pointing. Antenna less than 36: of forewing length, entirely scaled, cilia in male short. Thorax. With posterior crest. Legs with hind tibia in male modified, with long loose scales along dorsal margin. Wings (Figs 109, 905). Forewing index: 0.39– 0.42; forewing elongate subtriangular, with nearly straight costa, acute apex and oblique, weakly curved to nearly straight termen; chorda and Mstem both present, accessory cell very narrow, from below R2 to between R4 and R5; all veins separate from cell; R4, R5 and M1 about equidistant; M3 closer to M2 than to CuA1; CuA2 from 36: cell; CuP present at margin; wing pattern mottled dark grey to blackish (with brownish tinge in older specimens), with pale, long strigulae along costa, with intermittent, narrow, wavy black lines across wing in distal half, with a dark blotch halfway between distal margin of cell and apex and an outwardly oblique dark band beyond 56: dorsum, often forked or angled in middle of wing and running to 56: costa; rarely with a faint indication of dorsal blotch as two parallel streaks of paler grey scales beyond dark band from dorsum. Hindwing with long hairscales from upper surface of anal region in male but without anal fold; cubital pecten and anal tuft well developed; Rs and M1 closely approximated and parallel near base; M2 and M3 distant and nearly subparallel towards base, only gradually approaching each other; M3 and CuA1 very shortly stalked, CuA1 from below 36: cell; CuP present at wing margin; 1A+2A and 3A well developed; hindwing grey to dark grey. Pregenital abdomen (Fig. 907). Sternum 2 with well-developed anterolateral processes. Coremata reduced: eighth sternite with arched hind margin, intersegmental ventral sclerite weak, scale tufts indicated by sparse hair scales. Male genitalia (Figs 906, 908). Vinculum a ventrally wider, V-shaped band. Tegumen high and narrow, inverted U to V-shaped, a wide, moderately sclerotised band; tip of tegumen projecting as a subtriangular, sclerotised lobe, treated as not homologous with uncus and termed ‘scaphial ridge’ by Adamski and Brown (2001), variably set with short bristles on posteroventral surface. Gnathos arms weakly sclerotised, medially upcurved and
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parallel along subscaphium. Valva long and slender, ventral margin angled before 56:, middle section narrowest with costa nearly parallel to ventral margin, cucullus rather small, roundish, inner surface entirely covered with dense bristles; costal process membranous; outer surface of valva base with a large depression with a conspicuous, semicircular, strongly projecting dorsal margin. Juxta small, triangular, wide and low; caulis long. Aedeagus long, slender, strongly curved so that base and tip are nearly at a right angle, very lightly bulbous basally, otherwise parallel-sided; vesica with a large sheaf of lanceolate cornuti. No females known of Australian Gymnandrosoma species.
Distribution Gymnandrosoma is widely distributed across the New World, from southern Canada to northern Argentina. In Australia, the genus is only known from four specimens of G. gonomela (Lower) from Adelaide in South Australia, and a single specimen of an unnamed species from near Kalbarri in Western Australia.
Biology New World species of Gymnandrosoma are all borers, boring in fruit and nuts, with G. aurantianum Lima, a pest of citrus with the common name ‘orange worm’, reported from many different families including Citrus sp. (Rutaceae), Psidium guava (Myrtaceae) and Macadamia integrifolia (Proteaceae) (Adamski and Brown 2001). No hosts are known for the Australian species, but Lower (1899) stated for the type series from Blackwood near Adelaide ‘three specimens dislodged from Banksia sp. in September’, that strongly suggests that Banksia (Proteaceae) is the foodplant of G. gonomela.
Remarks Adamski and Brown (2001) provided a comprehensive, well-illustrated revision of Gymnandrosoma. The genus is represented by only five males from Australia, one recently collected and well-preserved male and three damaged males from the type series of G. gonomela (Lower) from Adelaide, and a single male of an undescribed species from Western Australia. According to the hypothesised phylogeny in Adamski and Brown (2001), Gymnandrosoma is
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characterised by: 1, a male antennae with the basal portion slightly flattened, with conspicuous cilia; 2, a relatively wide forewing with length 1.98– 2.08× width; 3, a male hindtibia with dense, modified scaling; 4, an unmodified, not swollen valva, and 5, numerous, relatively short cornuti arranged horizontally in vesica. The two Australian species are tentatively included in the genus, pending discovery of females. As in Gymnandrosoma, they have a hindtibia with modified scaling, an unmodified and not swollen valva and numerous cornuti set obliquely to the aedeagus axis, but they lack the antennal modification and have narrow forewings (length 2.4–2.6× width). The overall similarity of the tegumen apex with scattered bristles in its posteroventral surface is the most compelling reason to include the Australian species in Gymnandrosoma, though the valva is strongly derived with a conspicuous semicircular depression on its outer surface.
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Hindwing with short cell, dark grey, often yellow at least in basal half; with M2 at base bent quite close to short stalk of M3 and CuA1. Thorax with conspicuous posterior crest. Hindtibia in male with a dorsal fringe of long scales. Tergum 8 in male a semicircular or subrectangular, lightly sclerotised plate with long scales; S8 unmodified, a small semicircular plate. The strongly sclerotised male genitalia are characterised by a clavate or ovate cucullus densely bristled with some strong spines along ventral margin on inner surface and densely covered with very long hair-like scales on outer surface, and an almost straight or strongly sinuate aedeagus with an elongate patch of very small, transverse cornuti. In the female genitalia, S7 has an unscaled medioanterior region, the ductus bursae is narrow and at least partially sclerotised in its posterior half, curved near the middle and then gradually widening into the corpus bursae that is not spinulose.
Description Constituent species There are seven described New World species, one named Australian species and an unnamed second species from Western Australia. gonomela (Lower, 1899), Proc. Linn. Soc. NSW 24: 94 (Arotrophora). [SA: Blackwood.] Comb. nov.
Archiphlebia Komai & Horak, gen. nov. (Figs 111, 112, 909–914) Type species: Argyroploce endophaga Meyrick, 1911b, Proc. Linn. Soc. NSW 36: 278, hereby designated. [Australia.]
Diagnosis Medium-sized moths with typical grapholitine wing pattern elements absent, forewing either leaden grey with short transverse blackish lines across basal 36: of wing as rows of raised scales on dorsum, indication of a blackish pretornal mark and a blackish subapical spot, or pale ochreous with a pale grey, ill-defined band from 36: costa to tornus and a large subtriangular blotch of orange or reddish ochreous scales in apical area. Forewing with a moderately large accessory cell with chorda from between R1 and R2 to between R4 and R5.
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Adults (Figs 909, 910) Wingspan. 12.5–19.0 mm; medium-sized. Head (Fig. 911). Ocellus well developed, chaetosema distinct. Frons in upper half with very dense, erect and long, forwardly projecting scales, nearly touching labial palpi, remainder with short, upwardly oriented scales. Proboscis rather short, not much longer than labial palpus. Labial palpus from rather short to moderately long, longer in female (1.6–2.4× diameter of eye), porrect, sinuate; second segment widened distally but scales appressed and rather short; terminal segment porrect, about 56: length of second, longer and more slender in female, with appressed scales, apex blunt. Antenna less than 36: length of forewing, entirely scaled. Thorax. With posterior crest. Legs with a dorsal fringe of long scales on male hind tibia (less conspicuous in A. rutilescens (Turner)), but otherwise unmodified. Wings (Figs 111, 112, 909, 910). Forewing index: 0.40–0.45; forewing subtriangular, costa nearly straight to gently curved throughout, termen almost straight, oblique; in A. endophaga (Meyrick) several parallel transverse interrupted rows of raised blackish scales on dorsum to fold; accessory cell moderately large, chorda from between R1 and R2 to R5 or between R4 and R5, M-stem to between M2 and M3; all veins present and separate beyond cell, R1 from 56O of cell, R4 to costa, R5 to termen, M2 approximated to M3 at base, CuA1 from
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posterior angle of cell, CuA2 from 36: of cell, CuP present at wing margin; wing without typical grapholitine pattern elements, colour and pattern variable but always with a darker mark or blotch in apical area, A. endophaga leaden grey with short transverse blackish lines across basal 36: of wing (as rows of raised scales in dorsal region), pretornal spot indicated by black scales and a small blackish subapical spot, A. rutilescens pale ochreous with a
pale grey ill-defined band from 36: costa to tornus and a large subtriangular blotch of orange or reddish ochreous in apical area. Hindwing with cubital pecten well developed; frenulum in female with three bristles; cell short; Rs and M1 parallel and closely approximated, sometimes anastomosing; M2 at base bent quite close to short stalk of M3 and CuA1, M3 and CuA1 short-stalked; CuA2 from 36: of cell; CuP present at wing margin,
Figs 909–914. Archiphlebia endophaga, adults, head, abdomen and genitalia. 909, 910, adults, Carnarvon, WA. 909, . 910, . 911, head, male. 912, 913, male, nr Carnarvon, WA, slide ANIC 16910. 912, 8th segment. 913, genitalia. 914, female genitalia, Carnarvon, WA, slide ANIC 2453. Scale = 2 mm.
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basal loop of 1A+2A only weakly indicated, 3A present; pale yellow at least in basal half in A. endophaga, dark grey in A. rutilescens. Pregenital abdomen (Fig. 912). Sternum 2 with well-developed anterolateral processes. Tergum 1– T8 of male without the modified scales present in Cryptophlebia Walsingham. Tergum 8 a subrectangular or semicircular, lightly sclerotised plate with long scales. Coremata: eighth sternite a crescentic (A. endophaga) or subrectangular (A. rutilescens) plate densely set with long hair scales along hind margin; intersegmental ventral sclerite lightly sclerotised, not modified; scale tufts absent. Male genitalia (Fig. 913). Vinculum a U- or V-shaped band, wider ventrally. Tegumen dorsally rounded, a simple narrow to moderately broad band. Uncus and socius absent. Gnathos a pair of lateral bands, continuous with tegumen margin (A. rutilescens) or not (A. endophaga). Valva strongly sclerotised, basal excavation well-defined; costal process small; cucullus clavate or ovate, densely bristled with some strong spines along ventral margin on inner surface, densely covered with very long hair-like scales on outer surface. Juxta small (A. endophaga) or large (A. rutilescens), triangular; caulis long (A. endophaga) or short (A. rutilescens). Aedeagus almost straight or strongly sinuate, slightly bulbous at base; vesica with an elongate patch of very small, transverse cornuti. Female genitalia (Fig. 914). Ovipositor lobes rather broad. Sternum 7 with excavated, U- or V-shaped hind margin, medioanterior region strongly sclerotised and unscaled, with large ovate scaled lateral portions, deeply concave in A. rutilescens. Sterigma variable, a small cup, a ring around ostium, or a projecting ring-shaped wide collar surrounding end of ductus bursae. Ductus bursae in posterior half narrow, partially sclerotised and curved, from there gradually widening into corpus bursae, inception of ductus seminalis from middle to posterior 56: of ductus bursae. Corpus bursae not spinulose; signa two large, flat, curved, pointed horns with a large basal opening.
Distribution Archiphlebia is endemic to Australia. The genus occurs in Queensland, the Northern Territory, Western Australia and South Australia. Archiphlebia endophaga and an allied unnamed species are centred on Western Australia south from Broome and in the mallee of eastern South Australia. Archiphlebia rutilescens occurs in a wide band along the coast from the Kimberley of Western Australia, the Northern Territory and
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Queensland from the Gulf of Carpentaria and Cape York Peninsula south to Chinchilla.
Biology Meyrick (1911b) recorded that the holotype of A. endophaga has been bred from a larvae feeding ‘on pods of Acacia sp., eating the seeds and ejecting refuse through a hole, in October’ (Mimosaceae). Turner (1946: 204) stated that the holotype of A. diaema (Turner) [ = rutilescens (Turner)] is from Queensland, ‘one specimen bred from fruit’, but then goes on to say ‘Unfortunately the record of locality and host plant has been lost’. The species are found in arid, semiarid or seasonally dry locations.
Remarks Horak, Common and Komai (1996) listed A. rutilescens and A. endophaga as members of Cryptophlebia. The large posterior crest on the thorax, the strongly sclerotised male genitalia with a tegumen with a round top, a cucullus densely covered with long curved hair scales on outer surface and T8 in the male set with long hair scales suggest that Archiphlebia is allied to Thaumatotibia Zacher, Cryptophlebia and Gymnandrosoma Dyar. Superficial appearance, wing pattern and signa shape confirm such a relationship. Apomorphies characterising Archiphlebia are: 1, the posterior half of ductus bursae narrow and curved near the middle, and 2, S7 with unscaled medio-anterior region.
Constituent species We are aware of three species including two named ones. rutilescens (Turner, 1945), Trans. R. Soc. S. Aust. 69: 70 (Cnephasia). [Qld: Chinchilla, Leichhardt, Injune.] Comb. nov. diaema (Turner, 1946), Trans. R. Soc. S. Aust. 70: 204 (Eucosma). [Qld.] endophaga (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 278 (Argyroploce). [WA: Carnarvon.] Comb. nov.
Cryptophlebia Walsingham (Figs 108, 110, 915–926) Cryptophlebia Walsingham, 1900, Indian Mus. Notes 4 (1899): 105. Type species: Cryptophlebia
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carpophaga Walsingham, 1900, ibid. 4: 106 [ = Arotrophora (?) ombrodelta Lower, 1898, Proc. Linn. Soc. NSW 23: 48], by original designation. [India.] Pogonozada Hampson, 1905, Ann. Mag. Nat. Hist. (7)16: 586. Type species: Pogonozada distorta Hampson, 1905, ibid. (7)16: 586, by original designation. [China.] Phanerophlebia Diakonoff, 1957b, Tijdschr. Entomol. 100: 142 [originally as subgenus of Cryptophlebia]. Type species: Cryptophlebia perfracta Diakonoff, 1957b, ibid. 100: 142. by original designation. [Java.]
Diagnosis Medium-sized to very large moths with broad forewings, often with a darker subtriangular pretornal patch and usually with a small accessory cell with chorda from between R2 and R3 to between R4 and R5. Hindwing with short cell, especially in male, and thorax with an often conspicuous crest. Males of Cryptophlebia often have secondary sexual structures such as a modified hindtibia with tufts of hair-like scales, a modified hindwing with sometimes an elongate, heavily sclerotised pocket filled with fine hairscales beneath in distal half of CuA2, long and mane-like scales dorsally on the abdomen. Males of all species have T8 with a medial subtriangular or Y-shaped plate and with paired tufts of long filiform scales arising from shallow membranous pockets on each side, and nearly always coremata with tufts of modified, short, swollen scales. The male genitalia are characterised by a strongly swollen, clavate valva with few sparse, very large spines on the inner surface of the cucullus, and with dense, very long, curled scales on the outer surface. In the female genitalia, the sterigma is very variable but fused with the deeply excavated hindmargin of S7, the corpus bursae is usually spinulose in anterior 56O–169 and sometimes has a diverticulum, and the signa are two curved horns.
Description Adults (Figs 915–918) Wingspan. 11.5–30.0 mm (Australian species); medium-sized to very large. Head (Fig. 920). Ocellus well developed, chaetosema distinct. Frons in upper half with very dense, erect and long, forwardly projecting scales, nearly touching labial palpi, remainder with short,
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upwardly oriented scales. Proboscis rather short, not much longer than labial palpus. Labial palpus from rather short to moderately long, longer in female (1.6–2.7× diameter of eye), porrect, sinuate; second segment variably widened distally but scales appressed and rather short; terminal segment porrect, about 56: length of second, slender, with appressed scales, apex blunt. Antenna less than 36: length of forewing, entirely scaled or ventral band unscaled; cilia in male small. Thorax. With posterior crest. Legs with hindtibia and first hind tarsal segment of male sometimes modified, short and wide, with tufts of hair-like scales, inner surface of hindtibia often with naked area. Wings (Figs 108, 110, 915–918). Forewing index: 0.41–0.52; forewing oblong subtriangular, costa gently curved throughout, apex rounded, termen almost straight, oblique; accessory cell small or absent, chorda from between R2 and R3 (often closer to R3) to between R4 and R5, or chorda absent, Mstem to M2 or between M2 and M3; all veins present and separate beyond cell, R1 from 56O of cell, R2 arising nearer to R1 than to R3, R4 to costa, R5 to termen, M2 approximated to M3 at base, CuA1 from posterior angle of cell, CuA2 from 36: of cell, CuP present at wing margin; wing pattern often sexually dimorphic, either of greyish ochreous, pinkish, brownish to dark brown or reddish to dark red ground colour with a darker apex and usually a dark subtriangular pretornal mark followed by paler band from 36: costa to tornus, or brown with a paler elliptic mark on termen, or pale ochreous and either speckled with red-brown or with a large, bipartite dark mark below distal half of costa; with typical grapholitine pattern elements (ocelloid patch and dorsal blotch) reduced or absent. Hindwing sometimes sexually dimorphic in shape: subtrapezoidal to triangular in male, subtrapezoidal in female; male sometimes with elongate androconial pocket filled with fine hair-scales beneath in distal half of CuA2; cubital pecten well developed, frenulum in female with three bristles; cell short, especially in male; Rs and M1 clearly separate at base though approximated; M2 arising near M3, M3 and CuA1 connate or short-stalked; CuA2 from 36: of cell; CuA1 disappearing distally and CuA2 curved due to formation of androconial pocket in male of some species; CuP present at wing margin, basal loop of 1A+2A only weakly indicated, 3A present; pale to dark grey, greyish brown, sometimes whitish near base. Pregenital abdomen (Figs 925, 926). Sternum 2 with well-developed anterolateral processes.
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AUSTRALIAN OLETHREUTINE GENERA
Tergum 1–T8 of male with mane-like patch of long, easily removable scales. Tergum 8 of male with a sclerotised Y-shaped or subtriangular plate and with paired tufts of long filiform scales arising from shallow membranous lateral pockets near hindmargin. Coremata: eighth sternite a crescentic plate usually with short lateral projections; intersegmental ventral sclerite with a pair of short levers distally each associated with a tuft of modified, short, inflated scales; in some species levers and scale tufts absent. Male genitalia (Figs 919, 923, 924). Vinculum a V-shaped band, wider ventrally. Tegumen hoodshaped, a wide, dorsally strongly widened band, anterior edge sometimes concave in middle, with blunt top. Uncus absent, at most a rudimentary indication. Gnathos a pair of weakly sclerotised
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horizontal bars, articulated with tegumen. Valva swollen, clavate; basal excavation reaching base of cucullus; costal process well developed; cucullus sparsely bristled or lightly hairy but with a few very large spines on inner surface, usually along margin, and densely covered with very long hair-like scales on outer surface. Juxta triangular or trapezoidal; caulis rather long. Aedeagus bulbous basally, curved; vesica with two to a series of spiniform cornuti. Female genitalia (Figs 921, 922). Ovipositor lobes rather broad. Apophyses anteriores longer than apophyses posteriores. Hindmargin of S7 deeply concave, usually fused with sterigma. Sterigma variable, from a ventrally strongly sclerotised shallow pocket or funnel to a longitudinal shallow depression behind entrance to ostium, often with two longitudinal, sclerotised bands. Ductus bursae long,
Figs 915–918. Cryptophlebia, adults. 915, 916, C. rhynchias. 915, , nr Cooktown, Q. 916, , nr Hope Vale Mission, Q. 917, 918, C. ombrodelta. 917, , Daintree, Q. 918, , Cooktown, Q. Scale = 2 mm.
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Figs 919–926. Cryptophlebia, genitalia, head and abdomen. 919, C. rhynchias, male genitalia, Vanuatu, slide ANIC 2461. 920, head, C. ombrodelta, male. 921, C. rhynchias, female genitalia, Mt Bellenden-Ker, slide ANIC 2456. 922, C. ombrodelta, female genitalia, Moreton, Q, slide ANIC 16731. 923, 924, C. ombrodelta, aedeagus and male genitalia, Nambour, Q, slide ANIC 2109. 925, C. ombrodelta, male, coremata. 926, C. rhynchias, male, coremata.
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AUSTRALIAN OLETHREUTINE GENERA
widening anteriorly, membranous except for bipartite colliculum (sometimes indicated only by constriction) and an ill-defined sclerotisation in middle part; origin of ductus seminalis in anterior third of ductus bursae; bulla seminalis present; corpus bursae from globose to pyriform, usually spinulose in anterior 56O–169, sometimes with a diverticulum, with two usually large, curved, often laterally flattened, horn-shaped signa.
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margin; 2, male 8S with a pair of short projections laterally (absent in two undescribed Australian species with reduced coremata); 3, a greatly swollen, clavate valva with sparse strong spines on the inner surface of the cucullus, and, 4, the corpus bursae spinulose in anterior 56O–169 (absent in two undescribed Australian species).
Constituent species Distribution Worldwide except for the Nearctic region, well represented in the Indo-Pacific region. In Australia, the genus occurs in the Northern Territory, Western Australia, Queensland, and New South Wales.
Biology Taxonomically diverse plant families such as Anacardiaceae, Hippocastanaceae, Fabaceae, Proteaceae, Rhizophoraceae, Rutaceae, and Sapindaceae are recorded as hostplants for Cryptophlebia, but the genus is essentially associated with Fabaceae (Komai 1999). The larvae are borers of fruit (seeds, nuts, pods), galls or twigs. In Australia, C. ombrodelta is a pest of Macadamia spp. (Proteaceae) and Litchi chinensis (Sapindaceae). It has been reared from Buckinghamia celsissima (Proteaceae), Cupaniopsis anacardioides (Sapindaceae), Acacia spp., Adenanthera pavonina (Mimosaceae), Sesbania (Fabaceae), Bauhinia, Cassia, Poincianna and Delonix (Caesalpiniaceae). Sex pheromone components of Cryptophlebia have been reviewed by Komai (1999).
There are some 50 described species (Brown 2005), of which nine species including four described ones are found in Australia: ombrodelta (Lower, 1898), Proc. Linn. Soc. NSW 23: 48 (Arotrophora). [NSW: Sydney.] carpophaga Walsingham, 1900, Indian Mus. Notes 4 (1899): 106 (Cryptophlebia). [India.] iridosoma (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 264 (Argyroploce). [Qld: Brisbane.] ceratodes (Turner, 1946), Trans. R. Soc. S. Aust. 70: 209 (Eucosma). [Qld: Cape York.] rhynchias (Meyrick, 1905), J. Bombay Nat. Hist. Soc. 16: 586 (Platypeplus). [Sri Lanka.] pallifimbriana Bradley, 1953, Bull. Entomol. Res. 43: 688 (Cryptophlebia). [Fiji.]
Acanthoclita Diakonoff (Figs 113, 115, 927–935) Acanthoclita Diakonoff, 1982, Zool. Verh. (Leiden) 193: 27. Type species: Eucosma balanoptycha Meyrick, 1910, Rec. Indian Mus. 5: 218, by original designation. [India.]
Remarks
Diagnosis
Comprehensive reviews of Cryptophlebia have been given by Bradley (1953), Diakonoff (1957b), Razowski (1989), and Komai (1999). Adamski and Brown (2001) revised New World Ecdytolopha Zeller, Gymnandrosoma Dyar, and Pseudogalleria Ragonot, and discussed the phylogenetic relationships among these genera and the Old World Cryptophlebia Walsingham and Thaumatotibia Zacher. Komai (1999) recognised the following character states supporting the monophyly of Cryptophlebia: 1, male T8 subtriangular or Y-shaped with a pair of tufts of long filiform scales arising from shallow membranous pockets laterally near the hind
Smallish, rather narrow-winged, usually intricately patterned moths of eucosmine appearance with dense, fine strigulation along entire costa and a dark roundish spot in apex between two white strigulae, and with the hindwings usually translucent towards base and in male with modified scales on anal area above and/or below. However, one Australian species has the wing pattern and wing venation of a species of Matsumuraeses (Fig. 929), with only the genitalia referring the taxon to Acanthoclita. Acanthoclita has large, porrect, strongly sinuate labial palpi, with the second segment distally widened by long scales. The venation is diverse with R4 and R5 in the forewing
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separate or stalked and the hindwing usually with M3 and CuA1 stalked and distant from the base of M2 but in one species of Matsumuraeses facies with M3 and CuA1 connate and the base of M2 strongly bent towards M3. The male genitalia are diagnostic, with a very broad, short vinculum and a weakly sclerotised tegumen with the base of the gnathos arms often conspicuously fused with the tegumen, with long and very slender valvae with the cucullus usually crescent-shaped and often upcurved and, in addition to a band of spines along its ventral margin, nearly always with a single strong spine at its base somewhat distant from the margin. The juxta is subtriangular, dorsoventrally compressed, the band-shaped caulis curved and very long, and the long, curved or bent aedeagus has a sheaf of lanceolate deciduous cornuti and a more distal series of spiniform, fixed cornuti. The female genitalia of Acanthoclita are quite characteristic in the derived species with a complex sterigma surrounding the ostium and fused with the sclerotised, funnel-shaped end of the ductus bursae, but the simple transverse sclerite behind the ostium with a patch of modified scales on each side found in less derived species is shared with Matsumuraeses Issiki. The ductus bursae is membranous or variably sclerotised, with the origin of the ductus seminalis variable, and the ovate corpus bursae has two usually curved and blade-shaped, distally pointed signa, but sometimes their tip is roundedtruncate or the signa are distally much wider, paddle-like blades.
Description Adults (Figs 927–929) Wingspan. 11–14 mm (Australian species); small to medium-sized. Head (Fig. 935). Ocellus large, chaetosema very obvious. Maxillary palpi minute, scaled. Proboscis rather short, not much longer than labial palpus. Labial palpus large (1.5–2.1× diameter of eye), porrect and strongly sinuate in both sexes, with second segment sinuate and widened to a broadly triangular outline by long, loose scales along ventral and distal part of dorsal margin, partly concealing the long, slender, downward pointing distal segment with short, appressed scales. Antenna to beyond middle in male, no cilia visible in male. Thorax. Without crest, with moderate scale tuft from anterior corner of tegula. Legs unmodified.
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Figs 927–929. Acanthoclita, adult males. 927, A. trichograpta, Mt Webb Nat. Park, Q. 928, A. sp., Darwin, NT. 929, A. sp., Richmond R., NSW. Scale = 2 mm.
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AUSTRALIAN OLETHREUTINE GENERA
Wings (Figs 113, 115, 927–929). Forewing index: 0.32–0.38 (Australian species); forewing of variable shape, subrectangular, apex acute and protruding to rectangular, termen usually with notch below apex and with remainder of termen convex and often protruding beyond apex (not in Australian species); with chorda and M-stem well developed, chorda from between R1 and R2 to base of R5 or to between stalk of R4 and R5 and M1, closing vein weak; all veins present beyond cell, R1 from about middle of cell, R4 and R5 separate or stalked, M3 and CuA1 distally usually curved towards apex and M2 and M3 often closer on margin than at base, CuA2 from beyond middle to 36: cell; CuP present at margin; forewing pattern diverse and complex, greyish brownish, usually with very fine pale strigulae separated by fine dark marks along entire costa and with a conspicuous roundish dark spot at apex edged with a white strigula before and below apex, with ocelloid patch at most weakly indicated and row of parallel black lines sometimes extending in a curve towards 6M costa, dorsal blotch a variably developed curved brownish mark tapering towards centre of wing; one Australian species of Matsumuraeses-like appearance (Fig. 929) with wing pattern entirely or in large blotchy areas suffused by uniformly brown or whitish ochreous scales. Hindwing with well-developed cubital pecten and males sometimes with enlarged anal area (not in Australian species) and always with modified scaling on upper and/or lower surface, often with a patch of dense, short, modified scales on lower surface towards margin between CuP and 1A+2A, sometimes combined with a dense fringe of long scales along anal margin, or with a dense cover of modified lanceolate scales on the upperside behind 1A+2A or with modified scaling on both sides with a combination of orange scaling and a large bolster of blackish scales below; in female with a frenulum of three bristles; Rs and M1 closely approximated and parallel at base, usually M3 and CuA1 stalked and distant at base from M2, but in the species with Matsumuraeses facies M2 is bent towards M3 at base and M3 and CuA1 are connate to barely stalked; CuA2 from middle of cell in species with Matsumuraeses facies but usually from further distally; basal loop of 1A+2A only weakly indicated; 3A present; greybrown to blackish brown with basal half of wing usually very sparsely scaled, translucent, especially behind base of Cu.
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Pregenital abdomen (Fig. 934). Well-developed anterolateral processes on S2. Coremata: eighth sternite a small, subrectangular to pentagonal or crescentic plate, intersegmental ventral sclerite large, crescentic or bilobed, sometimes with a posterior V-shaped rib, with short lateral levers associated with small to very large tufts of long, broad, distally rounded scales, in one Australian species scale tufts absent; T8 in male usually with strongly sclerotised T-shaped anterior margin, hindmargin rounded or bilobed; in males of some species very long scales with modified (?glandular) sockets laterally on hindmargin of 7th or 6th and 7th segments and/or a band of very short, dense scales along anterior margin of T3 and T4, with lanceolate scales on remainder of these two tergites. Male genitalia (Figs 930, 931). Vinculum a very broad, lightly sclerotised band. Tegumen high and narrow, subtriangular to ovate, quite deep but very weakly sclerotised except for anterior margin. Uncus not differentiated. Gnathos in some species very obvious as two horizontal arms apparently fused with tegumen, rising medially and fading. Valva long and very slender, constricted at or before midlength; basal excavation large; costal process developed; basal half with an angular sacculus or elongate and distally gradually tapering; cucullus from crescentic to falcate, rarely clavate, usually very narrow and upcurved, with a row of heavy spines along ventral margin and nearly always with one or two large spines isolated and more distant from margin at base of cucullus; juxta subtriangular or a dorsoventrally compressed rectangle; caulis a very long, curved band; aedeagus long, slender, tapering towards tip, strongly bent or curved and apically weakly sinuate; vesica with a sheaf of deciduous, lanceolate cornuti and a more distal series of minute to strong, spiniform, fixed cornuti. Female genitalia (Figs 932, 933). Ovipositor lobes elongate. Apophyses long, slender, nearly straight. Hindmargin of S7 rounded-concave to V-shaped. Sterigma at its simplest a small transverse sclerite behind ostium carrying a patch of modified scales on each side, in the more derived condition extended in a smooth semicircle around ostium with the entrance into the ostium a sclerotised funnel and the ventral margin sometimes forming a raised rim. Ductus bursae membranous or variably sclerotised, with either the posterior half with a strongly sclerotised, curved band or with a weak sclerotisation just below ostium or below inception
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of ductus seminalis; origin of ductus seminalis variable, from 56: below ostium to close to corpus bursae; bulla seminalis present; corpus bursae ovate, with two curved, blade-shaped signa, usually pointed towards apex, more rarely with roundedtruncate apex or signa resembling distally widened, paddle-shaped blades.
Distribution Palaearctic, Ethiopian, Oriental and Australian regions, to Fiji in the Pacific. In Australia the genus occurs in the northern, wetter areas of the Northern Territory and in north-east Queensland, with one taxonomically isolated species of Matsumuraeses-like appearance on the eastern coast
Figs 930–935. Acanthoclita, genitalia, abdomen and head. 930, 931, male genitalia with aedeagus. 930, A. trichograpta, nr Cooktown, Q, slide T2322. 931, A. sp., East Point, Darwin, NT, slide ANIC 8198. 932, 933, female genitalia. 932, A. trichograpta, Mt Webb Nat. Park, Q, slide ANIC 1297. 933, A. sp., Richmond R., NSW, slide T2300. 934, male 8th segment, A. trichograpta. 935, head, A. sp., male.
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from Richmond R. in New South Wales to Toowoomba in Queensland.
Biology The type species A. balanoptycha has been reared from larvae feeding on leaves and galls on leaves of Pongamia glabra in India (Fletcher [1921]) and from larvae folding the leaves of Derris elliptica in the Malay Peninsula (Fletcher 1932). Fletcher ([1921], 1932) described and figured the biology of A. conciliata (Meyrick) in India, feeding on flowers, leaves and shoots of Butea frondosa, sometimes with the bases of the flowers and the shoot partially bored. Meyrick (1927a) recorded A. phaulomorpha (Meyrick) bred from leaves of Sesbania aegyptiaca. Clarke (1976) reported Pongamia glabra and Derris elliptica as foodplants for A. defensa (Meyrick). Komai reared two species of Acanthoclita from Desmodium spp. in the Philippines and one species from Derris trifoliata in the Ryukyu Is. All the hostplants reported for Acanthoclita belong to the Fabaceae and the larvae are primarily leafrollers though A. conciliata is reported to bore into the fleshy parts of flowers and into shoots. No biological information is available on the Australian species, but Pongamia, Desmodium and Derris all occur in Australia. Males of A. phaulomorpha were captured by field traps baited with a 1: 1 mixture of (Z)–8-Dodecen1-ol acetate and (E)-8-Dodecen-1-ol acetate in Senegal (Komai 1999).
Remarks Diakonoff (1982) figured head, wing venation and genitalia of both sexes with his original description of the genus. Clarke (1958) provided photos of adult and genitalia of the types of several Meyrick species. Though the majority of Acanthoclita species are easily recognised and characterised by several obvious apomorphies such as the wing shape and venation, a distinctive forewing pattern, a hindwing with a translucent base and modified anal region, the fusion between gnathos and tegumen and the characteristic valva with one or two isolated spines at the base of the cucullus, there is no convincing synapomorphy apart from the shape of the cucullus to link the most plesiomorphic, Matsumuraeses-like Australian species with all the other species. Matsumuraeses clearly is the sister group of Acanthoclita, with similar labial palpi, a similar sterigma in the more plesiomorphic taxa and a
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Matsumuraeses-type wing shape, venation and forewing pattern in the least derived Australian Acanthoclita (Fig. 929). A patch of dense, modified scales usually on both sides of the hindwing between CuP and 1A+2A present in Acanthoclita is shared with almost all Matsumuraeses species. The following non-Australian species are here transferred to Acanthoclita: argyrotorna (Diakonoff, 1984a), Entomol. Basil. 9: 377 (Grapholita (Aspila)). [Sumba.] Comb. nov. bidenticulana (Bradley, 1957), Nat. Hist. Rennell Island, British Solomon Islands 2(19): 93 (Eucosma). [Rennell I.] Comb. nov. conciliata (Meyrick, 1920), Exot. Microlepid. 2: 245 (Eucosma). [India.] Comb. nov. hilarocrossa (Meyrick, 1931), in de Joannis, Ann. Soc. Entomol. Fr. 98: 716 (Eucosma). [Vietnam.] Comb. nov.
Constituent species Apart from the four species referred to Acanthoclita above, we are aware of six non-Australian species previously included in Acanthoclita: A. phaulomorpha (Meyrick) from Egypt, A. balanoptycha (Meyrick) from India, A. defensa (Meyrick) from Fiji, A. iridorphna (Meyrick) from Taiwan and A. acrocroca Diakonoff from Sri Lanka, with the position of A. balia Diakonoff from Sri Lanka (Diakonoff 1982) to be confirmed. At least six unidentified and one named species are known from Australia. trichograpta (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 275 (Argyroploce). [Qld: Cairns, Kuranda.] taophanes (Meyrick, 1921), Exot. Microlepid. 2: 447 (Eucosma). [Qld: Cairns.]
Pammenopsis Kuznetzov (Figs 117, 936–944) Pammenopsis Kuznetzov, 2003, Entomol. Obozr. 82: 740. Type species: Eucelis critica Meyrick, 1905, J. Bombay Nat. Hist. Soc. 16: 587, by original designation. [India.]
Diagnosis Medium-sized, rather narrow-winged, pale greyish ochreous (P. barbata, sp. nov.) or dark grey-brown
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moths of Bactra-like appearance with forewing markings mainly of very fine, dark, largely longitudinal lines, with dorsal half paler. Males of Pammenopsis are immediately recognisable by the conspicuous fringe of very long black hair scales along the anal margin of the hindwing and the band of modified scales on the lower surface behind M1. Pammenopsis has large, porrect and strongly sinuate labial palpi, with the second segment distally widened by long scales, often nearly concealing the long, slender distal segment. Hindwing with M3 and CuA1 long-stalked and 3A a trace only in female and absent in male, frenulum with three bristles in female. Male with a long scale pencil from lower angle of tegulae, two transverse dorsal bands of modified scales on T4 and T5 and two deeply invaginated, lateral bundles of long scales between A6 and A7, which are diagnostic for Pammenopsis. The male genitalia have a simple, band-like tegumen, hardly widened across the faintly pointed apex, long and slender valvae with a shallow (P. barbata, sp. nov.) to very deep ventral emargination near middle, a long and distally reflexed caulis and a sharply bent aedeagus widened in basal part with a sheaf of sinuate, lanceolate cornuti. The modified, sclerotised and erect ovipositor lobes in the female genitalia are diagnostic for Pammenopsis, the ductus and corpus bursae are membranous, with two large, hornshaped signa, flattened except for hollow base.
Description Adults (Fig. 936) Wingspan. 11.5–15.0 mm; medium-sized. Head (Figs 938). Ocellus well developed, chaetosema present. Proboscis short, about length of labial palpus. Labial palpus large (1.8× diameter of eye), porrect and strongly sinuate in both sexes, with second segment sinuate and greatly widened in distal half with very long, loose scales along dorsal and especially ventral margin, often nearly concealing the very long, slender, distal segment. Antenna to middle of forewing, no cilia visible in male. Thorax. Without crest, but scale tuft from anterior corner of tegulae as long as fore coxa in male, shorter in females. Legs unmodified. Wings (Figs 117, 936, 937, 939). Forewing index: 0.33–0.35; forewing long and narrow, nearly parallelsided with acute apex and oblique termen; cell 0.66 length of wing, with chorda and M-stem well developed but closing vein weak; all veins present and separate beyond cell, R1 from just beyond middle of cell, R3 from somewhat closer to R4 than
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to R2, base of M3 from slightly closer to M2 than to CuA1, M3 and CuA1 distally both strongly curved towards apex, CuA2 from 36: cell; CuP present at margin; forewing pattern of Bactra-like appearance, longitudinally finely striate with brownish grey on creamy or ochreous ground colour, with fine greybrown strigulae along costa, giving rise to several outwardly oblique silvery lines before apex, with some silvery marks parallel along dorsal half of termen, preceded by a few irregular, longitudinal, black dashes; dorsal half of wing paler, with a brownish suffusion on lower angle of cell and a patch of dark brown scales on dorsum below fold before middle. Hindwing with weakly developed cubital pecten and axillary tuft, in male with a fringe of very long, conspicuous, stiff, black hairscales from anal margin (Fig. 937) and with a band of modified, spatulate scales on the lower surface behind M1 (Fig. 939), frenulum in female of three bristles; cell short (about 0.38 length of wing in male, about 0. 45 in female); Rs and M1 closely approximated and parallel at base, M2 widely distant from base of longstalked M3 and CuA1, CuA2 from 6M cell; basal loop of 1A+2A weak; 3A a weak trace in female, not discernible in male; pale brownish grey, with pale fringe and the very long, modified blackish scales mentioned above along anal margin. Pregenital abdomen (Figs 942, 944). Sternum 2 with well-developed anterolateral processes and long, slender, straight apodemes. In male two large, deeply invaginated bundles of modified scales laterally between 6th and 7th segment, and two bands of modified, broad scales anteriorly on T4 and T5. Coremata reduced: eighth sternite a semioval plate with bilobed hindmargin (P. barbata, sp. nov.) or a trapezoidal plate (P. critica); intersegmental ventral sclerite and scale tufts not developed. Male genitalia (Figs 940, 941). Vinculum a moderate band. Tegumen bent at a right angle at base and articulating with anterior edge of vinculum, high and narrow, a narrow band sometimes with a slightly widened and weakly pointed apex, with a dorsally projecting membranous lobe with long, thin scales. Gnathos two near-membranous, curved, long bands, medially rising. Valva long and slender, with a longitudinal ridge or groove in basal half of outer surface, with ventral margin weakly to strongly rounded in basal half, with a moderate to very deep emargination in middle, cucullus ovate to subtriangular, with relatively few but very strong bristles along ventral margin on inner surface. Caulis long and deeply reflexed below aedeagus; aedeagus long, sharply
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bent, basal third with at least twice the diameter of the lightly curved distal portion beyond bend; vesica with a sheaf of sinuate, lanceolate cornuti. Female genitalia (Fig. 943). Ovipositor lobes modified, erect and sclerotised, with curved distal margins approximated, smooth and with only few large bristles on outer surface. Apophyses moderately long, slender; anterior margin of T8 conspicuously bilobed. Ostium lightly invaginated behind weakly concave hindmargin of S7, and with two patches of large, modified scales in membrane posterior to ostium. Transition from ductus to corpus bursae gradual, ductus bursae short and membranous, without colliculum, smooth and narrow to origin of ductus seminalis, followed by a portion of equal length widening gradually towards
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corpus bursae proper and finely dimpled as corpus bursa itself; corpus bursa teardrop-shaped to ovate, with two large, curved, distally flattened, hornshaped signa with hollow base.
Distribution In Australia known from widely scattered localities across the tropical north, from the Kimberley in Western Australia, the Darwin and Borroloola areas in the Northern Territory, and from the islands of the Torres Strait to Rockhampton in Queensland. Pammenopsis critica has been reported from India, Sri Lanka, Bangladesh, Thailand, and Vietnam (Diakonoff 1982; Lateef and Reed 1990; Kuznetzov 2003).
Figs 936–939. Pammenopsis barbata, adult, hindwing and head. 936, adult , paratype, Sue (Warraber) I., Q. 937, fringe along hindwing anal margin, male. 938, head, male. 939, scale fringe along M1, male hindwing. Scale = 2 mm.
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Biology Fletcher ([1921]) provided detailed biological information on P. critica, which he described as a minor pest on Cajanus indicus (Pigeon Pea) (Fabaceae) in India, with ‘the larvae rolling and webbing together the top-leaves of the foodplant and also eating into flower-buds and boring into
pods and devouring the seeds’. Lateef and Reed (1990) reviewed the biology of this species as a pest on Pigeon Pea. The species is also reported to feed on pods of Crotalaria juncea (Fabaceae) (Fletcher 1932). Nothing is known about the biology of P. barbata, sp. nov., and the very diverse localities give no indication of a possible host.
Figs 940–944. Pammenopsis barbata, genitalia and abdomen. 940, 941, male genitalia and aedeagus, holotype, slide T2314. 942, male abdomen, segments 6 and 7 with scale bundles. 943, female genitalia, nr Borroloola, NT, slide T2393. 944, male abdomen (ap, anterolateral processes).
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Remarks Clarke (1958) provided photos of adults and genitalia of both sexes of types of P. critica and its synonyms, and Diakonoff (1982) referred the species to Pammene Hübner and figured the male genitalia and abdomen from Sri Lankan material. Kuznetzov (2003) provided figures of wing venation, genitalia of both sexes, and abdomen of P. critica from Vietnamese material, and mentioned that Pammenopsis is similar to Pammene in abdominal androconial organs although their configurations are different, and similar to Matsumuraeses Issiki in copulatory organs of both sexes and in host plants. Pammenopsis is very distinct superficially and also characterised by a considerable number of apomorphies. M3 and CuA1 in the hindwing are long-stalked and 3A is very weak in the female and absent in the male. The male has several quite unique secondary sexual structures: on the hindwing a fringe of very long, stiff, blackish hairscales along anal margin and a band of modified, spatulate scales on the lower surface behind M1, and on the abdomen two transverse bands of modified scales and two large, lateral, deeply invaginated bundles of eversible scales. The modified ovipositor lobes in the females are highly unusual for a tortricid.
Constituent species A single named species, P. critica with three junior synonyms is known from outside Australia, and one species occurs in Australia, described below: barbata, sp. nov.
Pammenopsis barbata Komai & Horak, sp. nov. (Figs 117, 936–944)
Material examined Holotype. , ‘Carnarvon Gorge, Q[ueensland], 31 Mar. 1957, I.F.B. Common’, ‘Genitalia slide T2314’ (ANIC). Paratypes. Western Australia: 1 , 14°39´S 126°57´E, Drysdale R., 18–21.viii.1975, I. F. B. C. & M. S. U.; 1 , Mt Barnett Stn, 19.vi.1980, L. & V. J. Robinson. Northern Territory: 1, 13°12´S 130°43´E, Tolmer Falls Turn Off, 10.iv.1991, M. Horak & A. Wells; 1, 12°42´S 131°59´E, Berry Springs, 8.iv.1991, M. Horak & A. Wells; 1, 16°27´S 136°05´E, Mc Arthur R., 48 km SW by S Borroloola, 29.x.1975, E. D. E.; 1, 16°10´S 136°15´E,
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Goose Lagoon, 11 km SE by S Borroloola, 31.x.1975, E. D. E. Queensland: 3 , 10°12´S 142°49´E; Sue (Warraber) I., 6–10.xii.1977, E. D. E; 1 , 17°16´S 145°54´E, Base Cableway Mt Bellenden-Ker, 80 m, 19.x.1981. E. D. E.; 1 , same label data as holotype; 1 , Rockhampton, 3.ix.1954, I. F. B. C. (ANIC, MQ, NTM, BMNH).
Diagnosis Pammenopsis barbata has the same wing pattern as P. critica, but the Australian species is lighter in colour. In the Australian fauna, the male is unmistakable with the conspicuous black scale fringe along the anal margin of the hindwing, and the female by the combination of its wing pattern and the modified ovipositor lobes. The male genitalia of P. barbata are characterised by a weakly constricted valva with a much wider cucullus ‘neck’ than P. critica, and a weakly rounded rather than angled and projecting sacculus.
Description Male (Fig. 936) Wingspan 11.5–14.5 mm. Head (Fig. 938), thorax and tegulae pale yellowish ochreous, inner surface of palpus whitish. Forewing ground colour creamy to pale greyish ochreous, longitudinally finely striate with pale greyish brown, especially in distal area beyond cell; with fine, long, brown costal strigulae along entire costa, distally giving rise to strongly oblique silvery lines running to below apex and interspersed with some white scales just before apex; with a series of silver spots parallel along dorsal half of termen preceded by a few irregular, longitudinal black dashes; dorsal half of wing paler with white scales between black dashes, a brownish suffusion on lower angle of cell, a dark brown spot on dorsum near fold below middle and often further dark brown scales along fold towards base of wing; fringe whitish, grey-brown just below apex. Hindwing pale greyish ochreous, with slightly darker marks in apex; fringe whitish ochreous, ochreous in apex and with a conspicuous fringe of very long blackish scales along anal margin (Fig. 937). Abdomen pale yellowish ochreous, with a transverse band of persistent blackish scales along anterior margin of T4 and T5 and a deeply invaginated large bundle of yellowish scales laterally between 6th and 7th segments (Figs 942, 944); hindmargin of eighth sternite bilobed. Male genitalia (Figs 940, 941). Tegumen a narrow band with a slightly widened, pointed apex giving rise to a triangular membranous apical lobe
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set with narrow scales. Valva long and slender, sacculus only weakly projecting and ventral emargination below cucullus shallow; cucullus roughly semicircular, with a band of strong spines along ventrodistal margin. Aedeagus sinuate, bent at nearly right angle beyond bulbous base and tapering tip lightly upcurved; vesica with a group of lanceolate cornuti. Female Wingspan 12–14 mm. Wing pattern as for male, but hindwing slightly darker and without the fringe of black scales along anal margin. Female genitalia (Fig. 943). Ovipositor lobes modified, erect, sclerotised, smooth with few bristles, with curved distal margins approximated. Sterigma two patches of large, modified scales posterior to ostium. Ductus bursae membranous, posterior half narrow, smooth, anterior half below inception of ductus seminalis finely dimpled and gradually widening towards corpus bursae. Signa two large curved rugose horns with sharp points, with round hollow base and apically flattened.
Distribution and biology Pammenopsis barbata is known across the top end of Australia, from northern Western Australia across the Northern Territory to northern Queensland and south to Central Queensland at Rockhampton. It has been collected in tropical monsoon forest and tropical eucalypt savannah. No host information is available for P. barbata, but a similarly modified ovipositor suggests a biology similar to that of P. critica.
Grapholita Treitschke (Figs 114, 945–955) The genus Grapholita comprises at least two possible monophyletic groups, the subgenera Grapholita with the majority of Australian species and Aspila Stephens that includes Grapholita (Aspila) molesta (Busck). The monophyly of the genus, however, has not been well established. Diagnostic characters used by previous authors to unite them (e.g. presence of coremata) are plesiomorphic. Komai (1999) tentatively proposed a synapomorphic character linking the two subgenera: coremata with eighth sternite with a pair of short projections laterally.
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Subgenus Grapholita Treitschke (Figs 114, 945–949) Grapholita Treitschke, 1829, in Ochsenheimer, Schmett. Eur. 7: 232. Type species: Pyralis dorsana Fabricius sensu Treitschke, 1829 [ = Tortrix lunulana [Denis & Schiffermüller], 1775, Ankündung syst. Werkes Schmett. Wienergegend: 127], by subsequent designation by Walsingham, 1895, Trans. Entomol. Soc. Lond. 1895: 517. [Europe.] Grapholitha Treitschke, 1830, in Ochsenheimer, Schmett. Eur. 8: 203 (an unjustified emendation of Grapholita Treitschke, 1929). Euspila Stephens, 1834, Illust. Br. Entomol. (Haustellata) 4: 103. Type species: Tinea compositella Fabricius, 1775, Syst. Entomol.: 663, by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 27, 57. [Europe.] Ephippiphora Duponchel, 1834, Ann. Soc. Entomol. Fr. 3: 446. Type species: Pyralis dorsana Fabricius sensu Duponchel, 1834, [ = Phalaena jungiella Clerck, 1759, Icon. Insect. rariorum 1: pl. 12, fig. 9], by monotypy. [Europe.] Stigmonota Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 182. Type species: Phalaena jungiella Clerck, 1759, Icon. Insect. rariorum 1: pl. 12, fig. 9, by subsequent designation by Desmarest, 1857, in Chenu, Encycl. Hist. Nat. (Papillons nocturnes): 224 (as jungiana, an unjustified emendation by Guenée, 1854, ibid. (2)3: 183). [Europe.] Endopisa Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 182. Type species: Grapholitha nebritana Treitschke, 1830, in Ochsenheimer, Schmett. Eur. 8: 208, by subsequent designation by Desmarest, 1857, in Chenu, Encycl. Hist. Nat. (Papillons nocturnes): 224 (misspelled as Eudopisa Guenée). [Europe.] Ebisma Walker, 1866, List Specimens Lepid. Insects Colln Br. Mus. 35: 1803. Type species: Ebisma seclusana Walker, 1866, ibid. 35: 1804, by monotypy. [New Guinea.]
Diagnosis Small moths with grapholitine wing pattern with variably developed dorsal blotch consisting of one, two, or four whitish lines from middle of dorsum, rarely without. Grapholita is characterised by rather
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short, upcurved labial palpi with long, spreading scales along ventral margin, a hindwing without sexual dimorphism in venation and with M1, M2 and M3 widely separate and parallel. The male genitalia are characterised by a weakly sclerotised tegumen except for its anterior margin, absence of uncus and socius, the valva constricted near middle and a rounded or ovate cucullus. The female genitalia are characterised by a rather short and wide ductus bursae and the absence of a diverticulum to the corpus bursae (Australian species). Coremata usually well developed, with paired tufts of filiform and sometimes also lanceolate scales, with intersegmental ventral sclerite usually with a pair of long levers, and with a small crescent-shaped eighth sternite with a pair of short projections laterally.
Description Adults (Fig. 945) Wingspan. 7.5–12.5 mm (Australian species); small. Head (Fig. 949). Ocellus large, chaetosema distinct. Proboscis rather short, not much longer than labial palpus. Labial palpus ascending, rather short (1.6–2.1× diameter of eye); second segment smooth above but with long, spreading scales beneath; terminal segment slender, with appressed scales, hardly bent forward; antenna rather long, in larger species exceeding 56O length of forewing; without scales along a narrow anterior band, cilia minute. Thorax. Without crest. Legs unmodified. Wings (Figs 114, 945). Forewing index: 0.38– 0.46 (Australian species); forewing subtriangular, costa curved, apex rounded, termen slightly notched below apex; chorda from between R1 and R2 to R5 or to between R4 and R5, M-stem to between M2 and M3 (chorda and M-stem both absent in G. tornosticha); all veins present and separate beyond cell; R1 arising before middle of cell; R2 nearer to R3 than R1; R4 to costa; R5 to termen; M1 and M2 subparallel; CuA1 from posterior angle of cell; CuA2 from about 36: of cell; CuP evident at margin; grey or greyish brown to dark brown, with five (sometimes six) costal strigulae beyond middle of costa, with variably defined dorsal blotch consisting of one, two, or four whitish lines from middle of dorsum or rarely without, with typical ocelloid patch of a pair of vertical leaden lines enclosing some black dots or dashes or ocelloid patch only indicated by a vertical leaden line above tornus (compositella-group), and
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with whitish stria on termen below apex. Hindwing subtrapezoidal, costa curved, apex rounded, with well-developed cubital pecten; frenulum of female consisting of three bristles; Rs and M1 approximated towards base; M1, M2 and M3 parallel; M3 and CuA1 connate or very shortstalked; CuA2 arising beyond middle of cell; CuP present at margin; grey to greyish brown, rarely white or subhyaline. Pregenital abdomen (Fig. 948). Sternum 2 with well-developed anterolateral processes or sometimes anterolateral processes reduced (the conficitana-group as defined below). Coremata: eighth sternite a small crescentic plate with a pair of short projections laterally; intersegmental ventral sclerite with a pair of long levers associated with tufts of long filiform scales and sometimes also broad lanceolate scales (eighth sternite and intersegmental ventral sclerite not fused, see Komai (1999: 24)); T8 in male a sclerotised hood-like plate with a more strongly sclerotised anterior margin. T5 and T6 in male with patch of modified scales in the conficitana-group; the non-Australian scintillana-group with anterior corners of T8 with paired pockets containing tufts of filiform scales (Komai 1999: 86). Male genitalia (Fig. 946). Vinculum U- or V-shaped, wider ventrally. Tegumen subtriangular to ovate, hood-shaped, weakly sclerotised except for anterior margin, sometimes with hairs laterally, or the top emarginated in lateral view. Uncus not differentiated. Anal tube membranous or with sclerotised subscaphium. Gnathos variable, from weakly sclerotised horizontal arms fused with tegumen to medially fused and expanded to a broad lobe (G. zapyrana), to a pair of wide sclerotised plates. Valva constricted near middle; basal excavation large; costal process small; sacculus rounded or angular; its outer surface sometimes with a patch of scales (easily removable). Juxta triangular or trapezoidal, wide and low, caulis rather long and wide, band-shaped, in the same plane as the aedeagus. Aedeagus moderately long, often with bulbous base, gradually narrowed or abruptly constricted before middle; vesica with a band of cornuti, numerous deciduous ones near base and a group of fixed, spiniform ones more distally. Female genitalia (Fig. 947). Ovipositor lobes subreniform. Sternum 7 usually weakly sclerotised, rounded-concave in posterior margin. Sterigma an often weakly sclerotised plate with a narrow, protruding ventral lip. Ductus bursae short and usually rather wide, widening anteriorly; distal end
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often with a cup- or funnel-shaped sclerotisation, followed by a small, narrow colliculum, anterior 36: variably sclerotised, either ill-defined areas or a transverse band or a spiny patch; ductus seminalis usually wide, arising from about middle of ductus bursae, rarely from neck of corpus bursae; bulla seminalis present, sometimes almost directly connected with corpus bursae (G. tetrazancla); corpus bursae with two, sometimes very long, horn-shaped signa.
Distribution Two species-groups occur in Australia (see Remarks). The compositella-group, known from the Holarctic and the Oriental regions, is restricted to northernmost Western Australia, the Northern Territory and to north-eastern Queensland. The conficitana-group, endemic to Australia, is known from all States and Territories.
There are two species-groups in Australia: the compositella-group, one of eight species-groups that Komai (1999) recognised in the Palaearctic region, and the conficitana-group that is endemic to Australia.
The compositella-group Antenna long, 36:– 6M length of forewing; forewing usually with well-defined dorsal blotch or rarely without (G. caecana Schläger) and with ocelloid patch indicated by a single vertical leaden line; S2 with well-developed anterolateral processes; tegumen sometimes with hairs laterally; valva usually shallowly constricted, sacculus rarely angulate and never with an acute angle, cucullus elliptic to ovate, aedeagus curved or straight, gradually tapering distally; vesica with deciduous needle-like and short fixed spiniform cornuti; anal tube membranous; T5 and T6 of male without patch of modified scales; ductus bursa posteriorly with funnel- or cup-shaped sclerotisation.
Biology This group is mainly associated with Fabaceae in the Palaearctic Region (Komai 1999). Only scant biological information is available on the Australian species, but all known larval host plants are Fabaceae. Grapholita zapyrana has been reared from larvae feeding on seed of native Glycine spp. grown in a shade house at CSIRO Black Mt, Canberra, and Meyrick (1881) reported the moths flying briskly in the hot sunshine round the purple blossoms of Hardenbergia violacea (as H. monophylla). Turner (1925) noted the larvae of G. tetrazancla tunnelling in leaves of ‘seven-year bean [Phaseolus lunatus]’ and Komai has reared this species from leaves of Alysicarpus vaginalis in Darwin, Northern Territory. Meyrick reported the moths of G. floricolana frequenting the flowers of Bursaria spinosa (Pittosporaceae), sitting on them in the hot sunshine. Sex pheromone components of Grapholita have been reviewed by Komai (1999).
Remarks Comprehensive reviews of Grapholita have been given by Heinrich (1926), Obraztsov (1959, as Grapholitha), Danilevsky and Kuznetzov (1968), Razowski (1989), and Komai (1999). The monophyly of the subgenus Grapholita has not been well supported. Komai (1999) tentatively lists two possible autapomorphies for the subgenus: pheromone component and setal pattern of larvae.
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The conficitana-group Antenna rather short, 56O length of forewing; forewing with ill-defined dorsal blotch and with typical ocelloid patch indicated by a pair of vertical leaden lines enclosing some black dots or dashes; S2 with anterolateral processes reduced; valva deeply constricted, sacculus angled, sometimes acute, cucullus ovate to subcircular; anal tube with sclerotised subscaphium; aedeagus curved, bulbous in basal 56: to 369 then abruptly narrowed distally; vesica with deciduous needle-like and short fixed spiniform cornuti, T5 and T6 of male with patch of modified scales; sterigma a rectangular or ovate plate, ductus bursae rather wide, gradually widening anteriorly, largely membranous.
Constitutent species The subgenus Grapholita has some 100 described species world wide (Brown 2005). 1. The conficitana-group antitheta (Meyrick, 1911b), Proc. Linn. Soc. NSW 36: 288 (Laspeyresia). [WA: Albany.] conficitana (Walker, 1863), List Specimens Lepid. Insects Colln Br. Mus. 28: 412 (Carpocapsa). [Qld: Moreton Bay.] amphitorna (Turner, 1916), Trans. R. Soc. S. Aust. 40: 534 (Laspeyresia). [Qld: Brisbane.] parvisignana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 654 (Stigmonota). [NSW: Sydney.]
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argyrocyrta (Turner, 1939), Pap. Proc. R. Soc. Tasm. 1938: 80 (Laspeyresia). [Tas.: Port Arthur on the Tasman Peninsula.] argyroela (Turner, 1946), Trans. R. Soc. S. Aust. 70: 219 (Laspeyresia). [Qld:
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Coolangatta [as Tweed Heads]; SA: Flinders I.] iridescens (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 655 (Stigmonota). [NSW: Parramatta and Murrurundi.]
Figs 945–949. Grapholita (Grapholita), adult genitalia, abdomen and head. 945, G. zapyrana, , Black Mt, ACT. 946, G. zapyrana, male genitalia, Brisbane, Q, slide ANIC 3142. 947, G. parvisignana, female genitalia, Richmond R., NSW, slide ANIC 8173. 948, G. zapyrana, 8th segment with coremata, male (iv, intersegemental ventral sclerite; le, levers for scale tufts of coremata. 949, head, G. zapyrana, male. Scale = 2 mm.
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zapyrana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 653 (Stigmonota). [Qld: Brisbane, Helidon, and Toowoomba; NSW: Sydney, Parramatta, and Bulli; Vic.: Melbourne.] floricolana (Meyrick, 1881), Proc. Linn. Soc. NSW 6: 656 (Stigmonota). [NSW: Parramatta, Bowenfels.] 2. The compositella-group tetrazancla (Turner, 1925), Trans. R. Soc. S. Aust. 49: 60 (Laspeyresia). [NT: Darwin and Melville I.] tornosticha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 220 (Laspeyresia). [Qld: Cape York.]
Subgenus Aspila Stephens (Figs 950–955) Aspila Stephens, 1834, Illust. Br. Entomol. (Haustellata) 4: 104. Type species: Phalaena lediana Linnaeus sensu Haworth, 1811 [ = Coccyx janthinana Duponchel, 1835, in Godart and Duponchel, 1835, Hist. Nat. Lepid. Papillons Fr. 9: 245], by subsequent designation by Fernald, 1908, Genera Tortricidae Types: 27, 58. [France.] Opadia Guenée, 1845, Ann. Soc. Entomol. Fr. (2)3: 182. Type species: Grapholitha funebrana Treitschke, 1835, in Ochsenheimer, Schmett. Eur. 10: 3: 116, by monotypy. [Europe.] Coptoloma Lederer, 1859, Wien Entomol. Montschr. 3: 124, 370. Type species: Coccyx janthinana Duponchel, 1835, in Godart and Duponchel, Hist. Nat. Lepid. Papillons Fr. 9: 245, by monotypy. [France.]
Diagnosis Dark grey-brown to blackish, small to mediumsized moths with little apparent grapholitine wing pattern with or without a barely discernible whitish dorsal blotch. Hindwing sometimes sexually dimorphic in shape and in male sometimes with fringe on parts of termen consisting of modified scales. Venation similar to that of Grapholita s. str. Coremata with a pair of tufts of filiform or lanceolate scales, with eighth sternite reduced or with both eighth sternite and intersegmental ventral sclerite reduced. The male genitalia are characterised by stout and pincers-shaped valvae in dorsal view, often with reticulate wrinkles on outer surface, a stout, short, basally swollen aedeagus with bulbus ejaculatorius dorso-proximally. The female are characterised by a heavily sclerotised
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sterigma, a very short ductus bursae without a typical colliculum but with a dark polygonal, ovate or conical sclerite, and a usually heavily sclerotised seventh sternite with pointed posterior corners.
Description Adults (Figs 950, 951) Wingspan. 9.5–16.0 mm (Australian species: 9.5–13.0 mm); small to medium-sized. Head (Fig. 952). Ocellus large, chaetosema very distinct. Proboscis rather short, longer than labial palpus. Labial palpus ascending, short (1.8–1.9× diameter of eye); second segment rough-scaled beneath; terminal segment short, about 56M length of second, with appressed scales, apex blunt. Antenna relatively short, less than 56O length of forewing; without scales along anterior band, cilia small. Thorax. Without crest. Legs unmodified. Wings (Figs 950, 951). Forewing index: 0.44– 0.47 (Australian species); forewing subtriangular, costa curved, apex rounded, termen slightly notched below apex or gently arched; M-stem to M2, chorda from between R1 and R2 to R5 or between R4 and R5; all veins present and separate beyond cell; R1 arising before middle of cell; R2 nearer to R3 than R1; R4 to costa; R5 to termen; M1 and M2 subparallel; CuA1 from posterior angle of discal cell; CuA2 from about 36: of cell; CuP evident at wing margin; variously dark greybrown, with five (sometimes six) costal strigulae beyond middle of costa, with or without illdefined dorsal blotch consisting of four whitish, irregular, barely discernible lines from middle of dorsum, or without dorsal blotch; ocelloid patch consisting of a pair of vertical leaden lines enclosing some blackish dots; whitish stria below apex usually absent. Hindwing sometimes sexually dimorphic in shape: in male trapezoidal or triangular and with a wide band of modified scales above along margin, with modified fringe (consisting of filiform scales or scales with triangular apex) on margin between CuP and 1A+2A; in female hindwing normal, semiovate with normal scaling and fringe; well-developed cubital pecten present; frenulum of female consisting of three bristles; Rs and M1 approximated towards base, M1, M2 and M3 parallel; M3 and CuA1 connate, CuA2 arising from 169 of discal cell, CuP present at wing margin, basal loop of 1A+2A only weakly indicated, 3A present; grey to greyish brown. Pregenital abdomen (Fig. 954). Sternum 2 with well-developed anterolateral processes. Coremata:
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eighth sternite a large, sclerotised semiovate plate with a pair of short projections laterally; intersegmental ventral sclerite with a pair of levers distally associated with tufts of long, filiform scales, or eighth sternite a less sclerotised rectangular plate often concave on posterior margin and a less sclerotised intersegmental ventral sclerite with short levers associated with tufts of lanceolate scales; T8 a hood-like plate. Male genitalia (Fig. 953). Vinculum U-shaped, wider ventrally. Tegumen subtriangular, hoodshaped, weakly sclerotised except for anterior margin, top sometimes emarginated in lateral view. Uncus not differentiated. Gnathos a pair of sclerotised horizontal arms completely fused with tegumen. Valva usually stout, heavily sclerotised, in
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dorsal view pincers-shaped (valva strongly bent at neck), the outer surface often with reticulate wrinkles; basal excavation well-defined; costal process small. Juxta triangular or trapezoidal, caulis a wide, rather long band. Aedeagus stout, usually short, swollen basally, often with a concavity dorso-posteriorly, connecting with bulbus ejaculatorius dorso-anteriorly; vesica with or without slender, deciduous cornuti, usually 1–2, rarely many (G. (A.) molesta). Female genitalia (Fig. 955). Ovipositor lobes subreniform. Sternum 7 usually heavily sclerotised, hindmargin produced at both corners. Sterigma a heavily sclerotised structure in the form of a ring or a transverse rectangle. Ductus bursae very short and wide, apparently with colliculum reduced but
Figs 950–955. Grapholita (Aspila) molesta, adults, head, genitalia and abdomen. 950, 951, adults, Canberra, ACT. 950, . 951, . 952, head, male. 953, 954, male, Canberra, ACT, slide ANIC 8211. 953, genitalia. 954, 8th segment. 955, female genitalia, Canberra, ACT, slide ANIC 8212. Scale = 2 mm.
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with a polygonal, ovate, or conical, heavily sclerotised, concave sclerite; ductus seminalis narrow or wide, arising laterally anteriorly to the sclerotised ring; bulla seminalis sometimes large and with a broad connection to bursa copulatrix.
shaped in dorsal view, and 4, the reduced coremata. Grapholita (A.) dysaethria, described from Sri Lanka (Diakonoff 1982) and recorded from the Philippines (Komai 1999), is known from one Australian specimen, a male from Mission Beach, Queensland.
Distribution Holarctic, Oriental and Australian regions. In Australia, Aspila is known only from Mission Beach in northern Queensland except for G. (A.) molesta, the Oriental fruit moth, accidentally introduced into Australia in the 1930s and now occurring in Tasmania, South Australia, New South Wales, the Australian Capital Territory, and Queensland.
Biology Recorded hosts are almost exclusively rosaceous plants, with the larvae feeding as shoot- or fruitborers, but no biological information is available on the Australian species apart from G. (A.) molesta. Heinrich (1926) reported larvae of G. (A.) prunivora (Walsh) feeding on ‘black knot’ fungus and aphid galls on oak and elm as well as rosaceous plants such apple, plum, peach and cherry. Rothschild and Vickers (1991) reported G. (A.) molesta has been found infesting shoots of Eugenia myrianthus (Myrtaceae) in Argentina. Park (1983) recorded Diospyros kaki (Ebenaceae) in addition to Rosaceae as host plants for G. (A.) molesta. In Australia, G. (A.) molesta, the Oriental fruit moth, is a serious pest of stone fruit, in particular peach, and has recently also become a pest in adjacent pear orchards (Sexton and Il’ichev 2000). Mating disruption by pheromone, developed at CSIRO Entomology (Vickers et al. 1985), is successfully used to control G. (A.) molesta in Australia. Sex pheromone components of Aspila have been reviewed by Komai (1999).
Remarks The Palaearctic species of Aspila were revised by Danilevsky and Kuznetzov (1968). Komai (1999) reviewed the subgenus and discussed its monophyly, that is supported by: 1, a ductus bursae with a polygonal, ovate, or conical, heavily sclerotised, concave sclerite; 2, the aedeagus connecting dorso-anteriorly with the bulbus ejaculatorius; 3, the stout valva that is pincers-
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Constituent species The subgenus Aspila has 17 described species, of which two occur in Australia. dysaethria Diakonoff, 1982, Zool. Verh. (Leiden) 193: 16 (Grapholita). [Sri Lanka.] molesta (Busck 1916), J. Agric. Res. 7: 373 (Laspeyresia). [N. America.]
Microsarotis Diakonoff (Figs 116, 956–963) Microsarotis Diakonoff, 1982, Zool. Verh. (Leiden) 193: 10. Type species: Laspeyresia palamedes Meyrick, 1916, Exot. Microlepid 1: 564, by original designation. [India.]
Diagnosis Species of Microsarotis are small, rather broadwinged dark grey moths with, at higher magnification, iridescent scales with light blue to golden gloss on body and forewings and with a conspicuous grapholitine pattern on a grey background, with large black and white costal strigulae, a dorsal blotch of bent, parallel, silvery lines framed by black, the ocelloid patch a broad leaden band without black dashes and the termen without notch. Microsarotis has a rather small head with smallish, upcurved, weakly sinuate, yellowish white labial palpi with a crescentic second segment. The venation is characterised by a forewing with chorda and M-stem well developed, all veins present and distant from each other at base, and a hindwing with Sc+R1 and Rs close to end of cell, Rs and M1 distant from base, and M2 widely distant from stalk of M3 and CuA1. The male has unique scent organs consisting of two large scale tufts deeply invaginated between a curved, very narrow T8 and a modified T7 with scaled pockets on outer surface and a small bipronged median sclerite on inner surface. The male genitalia are diagnostic with a narrow, distally tapering tegumen topped by
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a large, round or transversely ovate disc, strongly swollen, short, ovate to triangular valvae with small basal excavation, a strongly convex outer surface and the inner surface with a dense patch of strong bristles and a rounded-triangular naked apex except for deciduous, flattened bristles, the aedeagus tapering toward apex with vesica irregularly sclerotised in proximal half. The female genitalia also are diagnostic with the cup to funnelshaped sterigma fused with S7 beneath its posterior apex, the ductus bursae bipartite, narrow posteriorly and widened anteriorly, with two large, confluent patches of dorsal sclerotisation below ductus seminalis each with a short lateral membranous projection from outer wall, with a longitudinal line of single spinules between them, sometimes also with a median longitudinal line of spinules on ventral surface, and the corpus bursae with two long, slender, curved, horn-shaped signa.
Description Adults (Figs 956, 957) Wingspan. 7–13 mm; very small to small. Head (Fig. 958). Small. Ocellus moderate. Labial palpus lightly sinuate, upcurved, slender but medially weakly dilated, rather short (1.3–1.5× diameter of eye), white with yellowish tinge; second segment curved, slightly crescentic with loose scales along ventral margin, longest in middle; terminal segment rather long, cone-shaped with appressed scales. Proboscis longer than palpus. Antenna to about middle of wing, thicker in male, dorsally with appressed scales and with a narrow band of very short cilia. Thorax. Smooth, like wing and rest of body scales appressed and iridescent, with pale bluish and golden sheen. Legs unmodified. Wings (Figs 116, 956, 957). Forewing index 0.39–0.46. Forewing subtriangular, costa weakly curved towards apex, apex rounded, termen oblique and nearly straight, never notched, dorsum strongly curved at base; cell widening distally, closing vein fully developed only between M-stem and CuA1, chorda from closer to R1 than R2 to just beyond R5, M-stem to between M2 and M3; all veins separate and distant from each other at base; CuP well developed near margin; anal loop 0.31× length of 1A+2A; wing pattern of Grapholita-like appearance, very similar to some species, e.g. G. delineana Walker, with iridescent scales with a metallic gloss and a well-defined pattern on silvery grey ground with very large, alternating white and black costal strigulae, a dorsal blotch of four well-defined,
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outwardly bent, parallel, silvery lines separated and framed by blackish lines, the ocelloid patch a broad band of leaden scales from near tornus to beyond middle of wing without any trace of black dashes, and often with a row of dark scales around apex and along termen. Hindwing with very weak cubital pecten and anal tuft, each of a few hairs only; frenulum with two bristles in female; closing vein of cell strongly oblique but anteriorly only a trace in membrane, lower surface in male with a band of long and narrow, modified scales along costal edge of cell, appearing darker or lighter depending on illumination; venation not sexually dimorphic, with Sc+R1 and Rs closely approximated even near end of cell but Rs and M1 subparallel and widely distant from base, M2 widely distant from base of M3 and CuA1, which are stalked to 56:–56O; CuA2 from about 6M cell; cell with oblique outer margin, triangular; CuP present at wing margin; 1A+2A and 3A well developed; dark grey, fringe whitish to white with dark basal line. Pregenital abdomen (Fig. 962). Sternum 2 with well-developed anterolateral processes. Tergum 8 of male hood-shaped, membranous, with a narrow anterior edge. Tergum 7 of male with strongly sclerotised posterior edge and with a median bipronged or horseshoe-shaped process from inner side, in some species with paired shallow pockets filled with moderately long scales along anterior edge; two very large scale bundles deeply invaginated in membrane between T7 and T8. Coremata: eighth sternite subrectangular plate, scale-tufts and intersegmental ventral sclerite not developed. Male genitalia (Figs 959–961). Vinculum U-shaped, wide ventrally, becoming narrow dorsally. Tegumen a narrow band, articulated with anterior edge of vinculum, narrow and high, evenly tapering from base to about 36:; apex of tegumen abruptly widened into a large, flat, round or transversely ovate disc. No trace of uncus or socii. Gnathos indicated by membranous connection to anal tube. Valva strongly swollen, short, ovate to triangular, with very small basal excavation and no apparent costal process, outer surface strongly convex, apical third of inner surface with dense, long bristles but not to margin, leaving a roundedtriangular naked apical lobe, sometimes with some deciduous, flattened bristles along ventral margin. Juxta triangular with concave sides, caulis connected to aedeagus near middle. Aedeagus large, widest near base, tapering towards apex; vesica with a sheaf of deciduous cornuti (distally broadly
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rounded in M. sanderyi, sp. nov.) associated with a long, irregularly sclerotised portion of vesica. Female genitalia (Fig. 963). Ovipositor lobes slender, subreniform. Sternum 7 small, triangular with apex projecting as lobe over cup- to funnelshaped sterigma that is fused with S7. Posterior portion of ductus bursae a narrow, membranous tube, with abruptly thickened and partially split wall some way below ostium, anterior third abruptly widening, giving impression of pearshaped corpus bursae; the widening section just anterior to origin of ductus seminalis with large, paired, more or less confluent dorsal patches of sclerotisation each with a short membranous projection anterolaterally and with a longitudinal line of single spinules between them, sometimes also with a median longitudinal line of spinules on ventral surface. Corpus bursae round, signa two long, slender, curved spines or horns.
Distribution Microsarotis is known from Madagascar, La Réunion, Sri Lanka, India, Nepal, Indonesia and Australia, where the genus has been collected at Darwin, Northern Territory, and at Townsville and Bucasia near Mackay, Queensland.
Biology Fletcher (1932) reported that M. palamedes had been reared in India from larvae on tamarind leaves (Tamarindus, Caesalpiniaceae) at Coimbatore and Pusa and from larvae in flower-buds of Bauhinia purpurea (Caesalpiniaceae) at Nagpur.
scaling of body and wings, a venation with Sc+R1 and Rs close to end of cell and Rs and M1 widely distant from base, two large scale bundles invaginated between modified T7 and T8, tegumen tapering distally and topped by a large, disc-shaped apex, swollen valvae with small basal excavation, caulis attached to middle of aedeagus, sterigma fused with and beneath apex of S7, ductus bursae strongly widened in anterior third with two large, dorsal, confluent patches of sclerotisation each with a short apodeme from outer wall. The subrectangular eighth sternite in male that could be regarded as one of the elements of the coremata (see Komai 1999: 24–25) and the tegumen strongly curved at base suggest that this genus is a member of the Grapholita genus-group (Komai 1999). The position of Microsarotis within the group is not clear. The following two species are here referred to Microsarotis: bicincta (Diakonoff, 1976), Zool. Verh. (Leiden) 144: 10 (Grapholitha). [Nepal.] Comb. nov. lygistis (Diakonoff, 1977), Ann. Soc. Ent Fr. (N. S.) 13: 101 (Laspeyresia). [La Réunion.] Comb. nov.
Constituent species Four species of Microsarotis have been named from outside Australia, the two above and M. palamedes (Meyrick, 1916) and M. pauliani Diakonoff, 1988b, with M. lucida sensu Diakonoff, 1982 a possible fifth species. Two species are known from Australia, an unnamed one with a few specimens from Darwin, and a very long series of a different species named below. sanderyi, sp. nov.
Remarks Clarke (1958) provided photos of adult and female genitalia of the female holotype of M. palamedes and Diakonoff (1982) figured head, venation and genitalia of both sexes of M. palamedes and M. lucida (Meyrick) from Sri Lanka. However, the wing pattern and female genitalia of M. lucida sensu Diakonoff are very different from those of the type figured by Clarke (1958); the latter appears not to be a Microsarotis species. Diakonoff (1976, 1977, 1988b) provided illustrations of genitalia and in one case of the adult of further Microsarotis species, some described as Grapholita or Laspeyresia, and figured M. palamedes from Sumba (Diakonoff 1984a). The monophyly of Microsarotis is based on several conspicuous apomorphies: the iridescent
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Microsarotis sanderyi Komai & Horak, sp. nov. (Figs 956, 959, 960, 962, 963)
Material examined Holotype. , ‘Bucasia, N[or]th Q[ueens]l[an]d, Ken J. Sandery, 24 Jan. 2003’, ‘ANIC genitalia slide No. 18716 ’ (ANIC). Paratypes. Queensland: same label data as holotype: 1 , 15.xii.2001; 1 , 10.i.2002; 1, 21.i.2002; 1 , 3.x.2002; 1 , 20.x.2002; 1, 31.x.1002; 1, 22.xii.2002; 56 , 7, 10.–31.i.2003; 13 , 1, 2.–7.ii.2003; 1, 13.ii.2003; 1 ,
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457
Figs 956–963. Microsarotis, adults, head, genitalia and abdomen. 956, M. sanderyi, paratype, Bucasia, Q. 957. M. sp., , Darwin, NT. 958, head, M. sp., Thailand. 959, 960, M. sanderyi, male genitalia and aedeagus, holotype, slide ANIC 18716. 961, M. sp., male genitalia, Darwin, NT, slide ANIC 8174. 962, M. sanderyi, 8th segment with invaginated scale tufts, male. 963, M. sanderyi, female genitalia, paratype, Bucasia, Q, slide ANIC 18773. Scale = 2 mm.
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14.ii.2003; 1, 23.ii.2003; 1 , 22.iii.2003; 74 , 2, 4.– 26.i.2004; 1 , 22.ii.2004 (ANIC, QMBC, BMNH).
Diagnosis Microsarotis sanderyi has a silvery grey head, thorax and forewing base without brownish tinge and the distal portion of the forewing with a coppery sheen but only a very weak brownish tinge, and a uniformly leaden band from tornus to beyond middle of wing. The diagnostic feature is the conspicuous and uninterrupted line of blackish scales along the termen that is lacking in M. palamedes. The male genitalia of M. sanderyi are characterised by the horseshoe-shaped process from the hindmargin of T7, the lightly transversely ovate tegumen top, a valva with its outer surface greatly projecting as a ventrodistal, rounded, cone-shaped bulge, and a long, slender, irregularly tapering aedeagus. The female genitalia are characterised by a lamella antevaginalis with a deeply sinuate hindmargin, the ductus bursae with a rather short, thin-walled section below the ostium, a single longitudinal line of evenly small spinules on both dorsal and ventral wall of widened anterior portion and large signa from sclerotised rings. M. palamedes has larger signa and the longitudinal line of spinules in the ductus bursae is longer.
Description Male (Fig. 956). Wingspan 7–9 mm. Head, thorax and tegulae silvery grey, scales smoothly appressed with bluish iridescence; frons, labial palpi and scales on anterior surface of antenna creamy white. Forewing uniformly silvery grey in basal third, scales smoothly appressed with bluish iridescence; very large white costal strigulae distally increasing in size and alternating with black with first indistinct pair at 56M costa, separated from next pair by a long strongly oblique black streak; small second pair at middle of costa continued as a strongly oblique silvery grey streak; distal five white strigulae large, elongate, triangular, separated by black scales and less oblique towards apex. Distal half of wing blackish with an uninterrupted line of black scales along termen; ocelloid patch indicated by an erect, lightly inwardly curved band of uniformly leaden scales from tornus to beyond middle of wing, edged with black scales but without any black bars; dorsal blotch of two pairs of distinct, outwardly curved silvery lines with scattered whitish scales, each pair separated by a line of greyish black scales and
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central line between the two pairs and lateral edging deep black. Hindwing dark grey, darker towards tip, with hardly a brownish tinge; fringe whitish to grey in apex with dark basal line. Abdomen silver grey. T7 with narrowly, strongly sclerotised anterior margin and adjacent eversible small loose scale tuft in shallow pocket on each side, a strongly sclerotised, horseshoe-shaped process with long, slender prongs from inner surface of broadly sclerotised hindmargin; two very large scale bundles invaginated between T7 and T8. Male genitalia (Figs 959, 960). Tegumen moderately constricted below lightly transversely ovate top. Valva strongly inflated with convex outer surface, flat inner surface with patch of dense, long spines in distal half not reaching to tip, leaving the ventro-distal rounded-triangular apex naked except for some deciduous, flattened bristles. Juxta triangular, wide. Aedeagus tapering apically, with a sheaf of short, broadly rounded, deciduous cornuti, with a pointed base and subbasal attachment point. Female Wingspan 7.5–9.5 mm. Wing pattern as in male. Female genitalia (Fig. 963). Ovipositor lobes slender. Sterigma with lamella antevaginalis deeply sinuate, laterally excavated and medially projecting as a blunt triangle. Distal, slender part of ductus bursae long and anterior 6M with thicker wall and a partial, longitudinal split; widened portion anterior to ductus seminalis with two large, longitudinally ovate sclerotised patches and a single longitudinal line of 10–20 evenly small spinules on both dorsal and ventral wall. Signa two long, fairly stout horns from a sclerotised ring.
Distribution and biology Microsarotis sanderyi has been collected in large numbers at light in a suburban garden at Bucasia, Queensland.
Remarks A single male from Townsville is probably also M. sanderyi but has not been included in the type series. The much larger signa of M. sanderyi and the spinules in the ductus bursae are the crucial diagnostic characters separating it from an unnamed species from the Northern Territory. This species is named for Ken J. Sandery who for years has been making a significant contribution to the ANIC with his collecting at
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Bucasia, and who caught and set all the material of M. sanderyi.
Commoneria Komai & Horak, gen. nov. (Figs 118, 964–970) Type species: Laspeyresia cyanosticha Turner, 1946, Trans. R. Soc. S. Aust. 70: 218, hereby designated. [Australia.]
Diagnosis Small to medium-sized, evenly grey to dark grey moths with typically grapholitine markings, with distally subrectangular forewings with a deep notch below apex and the central portion of the termen projecting further than the blunt, roundedrectangular apex. The frons and the long, slender, upcurved labial palpi are conspicuously pale, yellowish, contrasting with the dark grey vertex scaling. Hindwing with Rs and M1 distant at base, M3 and CuA1 connate. The large anterolateral processes on S2 and in the male the scent organ on T8 and the bifurcate hindmargin of eighth sternite are diagnostic for Commoneria. The male genitalia have a simplified, weakly sclerotised, high and narrow tegumen, long, slender valvae with a deep emargination at 56: of ventral margin, a long caulis and an aedeagus with a bulbous base. The female genitalia are characterised by a deeply V-shaped hindmargin to S7, a lightly and unevenly sclerotised, narrow ductus bursae, a corpus bursae with two thorn-shaped signa, and a ductus seminalis originating from the ductus bursae close to the corpus bursae.
Description Adults (Fig. 964) Wingspan. 10.0–13.5 mm; small to mediumsized. Head (Fig. 970). Ocellus large, chaetosema with few long bristles. Vertex with two large lateral tufts of long, loose, spreading, dark grey scales, their tips medially converging and anteriorly protruding between antennae. Uppermost part of frons with downward oriented, moderately long, rather loose yellowish scales, remainder with minute, appressed, medially and upwardly directed yellowish scales. Proboscis short, hardly longer
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than labial palpus. Labial palpus slender, moderately long (1.5–1.7× diameter of eye) and upcurved, yellowish, second segment strongly and evenly curved, with shortish but rough scaling, especially along ventral margin, third segment long (nearly half length of second segment), straight and apically tapered, with short, appressed scales. Antennae short, not reaching to middle of forewing, cilia in male barely discernible. Thorax. Without crest. Legs unmodified. Wings (Figs 118, 964). Forewing index: 0.43; forewing dilated, distally subrectangular, termen with a deep notch below rounded-rectangular apex and with central portion from M1 to M3 protruding further than apex; discal cell 0.63× length of wing, with chorda and M-stem well developed but closing vein weak; all veins present and separate beyond cell, R1 from middle of cell, R3 from much closer to R4 than to R2, base of M3 equidistant to M2 and CuA1, CuA2 from 36: cell; CuP present at margin; forewing evenly grey with ochreous sheen to dark grey-brown, with typically grapholitine markings: four pairs of white strigulae from apex to middle of costa and some indistinct ones before middle; with lines of leaden scales with metallic blue sheen originating from some strigulae, one from 2nd pair of strigulae from apex running to notch on termen and a second, inwardly curved one from 4th pair of strigulae first running towards termen and then to tornus, bordering an apical area with cinnamon-tipped scales that contains 2–3 longitudinal, parallel black dashes between M2 and CuA2; with a conspicuous row of dark scales along notched termen and with dorsal blotch ranging from a series of 3–4 fairly welldefined, outwardly curved yellow lines to faint traces only or an indistinct whitish smudge. Hindwing with moderate cubital pecten and scattered long hairscales across anal area; frenulum of two (C. cyanosticha) or three (unnamed species) bristles in female; Rs and M1 distant at base, M2 distant and nearly parallel to M3, M3 and CuA1 connate, basal loop of 1A+2A only a weak trace, 3A well developed; dark grey-brown with bronze sheen to blackish, paler towards base, with pale fringe. Pregenital abdomen (Fig. 965). Sternum 2 with large and conspicuous anterolateral processes, longer than the short, curved apodemes with their broadened apex. Tergum 8 in male with a yokeshaped sclerite along anterior margin with a large tuft of large, modified, inflated scales at each end, with a dense fringe of very long, slender scales along curved hindmargin, and with an area of large scale sockets in the center of T8, connected by indistinct
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medial sclerotisation to yoke-shaped sclerite. Coremata reduced: eighth sternite with a medially concave hind margin; intersegmental ventral sclerite sometimes present, scale tufts always absent. Male genitalia (Figs 968, 969). Vinculum a narrowish band. Tegumen a narrow band, curved abruptly at base and articulating with anterior edge of vinculum, high, narrow, subtriangular and
tapering towards rounded apex, very weakly sclerotised. No trace of uncus. Gnathos indicated by very weakly sclerotised band. Valva long and slender with nearly straight costa; ventral margin in basal third projecting strongly as a rounded angle followed by a deep emargination just beyond 56: lined with a flange towards both ends; basal opening large; costal process absent; cucullus
Figs 964–970. Commoneria, gen. nov., adult, genitalia and abdomen. 964, C. sp., , nr Cooktown, Q. 965, C. cyanosticha, 8th segment, male. 966, C. cyanosticha, signa, detail of 967. 967, C. cyanosticha, female genitalia, syntype, Koah, Q, slide ANIC 8183. 968, 969, C. cyanosticha, male genitalia and aedeagus, syntype, Koah, Q, slide ANIC 8109. 970, C. cyanosticha, head, male. Scale = 2 mm.
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elongate, semiovate, with numerous, dense bristles ventrally on inner surface and densely covered with long hairscales on outer surface. Caulis long and curved; aedeagus sinuate in lateral view, with a bulbous basal half and a narrow, parallel-sided distal part in ventral view; vesica without cornuti. Female genitalia (Figs 966, 967). Ovipositor lobes broad and flat, subelliptic. Apophyses short to moderately long, slender. Sternum 7 with a deeply V-shaped hindmargin with ostium at its lowest point. Ductus bursae narrow, shorter than corpus bursae to very short, lightly and unevenly sclerotised with a short, partly membranous, sometimes constricted portion at 56: below ostium. Corpus bursae roundish to teardrop-shaped, with ductus seminalis originating from posterior end close to ductus bursae; bulla seminalis present; signa two very small thorns (C. cyanosticha) (Fig. 966).
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can be regarded as coremata elements regardless of the absence of scale tufts. Furthermore, the tegumen that is curved at base and articulating with the anterior edge of the vinculum suggests the genus is a member of the Grapholita-group. This most attractive genus is named in honour of Ian Common who inspired the authors’ work on Australian Tortricidae.
Constituent species One named and possibly one unnamed species are known from Australia: cyanosticha (Turner, 1946), Trans. R. Soc. S. Aust. 70: 218 (Laspeyresia). [Qld: Koah.] Comb. nov.
Parapammene Obraztsov Distribution Known from north-eastern Queensland only, northward from near Ingham to near Batavia Downs on Cape York Peninsula, often from transition zones of rainforest to eucalypt forest or from riverine associations.
Biology Turner (1946) reported that the type series of C. cyanosticha had been reared in December from fruit of Parinari nonda (Chrysobalanaceae) from Koah near Cairns.
Remarks The deep notch below the apex on the forewing termen and the position of the male scent organ dorsally, on T8, are the two most obvious generic apomorphies. Two possible further apomorphies are found in the female genitalia, the origin of the ductus seminalis from the posterior part of the corpus bursae close to the ductus bursae and the deeply V-shaped hindmargin of S7. Microsarotis Diakonoff shares the dorsal scent organ associated with T8 and also has Rs and M1 distant at base, but the genitalia do not suggest a close relationship between the two genera. Sternum 8 is as long as T8 and with a concave hind margin that is unusual among Grapholitini except for Loranthacydia Horak, Common & Komai, but the configuration of the genitalia and the wing venation clearly refer the genus to the Grapholitini. The compact eighth sternite and large intersegmental ventral sclerite
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(Figs 119, 120, 971–978) Parapammene Obraztsov, 1960, Tijdschr. Entomol. 103: 125. Type species: Phthoroblastis selectana Christoph, 1882, Bull. Soc. Imp. Nat. Moscou 56 1881: 426, by original designation. [Russia: Amur Region.] Diamphidia Obraztsov, 1961, Tijdschr. Entomol. 104: 51. Type species: Pammene petulantana Kennel, 1901, Dtsche Entomol. Z. 13: 301, by original designation. [Russia: Amur Region.] A junior homonym of Diamphidia Gerstaecker, 1855, Ber. Akad. Wiss. Berlin 1855: 638 (Coleoptera). Pammenodes Danilevsky & Kuznetzov, 1968, Fauna SSSR (N. S.) 98 (Lepid. 5(1)): 334. Type species: Pammene glaucana Kennel, 1901, Dtsche Entomol. Z. 13: 302, by original designation. [Russia: Primorsky Krai.] Mimarsinania Koçak, 1981, Priamus 1: 116. Replacement name for Diamphidia Obraztsov, 1961. Diplosemaphora Diakonoff, 1982, Zool. Verh. (Leiden) 193: 35. Type species: Diplosemaphora amphibola Diakonoff, 1982, ibid. 193: 36, by original designation. [Sri Lanka.]
Diagnosis Very small to small moths with a brownish grey to dark brownish forewing with ochreous irroration, with median fascia usually developed, with an
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ocelloid patch with reduced or absent outer edge, and without or with ill-defined whitish stria below apex. Hindwing venation sexually dimorphic; male hindwing with variable venation: with Sc+R1 and Rs fused beyond outer edge of cell in all Australian species, or without trace of Rs, or with Sc+R1 and Rs separate to margin as in females. Upper surface of male hindwing usually with narrow or hair-like, modified scales along Sc+R1 and Rs, sometimes central portion of wing with very small, modified scales. Sternum 2 without anterolateral processes, and T4 and T5 in male sometimes with a transverse band of modified scales or granular spots. In the male genitalia, the tegumen is a broad band with its top laterally emarginate, and the curved or sinuate aedeagus is tapering gradually or abruptly narrowed in distal 56:–56O. The female genitalia are characterised by S7 a convex rather than a flat plate and by a narrow, band-like longitudinal sclerite in the proximal part of the ductus bursae.
Description Adults (Figs 971, 972) Wingspan. 6.5–17 mm (Australian species: 6.5–9.5 mm); very small to small. Head (Fig. 975). Ocellus well developed, chaetosema very obvious. Proboscis rather short, not much longer than labial palpus. Labial palpus from rather short to moderately long (1.6–2.9× diameter of eye); either short, ascending, with second segment roughly scaled beneath and with terminal segment very short, with obtuse apex, or palpus longer, porrect, sinuate, with second segment distally strongly widened, triangular, and with rather long terminal segment with pointed apex. Antenna short, less than 56O forewing length; anterior surface without scales; cilia in male at most 56M width of flagellar segment. Thorax. Without crest. Legs unmodified. Wings (Figs 119, 120, 974, 971, 972). Forewing index: 0.38–0.49 (Australian species: 0.42–0.46); forewing oblong subtriangular, costa gently curved, apex rounded or obtusely pointed, termen almost straight or convex, oblique; chorda and M-stem present or absent, chorda if present from between R1 and R2 to between R4 to R5, M-stem if present to M2 or between M2 and M3; all veins present and separate beyond cell; R1 arising before middle of cell; R2 nearer to R3 than R1; R4 from anterior angle of cell, to costa; R5 to termen; M1 and M2 parallel; CuA1 from posterior angle of cell; CuA2 arising beyond middle of cell; CuP present at wing
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margin; brownish grey to dark brownish grey with whitish ochreous to ochreous irroration, with or without median fascia, with five costal strigulae beyond middle of costa, with subcostal leaden streaks, and with ocelloid patch (the outer edge absent or having a tendency to be reduced), without whitish stria below apex (if present, very illdefined). Hindwing semiovate, costa gently arched, apex rounded; cubital pecten present; frenulum of female consisting of three bristles; on upper surface in males very narrow or hair-like modified scales along Sc+R1 and Rs in about distal 56O of cell; venation in male variable, of three types, in most species Sc+R1 and Rs fused beyond outer edge of cell, or trace of Rs absent (not in Australian species), or Sc+R1 and Rs separate to margin (not in Australian species), while in female Sc+R1 and Rs separate to margin; Rs to costa; M3 and CuA1 slightly separate at base or connate or short-stalked; CuA2 arising beyond middle of cell, basal loop of 1A+2A only weakly indicated, 3A present; brownish grey, sometimes with a strong greenish reflection, in male of some Australian species central wing portion with very small, modified scaling, in some non-Australian species with a narrow patch of black scales along 3A. Pregenital abdomen (Figs 977, 978). Sternum 2 without anterolateral processes (Fig. 977). Tergum 4 and T5 of male sometimes with a transverse band of narrow or hair-like scales and/or one of blackish scales along anterior margin respectively or T4 and T5 with a transverse band of granular spots along anterior margin (not in Australian species). Coremata: eighth sternite a crescentic plate; intersegmental ventral sclerite with a pair of short levers; paired scale-tufts well developed or rarely absent; T8 sclerotised, hood-shaped, with a subtriangular anterior ridge. Male genitalia (Fig. 973). Vinculum a Ushaped band, broad ventrally and becoming narrower dorsally. Tegumen a rather wide, subtriangular band with top emarginate in lateral aspect. Uncus not differentiated. Gnathos an inconspicuous pair of narrow sclerotised arms fused with tegumen. Valva constricted before middle; cucullus subtriangular or subrectangular, basal excavation large; costal process reduced. Juxta triangular or trapezoidal, caulis rather long. Aedeagus sinuate or curved, abruptly narrowed in distal 56:–56O or gradually narrowed; vesica with a group of spiculate cornuti. Female genitalia (Fig. 976). Ovipositor lobes elongate. Sternum 7 a strongly convex plate, with ostium in excavation of posterior margin. Sterigma
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Figs 971–978. Parapammene, adults, genitalia, hindwing, head and abdomen. 971, P. sp. A, , Smith Point, Cobourg Peninsula, NT [image reversed]. 972, P. sp. C, , Black Point, Cobourg Peninsula, NT [image reversed]. 973, P. sp. C, male genitalia, Black Point, Cobourg Peninsula, NT, slide ANIC 8097. 974, detail, male hindwing, P. sp. A. 975, head, P. selectana, Japan. 976, P. dyserasta, female genitalia, holotype, slide ANIC 8178. 977, 978, male abdomen, P. sp. D, Black Mt, ACT, slide ANIC 8208. 977, 2nd sternite. 978, 8th segment. Scale = 2 mm.
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a sclerotised plate or not developed. Ductus bursae with cup- or tube-shaped sclerotisation at posterior end and with an irregular, sclerotised ring in anterior 56: to 56O, from dorsal wall of which a narrow, band-like sclerite extends in a spiral to the ventral wall at the juncture of ductus and corpus bursae; ductus seminalis arising laterally, from anterior 56:–56O of ductus bursae; bulla seminalis present; corpus bursae ovate, in some nonAustralian species constricted in anterior 56M; with two horn-shaped signa.
Distribution Palaearctic, Oriental, and Australian regions. In Australia, Parapammene has been recorded from: the northernmost parts of the Northern Territory; northern Queensland from the Cape York Peninsula, the Cairns area and from South Queensland at Warwick; New South Wales at Bathurst, and the Australian Capital Territory. These small moths no doubt are undercollected.
Biology The known larvae of Parapammene are borers in stems and seeds. Hostplants recorded in the Palaearctic region chiefly belong to the Fagaceae, and a few species use Tiliaceae and Aceraceae as larval hosts (Komai 1999). In Australia, one undescribed species has been reared from larvae feeding in seed capsules of Dodonaea viscosa (Sapindaceae) on Black Mt, ACT.
Remarks Comprehensive reviews of Parapammene have been given by Obraztsov (1960), Danilevsky and Kuznetzov (1968), Razowski (1989), and Komai (1999). The following character states support the monophyly of Parapammene (Komai 1999): 1, ductus bursae with a narrow, band-like sclerite; 2, seventh sternite of female a convex (not flat) plate, and, 3, S2 without anterolateral processes.
Ixonympha Komai & Horak, gen. nov. (Figs 121, 979–984) Type species: Hyphantidium hyposcopa Lower, 1905, Trans. R. Soc. S. Aust. 29: 105, hereby designated. [Australia.]
Diagnosis Small, narrow-winged, very characteristic-looking moths with blackish forewings with a broad whitish band from 6M costa to tornus, dorsally variously obliterated, and with conspicuous orange hindwings, at first look not suggesting a tortricid. Labial palpi small, slender and lightly upcurved, with fairly rough scaling, especially beneath. Forewing with chorda absent and M-stem at best very weak; hindwing with Rs and M1 stalked in female and Sc+R1 short-stalked with stem of Rs and M1, in male no trace of Rs as result of fusion of Sc+R1, and Rs and M3 and CuA1 connate to barely stalked in female but stalked to nearly half in male. Abdomen with anterolateral processes of S2 very weakly developed, coremata two eversible bundles of large scales at each end of a curved rod. The male genitalia have a hood-like tegumen with a pair of curved, finger-like processes laterally on each side, and the shape of the valva is diagnostic for Ixonympha, with a large, outwardly curved, projecting point on ventral margin at base of cucullus and a low, triangular projection halfway to apex. The aedeagus is sinuate, robust to very stout, with 6–12 stout, spine-like cornuti. The female genitalia are highly derived and diagnostic for the genus, with a complex sterigma, an inverted funnelshaped and sclerotised posterior portion (ductus bursae) giving rise to a small, round corpus bursae with a small, scobinate signum and adjacent to the larger, ovate, flimsy bulla seminalis that is apparently joined to the corpus bursae without intermediate connection.
Description Constituent species Sixteen Parapammene species have been named, including one Australian species (Brown 2005). We are aware of four unnamed Australian species. dyserasta (Turner, 1916), Trans. R. Soc. S. Aust. 40: 535 (Laspeyresia). [Qld: Kuranda.]
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Adults (Fig. 979) Wingspan. 9–11 mm. Small. Head (Fig. 980). Ocellus large, chaetosema with short, barely visible bristles. Proboscis short, slightly longer than labial palpus. Labial palpus small (1.2– 1.4× diameter of eye), slender, straight to lightly upcurved, second segment rough-scaled, especially
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below, terminal segment short and relatively stout, with appressed scales. Antennae to beyond middle of forewing, cilia in male barely discernible. Thorax. Without crest. Legs unmodified. Wings (Figs 121, 979). Forewing index: 0.37– 0.38; forewing subtriangular with straight costa, acute apex, strongly oblique termen and curved
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dorsum; cell 0.60× length of wing, with chorda absent and M-stem at most a faint trace in wing membrane, closing vein weak; all veins present and separate beyond cell, R1 from middle of cell, R2 from closer to R3 than to R1, CuA2 from just beyond 36: cell; CuP present at margin; forewing blackish, with an irregular, broad, whitish band
Figs 979–984. Ixonympha, gen. nov., adult, head, genitalia and abdomen. 979, I. sp., , nr Kulgera, NT. 980, head, I. sp., Alice Springs, NT. 981, I. sp., male 8th sternite, slide T2309. 982, I. sp., female genitalia, nr Mataranka, NT, ANIC 3144. 983, 984, I. sp., male genitalia and aedeagus, Alice Springs, NT, slide T2309. Scale = 2 mm.
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OLETHREUTINE MOTHS OF AUSTRALIA
from 6M costa to dorsum before tornus, variably interrupted towards dorsum with blackish scales; strigulae along costa variably developed, and distal area beyond whitish band in paler species with transverse blackish and some leaden-metallic strigulae, and some scattered whitish scales. Hindwing narrow, with sparse cubital pecten and axillary tuft; frenulum long, of two bristles in female; closing vein weak; in male no trace of Rs as result of complete fusion of Sc+R1 and Rs, in female Rs and M1 stalked, Sc+R1 short-stalked with stalk of Rs and M1, M3 and CuA1 connate to barely stalked in female but stalked to nearly half in male, CuA2 from 36: cell in female, from beyond 36: in male; CuP only a weak trace at margin; 1A+2A and 3A present but not tubular veins; conspicuously orange, some species with bronze or greyish scales in apex. Pregenital abdomen (Fig. 981). Sternum 2 with anterolateral processes hardly developed. Coremata: eighth sternite a small crescentic sclerite; intersegmental ventral sclerite with paired long levers associated with large eversible bundles of narrow scales at each end. Male genitalia (Figs 983, 984). Vinculum a moderately broad band. Tegumen strongly bent at base, articulating with anterior margin of vinculum, high, triangular in outline, hoodshaped, with a pair of digital, curved processes laterally (Fig. 983). No trace of uncus. Gnathos at most membranous, upcurved bands. Valva strongly sclerotised, long, lightly curved and medially narrowed, ventral margin lined with a broad flange and with a large, outwardly curved, projecting point at base of cucullus and a second, low, triangular projection halfway to apex; costal process absent. Caulis stout and short; aedeagus sinuate, tapering towards apex from bulbous base, moderately long and robust to short and very stout; vesica with 6–12 very stout spine-shaped cornuti. Female genitalia (Fig. 982). Ovipositor lobes elongate-ovate; hair short and sparse. Apophyses moderately long, slender. Sternum 7 with a deeply emarginate hindmargin, with a subquadrate central sclerite and two lateral, fan-shaped, sclerotised and densely scaled areas. Sterigma separate from S7, a complex, strongly sclerotised, posteriorly narrowed groove-shaped dorsal process from a collar-like ring around ostium. Ductus and corpus bursae highly modified, with bulla seminalis apparently incorporated into the one complex structure: posteriormost portion below ostium heavily sclerotised, inverted funnel-shaped with a longitudinal, somewhat crumpled fold ventrally,
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with scattered heavy spines in anterior half, bifurcating below wide end of funnel into a small, globose corpus bursae and a larger but flimsier, ovate bulla seminalis [?] with the ductus seminalis originating from its neck; corpus bursae with a single signum, either a small scobinate plate or a thorn with a small domed base.
Distribution Ixonympha is known from ten mostly single specimens in the ANIC collected throughout Australia including Toowoomba in Queensland, Melbourne, South Australia, Geraldton and the Kimberley in Western Australia, and central Australia and northern parts of the Northern Territory. These scattered and extremely diverse localities suggest that Ixonympha occurs wherever its host, mistletoe, occurs. Mistletoe is absent from Tasmania and Ixonympha therefore is not expected to be found there.
Biology One specimen of Ixonympha has been reared from a deployed seed of the mistletoe Amyema quandang (Loranthaceae) on Acacia papyrocarpa during a study of mistletoe ecology in South Australia by N. Reid. The larvae bore into both newly deployed seeds and holdfast phase seedlings, forming a small waxy, frass-covered dome alongside the remnant shell of the seed or seedling (Reid 1984). The seed with the larva was collected 1–3 June on Middleback Station, SA, the adult emerged 24 October. Reid (1987) reported that 7–36% of deployed seeds or seedlings were destroyed by Ixonympha larvae. This unusual life history could explain the occurrence of Ixonympha across a surprisingly wide range of biotopes, with the presence of mistletoes the crucial link between the localities. The small size of the moths and the many single specimens also tally with such a biology.
Remarks Several apomorphies demonstrate the monophyly of Ixonympha, among them the very unusual colour of fore- and hindwings. The genitalia in both sexes of Ixonympha are highly derived, with the two projections from the cucullus characterising the male and the inverted funnel-shaped and heavily sclerotised ductus bursae and the single plateshaped signum the female. Ixonympha is closely related to Andrioplecta Obraztsov and Strophedra
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AUSTRALIAN OLETHREUTINE GENERA
Herrich-Schäffer, with the three genera sharing the following synapomorphies: 1, Sc+R1 and Rs entirely fused in the male; 2, two frenulum bristles in female; 3, M3 and CuA1 stalked in hindwing. Furthermore, a pair of digitiform or ear-shaped processes or flaps is present in the lateral tegumen wall of Ixonympha and several species of Strophedra and Andrioplecta (Komai 1999: figs 338, 344, 365). Andrioplecta is possibly the sister group of Ixonympha, sharing a bulla seminalis broadly connected to or continuous with the corpus bursae. The ten specimens of Ixonympha in the ANIC represent at least three different species superficially recognisable by the shape of the white band on the forewing and the colour of the hindwing, in particular the presence of darker scales on its apex.
Constituent species One named and at least two unnamed species are known from Australia. hyposcopa (Lower, 1905), Trans. R. Soc. S. Aust. 29: 105 (Hyphantidium). [Vic.: Melbourne.] Comb. nov.
Olethreutine Moths Final.indd 467
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UNPLACED TAXA The types of the following three species could not be located either in the ANIC or the BMNH, and the descriptions alone are insufficient to link them with species known from the type localities listed. Argyroploce angustifascia Turner was erroneously omitted from the Checklist (Nielsen et al. 1996). An ANIC file note states that according to I. F. B. Common the species ‘may be an Oecophorine’. The two Walker types in the BMNH are considered lost and marked as such on the museum’s card index, in the case of A. dentatana with the ‘lost’ label in Walker’s own handwriting glued to the card. angustifascia Turner, 1925, Trans. R. Soc. S. Aust. 49: 58 (Argyroploce). [Qld: Toowoomba.] dentatana Walker, 1864, List Specimens Lepid. Insects Colln Br. Mus. 30: 991 (Grapholita). [NSW: Sydney.] diffusana Walker, 1864 List Specimens Lepid. Insects Colln Br. Mus. 30: 986 (Sciaphila ?). [South Australia.]
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APPENDIX 1 Character Matrix
120503•Olethreutine Moths 4pp.in477 477
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478
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Characters (Table 1.)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Williella sauteri Horak
1
1
0
0
0
1
1
0
-
1
3
1
0
0
0
Templemania animosana Busck
1
1
0/1
0
0
0
2
0
-
1
3
0
0
2
0
Cryptaspasma Walsingham
1
1
0/1
0/1
0/1
0
2
0
-
1
0/1
0/1
0
1
0
Collogenes Meyrick
1
0
0
0
0
2
0/1
0
-
1
1
1
0
1
0
Oxysemaphora Diakonoff
1/2
1
0/1
0
1
0/1
1
0
-
1
0
1
0
0
0
Cnecidophora, gen. nov.
1/2
1
0
1
1
1
1
0
-
1
0
1
0
0
0
Sorolopha Lower
1
1
0/1
0/1
0/1
0
1
0
-
1
1/2
1
0
0
1
Sycacantha Diakonoff
1
2
0/1
0
0
0
1
0
-
1
1
1
0
0
0/1
Atriscripta, gen. nov.
1
2
1
0
0
0
1
0
-
1
1
0
0
0
0
Demeijerella Diakonoff
1
2
0
0/1
1
1
1
0
-
0
0
1
0
0
0
Lobesia Guenée
1
1
0
1
1
0/1
1/2
0
-
0
0
1
0
0
0/1
Gnathmocerodes Diakonoff
1
1
0
1
1
1
1
0
-
1
1/2
1
0
0
0/1
Rhectogonia Diakonoff
1/2
1
0
1
1
1/2
1
0
-
1
0
1
0
0
1
Ophiorrhabda Diakonoff
1
1
0/1
0/1
1
1
1/2
0
-
1
1
1
0
0
1
Dudua Walker
1/2
2
1
0
0
0
1
0
-
1
1
1
0
0
1
Trachyschistis Meyrick
1
2
0/1
0
0/1
1/2
2
0
-
1
1
1
0
0
1
Archilobesia Diakonoff
1
2
0
1
1
1/2
1
0
-
1
1
1
0
0
1
Euobraztsovia Diakonoff
1
2
1
0
0
0
1
0
-
1
1/2
1
0
0
1
Eremas Turner
0/1
2
0/1
0
0/1
1
1
0
-
1
1
1
0
0
1
Zomariana Diakonoff
1
1
0
1
1
1/2
0/1
0
-
1
1
1
0
0
0
Podognatha Diakonoff
0/1
1
0/1
1
1
1/2
1
0
-
1
3
1
0
0
1
Temnolopha Lower
1
1
0
0/1
1
1/2
0/1
0
-
1
2
1
0
0
0
Diakonoffiana Koçak
1/2
2
0
0/1
0/1
1
1
0
-
1
1
1
0
0
1
Megalota Diakonoff
1
1
0
1
1
1
0/1
0
-
1
1
1
0
0
0/1
Costosa Diakonoff
1
1
0
1
1
1
1
0
-
1
1
1
0
0
1
APPENDIX 1
120503•Olethreutine Moths 4pp.in478 478
Appendix 1. Character matrix.
120503•Olethreutine Moths 4pp.in479 479
17
18
19
20
21
22
23
24
25
26
27
28
29
30
Williella sauteri Horak
0
0
0/1
0
1
2
0
0
0
2
1
-
-
0
0
Templemania animosana Busck
1
0
1
0
1
1
0
0
0
0
0
0
2
0
0
Cryptaspasma Walsingham
0
0/1
0
0
1
0/1
0
0
0
0/1
0
0
1
0/1
0
Collogenes Meyrick
2
0
0
0
1
2
0
0
0
0/1
1
-
-
3
0
Oxysemaphora Diakonoff
0
0
0
0
1
1
0
0/1
1
0
0
0
1
0
0
Cnecidophora, gen. nov.
1
0
0
0
1
1
0
0
0
0
0
0
1/2
0/1
0
Sorolopha Lower
1
0
0/1
0
1
2
0
0
0
0
0
0
2/3
0
0
Sycacantha Diakonoff
1
0
2
0
1
1
1
1
0
0/1
0/1
1/2
0
0
0
Atriscripta, gen. nov.
1
0
2
0
1
1
1
1/2
0/1
0
0
2
1
0
0
Demeijerella Diakonoff
0
0
0/1
0
1
2
0
0
0
0
1
-
-
3
0
Lobesia Guenée
0
0
0/1
0
1
0/4
0
0
0
1/2
0/1
0/2
1
0/2
0
Gnathmocerodes Diakonoff
0
0
0
0
1
1
0
0
0
0/1
0
0
2
0
0
Rhectogonia Diakonoff
0
0
1
0
1
3
1
1
1
0/1
0
0/1
1
0
0
Ophiorrhabda Diakonoff
1
0
0/1
0
1
1
0
0
0
0
0
0
1/2
0
0
Dudua Walker
0
0
0/2
0
1
1/2
0
0
0
0/1
0
0
3
0/1
0
Trachyschistis Meyrick
1
0
0
0
1
0/4
1
1
0
0/1
0
0
1
0
0
Archilobesia Diakonoff
0
0
1
0
1
1/2
0
0
0
0
0
0
2/3
0/1
0
Euobraztsovia Diakonoff
0
0
1
0
1
1/4
1
0/1
0/1
0
0
0
1
1
0
Eremas Turner
0
0
0/1
0
1
3
0
0
0
0
0
0
2
0
0
Zomariana Diakonoff
0
0
0/1
0
1
0
0
0
0
0/1
0
2
2
0
0
Podognatha Diakonoff
0
0
1/2
0
1
2/5
0
0
0
0
0
2
1
0/1
0
Temnolopha Lower
0
0
1
0
1
1
0
1
0
0
0
2
2
1
0
Diakonoffiana Koçak
1
0
1
0
1
1/4
0/1
1
1
0
0/1
0
3
0/1
0
Megalota Diakonoff
1
0
0/1
0
1
1
0
0/1
0/1
0
0
2
2
0/1
0
Costosa Diakonoff
1
0
1
0
1
2
0
0
0
0/1
0/1
0
2
0/1
0
APPENDIX 1 479
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16
480
5/5/06 2:45:01 PM
32
33
34
35
36
37
38
39
40
41
42
43
44
45
Williella sauteri Horak
2
0
0
0
0
0
0
0
0
0
0
0
1
0
1
Templemania animosana Busck
1
0
0
0
0
0
0
0
0
0
0
0
4
0
1
Cryptaspasma Walsingham
1
0
0
0
1
0
0
2
0/1/2 0
0
0
2
0
1
Collogenes Meyrick
2
0
0
1
0
0
0
0
0
0
0
0
2/3
0
0
Oxysemaphora Diakonoff
1
0
0
0
0
0
0
0
0/1
0
0
0
2
0
1
Cnecidophora, gen. nov.
2
0
0
0
0
0
0/1
0/1
0
0/2
0
0
4
0
0/1
Sorolopha Lower
1
0
0
0
0/1
0
0
0
0/3
0/1
0
0
2
0
2
Sycacantha Diakonoff
2
0
0/1
0
0
0
1
0
0
0
0
0
2
0
1/2
Atriscripta, gen. nov.
1
0
0
0
0
0
0
0
0
0
0
0
2
0
2
Demeijerella Diakonoff
1
0
0
0
0
0
0
0
0
0
0
0
2
0
1
Lobesia Guenée
2
0
0
0
0/1/2 0
0/1
0
4
0
0
0
2/4
0
0/1
Gnathmocerodes Diakonoff
1
0
0/1
0
0/1
0
0
1
4
0
0
0
2
0
2
Rhectogonia Diakonoff
2
0
0
0
1
0
0
2
0
0
0
0
2
0
2
Ophiorrhabda Diakonoff
1
0
1
0
0
0
0/1
2
4/5
0
0
0
2
0
2
Dudua Walker
2
0
1
0
2
0
1
1
0
2
0
0
2
0
2
Trachyschistis Meyrick
1
0
1
0
1
0
1
1
5
0
0
0
2
0
1/2
Archilobesia Diakonoff
0
0
0
0
2
0
1
1
4
0
0
0
2
0
2
Euobraztsovia Diakonoff
2
0
1
0
0
0
1
1
0
1
0
0
2
0
2
Eremas Turner
2
0
0
0
0
0
1
0
4
0
0
0
1
0
1
Zomariana Diakonoff
1
0
0
0
0
0
0
2
3
0
0
0
2
0
2
Podognatha Diakonoff
2
0
0/1
0
2
0
0
2
2
0
0
0
2
0
2
Temnolopha Lower
2
0
1
0
1
0
1
2
5
0
0
0
2
0
2
Diakonoffiana Koçak
1
0
1
0
0
0
1
0
1
0
0
0
2
0
2
Megalota Diakonoff
2
0
0
0
1
0
0/1
0/2
0/4
0
0
0
2/3
0
2
Costosa Diakonoff
2
0
0
0
2
0
0
2
0
1
0
0
2
0
2
APPENDIX 1
120503•Olethreutine Moths 4pp.in480 480
31
120503•Olethreutine Moths 4pp.in481 481
47
48
49
50
51
52
53
54
55
56
57
58
59
60
Williella sauteri Horak
4
0
0
1
1
0
0
0
0
0
2
1
0
0
0
Templemania animosana Busck
0
0
0
2
1
0
0
0
0
0
2
1
0
1
0
Cryptaspasma Walsingham
0/1/2 0
0
1
1
0
0
0
0
0
2
0
0
0
0
Collogenes Meyrick
1
0
0
0
1
0
0
0
0
1
2
0
0
0
0
Oxysemaphora Diakonoff
1
0
0
1
1
0
1
0/1
0
2
2
0
2
0
0
Cnecidophora, gen. nov.
1/2
0
0
1
1
0
0/1
0
0
2
2
0
0
0
0
Sorolopha Lower
1
0
0
1
1
0
0
0
0
1/3
2
0
0
0
0
Sycacantha Diakonoff
1
0
0/1
1
1
0
0
0/1
0
2
2
0
0
0
0
Atriscripta, gen. nov.
1
0
0/1
0
1
0
0
0
0
5
2
0
0
0
2
Demeijerella Diakonoff
1
0
1
1
1
0
0
0
0
1
2
0
0
0
0
Lobesia Guenée
0/2/3 0
0/1
0/2
1
0
0
1
0
0/1
2
0
0
0
0
Gnathmocerodes Diakonoff
1/2
0
0
1
1
0
0
0
0
1/2
2
0
0
0
0
Rhectogonia Diakonoff
2
0
0
1
1
0
0
1
0
1
2
0
0
0
0
Ophiorrhabda Diakonoff
0/1
0
0
1
1
1
0
0
0
1
2
0
0
0
0
Dudua Walker
0/1/2 0
0
1
1
0
0/1
0
0
1
2
0
0
0
0
Trachyschistis Meyrick
0
0
0
1
1
0
0
0
0
1
2
0
0
0
0
Archilobesia Diakonoff
1/2
0
0
1
1
0/1
0
0
0
1
2
0
0
0
0
Euobraztsovia Diakonoff
1
0
0
1
1
0
0
0
0
1
2
0
0
0
0
Eremas Turner
0/2
0
0
0
1
0
0
0
0
1
2
0
0
0
3
Zomariana Diakonoff
1/2
0
0
1
1
1
0
0
0
1
2
0
0
0
0
Podognatha Diakonoff
1
0
0
1
1
0
0
0
0
2
2
0
0
0
0
Temnolopha Lower
1
0
0
1
1
1
0
0
0
1
2
0
0
0
0
Diakonoffiana Koçak
2
0
0
1
1
1
0
1
0
1
2
0
0
0
0
Megalota Diakonoff
0
0
0
1
1
0
0
0
0
1
2
0
0
0
0
Costosa Diakonoff
1/2
0
0
1
1
0
0
0
0
1
2
0
0
0
0
APPENDIX 1 481
5/5/06 2:45:03 PM
46
482
5/5/06 2:45:04 PM
62
63
64
65
66
67
68
69
70
71
72
73
74
75
Williella sauteri Horak
1
2
1
1
2
1
0
0
1
4
2
1
0
0
2
Templemania animosana Busck
1
2
0
0
0
1
0
0
1
1
2
1
0
0
2
Cryptaspasma Walsingham
1
2
0
0
2
0
0
0
1
0
1
1
0/1
0
0
Collogenes Meyrick
0/1
2
0
0
1
0
0
1
-
-
-
-
0
0
0
Oxysemaphora Diakonoff
0
2
0/1
0
1
1
0
0
1
5
0
1
0
0
1/2
Cnecidophora, gen. nov.
0
2
0
0
1
1
0
0
1
0
5
1
0
0
1/2
Sorolopha Lower
0
0
1
0
0
0
0
0
1
2
5
0
0
1
2
Sycacantha Diakonoff
0
0
0
0
0
0
0
0/1
1
2
0
0
0
1
1/2
Atriscripta, gen. nov.
0/1
0/2
1
0
0
0
0
1
-
-
-
-
0
1
2
Demeijerella Diakonoff
0
2
0/1
0
1
0
0
1
-
-
-
-
0
1
2
Lobesia Guenée
0/1
0/2
0/1
0
0/1
0
0
0/1
1
2/5
0/5
0
0/1
0/1
0/1/2
Gnathmocerodes Diakonoff
0
2
1
0/1
1
0
0
0
1
3
3
0
0
0
1
Rhectogonia Diakonoff
0
2
1
0
0
1
0
1
-
-
-
-
0
0/1
1
Ophiorrhabda Diakonoff
0
0
1
0/1
1
0/1
0
0
1
1/3/4 2
1
0
0
1
Dudua Walker
0
0
1
0/1
1
0
0
0
1
1/3/4 2/5
1
0
0
1
Trachyschistis Meyrick
0
0
1
1
1
0
0
0
1
4
2
1
0
0
1
Archilobesia Diakonoff
0
0
1
1
1
0
0
0
1
4
2
1
0
0
1
Euobraztsovia Diakonoff
0
0
1
0
1
0
0
0
1
3
2
0/1
0
0
1
Eremas Turner
1
2
0
1
1
0
0
0
1
0
5
1
0
0
1
Zomariana Diakonoff
0/1
0
0
1
1
0
0
1
-
-
-
-
0
0
2
Podognatha Diakonoff
0/1
0
1
1
0
0
0
0
1
1/5
2
1
0
1
0/2
Temnolopha Lower
0
0
1
1
0
0
0
0
1
1/4
2
1
0
0
1/2
Diakonoffiana Koçak
0
0
0/1
0/1
1
0
0
0
1
0/1/2 2
1
0/1
0
0/1
Megalota Diakonoff
0
1
1
0
1
1
0
0
0/1
5
2
1
0/1
0
0
Costosa Diakonoff
1
0
0
0
1
0
0
0
0/1
5
2
0/1
1
-
-
APPENDIX 1
120503•Olethreutine Moths 4pp.in482 482
61
120503•Olethreutine Moths 4pp.in483 483
77
78
79
80
81
82
83
84
85
86
87
88
89
90
Williella sauteri Horak
2
0
0
1
2
0
0
0
0
0
0
0
0
0
0
Templemania animosana Busck
2
0
0
1
2
0
0
0
0
0
0
0
0
0
0/1
Cryptaspasma Walsingham
2
0
0
0/1
0/2
0
0
0
1/2
0
0
0/3
0
0
0
Collogenes Meyrick
2/4
0
0
0
0/2
0
0
0
3
0
1
0/1/7 0/1
0
0
Oxysemaphora Diakonoff
2
0
1
-
-
-
0
0
3
0
1
2
2
0
0
Cnecidophora, gen. nov.
1/2
0
0
0
3
0
2
0
3
0
1
2
2
0
0
Sorolopha Lower
0
0
0
0/1
3
0
1
0
2
0
1
0/1
0/1/2 0
0/1
Sycacantha Diakonoff
5
0
0
1
4
0
0
0
1
0
1
0
0/1
0
0/1
Atriscripta, gen. nov.
2
0
0
1
3/4
0
1
0
1
0
0
0
1
0
0
Demeijerella Diakonoff
2
0
0
0
0
0
0
0
2
0
0
0
0
0
0
Lobesia Guenée
2
0
0
1
0/2
0
0
0
3
0
0/1
0
0/1
0
1
Gnathmocerodes Diakonoff
1
0
0
0/1
0/3
0
0/1
0/1
3
1
1
0/5
0/1
0
1
Rhectogonia Diakonoff
0/2
0
0
1
3
0
1
0
2
1
1
7
1
0
0
Ophiorrhabda Diakonoff
2
0
0
0
3
0/1
1
0/1
2
1
1
2
0/1
0
0
Dudua Walker
2
0
0
0
1
1
0
0
1/2
0
0/1
0/2
0/1
0
0
Trachyschistis Meyrick
2
0
0
0
1
1
0
0
1/2
0
0
0/2
0
0
0
Archilobesia Diakonoff
2
0
0
0
1
1
0
0
2
0
0/1
2
0
0
0
Euobraztsovia Diakonoff
2
0
0
1
1/3
0/1
1
0
2/3
1
1
0/2
1
0
1
Eremas Turner
2
0
0
0
3
0
1
0
1
0
0/1
0/2
2
0
0
Zomariana Diakonoff
2
1
0
1
2
0
0
0
1
0
1
0
1
0
1
Podognatha Diakonoff
0/1/4 0
0
0
0
0
0/1
0
2
0
1
0/1
0/1
0
0
Temnolopha Lower
2
1
0
0/1
0
0
0
1
1
0
0
0
0
0
1
Diakonoffiana Koçak
0/1
0
0
1
3
0
1
0
1
1
0
0
0/1
0
0
Megalota Diakonoff
3/4
0
0
0/1
3
0
1
0
2
0
0/1
0/2
0
0
0
Costosa Diakonoff
-
-
0
0/1
0
0
0
0
1
0
1
0/2
0
0
1
APPENDIX 1 483
5/5/06 2:45:05 PM
76
484
5/5/06 2:45:06 PM
92
93
94
95
96
97
98
99
100
101
102
103
104
105
Williella sauteri Horak
0
0
0
0
-
0
-
0
0
0
0
0
0
0
0
Templemania animosana Busck
0
0
0
0
-
0
-
0
0
0
0
0
0
0
0
Cryptaspasma Walsingham
0
1
1
0
-
1
0
0
0
0
0/1
2
1
2
-
Collogenes Meyrick
0
1
1
0
-
0
-
0
0
0
0
3
1
1/2
2
Oxysemaphora Diakonoff
0
1
2
0
-
0
-
0
0
0
0
4
1
0
0
Cnecidophora, gen. nov.
0
1
2
0
-
0
-
0
0
0
0
4
1
0
0
Sorolopha Lower
0/1
1
1/2
0
-
0
-
0
0
0/1
0
3
7
2
-
Sycacantha Diakonoff
0
1
2
0
-
0
-
0
1
0/1
0
3/4
1
0
0/1
Atriscripta, gen. nov.
0
4
2
0
-
0
-
0
1
1
0
3
1
0
1
Demeijerella Diakonoff
0
2
2
0
-
0
-
0
1
0
0
2/3
1
0
0
Lobesia Guenée
1
1
2
0/1
1
0
-
0
0
0
0
2/3/4 1
2
-
Gnathmocerodes Diakonoff
1
1
1/2
1
1
0
-
0
0
0
0
2/3
4
2
-
Rhectogonia Diakonoff
0
1
1
0
-
0
-
0
0
1
0
2
6
2
-
Ophiorrhabda Diakonoff
0/1
1
2
0
-
0
-
0
0
0/1
0
3
1
2
-
Dudua Walker
0/1
1
2
0
-
0
-
0
0
0/2
1
2
1
0/2
0
Trachyschistis Meyrick
0
1
2
0
-
0
-
0
0
0/2
1
2
1
0
0
Archilobesia Diakonoff
0
1
2
0
-
0
-
0/3
0
2
1
2
1
2
-
Euobraztsovia Diakonoff
0
1/2
2
1
1
0
-
2
1
0
0
2
2
2
-
Eremas Turner
0
1
1
0
-
1
2
0
0
0
0
1/2
3
2
-
Zomariana Diakonoff
0
2
1
1
0
0
-
1
0
0
0
2
1
2
-
Podognatha Diakonoff
0
1
1
0
-
0
-
2
0
0
0
3
3
0
0
Temnolopha Lower
1
1
2
0
-
1
1
0
0
0/1
0
2
5
2
-
Diakonoffiana Koçak
0/1
1
2
0/1
2
0
-
0/3
1
0
0
2
1
2
-
Megalota Diakonoff
0
1
1
0
-
1
2
0
0
0
0
3
1
1
2
Costosa Diakonoff
0
4
2
0
-
1
1
0
0
0
0
2
1
0
0
APPENDIX 1
120503•Olethreutine Moths 4pp.in484 484
91
120503•Olethreutine Moths 4pp.in485 485
107
108
109
110
111
112
113
114
115
116
117
118
119
120
Williella sauteri Horak
0
0
1
0
2
0
1
2
1
0
0
-
0
3
0
Templemania animosana Busck
0
2
1
0
4
0
1
2
1
0
1
0
0
2
0
Cryptaspasma Walsingham
-
0
1
0
1
0
1
0/1
1/2
4
1
0
0
2
0
Collogenes Meyrick
1
0
1
0
0/3
0/1
0/1
2
0/1
4
0/1
0
0
2/3
0
Oxysemaphora Diakonoff
0
0
0
0
2
1/2
1
1
0
0
1
1
0
2
0
Cnecidophora, gen. nov.
0
0
0
0
2
0/1
1
1
1
0
1
1
0
2
0
Sorolopha Lower
-
0
0
1
1
2
1
1/2
0
1
0
-
0
1/2
0
Sycacantha Diakonoff
0
1
0
1
2/4
2
1
1/2
0/1
0
0
-
2
2
0
Atriscripta, gen. nov.
0
1
0
1
3
2
1
1
1
0
0
-
2
2/3
0
Demeijerella Diakonoff
0/1
0
0
0
3
0
1
1
0
4
1
0
2
0/1/2/3 0
Lobesia Guenée
-
0
0
0
1/3/4 1/2
1
0/1/2 0/1
3/4
0
-
0
0/1/2
0
Gnathmocerodes Diakonoff
-
0/1
0
0
3
1/2
1
2
0
1
0
-
0
1/2
0
Rhectogonia Diakonoff
-
0
0
0
2/4
2
1
2
0
1
0
-
0
2
0
Ophiorrhabda Diakonoff
-
0
0
0
2/3
2
1
1/2
0
0
0
-
0
1
0
Dudua Walker
1
0/1
0
0
2/3
2
1
2
0
1
0
-
0
2
0
Trachyschistis Meyrick
1
0
0
0
1/3
2
1
2
0
1
0
-
0
2
0
Archilobesia Diakonoff
-
0
0
0
1/3
2
1
1
0
1
0
-
0
2/3
0
Euobraztsovia Diakonoff
-
0
0
0
2/3
2
1
1
0
1
0
-
0
2
0
Eremas Turner
-
0
0
0
1
1
1
1
0
0
0
-
0
3
0
Zomariana Diakonoff
-
0/1
0
0
3
1
1
0
1
1
0
-
0
0
0
Podognatha Diakonoff
0
0
0
0
2
1/2
1
1/2
0
1
0/1
0
0
2
0
Temnolopha Lower
-
0/1
0
0
1/3/4 2
1
1
0
1
1
0
0
2
0
Diakonoffiana Koçak
-
0
0
0
2/3/4 2
1
1
0
1
0
-
0
0
0
Megalota Diakonoff
1
1
0
1
1
2
1
1
0/1
2
0
-
0
2
0
Costosa Diakonoff
1
0
0
0
3
2
1
1
0
0
0
-
0
3
0
APPENDIX 1 485
5/5/06 2:45:07 PM
106
486
5/5/06 2:45:08 PM
122
123
124
125
126
Williella sauteri Horak
0
0
1
0
-
0
Templemania animosana Busck
0
0
1
0
-
0
Cryptaspasma Walsingham
0
0
0
1
0
0
Collogenes Meyrick
0
0/1
0
1
0
0
Oxysemaphora Diakonoff
0
0
1
1
0
0
Cnecidophora, gen. nov.
0
0
0
1
1
0
Sorolopha Lower
0
0
1
0/1
1
0/3
Sycacantha Diakonoff
0
0/1
1
0/1
0/1
0/3
Atriscripta, gen. nov.
0
0
1
0
-
0
Demeijerella Diakonoff
0
0
1
1
0
1
Lobesia Guenée
0
0/1
0/1
0/1
0
3
Gnathmocerodes Diakonoff
0
0
1
0
-
0
Rhectogonia Diakonoff
0
0
1
0
-
0
Ophiorrhabda Diakonoff
0
0
1
0
-
0
Dudua Walker
0
0
1
1
0
0
Trachyschistis Meyrick
0
1
-
-
-
-
Archilobesia Diakonoff
0
0
1
1
0
0
Euobraztsovia Diakonoff
0
0
1
1
0
0
Eremas Turner
0
0
1
1
0
0
Zomariana Diakonoff
0
1
-
-
-
-
Podognatha Diakonoff
0
0
1
0
-
4
Temnolopha Lower
0
0
1
0
-
4
Diakonoffiana Koçak
0
0
1
1
0/1
4
Megalota Diakonoff
0
0
1
0
-
4
Costosa Diakonoff
0
0
1
1
1
4
APPENDIX 1
120503•Olethreutine Moths 4pp.in486 486
121
120503•Olethreutine Moths 4pp.in487 487
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Gatesclarkeana Diakonoff
1
1
0
1
1
1
1
0
-
1
1/2
1
0
0
0/1
Metrioglypha Diakonoff
1
1
0
1
1
1
1
0
-
1
1/2
1
0
0
0/1
Dactylioglypha Diakonoff
1
2
0/1
0
0
0
1
0
-
1
1
1
0
0
0/1
Statherotis Meyrick
1
1
0
0/1
0/1
1/2
0/1
0
-
1
2/3
0/1
0
0
1
Aterpia Guenée
1
1
0
1
1
1/2
1
0
-
1
3
0/1
0
0
0
Proschistis Diakonoff
1
1
0
1
1
1/2
1
0
-
1
3
1
0
0
0
Rhodacra Diakonoff
0/1
2
0/1
0/1
1
0
1
0
-
1
1
0/1
0
0
1
Bactra Stephens
1/2
1
0
1
1
1/2
0/1
0
-
1
1/2
0
0
0
0
Syntozyga Lower
1/2
1
0
1
1
1/2
0/1
0
-
1
1/2
0/1
0
0
0
Endothenia Stephens
2
1
0
1
1
1
1
0
-
1
2
0/1
0
0
0/1
Acroclita Lederer
2
0
0
1
1
2
0
0
-
0/1
0
0
0
0
0
Tritopterna Meyrick
2
0
0
1
1
2
0
0
-
0/1
0
0
0
0
0
Fibuloides Kuznetsov
2
1
0
1
1
2
0
0/1
0
1
0
0/1
0
0
0
Noduliferola Kuznetsov
1/2
1
0
1
1
2
0
0
-
0
0
1
0
0
0
Rhopobota Lederer
2
1
0
1
1
1
1
0
-
0
0
0
0
0
0
Melanodaedala, gen. nov.
1
2
0
1
1
0
2
0
-
1
0
0
0
0
0
Hermenias Meyrick
2
1
0
1
1
2
0
1
0
1
3
0
0
0
0
Eccoptocera Walsingham
2
1
0
1
1
2
0
1
0
1
1
0
0
0
0
Strepsicrates Meyrick
2
1
0
1
1
1/2
1
1
0/1
1
1
0
0
0
0
Spilonota Stephens
2
1
0
1
1
0/1
1/2
0/1
0
0/1
0/1
0
0
0
0
Holocola Meyrick
2
1
0
1
1
1
1
0/1
0/1
1
1/2
0
0
0
0
Epiblema Hübner
1
2
0
1
1
0
1
0
-
0
0
0
0
0
0
Coenobiodes Kuznetsov
1
1
0
1
1
1
1
0
-
0
0
0
0
0
0
absconditana-group
2
0
0
1
1
2
0
0
-
0
0
0
0
0
0
Crocidosema Zeller
2
1
0/1
0/1
0/1
2
0
0
-
1
1/2
0
0
0
0
APPENDIX 1 487
5/5/06 2:45:09 PM
1
488
5/5/06 2:45:10 PM
17
18
19
20
21
22
23
24
25
26
27
28
29
30
Gatesclarkeana Diakonoff
1
0
0/2
0
1
1
0
0/1
0/1
0
0
0
2
0/1
0
Metrioglypha Diakonoff
0
0
1/2
0
1
1
0
0
1
0
0
0
3
0/1
0
Dactylioglypha Diakonoff
0
0
0/1
0
1
5
0/1
1
1
0/1
0
0
2/3
1
0
Statherotis Meyrick
0
0
1
0
1
0/1
0/1
1
0/1
0
0
0
1/2/3
1/2/3 0
Aterpia Guenée
0
0
0
0
1
1
0
0
0/1
0
0
0
2/3
1/2/3 0
Proschistis Diakonoff
0
0
0/1
0
1
1/4
0
0/1
0
0
0
0
1/2/3
1
0
Rhodacra Diakonoff
0
0
0/1
0
1
0/1
1
1
0
0
0
0
3
0/1
0
Bactra Stephens
0
0
0
0
1
1
0
0
0/1
1
0
0
1
0
0
Syntozyga Lower
0
0
0/1
0
1
1
0
0
0/1
2
0
0/2
1
0/2
0
Endothenia Stephens
0
0
0/1
0
1
1/2
0
0
0
1
0
0
1
0
0
Acroclita Lederer
0
0
1
1
1
3
0
0
1
1
0
0
3
1
2
Tritopterna Meyrick
0
0
1
1
1
3
0
0
1
1/2
1
-
-
1
2
Fibuloides Kuznetsov
0
0/1
1/2
0/1
1
0/1
0
0/1
1
1/2
0/1
0
2
1
0/1/2
Noduliferola Kuznetsov
0
1
1
1
1
1
0
1
0
2
1
-
-
1
2
Rhopobota Lederer
0
0/1
2
0
0/1
1
0
0/1
1
0
1
-
-
1
0/1/2/3
Melanodaedala, gen. nov.
0
0
1
0
1
1
0
1
1
0
0
0
3
0
0
Hermenias Meyrick
0
1
1/2
0
1
1
0
0
0/1
1
0/1
0
3
1
2
Eccoptocera Walsingham
0
1
1
0
0
1
0
0
1
2
1
-
-
0
4
Strepsicrates Meyrick
0
1
0
0/1
0/1
1
0
0
0
1/2
1
-
-
0
0
Spilonota Stephens
0
0/1
1
0
0/1
1
0
0
0/1
0/1
0/1
0
0/1/2/3 0/1
0/1/2
Holocola Meyrick
0
0/1
1
0
0/1
3/5
0
0
1
0/1
0/1
0
2
0
0/2
Epiblema Hübner
0
1
1
0
0
1
0
0
0
1
0
0
1
0
0
Coenobiodes Kuznetsov
0
0
0/1
0
1
1
0
0
0
0
0
0
1
1
0
absconditana-group
0
0
1
0
1
1
0
0
0/1
0
0
0
3
0
0
Crocidosema Zeller
0
0/1
1
0
0/1
2
0
0
1
1
0
0
3
0
0/1/2
APPENDIX 1
120503•Olethreutine Moths 4pp.in488 488
16
120503•Olethreutine Moths 4pp.in489 489
32
33
34
35
36
37
38
39
40
41
42
43
44
45
Gatesclarkeana Diakonoff
2
0
1
3
0/2
0
0
1
0/4
1
0
0
2
0
2
Metrioglypha Diakonoff
2
0
1
0
0
0
0
0
0
0
0
0
2/4
0
2
Dactylioglypha Diakonoff
2
0
1
0/3
0
0
0
0
0
0
0
0
2
0
2
Statherotis Meyrick
2
0
0
0
0
0
0
0
0
0/1
0/1
0
2
0
2
Aterpia Guenée
2
0
0
1
0
0
0
0
5
0
0
0
2
0
2
Proschistis Diakonoff
2
0
0
1
0
0
0
0
5
0
0
0
2
0
1/2
Rhodacra Diakonoff
2
0
1
1
1
0
1
1
0
1
0
0
2
0
2
Bactra Stephens
2
0
0
0
0
0
0
0
0
0
0
0
3/4
0
2
Syntozyga Lower
2
0
0
0
0
0
0
0
0
0
0
0
4
0
1
Endothenia Stephens
2
0
1
3
0
0
0/1
2
4
0
0
0
0/1/2/3/4 0
1
Acroclita Lederer
2
1
0
0
0
0
0
1
0
0/3
1
0
4
1
1/2
Tritopterna Meyrick
2
1
0
0
0
0
0
0
0
0
1
0
2
1
1/2
Fibuloides Kuznetsov
2
1
1
0
0/1
0
0
0
0/1/4 0
1
0
2/4
0
2
Noduliferola Kuznetsov
2
1
1
0
0
0
0
0
0
0
1
0
4
0
2/3
Rhopobota Lederer
2
1
0/1
0
0
0
0
0
0
0
0/1
0
2
0
2
Melanodaedala, gen. nov.
2
0
0
0
0
0
0
0
5
0
1
0
2
0
2
Hermenias Meyrick
0
0/1
1
0
1
0
0
2
1
0
0
0
2
0
2
Eccoptocera Walsingham
2
1
1
0
2
0
1
2
1
0
1
0
3/4
0
2
Strepsicrates Meyrick
1
0
0
0
1
0
0
2
1
0
0
0
2
0
1/2
Spilonota Stephens
2
1
1
0
0/2
0
0
0/2
1
0
0/1
0
2/4
0
2
Holocola Meyrick
0/1/2 1
1
0
2
0
1
2
1
0
0/1
0
2
0
2
Epiblema Hübner
1
1
1
0
0
0
0
0
0
0
0
0
2
0
1/2
Coenobiodes Kuznetsov
2
0
0
0
0
0
0
0
0
0
0
0
2/4
0
2
absconditana-group
0
0
0
0
0
0
0
0
0
0
0
0
2
0
2
Crocidosema Zeller
0
1
1
2
0
0
0
0
5
0
0/1
0
2/4
0
2
APPENDIX 1 489
5/5/06 2:45:11 PM
31
490
5/5/06 2:45:12 PM
47
48
49
50
51
52
53
54
55
56
57
58
59
60
Gatesclarkeana Diakonoff
1/2
0
0
1
1
1
0/1
0/1
0
0
2
0
0
3
0
Metrioglypha Diakonoff
1
0
0
0
1
0
0
0
0
2
2
0
1
0
0
Dactylioglypha Diakonoff
1/2
0
0/1
0
1
0
0
1
0
2
2
0
1
0
0
Statherotis Meyrick
1
0
0
0
1
0
0
0/1
0
2
2
0
0/1
0
0
Aterpia Guenée
0/1/2 0
0
0
1
0
0
0
0
1
2
0
0
1
0
Proschistis Diakonoff
1
0
0
0
1
0
0
0/1
0
1
2
0
0
1
0
Rhodacra Diakonoff
1
0
0
1
1
0
0
0
0
1
2
0
0
0/1
0
Bactra Stephens
0/1
0
0
0/1
0
0
0
0
0
2
2
0
0
2
0
Syntozyga Lower
0/1
0
0
0
0
0
0
0
0
1
2
0
0
0/2
0
Endothenia Stephens
1
0
0
1
1
0
0
0
0
2
2
0
0
0
0
Acroclita Lederer
2
1
0/1
0
0
0
0
0
0
2
2
1
0
0
0
Tritopterna Meyrick
1/2
1
0
0
0/1
0
0
0
0
2
2
1
0
0
0
Fibuloides Kuznetsov
2
1
0
1
1
0
0/1
0
0
2
2
0/1
0
0
0
Noduliferola Kuznetsov
2/3
1
0/1
2
1
0
0
0
0
2
2
1
3
0
4
Rhopobota Lederer
2
0
0
1
1
0
0
0
0
2
2
1
3
0
0
Melanodaedala, gen. nov.
2
1
0
1
0
0
0
0
0
1/2
2
1
0
0
0
Hermenias Meyrick
2
1
0
1
0
0
0
0
0
2
2
0/1
0
0
0
Eccoptocera Walsingham
3
1
0
1
0/1
0
0
0
0
2
2
1
0
0
0
Strepsicrates Meyrick
2
1
0
1
1
0
0
0
0
2
2
1
0
0
0
Spilonota Stephens
2/3
1
0
1
1
0
0
0
0
2
2
1
0
0
0
Holocola Meyrick
2/3
1
0
1
0/1
0
0
0
0
2
2
1
0
0
0
Epiblema Hübner
2
0
0/1
1
0/1
0
0
0
0
2
2
1
0
0
0
Coenobiodes Kuznetsov
2
1
0/1
1
1
0
0
0
0
2
2
1
0
0
0
absconditana-group
2
1
0/1
1
1
0
0
0
0
2
2
1
0
0
0
Crocidosema Zeller
2/3
0
0
1
1
0
0
0/1
0
2
2
1
0
0
0
APPENDIX 1
120503•Olethreutine Moths 4pp.in490 490
46
120503•Olethreutine Moths 4pp.in491 491
62
63
64
65
66
67
68
69
70
71
72
73
74
75
Gatesclarkeana Diakonoff
0
0
0/1
0
1
1
0
0
1
0/3
5
0
0
0
2
Metrioglypha Diakonoff
0
2
1
1
0
0
0
0
1
0/1
2/3
1
0
0
0/1/2
Dactylioglypha Diakonoff
0
0
1
1
0
0
0
0
1
0
5
1
0
1
1
Statherotis Meyrick
0
0/2
1
1
0
0
0
0
1
0/1/3/5 2
1
0
0
0/1
Aterpia Guenée
0
0
1
0
1
0
0
0
1
2/3
5
0/1
0
0
0
Proschistis Diakonoff
0
0/2
0/1
0
0
0
0
1
-
-
-
-
0
1
2
Rhodacra Diakonoff
0
2
1
1
1
0
0
0
1
1/4
0/2
1
0
0
0/1
Bactra Stephens
1
0/2
0/1
0/1
1
1
0/1
0
1
1
4
1
0
0
0/1
Syntozyga Lower
1
0/2
0/1
0
2
0/1
0
1
-
-
-
-
0
0
1
Endothenia Stephens
1
2
0
1
0
0
1
0
1
4
2
1
0
0
0/1/2
Acroclita Lederer
1
1
0
1
0
1
0
0/1
0
2/6
5
1
0
1
2
Tritopterna Meyrick
1
1
0
1
0
1
0
0
1
0
5
1
0
1
1/2
Fibuloides Kuznetsov
0
0
1
0/1
0/1
1
0
0
0/1
5
0
0
0
0
1
Noduliferola Kuznetsov
0
0
1
0
1
1
0
0/1
0
0
0
0
0
0
0
Rhopobota Lederer
0
1
0
0
0
1
0
0
0
0/3
0
0
0
0/1
1/2
Melanodaedala, gen. nov.
1
1
1
0
0
1
0
0
0/1
5
0
1
0
1
2
Hermenias Meyrick
0/1
1/2
1
0/1
0
1
0
0
1
0/1
0
0
0
1
0/1
Eccoptocera Walsingham
0/1
0/2
1
0
0
1
0
0
1
4
0
1
0
0
0
Strepsicrates Meyrick
0/1
0/1/2 0
0/1
0
1
0
0/1
0
0
3
0
0
0
0
Spilonota Stephens
1
0/1/2 1
1
0
1
0
0/1
0/1
0/3/5
0
1
0
1
2
Holocola Meyrick
0/1
0/1/2 1
0/1
0
1
0
0/1
0/1
0/1/5
0
0/1
0
0/1
2
Epiblema Hübner
0/1
0/1
0
1
0
1
0
0
1
2/3
3
0
0
0
1/2
Coenobiodes Kuznetsov
0/1
1
0/1
1
1
1
0
0
0/1
2/5
0
0
0
0/1
1/2
absconditana-group
0/1
0/2
0/1
1
1
1
0
0
1
5
0
0/1
0
0
0/1/2
Crocidosema Zeller
0/1
2
0
0
1
1
0
0
1
0/3
0/3
1
0
0
1/2
APPENDIX 1 491
5/5/06 2:45:13 PM
61
492
5/5/06 2:45:14 PM
77
78
79
80
81
82
83
84
85
86
87
88
89
90
Gatesclarkeana Diakonoff
2
0
0
0
0
0
0
1
1
0
1
0
0
0
0/1
Metrioglypha Diakonoff
2/4
0
0
0/1
0
0
0
0
2
0
0
0
0/1
0
0
Dactylioglypha Diakonoff
0/2
0
0
1
2/3
0
1/3
0
1/2
1
0
0
1
0
0
Statherotis Meyrick
2
0
0
1
0/2
0
0
0
2
0/1
0/1
0/7
1
0
0
Aterpia Guenée
2/4
0
0
1
2/3
0
1
0
1
0
0
0
1
0
0
Proschistis Diakonoff
0/1
0
0
1
3
0
1
0
2
0
0
0
1
0
0
Rhodacra Diakonoff
2
0
0
0/1
3
0
1
0
2
0
1
7
1
0
0
Bactra Stephens
2
0/1
0
0
0
0
0/1
0
2
0
1
0/3/5 0/1
0
1
Syntozyga Lower
4
0
0
0/1
0/3
0
0/1
0
2/3
0
0/1
0/3
0/1
0
1
Endothenia Stephens
2
0
0
0
0
0
0
0
1/2/3 0
0
2
0
0
0
Acroclita Lederer
5
0
0
0
1
0/1
0
0
2
0
1
0
0
0
0/1
Tritopterna Meyrick
0
0
0
0/1
1
0
0
0
2
0
1
6
0
0
0/1
Fibuloides Kuznetsov
2
0
0
1
5
0
0
0
3
0
1
2/5/7 0/1/2 0
1
Noduliferola Kuznetsov
3/4
0
0
0/1
1
0
0
0
2
1
1
0/5
1
0
0/1
Rhopobota Lederer
1/2
0
0
0/1
6
0
0
0
3
1
1
4/7
0
0
0
Melanodaedala, gen. nov.
2
0
0
1
5
0
0
0
3
0
1
5/6/7 0/1
0
1
Hermenias Meyrick
2
0
0
0
0
0
0
0
3
0
1
4/5/6 0/2
0
1
Eccoptocera Walsingham
2/4
0
0
0
0
0
0
0
3
0
1
0/4/7 2
0
1
Strepsicrates Meyrick
0/4
0
0
0/1
0/5
0
0
0
2
0
1
7
1
0/1
Spilonota Stephens
1/2
0
0
0
0
0
0
0
2/3
0
1
5/6/7 1/2
0
0/1
Holocola Meyrick
2/6
0
0
0/1
0
0
0
0
2/3
0
1
1/5
0/1
0
1
Epiblema Hübner
2
0
0
0/1
3
0
1
0
2
1
1
0
0
0
0
Coenobiodes Kuznetsov
0/1
0
0
0/1
0
0
0
0
2
1
1
0
0
0
0
absconditana-group
2/4
0
0
0/1
3
0
1
0
2
1
1
0
0
0
1
Crocidosema Zeller
2
0
1
-
-
-
0
0
2
0
1
0/5
0
0
1
2
APPENDIX 1
120503•Olethreutine Moths 4pp.in492 492
76
120503•Olethreutine Moths 4pp.in493 493
92
93
94
95
96
97
98
99
100
101
102
103
104
105
Gatesclarkeana Diakonoff
0
4
1
1
1
0
-
1
0
0
0
4
1
2
-
Metrioglypha Diakonoff
0/1
1
2
0
-
0
-
0
0
1
0
3
1
2
-
Dactylioglypha Diakonoff
0
1
2
0
-
0
-
0
0/1
0
0
3
1/2
2
-
Statherotis Meyrick
0/1
1
2
0
-
0
-
0/3
0/1
0
0
3/4
1
2
-
Aterpia Guenée
0
1
2
0
-
0
-
0
0
2
0
2
5
2
-
Proschistis Diakonoff
0
1
2
0/1
1
1
1
0
3
2
0
2
5
2
-
Rhodacra Diakonoff
0
1
2
0
-
0
-
0
0/1
0
0
2
1
2
-
Bactra Stephens
1
4
1
0/1
1
0
-
0
0
0
0
1/2
1
2
-
Syntozyga Lower
1
4
1
0/1
1
0
-
0
0
0
0/1
2/3/4 2
2
-
Endothenia Stephens
0
4
1
1
1
0
-
0
0
0
0
1/2
1
0/2
0/2
Acroclita Lederer
0
1
1
1
3
0
-
0
0
3
0
2
1/5
0
1
Tritopterna Meyrick
0
1
1
1
3
0
-
0
2
3
0
2
2
2
-
Fibuloides Kuznetsov
0/1
1
1
0/1
3
0
-
0
0
3
0
3
2
0
1
Noduliferola Kuznetsov
0
1
1
0
-
0
-
0
1/3
0
3
2
2
0
1
Rhopobota Lederer
0
3
1/2
0
-
0
-
0
0
0/3
0
3
2
0
1
Melanodaedala, gen. nov.
1
1
1
0
-
0
-
0
1
3
2
3
2
0
1
Hermenias Meyrick
0
1
1
0
-
0
-
0
0
0
0
3
1
0
1
Eccoptocera Walsingham
0
4
1
0
-
0
-
0
0
0
0
3
1
0
1
Strepsicrates Meyrick
0
1
1
0
-
0
-
0
0
0
0
2
2
0
1
Spilonota Stephens
0
1
1
0
-
0
-
0
0
0/4
0
2/3/4 1
0/2
1
Holocola Meyrick
0
1
1
0
-
0
-
0
0
0
0
3
4/7
0
1
Epiblema Hübner
0
1
1
0
-
0
-
0
2
0
0
2
2
0
1
Coenobiodes Kuznetsov
0
1
1
0
-
0
-
0
0
1
0
2
2
0
1
absconditana-group
0
1
1
0
-
0
-
0
0/2
0
0
2
1
0
1
Crocidosema Zeller
0
1
1
0
-
0
-
0
2
0/4
0
2
8
0
1
APPENDIX 1 493
5/5/06 2:45:16 PM
91
494
5/5/06 2:45:17 PM
107
108
109
110
111
112
113
114
115
116
117
118
119
120
Gatesclarkeana Diakonoff
-
0
0
0
4
2
1
1
0
3/4
0
-
0
1/2
0
Metrioglypha Diakonoff
-
0
0
0
3
1
1
2
0
1
0
-
0
3
0
Dactylioglypha Diakonoff
-
0
0
0/1
3
1
1
1
0
1
0
-
0
2/3
0
Statherotis Meyrick
-
0
0
0
2/3
1
1
2
0
1
0
-
0
2/3
0
Aterpia Guenée
-
0
0
0
2
1
1
2
0
0
1
0
0/1
3
0
Proschistis Diakonoff
-
0
0
0
1
0/1
1
2
0
0/3
1
0
1
2
0
Rhodacra Diakonoff
-
0
0
0
2/3
2
1
2
0
0/1
0/1
0
0
2
0
Bactra Stephens
-
0/1
0
1
1/4
0/1
0/1
1/2
0/1
3/4
0/1
0
0
2/3
0
Syntozyga Lower
-
0
0
0
4
0/1
1
2
0
4
0
-
0
2/3
0
Endothenia Stephens
1
0
0
0
0/2
0/1/2 0
2
0
0
0
-
0
2
0
Acroclita Lederer
0
0
0
0
3
1
1
1
1
4
1
0
0
2
0
Tritopterna Meyrick
-
1
0
1
3
1
1
1/2
1
4
1
0
0
2
0
Fibuloides Kuznetsov
0
1/3
0
1
2
1
1
1
1
0/2
1
2
0
2
0
Noduliferola Kuznetsov
0
0
0
0
2
2
1
0/1
0
0/2
1
0/2
0
0
0
Rhopobota Lederer
0/1
1
0
0/1
2/3
1
1
1/2
0/1
2/3
1
2
0
2/3
0
Melanodaedala, gen. nov.
0
1
0
1
1
2
1
1
1
3
1
0
0
2
2
Hermenias Meyrick
0
2
0
1
1
2
0
1
0
0/1
1
0
0
1
0
Eccoptocera Walsingham
0
1
0
1
1
2
0
0/1
0/1
0
1
0
0
2
3
Strepsicrates Meyrick
0
1
0
1
1/2
2
0
1
0
2
1
0
0
1/2
0
Spilonota Stephens
0
1
0
1
1
2
0
1
0
2
1
0
0
1/2
0/3
Holocola Meyrick
0
2
0
1
1
2
0
1/2
0
0/2
1
0
0
0/1/2 0/4
Epiblema Hübner
0
1
0
0
0/2
1
1
1
0
2
1
0
0
1
0
Coenobiodes Kuznetsov
0
1/3
0
0/1
2/3
2
1
1
0
2
1
0
0
0/1
0
absconditana-group
1
0
0
0/1
1/3
2
0/1
1/2
0/1
4
1
0
0
3
0
Crocidosema Zeller
0
0
0
1
1
2
1
1
0
2/4
1
0
0
2
0
APPENDIX 1
120503•Olethreutine Moths 4pp.in494 494
106
120503•Olethreutine Moths 4pp.in495 495
122
123
124
125
126
Gatesclarkeana Diakonoff
0
0
1
0
-
3/4
Metrioglypha Diakonoff
0
0
1
1
0
4
Dactylioglypha Diakonoff
0
0
1
1
0
4
Statherotis Meyrick
0
0
1
1
0/1
4
Aterpia Guenée
0
0
1
0/1
0/1
4
Proschistis Diakonoff
0
0
1
0
-
4
Rhodacra Diakonoff
0
0
1
1
0
4
Bactra Stephens
0
0
1
0
-
2
Syntozyga Lower
0
1
-
-
-
-
Endothenia Stephens
0
0
1
0
-
2
Acroclita Lederer
0
0
1
1
0
0
Tritopterna Meyrick
0
1
-
-
-
-
Fibuloides Kuznetsov
0
0
1
1
0
0
Noduliferola Kuznetsov
1
0
1
1
0/1
0
Rhopobota Lederer
0
0
1
0/1
1
0
Melanodaedala, gen. nov.
1
0
0/1
1
0
0
Hermenias Meyrick
1
1
-
-
-
-
Eccoptocera Walsingham
0
0
0
1
0
0
Strepsicrates Meyrick
1
1
-
-
-
-
Spilonota Stephens
1
0
1
1
0
0
Holocola Meyrick
1
0/1
1
1
0
0
Epiblema Hübner
0
0
1
1
0
0
Coenobiodes Kuznetsov
0
0
1
1
0
0
absconditana-group
0
0
1
1
0
0/1
Crocidosema Zeller
1
0
1
1
0
0
APPENDIX 1 495
5/5/06 2:45:17 PM
121
496
5/5/06 2:45:19 PM
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Heleanna Clarke
3
1
0
1
1
1
1
0
-
0
0
0
0
0
0
Icelita Bradley
3
2
1
0
0/1
0
2
0
-
1
1/2
0
0
0
0
Eucosmophyes Diakonoff
1
2
0
1
1
1
1
0
-
0
0
0
0
0
0
Herpystis Meyrick
1
1
0
0/1
0/1
1
1
0
-
0
0
0
0
0
0
Whittenella, gen. nov.
2
2
0
1
1
1
1
0
-
0
0
0
0
0
0
Aglaogonia, gen. nov.
2
1
0
1
1
1
1
0
-
0
0
0
1
0
0
Anathamna Meyrick
2
1
0
1
1
1
1
0
-
1
0
0/1
0
0
1
Ancylis Hübner
2
1
0
1
1
1
1
0
-
0/1
0
0
0
0
0
Ancylophyes Diakonoff
2
0
0
1
1
2
0
0
-
0
0
0
1
0
0
Anthozela Meyrick
1
2
0
0
0
1
1/2
0
-
1
1
0
0
0
0
Balbidomaga Diakonoff
2
0
0
1
1
1
1
0
-
0
0
0
0/1
0
0
Cyphophanes Meyrick
2
1
0
1
1
0/1
1
0
-
1
1
1
0
0
0
Eucosmogastra Diakonoff
1/2
1
0/1
0/1
0/1
0/2
1/2
0
-
1
1
1
0
0
0
Helictophanes Meyrick
2
1
0
1
1
2
0
0
-
0/1
1
1
0
0
0
Irianassa Diakonoff
0/1
2
1
0
0
2
2
0
-
1
1
0
0
0
0
Loboschiza Diakonoff
1/2
1
0/1
0/1
0/1
1
1
0
-
1
1/2
0
0
0
1
Metaselena Diakonoff
1
2
0/1
0/1
0/1
1
1
0
-
1
1
0
0
0
1
Oriodryas Turner
1
2
0/1
0
0
1
1
0
-
1
1
1
0
0
0
Periphoeba Bradley
2
0
0
1
1
2
0
0
-
0
0
0
1
0
0
Pseudancylis, gen. nov.
2
2
0
1
1
1
1
0
-
0
0
0
1
0
0
Pternidora Meyrick
0/1
2
0
1
1
0
2
0
-
1
0
0
0
0
0
Tetramoera Diakonoff
2
1
0
1
1
2
0
0
-
0
0
0
0
0
0
Thysanocrepis Diakonoff
1
1
0
1
1
1
1
0
-
1
1
0
0
0
0
Toonavora, gen. nov.
2
2
0
1
1
1
1
0
-
1
1
0
0
0
0
Grapholita zapyrana Meyrick
1
1
1
2
2
1
1
0
-
1
0
0
0/1
0
0
APPENDIX 1
120503•Olethreutine Moths 4pp.in496 496
1
120503•Olethreutine Moths 4pp.in497 497
17
18
19
20
21
22
23
24
25
26
27
28
29
30
Heleanna Clarke
0
0/1
1
1
1
3/5
1
1
1
1/2
0/1
0
3
1/3
2
Icelita Bradley
0
0
1/2
0
1
1
0/1
0/1
1
1
0/1
0
3
0/1
0/1/2
Eucosmophyes Diakonoff
0
0
0
0
1
0/1
0
0
1
1
0/1
0
3
0/1
0
Herpystis Meyrick
0
0
1
0
1
2
0
0
1
2
1
-
-
0/1
0
Whittenella, gen. nov.
0
0
1
1
1
0/1
0
0
1
0
1
-
-
0
2
Aglaogonia, gen. nov.
0
0
1
0
1
3
0
0
1
2
1
-
-
0
2
Anathamna Meyrick
0
0
1
0
1
3
1
1/2
1
0
1
-
-
0
2
Ancylis Hübner
0
0
1
0
0/1
3
0
0
1
1
0/1
0
2
0
1/2
Ancylophyes Diakonoff
0
0
1
0
1
3
0
1
1
0
1
-
-
1
2
Anthozela Meyrick
0
0
0/1
0
1
1
0/1
0/1
0
0
0
0
1
0
0
Balbidomaga Diakonoff
0
2
1
0
1
3
0
1
1
2
0
0
3
1
1/2
Cyphophanes Meyrick
0
0
1
0
1
1
2
0
0
0
0
0
0
0
0
Eucosmogastra Diakonoff
0
0
0/1
0
1
1/2
0
0
0/1
0
0
0
1
0
0
Helictophanes Meyrick
0
0
0/1
0
1
4
0/1
1
0
0/1
0/1
0
2
1
0
Irianassa Diakonoff
0
0
0
0
1
1
3
1
1
0
0
0
2
0
0
Loboschiza Diakonoff
0
0
0/1
0
1
1
1
0/1
0
0
0/1
0/2
3
0
0/1
Metaselena Diakonoff
0
0
1/2
0
1
3/5
1/2
1/2
0/1
0
1
-
-
4
2
Oriodryas Turner
0
0
0
0
1
4
0
0
0
0
0
0/1
2
1
0
Periphoeba Bradley
0
0
0/1
0
1
1
0
0
1
2
1
-
-
1
1
Pseudancylis, gen. nov.
0
0
1
0
1
5
0/3
1
1
2
1
-
-
0
2
Pternidora Meyrick
0
0
0
0
1
4
0
0
0
0
0
0
1
0
0
Tetramoera Diakonoff
0
0
1
0
1
1/3
0/1
1
1
0
0
0
3
1
0
Thysanocrepis Diakonoff
1
0
1
0
1
6
0
0
0/1
0
1
-
-
0
1/2
Toonavora, gen. nov.
0
0
0/1
0
0
1
0
0
1
0
0
0
2
0/1
0
Grapholita zapyrana Meyrick
1
0
0
0
0
4
0
0
1
0
0
0
1
0
0
APPENDIX 1 497
5/5/06 2:45:20 PM
16
498
5/5/06 2:45:21 PM
32
33
34
35
36
37
38
39
40
41
42
43
44
45
Heleanna Clarke
2
1
0
0
0
0
0
0
0
0
0/1
0
2
0
2
Icelita Bradley
1
1
1
0
2
0
0
0/2
1
0
1
0
2/4
0
2
Eucosmophyes Diakonoff
1
0
0
0
2
0
1
2
1
0
0
0
2/4
0
2
Herpystis Meyrick
2
0/1
1
0
0
0
0
0
0
0
0/1
0
1/2
0
3
Whittenella, gen. nov.
2
1
1
0
0
0
0
0
0
0
1
1
2
0
2
Aglaogonia, gen. nov.
5
1
1
0
0
0
0
0
0
0
0
0
2
0
2
Anathamna Meyrick
1/2
1
1
0
2
1
1
0
0
0
0
0
2
0
1/2
Ancylis Hübner
1/2
0/1
0
0
1/2
0/1
1
2
1
0
0
0
2
0
1/2
Ancylophyes Diakonoff
5
0
1
0
0
0
1
1
0
0
0
0
2
0
2
Anthozela Meyrick
2
0
0
0
0
0
0
0/2
0/1
0
0
0
3/4
0
0
Balbidomaga Diakonoff
2
0
0
0
0
0
0
0
6
1
0/1
0
2
0
2
Cyphophanes Meyrick
2
0
0
0
0
1
0
0
0
0
0
0
2
0
0/2
Eucosmogastra Diakonoff
2
0
0
3
0
0
0/1
0/2
1/7
0
0
0
1/2
0
2
Helictophanes Meyrick
0/1/2 0
0
0
0
0/1
0
0
0
0
0
0
2/3/4 0
2
Irianassa Diakonoff
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
Loboschiza Diakonoff
2
0
0
0
0
0
0
0
0
0
0
0
2
0
0/1
Metaselena Diakonoff
0
0/1
1
0
1/2
0
0/1
2
0
0
0
0
2
0
1
Oriodryas Turner
1
1
1
0
2
0
1
2
1
0
0
0
2
0
2
Periphoeba Bradley
2
0
0
0
0
0
0
2
1
0
0
0
2
0
2
Pseudancylis, gen. nov.
2
0
1
0
0
0
0
0
0
0
1
0
2
0
1
Pternidora Meyrick
2
0
0
0
0
0
0
0
0
0
0
0
0/2
0
0
Tetramoera Diakonoff
0/1
0
0
0
0
0
0
0
0
0
0
0
2
0
2
Thysanocrepis Diakonoff
0
0
0
0
1
0/1
1
2
1
0
0
0
2
0
2
Toonavora, gen. nov.
2
1
1
0
0
0
0
0
0
0
0
0
2
0
1
Grapholita zapyrana Meyrick
1
0
0
1
0
0
0
0
0
0
0
0
1/2
0
0
APPENDIX 1
120503•Olethreutine Moths 4pp.in498 498
31
120503•Olethreutine Moths 4pp.in499 499
47
48
49
50
51
52
53
54
55
56
57
58
59
60
Heleanna Clarke
2
1
0
1
1
0
0
0
0
2
2
1
3
0
0
Icelita Bradley
2/3
1
0
2
0/1
0
0
0
0
2
2
1
2
0
0
Eucosmophyes Diakonoff
2
1
0
1
1
0
0
0
0
2
2
1
0
0
0
Herpystis Meyrick
4
1
0
1
1
0
0
0
0
2
2
1
0
1
0
Whittenella, gen. nov.
2
1
0
3
0
0
0
0
0
2
2
1
0/3
0
0
Aglaogonia, gen. nov.
2
0
0
0
1
0
0
0
0
1
2
1
0
0
1
Anathamna Meyrick
1
0
0
1
1
0
0
0
1
2
2
1
0
0
0
Ancylis Hübner
2/3
0
0
1
0/1
0
0
0
0
0/1
2
1
0
0
0
Ancylophyes Diakonoff
2/3
0
0
0
0
0
0
0
0
2
2
1
0
0
0
Anthozela Meyrick
1/2/3 0
0
2
1
0
0
0
0
1
2
1
0
0
0/1
Balbidomaga Diakonoff
1/2
1
0
1
0/1
0
1
0
0
1
2
1
0
0
1
Cyphophanes Meyrick
1/2
0
0
1
1
0
0
0
0
1
2
1
0
0
0
Eucosmogastra Diakonoff
1
0
0
1
1
0
0/1
0
0
2
2
1
3
0
0/1
Helictophanes Meyrick
2
0
0
1
1
0
0
0
0
0/1
2
1
1
0
1
Irianassa Diakonoff
1
0
1
1
1
0
0
0
0
2
2
1
0
0
0/1
Loboschiza Diakonoff
1/2
0
0/1
1/2
0/1
0
0
0/1
0/1
2
2
1
0
0
0
Metaselena Diakonoff
0/2/4 0
0
2
1
0
0
0
1
2
2
0
0
0
0
Oriodryas Turner
0
0
0
1
1
0
0
2
0
2
2
1
0
0
0
Periphoeba Bradley
2
0
0
1
1
0
0
0
0
0/1
2
1
3
0
1
Pseudancylis, gen. nov.
2
0
0
0
0
0
0
0
0
1
2
1
3
1
0
Pternidora Meyrick
1
0
1
2
1
0
0
0
0
1
2
1
3
0
1
Tetramoera Diakonoff
0/1/2 0
0
1
1
0
0
0
0
2
2
0/1
0
0
0
Thysanocrepis Diakonoff
1
0
0
1
1
0
0
0
0
1
2
0
0
0
1
Toonavora, gen. nov.
2
0
0
1
1
0
0
0
0
2
2
1
0
1
0
Grapholita zapyrana Meyrick
2
0
0
0
1
0
0
0
0
4
0
1
0
1
0
APPENDIX 1 499
5/5/06 2:45:22 PM
46
500
62
Heleanna Clarke
0/1
Icelita Bradley
63
5/5/06 2:45:23 PM
64
65
66
67
68
69
70
71
72
73
74
75
0/1/2 0/1
0
0
1
0
1
-
-
-
-
0
1
2
0
0/2
1
0
2
1
0
1
-
-
-
-
0
0
1/2
Eucosmophyes Diakonoff
0
2
0/1
0
1
1
0
1
-
-
-
-
0
0
0
Herpystis Meyrick
1
2
0
0
1
1
0
1
-
-
-
-
0
0
0/1
Whittenella, gen. nov.
1
1
1
1
0
1
0
1
-
-
-
-
0
1
2
Aglaogonia, gen. nov.
1
2
1
0
1
0
0
1
-
-
-
-
0
0
2
Anathamna Meyrick
0
0/2
1
0
1
1
0
0/1
1
0
5
0
0
0
1
Ancylis Hübner
0
0/2
1
0
1
0
0
0/1
0/1
0/5
0
0/1
0
0
2
Ancylophyes Diakonoff
0
2
1
0
1
0
0
1
-
-
-
-
0
0
0/1
Anthozela Meyrick
0/1
2
0
1
1
0/1
0
0/1
0
6
0
0
0
0
0
Balbidomaga Diakonoff
0/1
0
1
0
0
0
0
0
1
4/5
0
1
0/1
0
0
Cyphophanes Meyrick
0
1
1
1
1
0
0
0
0/1
3/5
6
0
0
0
0
Eucosmogastra Diakonoff
0/1
0/2
0
0
1
0/1
0
0
1
2
2
1
0
0/1
1
Helictophanes Meyrick
0
0
0
0
2
0/1
0
1
-
-
-
-
0
0
0/1
Irianassa Diakonoff
0/1
0
0/1
0
0/1
0
0
1
-
-
-
-
0
0
0/1
Loboschiza Diakonoff
0
0/2
0/1
0
1
0
0
1
-
-
-
-
0
0
0/1
Metaselena Diakonoff
0
0
1
0
2
1
0
0/1
1
2/5
0
0
0
0
1
Oriodryas Turner
1
2
0
1
0
0
0
0
1
1
2
1
0
0
1
Periphoeba Bradley
0
2
1
0
1
1
0
0
1
5
0
0
0
0
0
Pseudancylis, gen. nov.
0
0/2
0
0
2
1
0
0/1
1
7
0
0
0
0/1
2
Pternidora Meyrick
0
0/2
1
0
2
1
0
0/1
1
2
0/3
0
1
-
-
Tetramoera Diakonoff
0
2
1
0
2
1
0
0
1
3/5
7
0/1
0
0
0
Thysanocrepis Diakonoff
1
2
0
0
2
0
0
0
1
5
7
0
0
0
1/2
Toonavora, gen. nov.
0
0/2
1
0
2
1
0
0
1
3/5
7
0
0/1
0
0
Grapholita zapyrana Meyrick
1
0
1
0
0
1
0
1
-
-
-
-
1
-
-
APPENDIX 1
120503•Olethreutine Moths 4pp.in500 500
61
120503•Olethreutine Moths 4pp.in501 501
77
78
79
80
81
82
83
84
85
86
87
88
89
90
Heleanna Clarke
0/1
0
0
0
0
0
0
0
1
0
0/1
0
0/1
0
0
Icelita Bradley
1/2
1
0
0/1
1
0
0
0
2
0
1
0/3
0
0
1
Eucosmophyes Diakonoff
3
0
0
0/1
3
0
0
0
3
0
1
5/6
0
0
1
Herpystis Meyrick
2
0
0
1
3
0
1
0
3
0
0/1
0
1
0
0/1
Whittenella, gen. nov.
0
0
0
0
3
0
1
0
2
0
0
0
0
0
0
Aglaogonia, gen. nov.
2
1
0
0
0
0
0/1
0
2
0
1
7
2
1
0
Anathamna Meyrick
2
0
0
0
0
0
0
0
3
0
1
5
3
2
1
Ancylis Hübner
2
0
0
0/1
0
0
0
0
2/3
0
1
2/5/7 0
0
1
Ancylophyes Diakonoff
4
0
0
1
3
0
0
0
2/3
0
1
0
0
0
1
Anthozela Meyrick
4
0
0
1
8
2
0
0
1
0
0/1
0
0
0
0
Balbidomaga Diakonoff
4
0
0
1
5
0
0
0
2
0
0/1
0
0
0
0/1
Cyphophanes Meyrick
2
0
0/1
0/1
0
0
2
0
3
0
0
0/3
3
1
0
Eucosmogastra Diakonoff
2
1
0
0/1
1
1
0
0
2/3
0
0/1
0/2
0
0
0/1
Helictophanes Meyrick
2/4
0
0
0
3
0
1
0
3
0
1
0
3
1/2
1
Irianassa Diakonoff
2
0
0
1
3
0
1
0
3
0
0/1
1/5
1/2
1
0
Loboschiza Diakonoff
2/3
0
0
0/1
3
0
1
0
3
0
0/1
0/7
0/2
0/1/2 0/1
Metaselena Diakonoff
2
0
0
0/1
0
0
0
0
3
0
1
2
3
1
1
Oriodryas Turner
2
0
0
1
8
2
3
0
3
0
1
0
0
0
1
Periphoeba Bradley
2
0
0
0/1
0
0
0
0
3
0
1
5
0
0
1
Pseudancylis, gen. nov.
2
0
1
-
-
-
0
0
3
0
1
1
2
1
1
Pternidora Meyrick
-
-
0
0/1
1/5
0
0
0
3
0
1
3
0
0
0
Tetramoera Diakonoff
0
0
0
1
0
0
1
0
3
0
1
7
0
0
1
Thysanocrepis Diakonoff
2
0
1
-
-
-
0
0
3
0
1
7
2
1
1
Toonavora, gen. nov.
4
0
0
0/1
0
0
0
0
3
0
1
7
2
1/2
0/1
Grapholita zapyrana Meyrick
-
-
0
1
3
0
1
0
2
0
1
4/5
0
0
1
APPENDIX 1 501
5/5/06 2:45:24 PM
76
502
5/5/06 2:45:25 PM
92
93
94
95
96
97
98
99
100
101
102
103
104
105
Heleanna Clarke
1
1
2
0/1
0
0
-
0
0
0
0
2/3
2
0
1
Icelita Bradley
0
4
1
1
1
0
-
0
4
4
0
3
1
0/1
1
Eucosmophyes Diakonoff
0
1/4
1
0
-
0
-
0
0
0
0
4
1
0
1
Herpystis Meyrick
0
4
1
0
-
0
-
0
0
0
0
4
1
0
1
Whittenella, gen. nov.
0
1
1
0
-
0
-
0
0
0
1
2
1
0
1
Aglaogonia, gen. nov.
0
1
2
0
-
0
-
0
0
0
0
3
1
2
-
Anathamna Meyrick
0/1
1
2
0
-
0
-
0
0
0
0
4
1
0
1
Ancylis Hübner
0
1
1
0
-
0
-
0
0
0
0
3/4
2
0
1
Ancylophyes Diakonoff
0
1
2
0
-
0
-
0
0
0
0
3
2
2
-
Anthozela Meyrick
0
2
1
1
0/2
0
-
0
0
0
0
1/5
7
0
0/1
Balbidomaga Diakonoff
0/1
2
1
0
-
0
-
0
0
0
0
3
1
2
-
Cyphophanes Meyrick
1
1
2
0
-
0
-
0
0
4
0
4
1
0
1
Eucosmogastra Diakonoff
0
1
1
1
3
0
-
0
0
0
0
3
1
0
1
Helictophanes Meyrick
0
1
1
0
-
0
-
0
0
0
0
3
1
0
1
Irianassa Diakonoff
1
1
1
0
-
0
-
0
0
0
0
3/4
1
2
-
Loboschiza Diakonoff
0/1
1
2
0
-
0
-
0
0
0/2
0
3/4
7
0
1
Metaselena Diakonoff
1
1
2
0
-
0
-
0
0
0
0
4
1
0
1
Oriodryas Turner
0
1
1
0
-
0
-
0
0
0
0
1/5
7
0
1
Periphoeba Bradley
1
1
1
0
-
0
-
0
0
4
0
1/5
1
0
3
Pseudancylis, gen. nov.
0
1
2
0
-
0
-
0
0
0
0
3
1
0
1
Pternidora Meyrick
0
2
2
0
-
0
-
0
0
0
4
4
1
0
0/2
Tetramoera Diakonoff
0
1
2
1
0
0
-
0
0
0
0
2
1
0
0
Thysanocrepis Diakonoff
0
1
1
0
-
0
-
0
0
0
0
2
1
0
0
Toonavora, gen. nov.
1
1
1
0
-
0
-
0
0
0
0
2
2
0
0
Grapholita zapyrana Meyrick
0
1
2
0
-
0
-
0
0
0
0
3
0
0/1
0
APPENDIX 1
120503•Olethreutine Moths 4pp.in502 502
91
120503•Olethreutine Moths 4pp.in503 503
106
107
108
109
110
Heleanna Clarke
0
0/1
0
0
Icelita Bradley
0
0/2
0
Eucosmophyes Diakonoff
0
1
Herpystis Meyrick
0
Whittenella, gen. nov.
111
112
113
114
115
0/1/3 0/2
1
1/2
0
0
4
1
1
1
0
1
1
1
0
3
0
1
1/3
2
0
2
0
1
1
Aglaogonia, gen. nov.
-
0
0
0
Anathamna Meyrick
0
0
0
Ancylis Hübner
0
0
Ancylophyes Diakonoff
-
Anthozela Meyrick
116
118
119
120
2/3/4 1
0
0
1/2
0
0
1/2/3 1
0
0
1/2
1
1
0
2
1
0
0
2
1
1
1
0
2
1
0
0
0
0
1
1
2
0
4
1
0
0
3
0
3
0/1
1
1
0
1/3
0
-
0
0/1
0
0
0/3
0/1
1
1
0
0/3
0
-
0
0
3
0
0
1
2
1
1
0
0
0
-
0
0
0
3
0
1
1
2
0
1
0
4
0
-
0
0/1
0
0
0
0
0
1/3
2
0
1/2
0
0/4
1
0
0
0
0
Balbidomaga Diakonoff
-
0/1
0
0
3
1
1
1
0
4
0
-
0
0/1
0
Cyphophanes Meyrick
1
0
0
0
0
0/1
1
1
0
1/3/4 0
-
0
1
3
Eucosmogastra Diakonoff
0
0
0
0
5
2
1
1
0
3
0/1
0
0
0
0
Helictophanes Meyrick
0
0
0
0
5
1
1
1
0
0/3
0
-
0
0
0
Irianassa Diakonoff
-
0
0
0
5
2
0
2
0
5
1
1
0
1
2
Loboschiza Diakonoff
0
0/2
0
0/1
0/1
0
1
1
0
5
0/1
0
0
1/2
0
Metaselena Diakonoff
0
0
0
0
0/5
0
1
1
0
4
1
0
0
0
3
Oriodryas Turner
0
0
0
0
2
1
1
1
0
0/1
0
-
0
0
0
Periphoeba Bradley
0
2
0
1
0
0
1
1
0
4
1
0
0
0
0
Pseudancylis, gen. nov.
0
0
0
0
5
1
1
1
0
4
1
1
0
1
2
Pternidora Meyrick
0
1
0
1
5
2
1
0
0
2/3
1
0/1
0
0
3
Tetramoera Diakonoff
0/1
0
0
0
0
0
1
1
0
4
0
-
0
2
0
Thysanocrepis Diakonoff
0
0
0
0
5
1
1
1
0
0
0
-
0
0
0
Toonavora, gen. nov.
0
1/3
0
1
5
0
1
1
0
4
0
-
0
0
0
Grapholita zapyrana Meyrick
0
0
0
0
2
2
0
1
1
0
1
0
0
2
0
APPENDIX 1 503
5/5/06 2:45:27 PM
117
504
5/5/06 2:45:27 PM
122
123
124
125
126
Heleanna Clarke
1
0
1
1
0
0
Icelita Bradley
1
0
1
1
0
0
Eucosmophyes Diakonoff
1
0
1
1
1
0
Herpystis Meyrick
0
0
1
1
0
0
Whittenella, gen. nov.
0
0
1
1
0
0
Aglaogonia, gen. nov.
0
0
1
0
-
3
Anathamna Meyrick
0
0
1
1
0
0
Ancylis Hübner
0
0
1
0/1
0
1
Ancylophyes Diakonoff
0
0
1
1
1
0/1
Anthozela Meyrick
0
0
1
0/1
0
0
Balbidomaga Diakonoff
0
0/1
0/1
0
-
3
Cyphophanes Meyrick
0
0
1
1
1
0
Eucosmogastra Diakonoff
0/1
0
1
1
0
0
Helictophanes Meyrick
0
0
0/1
1
1
1
Irianassa Diakonoff
2
1
-
-
-
-
Loboschiza Diakonoff
0
0
0
0/1
0
0/3
Metaselena Diakonoff
0
0
0
0
-
3
Oriodryas Turner
0
0
1
1
0
0
Periphoeba Bradley
0
0
0/1
0/1
1
3
Pseudancylis, gen. nov.
0
0
1
0
-
0
Pternidora Meyrick
0
0
1
1
0
0
Tetramoera Diakonoff
0
0
0/1
0
-
1
Thysanocrepis Diakonoff
0
0
0/1
1
1
1
Toonavora, gen. nov.
0
0
0/1
1
1
1
Grapholita zapyrana Meyrick
0
0
1
1
0
0
APPENDIX 1
120503•Olethreutine Moths 4pp.in504 504
121
Appendix 2 Host plants of olethreutine genera
120503•Olethreutine Moths 4pp.in505 505
5/5/06 2:45:27 PM
506
TRIBE/GENUS
HOSTPLANT FAMILIES CONFIRMED
UNCONFIRMED HOST FAMILIES
EXTERNAL FEEDING
INTERNAL FEEDING
MICROCORSINI Cryptaspasma Walsingham
Arecaceae, Fagaceae, Lauraceae, Monimiaceae, Myrtaceae, Proteaceae, Rosaceae
x
Collogenes Meyrick
Lauraceae
x
OLETHREUTINI Oxysemaphora Diakonoff
Myrsinaceae, Rutaceae
? Pittosporaceae
Cnecidophora, gen. nov.
Myrsinaceae
x
Sorolopha Lower
Lauraceae, Magnoliaceae, Myrtaceae, Pandanaceae
x
Sycacantha Diakonoff
Annonaceae
x
Atriscripta, gen. nov.
–
Demeijerella Diakonoff
–
Lobesia Guenée
Myrtaceae, Proteaceae, Sapindaceae, Verbenaceae, Vitaceae, Liliaceae
x
x
Gnathmocerodes Diakonoff
Lecythidaceae mostly; Annonaceae, Malvaceae
x
?x
Rhectogonia Diakonoff
–
Ophiorrhabda Diakonoff
Myrtaceae mostly; Sapindaceae, Caesalpiniaceae
x
x
Dudua Walker
Euphorbiaceae (Glochidion) mostly; aprobola polyphagous
x
Trachyschistis Meyrick
Ebenaceae
x
Archilobesia Diakonoff
–
Euobraztsovia Diakonoff
–
Eremas Turner
–
Zomariana Diakonoff
Caesalpiniacea, Mimosaceae, Oxalidaceae, Surianaceae
x
Podognatha Diakonoff
Alangiaceae, Annonaceae
x
? Glochidion
x
x
APPENDIX 2
120503•Olethreutine Moths 4pp.in506 506
Appendix 2. Host plants of olethreutine genera.
5/5/06 2:45:28 PM
120503•Olethreutine Moths 4pp.in507 507
TRIBE/GENUS
HOSTPLANT FAMILIES CONFIRMED
UNCONFIRMED HOST FAMILIES
EXTERNAL FEEDING
Temnolopha Lower
Euphorbiaceae
? Zingiberaceae
Diakonoffiana Koçak
unidentified tree
x
Megalota Diakonoff
Euphorbiaceae, Mimosaceae
x
Costosa Diakonoff
–
Gatesclarkeana Diakonoff
polyphagous
x
x
Metrioglypha Diakonoff
unidentified
Dactylioglypha Diakonoff
Elaeocarpaceae
x
Statherotis Meyrick
Myrtaceae, Proteaceae, Sapindaceae
x
Aterpia Guenée
Lytraceae, Ranunculaceae, Rosaceae (Holarctic records)
x
Proschistis Meyrick
–
Rhodacra Diakonoff
–
? Vitis
INTERNAL FEEDING
x
x
BACTRINI Bactra Stephens
Cyperaceae, Juncaceae, Poaceae
x
Syntozyga Lower
Commelinaceae
x
Endothenia Stephens
mostly polyphagous, non–monocots (Holarctic records)
x
ENARMONIINI Anathamna Meyrick
–
Ancylis Hübner
Betulaceae, Dilleniaceae, Ericaceae, Fagales, Proteaceae, Rhamnaceae, Rosaceae, Rutaceae, Salicaceae (incl. Holarctic records)
Ancylophyes Diakonoff
Moraceae
Anthozela Meyrick
Piperaceae
Balbidomaga Diakonoff
–
Cyphophanes Meyrick
Rutaceae
x
Eucosmogastra Diakonoff
Annonaceae
x
x
x (mining)
x
507
–
APPENDIX 2
5/5/06 2:45:28 PM
Aglaogonia, gen. nov.
508
HOSTPLANT FAMILIES CONFIRMED
Helictophanes Meyrick
Apocynaceae, Euphorbiaceae, Meliaceae
UNCONFIRMED HOST FAMILIES
EXTERNAL FEEDING
INTERNAL FEEDING x
Irianassa Diakonoff Loboschiza Diakonoff
Meliaceae mostly; Oleaceae, Malvaceae
Metaselena Diakonoff
–
Oriodryas Turner
–
Periphoeba Bradley
–
Pseudancylis, gen. nov.
–
Pternidora Meyrick
–
Tetramoera Diakonoff
Poaceae
Thysanocrepis Diakonoff
–
Toonavora gen. nov.
Meliaceae (Toona)
x
x
x x
x
EUCOSMINI Crocidosema Zeller
Malvaceae mostly; Verbenaceae, Fabaceae
x
Epiblema Hübner
Asteraceae
x
absconditana-group
Myrsinaceae
Coenobiodes Kuznetzov
Taxaceae
Melanodaedala, gen. nov.
Anacardiaceae
x
Rhopobota Lederer
Anacardiaceae, Aquifoliaceae, Araliaceae, Berberidaceae, Cornaceae, Dipsacaceae, Ericaceae, Fagaceae, Rhamnaceae, Rosaceae
x
Fibuloides Kuznetzov
Fagaceae, galls on Anacardiaceae (Holarctic records)
Acroclita Lederer
Euphorbiaceae, Elaeagnaceae (Holarctic records)
Tritopterna Meyrick
Euphorbiaceae
Noduliferola Kuznetzov
Lamiaceae, Myrtaceae
x x
x x x
APPENDIX 2
120503•Olethreutine Moths 4pp.in508 508
TRIBE/GENUS
5/5/06 2:45:28 PM
120503•Olethreutine Moths 4pp.in509 509
TRIBE/GENUS
HOSTPLANT FAMILIES CONFIRMED
Heleanna Clarke
UNCONFIRMED HOST FAMILIES
EXTERNAL FEEDING
INTERNAL FEEDING
Anacardiaceae, Borraginaceae, Clusiaceae, Euphorbiaceae, Lecythidaceae
x
x
Spilonota Stephens
Myrtaceae (in Australia)
x
Strepsicrates Meyrick
Myrtaceae (in Australia)
x
Holocola Meyrick
Myrtaceae mostly, Epacridaceae, Mimosaceae
x
Hermenias Meyrick
Myrtaceae
x
Eccoptocera Walsingham
Myrtaceae mostly, Araliaceae, Epacridaceae
x
Herpystis Meyrick
Cuscutaceae, Euphorbiaceae, Moraceae
Icelita Bradley
Caesalpiniaceae
Eucosmophyes Diakonoff
Rubiaceae
Whittenella, gen. nov.
–
x x
GRAPHOLITINI x
Loranthacydia Horak, Common & Loranthaceae Komai x
Pammenemima Diakonoff
Fabaceae
Leguminivora Obraztsov
Fabaceae mostly; Anacardiaceae
x
Notocydia, gen. nov.
Caesalpiniaceae
x
Fulcrifera Danilevsky & Kuznetzov Fabaceae
?x
x
–
x
Cydia Hübner
Fabaceae, Fagaceae, Rosaceae, Sapindaceae
x
Thaumatotibia Zacher
Hippocrateaceae, Lauraceae, Myrtaceae, Poaceae, Proteaceae, Rutaceae, Sterculiaceae, Theaceae
x
Gymnandrosoma Dyar
Myrtaceae, Proteaceae, Rutaceae
x
Archiphlebia, gen. nov.
Mimosaceae
x
509
5/5/06 2:45:29 PM
APPENDIX 2
Apocydia, gen. nov.
510
HOSTPLANT FAMILIES CONFIRMED
UNCONFIRMED HOST FAMILIES
EXTERNAL FEEDING
INTERNAL FEEDING
Cryptophlebia Walsingham
Anacardiaceae, Fabaceae, Hippocastaneaceae, Proteaceae, Rhizophoraceae, Rutaceae, Sapindaceae
Acanthoclita Diakonoff
Fabaceae
x
?x
Pammenopsis Kuznetzov
Fabaceae
x
x
Grapholita Treitschke
Fabaceae
x
Aspila Stephens
Rosaceae
x
Microsarotis Diakonoff
Caesalpiniaceae
x
Commoneria, gen. nov.
Chrysobalanaceae
x
Parapammene Obraztsov
Fagaceae mostly; Aceraceae, Tiliaceae, Sapindaceae
x
Ixonympha, gen. nov.
Loranthaceae
x
x
APPENDIX 2
120503•Olethreutine Moths 4pp.in510 510
TRIBE/GENUS
5/5/06 2:45:29 PM
Index Index to scientific names The index covers only scientific names of Lepidoptera and host plant genera and families. Authors are given for Lepidoptera but not for plant taxa. Valid names of Australian olethreutine species and genera are in boldface. Abelmoschus, 305 abietella Matsumura, 317 ablabes Turner, 210 absconditana Walker, 9, 17, 18, 40, 60, 307, 311, 312, 313 absconditana-group, 306, 310–313, 317 abstrusa Kuznetzov, 341, 343, 345 Abutilon, 305 Acacia, 86, 153, 154, 161, 173, 313, 349, 367, 370, 435, 439, 466 Acanthoclita Diakonoff, 47, 55, 58, 60, 66, 396, 397, 398, 426, 427, 439–443 Acanthothyspoda Lower, 93, 98 acceptana Kuznetzov, 313, 314, 315, 317 Aceraceae, 464, 510 Acharneodes Meyrick, 68 achatana Denis & Schiffermüller, 229 Achillea, 403 aclyta Turner, 428, 429, 430 Acmena, 71 Aconitum, 183, 197 acrocausta Turner, 93 Acroclita Lederer, 1, 18, 41, 51, 57, 58, 80, 184, 225, 235, 299, 300, 305, 313, 317, 320, 324, 330, 331, 334–338, 341, 345, 349, 355, 371, 388, 390, 394, 395, 396 Acroclita-group, 1, 80, 313, 317, 320, 324, 345, 349, 388, 390, 394 acrocroca Diakonoff, 443 acrogypsa Turner, 38, 282, 283, 284, 285 acromochla Turner, 235 Acronychia, 84, 87 acrosema Turner, 352, 353, 355 Actinocentra Diakonoff, 151, 152 Adenanthera, 439 Adenoneura Walsingham, 422 adluminana Bradley, 278, 282 Aegiceras, 92 aellaea Turner, 39, 295, 296, 297, 298 Aemulatrix Diakonoff, 228, 275 aeneana Villers, 426 aeraria Meyrick, 257 aestuosa Meyrick, 327, 330, 331 aethalostola Turner, 370 aetheria Turner, 263, 264, 265, 266 Afrocostosa Aarvik, 176 Afzelia, 387 agenjoi Obraztsov, 423 Aglaia, 269 Aglaogonia gen. nov., 1, 18, 25, 36, 58, 220, 221–225 Agnathmacera Diakonoff, 121, 124 Agriophanes Meyrick, 426 Alangiaceae, 159, 506
120503•Olethreutine Moths 4pp.in511 511
Alangium, 159 alboatra Diakonoff, 271 albocingulata sp. nov., 1, 74, 75, 76, 77, 78, 79 Alcina Clarke, 375, 378 Allobrachygonia Fernald, 68, 71, 73 allochroma Diakonoff, 173, 176 Alloendothenia, 216, 219 Allohermenias, 320, 362, 369 allophlebodes Horak & Sauter, 272, 273, 274, 275 alphestis Meyrick, 124, 125 Alphitonia, 233 Alpinia, 163 altheana Mann, 306 Alypeta Turner, 93, 98, 430 Alysicarpus, 450 Alyxia, 261 amaeboea Lower, 192, 193, 195, 196 Ambrosia, 309 ammopastea Turner, 210 ammostigma Turner, 371 amoebaea (missp.), 193 amphibola Diakonoff (Diplosemaphora), 461 amphibola Diakonoff (Proschistis), 199, 202 amphitorna Turner, 450 Amyema, 466, 475 Anacardiaceae, 138, 182, 320, 324, 330, 348, 408, 439, 508, 509, 510 anachastopa Meyrick, 340, 341 Analdes Turner, 111, 117, 120 anaphorana Walsingham, 68 Anaphorodes Diakonoff, 71, 73 anaprobola Bradley, 133, 138 anastrepta Meyrick, 340, 341 Anathamna Meyrick, 18, 36, 57, 79, 104, 220, 221, 222, 225–229, 250, 270, 271, 272, 273, 275, 288, 298 ancalota Meyrick, 126, 128 anceps Meyrick, 173 Anchylopera Stephens, 229, 234, 238, 239 anconia Meyrick, 35, 212, 213, 214, 215, 216 Ancylis Hübner, 18, 36, 49, 51, 54, 59, 60, 220, 221, 225, 229–235, 236, 238, 239, 246, 280, 282, 284, 285, 298, 301, 355 Ancyloides Diakonoff, 235, 236, 238 Ancyloides Kuznetzov, 236 Ancylopera Agassiz, 229 Ancylophyes Diakonoff, 1, 18, 25, 36, 58, 220, 234, 235–239 ancyrota Meyrick, 331 andereggana Guenée, 196, 198, 199 Andrioplecta Obraztsov, 397, 426, 466, 467 angelica Diakonoff, 355 anguillana Meyrick, 36, 230, 233, 235 angulicostana Walsingham, 68, 69 angustifascia Turner, 467 animosana Busck, 17, 18 anisoptera Clarke, 138 Annonaceae, 138, 159, 256, 506, 507 Anoecophysis Diakonoff, 275
5/5/06 2:45:29 PM
512
INDEX
antaxia Meyrick, 310 antecellana Kuznetzov, 388 anthochroa Diakonoff, 252, 253, 254, 255, 256, 257 Anthozela Meyrick, 17, 18, 36, 65, 220, 221, 239–242, 270, 275, 278, 288 anthracosema Turner, 210 anticipans Meyrick, 408 Anticlea Stephens, 229 Antithesia Stephens, 104, 139 antitheta Meyrick, 450 Apeleptera Diakonoff, 196, 197, 199 Aphelia Stephens, 206, 207 aphenia Diakonoff, 176 Aphiaris Kuznetzov, 177, 182 apicinota Turner, 305, 306 Apocydia gen. nov., 1, 45, 51, 54, 55, 65, 398, 406, 407, 408, 419–422, 426 Apocynaceae, 261, 508 aporema Walsingham, 305, 473 approximana Heinrich, 183, 196, 197, 198 aprobola Meyrick, 22, 50, 134, 135, 136, 137, 138, 139 Apsidophora Diakonoff, 168 Aquifoliaceae, 324, 508 Araliaceae, 324, 378, 508, 509 Archilobesia Diakonoff, 18, 23, 32, 50, 61, 62, 129, 136, 138, 139, 142–145 archimedias Meyrick, 94, 96, 98 Archiphlebia gen. nov., 1, 46, 54, 59, 398, 426, 433–435 Archips Hübner, 178 arctana Staudinger, 334, 335, 336 Arecaceae, 71, 506 arescophanes Turner, 118, 120 argillacea Clarke, 257, 261 argillacea Turner, 235 argyrocyrta Turner, 451 argyroela Turner, 451 Argyroploce Hübner, 1, 93, 98, 104, 108, 125, 128, 134, 139, 142, 144, 146, 148, 151, 152, 159, 160, 161, 170, 173, 183, 187, 190, 195, 196, 199, 203, 205, 250, 252, 257, 262, 266, 292, 295, 427, 430, 433, 435, 439, 443, 467 Argyroptocha Diakonoff, 250 argyrotorna Diakonoff, 443 argyrotypa Turner, 371 arithmetica Meyrick, 104, 106, 107, 108 aromatias Meyrick, 235 Arotrophora Meyrick, 433, 436, 439 Artemisia, 403 Articolla Meyrick, 250, 257, 261, 262, 294, 430 artifica Meyrick, 233, 235 Artiphanes Diakonoff, 257, 261 Asaphistis Meyrick, 196, 197, 198, 199 asema Diakonoff, 199 aspersa Turner, 422 aspersana Kuznetzov, 216 Aspila Stephens, 398, 443, 448, 452–454 assimulatana Kuznetzov, 383 Assulella Kuznetzov, 317, 320, 321 Asteraceae, 307, 309, 403, 469, 508 Asthenia Hübner, 111, 118, 330 Astrobola Diakonoff, 184, 186, 187 Asymmetrarcha Diakonoff, 177, 182, 187 atactodes Turner, 104 Aterpia Guenée, 18, 26, 34, 48, 50, 61, 68, 80, 165, 183, 196–199, 200, 202, 203 atmophanes Turner, 371
120503•Olethreutine Moths 4pp.in512 512
atrifracta Diakonoff, 190, 191, 193 atrinodis Meyrick, 68 atripunctis Turner, 44, 411, 412, 413 Atriscripta gen. nov., 1, 17, 18, 31, 64, 80, 99, 100, 104–108 auchmera Turner, 139 aulacodes Lower, 399, 402, 403 aulacota Turner, 313 aurantiana Pryer, 271 aurantianum Lima, 432 auriflora Diakonoff, 100, 102 aurita Meyrick, 93 austera Turner, 412, 413 australis sp. nov. (Costosa), 1, 33, 174, 175, 176, 177 australis sp. nov. (Eccoptocera), 1, 42, 377, 378, 379 autoberylla Meyrick, 98 Averrhoa, 155 avida Meyrick, 380, 381, 383 avita Diakonoff, 187, 188 axiologa Turner, 142, 144 Bactra Stephens, 17, 18, 19, 35, 50, 54, 61, 64, 66, 117, 177, 205, 206–212, 215, 217, 219, 288, 444 bactrana Kennel, 209 Bactrini, 1, 17, 19, 35, 49, 50, 54, 79, 81, 162, 177, 178, 205, 206, 219 baeodes Turner, 371 Bakia Koçak, 68 balanoptycha Meyrick, 439, 443 Balbidomaga Diakonoff, 1, 18, 36, 61, 220, 221, 242–248, 278, 279, 281 balia Diakonoff, 443 Banksia, 49, 193, 221, 234, 432 banosii Gozmany, 210 Barbara Heinrich, 306 barbata sp. nov., 1, 47, 443, 444, 445, 446, 447, 448 Barringtonia, 120, 124, 348, 476 Basigonia Diakonoff, 161 Bathrotoma Meyrick, 317, 320, 321, 350, 353, 355, 369, 370, 371 bathysema Diakonoff, 240, 242 batianensis Diakonoff, 180, 181, 182 batrachodes Meyrick, 191, 192, 194, 195, 196 batrachopa Meyrick, 430 Bauhinia, 132, 138, 439, 456 Beilschmiedia, 71 Berberidaceae, 324, 508 Betulaceae, 233, 507 bicincta Diakonoff, 456 bidenticulana Bradley, 443 biscissana Meyrick, 230, 235 biuncana Kuznetzov, 330, 334 blackburnii Butler, 305, 306 Blastesthia Obraztsov, 306, 310 blepharopis Meyrick, 208, 209, 210, 211 Borraginaceae, 509 bostrychodes Diakonoff, 306 botrana Denis & Schiffermüller, 111, 115 Brachygonia Kirby, 68 brachyptycha Meyrick, 68, 69, 70, 71, 72, 73 brachytycha Turner, 361 Breynia, 383 bryopa Meyrick, 41, 184, 334, 335, 336, 337, 338, 341 Bubonoxena Diakonoff, 212, 215, 216 Buckinghamia, 115, 439 Bursaria, 450
5/5/06 2:45:29 PM
INDEX
Butea, 443 Byrsoptera Lower, 111, 112, 118, 120 Cacochroea Lederer, 307, 310 caecana Schläger, 450 Caesalpiniaceae, 51, 132, 138, 152, 154, 305, 387, 412, 427, 439, 456, 506, 509, 510 Cajanus, 446 Calamus, 71 calceata Meyrick, 355 callichlora Meyrick, 98 callicratis Meyrick, 257 calligrapha Meyrick, 289, 290 callimita Turner, 370 Callistemon, 353 Calophyllum, 348 calvifrons Walsingham, 301 capnopepla Turner, 208, 210 Capua Stephens, 120, 211, 235 capyra Meyrick, 41, 331, 336, 338, 339, 340, 341 Caragana, 418 carnicolor Meyrick, 152, 155 carpalima Meyrick, 230, 231, 232, 233, 234, 235, 236 Carpocampa Harris, 422 Carpocapsa Treitschke, 422, 426, 450 carpophaga Walsingham, 436, 439 Cassia, 305, 439 Castanea, 330 castanicolor Turner, 102, 103, 104 catapasta Turner, 371 catarrhactopa Meyrick, 199 catharoptis Lower, 331, 341 Catoptria Guenée, 423 celebensis Diakonoff, 294, 295 Cellifera Diakonoff, 129, 133 cellifera Meyrick, 129, 133, 134, 139 Centroxena Diakonoff, 66, 398, 426, 427 Cephalophyes Diakonoff, 185, 199 Cerata Stephens, 422 ceratodes Turner, 439 chalcitis Meyrick, 370 chaomorpha Meyrick, 426, 427, 429, 430 charmera Turner, 306 Cheirodendron, 378 Chenopodium, 309 Chiloides Butler, 207, 208, 210 chionodelta Meyrick, 32, 146, 147, 148 chionolitha Meyrick, 82, 84 chionopyra Diakonoff, 228 chionostoma Meyrick, 338, 339, 340, 341 chlidana Turner, 363, 370 chlorantha Meyrick, 321, 324 chloreis Turner, 41, 345, 346, 347, 348, 349 chlorohygra Diakonoff, 139 Choganhia Razowski, 93 Chrosis Guenée, 216 Chrysanthemum, 403 Chrysobalanaceae, 461, 510 chrysoplea Diakonoff, 31, 108, 109, 110, 111 chrysosema Turner, 216 chrysoxantha Meyrick, 239, 240, 241, 242 Cimeliomorpha Diakonoff, 270, 275 cinnabaritis Meyrick, 271 Cinnamomum, 96 circulata Diakonoff, 184, 186, 187
120503•Olethreutine Moths 4pp.in513 513
513
cirrholepida Clarke, 388 citharistis Meyrick, 219 Citrus, 432 clastomochla Turner, 370 clivosa Meyrick, 322 Clusiaceae, 348, 509 clydonias Meyrick, 165 clytocarpa Meyrick, 266, 268, 269, 270 Cnecidophora gen. nov., 1, 18, 28, 30, 65, 80, 81, 82, 86, 89–93 Cnephasia Curtis, 89, 93, 120, 435 coccophaga Falkovitsh, 112 Coccyx Treitschke, 306, 422, 452 Coenobiodes Kuznetzov, 17, 18, 40, 60, 300, 306, 310, 313–317 Collicularia Obraztsov, 423 Collogenes Meyrick, 1, 17, 18, 27, 29, 50, 65, 67, 68, 73–81 colonota Meyrick, 235 comitana Denis & Schiffermüller, 350 commatica Turner, 371 Commelina, 210, 215 Commelinaceae, 50, 206, 215, 507 Commoneria gen. nov., 1, 47, 54, 55, 65, 397, 398, 426, 459–461 commoni sp. nov., 1, 43, 389, 390, 391, 392 compositella Fabricius, 66, 448, 449, 450, 452 conciliata Meyrick, 443 confertana Walker, 183, 186, 187 conficitana Walker, 66, 449, 450 confracta Diakonoff, 177 confusa Turner, 395 Coniostola Diakonoff, 426 constrictana Meyrick, 24, 42, 49, 349, 350, 351, 352, 353, 354, 355, 369 coppelia Clarke, 387 Coptoloma Lederer, 452 Cordia, 348 Corethrarcha Diakonoff, 203, 205 corinthia Meyrick, 330 Cornaceae, 324, 508 corollana Hübner, 422 coronopa Meyrick, 235 Correa, 233 corynetes Diakonoff, 306 Cosmetra Diakonoff, 349 Cosmopoda Diakonoff, 163 Cosmorrhyncha Meyrick, 161 Costosa Diakonoff, 1, 18, 33, 63, 80, 161, 172, 173–177 cretata Diakonoff, 199 critica Meyrick, 443, 444, 445, 446, 447, 448 Crobylophora Kennel, 422 Crocidosema Zeller, 18, 22, 23, 40, 51, 57, 60, 299, 300, 301–306 crocopa Meyrick, 418 crocoptila Diakonoff, 330, 345 crocospila Meyrick, 134 crossoleuca Meyrick, 143, 144 crossota Meyrick, 39, 156, 161, 292, 293, 294, 295 Crotalaria, 418, 419, 446 Croton, 173 Cryptaspasma Walsingham, 18, 29, 50, 62, 67, 68–73, 76, 196 Cryptophlebia Walsingham, 25, 26, 46, 51, 55, 57, 59, 396, 397, 398, 426, 427, 430, 435–439 Cryptoschesis Diakonoff, 426
5/5/06 2:45:30 PM
514
INDEX
cryptosema Diakonoff, 361 cunninghamiacola Liu & Bai, 112, 468 Cupaniopsis, 439 Cuscuta, 383, 468 Cuscutaceae, 383, 509 cuscutae Bradley, 383 cyanitis Diakonoff, 165, 166 cyanopis Meyrick, 329, 330 cyanosticha Clarke, 168, cyanosticha Turner, 47, 459, 460, 461 Cyclacanthina Diakonoff, 212, 215, 216 cyclidias Meyrick, 261 cyclotoma Lower, 30, 93, 94, 95, 96, 97, 98 Cydia Hübner, 22, 45, 50, 57, 66, 165, 397, 398, 403, 406, 407, 416, 421, 422–426, 427 Cydia-group, 50, 397, 398, 403, 406, 407, 416, 427 Cymolomia Lederer, 129, 134, 142, 144, 168 Cynoglossum, 182 Cyperaceae, 50, 206, 209, 507 Cyperus, 209 Cyphophanes Meyrick, 1, 18, 37, 66, 220, 221, 225, 228, 248–252, 262, 275, 288, 295 Dactylioglypha Diakonoff, 18, 34, 64, 187–190, 203 dadionopa Diakonoff, 426 Danilevskia Kuznetzov, 423 dascia Bradley, 76, 78 deceptor Diakonoff, 133 decorata Meyrick, 190 decorigera Diakonoff, 108 defensa Meyrick, 443 delineana Walker, 455 delochlora Turner, 93, 94, 95, 96, 97, 98 delomilta Turner, 267, 268, 269, 270, 271 Delonix, 439 deloschema Turner, 42, 364, 366, 368, 371 deloxantha Turner, 269, 271 deltostoma Diakonoff, 327, 330, 331 Demeijerella Diakonoff, 17, 18, 31, 57, 80, 99, 108–111, 119 dentatana Walker, 467 derasana Hübner, 229 Derris, 443 Desmodium, 403, 405, 406, 443 diachorda Meyrick, 207, 210 diaema Turner, 435 Diakonoffiana Koçak, 18, 33, 59, 61, 62, 80, 161, 165–170, 172 Diakonoffiella Kuznetzov, 236 Diamphidia Obraztsov, 461 Dicephalarcha Diakonoff, 191, 193 Dichelia Guenée, 196 Dichrorampha Guenée, 50, 397, 398, 403, 406 Dichrorampha-group, 50, 397, 398, 403 Dicraniana Diakonoff, 423, 470 Didrimys Diakonoff, 119, 129, 132, 133, 134 Dierlia Diakonoff, 397, 426 dietziana Kearfott, 321 diffusa Bradley, 320 diffusana Walker, 467 Dilleniaceae, 233, 234, 507 Diospyros, 141, 454 Diplonearcha Meyrick, 380, 383 Diplosemaphora Diakonoff, 461 diplostigma Turner, 371 Dipsacaceae, 324, 508
120503•Olethreutine Moths 4pp.in514 514
Dipterina Meyrick, 187 discana Felder & Rogenhofer, 193 discotypa Turner, 120 dissias Meyrick, 406 distorta Hampson, 436 Dodonaea, 406, 425, 464 Dolichos, 418 dolichosticha Turner, 422 domestica Diakonoff, 182 dorophora Diakonoff, 242, 244, 246 dorsana Fabricius, 448 doxasticana Meyrick, 33, 57, 152, 153, 154, 155, 156 Dracontogena Diakonoff, 426 Dryas, 183, 197 drymoptila Lower, 32, 142, 144, 145 dryocoma Meyrick, 250, 252, 262 dryoptycha Meyrick, 134 Drypetes, 161, 163 Dudua Walker, 18, 22, 32, 50, 59, 63, 129, 133, 134–139, 142, 144, 146, 148 Duessa Clarke, 341, 343, 345 Dynatorhabda Diakonoff, 81, 82, 86, 89, 92, 120, 196 dysaethria Diakonoff, 454 dyscheranta Meyrick, 248, 249, 250, 251, 252, 295 dyschima Diakonoff, 128 dyselia Turner, 361 dyserasta Turner, 463, 464 Ebenaceae, 141, 454, 506 ebenocosma Turner, 361 ebenostigma Turner, 371 Ebisma Walker, 448 Eccopsis Zeller, 128, 134, 139, 161, 190 Eccoptocera Walsingham, 1, 18, 42, 54, 58, 299, 349, 375–379 Ecdytolopha Zeller, 397, 426, 427, 430, 439, 468 ejectana Walker, 356, 357, 358, 359, 360, 361 Elaeagnaceae, 51, 336, 508 Elaeagnus, 336 Elaeocarpaceae, 183, 190, 507 Elaeocarpus, 183, 190 elaeodes Lower, 93, 94, 95, 97, 98 electrosema Diakonoff, 31, 125, 126, 127, 128 Eleuthodema Bradley, 212 Embolostoma Diakonoff, 228 emmilta Diakonoff, 168 empalinopa Diakonoff, 187 empherana Bradley, 134 empyra Turner, 139 Enarmonia Hübner, 126, 219, 220, 221, 241, 262, 290 Enarmoniini, 1, 8, 11, 17, 25, 49, 54, 58, 79, 80, 81, 82, 92, 151, 161, 219, 220, 221, 234, 264, 270, 280, 295, 298, 299, 396 Enarmonodes Danilevsky & Kuznetzov, 220 endaphana Diakonoff, 215, 216 Endiandra, 71, 76 endophaga Meyrick, 46, 433, 434, 435 Endopisa Guenée, 426, 448 Endothenia Stephens, 17, 18, 19, 35, 50, 63, 66, 79, 81, 177, 178, 206, 207, 216–219 Engelana Diakonoff, 119 engone Diakonoff, 219 Enveryucelia Koçak, 142 eocnephaea Meyrick, 340, 341 Epacridaceae, 367, 378, 509
5/5/06 2:45:30 PM
INDEX
ephippias Meyrick, 206, 215, 216 Ephippiphora Duponchel, 448 Epiblema Hübner, 17, 18, 40, 60, 299, 300, 306, 307–310, 313, 317, 331 Epiblema-group, 300, 306, 307, 310, 313 Epicharis Hübner, 229 Epichorista Meyrick, 216 epidola Meyrick, 1, 371, 372, 374, 375 epidolana Lower, 375 Epinotia Hübner, 40, 49, 50, 234, 246, 299, 300, 301, 305, 306, 307, 311, 312, 313, 321, 324, 330 Epinotia-group, 300, 301 Epiphyas Turner, 22, 370 episema Diakonoff, 212, 213, 216 Episimoides Diakonoff, 81, 82, 92 Episimus Dyar, 81, 82, 86 Eremas Turner, 18, 32, 54, 59, 63, 80, 148–151, 190, 195 ergasima Meyrick, 129, 134 Ericaceae, 152, 233, 324, 507, 508 eridela Turner, 306 Erinaea Meyrick, 321, 322, 324 Eriopsela Guenée, 300 Erminea Kirby & Spence, 422 erotias Meyrick, 178, 182, 183 erythrana Meyrick, 230, 231, 232, 233, 235 erythrosema Turner, 235 erythrotypa Turner, 235 Esia Heinrich, 196, 198 Eubrochoneura Diakonoff, 100, 151, 152, 155, 156, 159 Eucalyptus, 357, 367, 374 Eucelis Hübner, 271, 443 Eucoenogenes Meyrick, 326, 330, 331 Eucosma Hübner, 73, 93, 156, 170, 173, 196, 210, 212, 216, 257, 282, 285, 290, 292, 295, 298, 299, 300, 306, 310, 311, 312, 313, 317, 320, 321, 330, 355, 361, 370, 371, 411, 412, 413, 419, 422, 427, 435, 439, 443 Eucosmini, 1, 3, 11, 17, 21, 25, 26, 28, 49, 54, 58, 60, 79, 80, 81, 82, 92, 109, 126, 151, 220, 221, 282, 285, 298, 299, 300, 307, 313, 317, 341, 376, 383, 396, 397, 399 Eucosmocydia Diakonoff, 426 Eucosmogastra Diakonoff, 17, 18, 37, 61, 221, 239, 252–257, 275, 278, 284 Eucosmomorpha Obraztsov, 220, 221, 238, 270 Eucosmophyes Diakonoff, 1, 17, 18, 28, 43, 54, 60, 300, 384, 388–392 Eudemis Hübner, 93, 94, 97 Eudemis-group, 93 Eudemopsis Falkovitsh, 93, 94, 97 Eugenia, 193, 330, 343, 357, 454 Eumarissa Clarke, 321, 324 Euobraztsovia Diakonoff, 18, 32, 63, 129, 134, 142, 144, 146–148 eupena Turner, 221, 222, 223, 224, 225 euphlebia Kawabe, 330 Euphorbia, 336 Euphorbiaceae, 50, 51, 138, 144, 161, 163, 173, 261, 336, 340, 341, 348, 383, 506, 507, 508, 509 euplectra Lower, 31, 121, 122, 123, 124 euploca Turner, 361 euprepes Turner, 313 eurymochla Turner, 216 euryochra Bradley, 317 euryphaea Turner, 191, 192, 193, 194, 195 eurypolia Turner (Sycacantha), 104 eurypolia Turner (Zomariana), 156
120503•Olethreutine Moths 4pp.in515 515
515
euryptycha Turner, 370 Euryptychia Clemens, 307 Euspila Stephens, 448 euthytoma Turner, 370 excitana Möschler, 306 exedra Turner, 31, 101, 102, 103, 104 exemplaris Meyrick, 266, 267, 268, 271 exocentra Meyrick, 405, 406 Exoria Meyrick, 353 extrusana Walker, 111, 112, 113, 114, 115, 117, 118, 119, 120 exvagana Walker, 310 Fabaceae, 51, 305, 403, 405, 408, 418, 419, 425, 427, 439, 443, 446, 450, 508, 509, 510 Fagaceae, 71, 324, 330, 425, 427, 464, 506, 508, 509, 510 fallax Meyrick, 170, 173 falsifalcellum Walsingham, 422 favillosa Diakonoff, 134 ferraria Turner, 422 Fibuloides Kuznetzov, 1, 18, 28, 41, 58, 60, 317, 326–334, 341, 342, 345, 355 Ficus, 238, 383 fidana Meyrick, 235 fimbriata Walsingham, 163, 164 finitimana Heinrich, 321 flammeata Kuznetzov, 221, 290 flavescens Kuznetzov, 290 flavocellana Clemens, 310 floricolana Meyrick, 450, 452 fluidana Meyrick, 367, 371 foenella Linnaeus, 307 foetorivorans Butler, 375, 378, 379 formosana Scopoli, 221 frustulosa Diakonoff, 159 fukugi Nasu, 345, 346, 348, 349 Fulcrifera Danilevsky & Kuznetzov, 1, 29, 45, 55, 66, 397, 398, 406, 407, 408, 415–419, 426 fulva Turner, 216 fulvoflua Diakonoff, 168 funebrana Treitschke, 452 fungiferana Meyrick, 170 furiosa Meyrick, 266, 267, 268, 269, 270, 271 galena Clarke, 340 Garcinia, 348, 474 Gatesclarkeana Diakonoff, 1, 18, 35, 49, 50, 62, 79, 129, 152, 156, 162, 177, 178–183, 206, 219 Gatesclarkeana-group, 1, 79, 162, 177, 178, 206 Geita Aarvick, 161 gemmarius Diakonoff, 186 gentianaeana Hübner, 216 Gephyroneura Obraztsov, 239, 242 geraropa Meyrick, 210 Gibberifera Obraztsov, 300 glaphyrana Rebel, 426 glaucana Kennel, 461 Glochidion, 50, 138, 144, 340, 506 glyceranthes Meyrick, 169 Glycine, 305, 408, 450, 473 glycinivorella Matsumura, 407, 408, 409 glycypolia Turner, 235 Gnathmocerodes Diakonoff, 18, 31, 49, 64, 86, 93, 120, 121–125, 128, 196 Gnathmocerodes-group, 93, 120 gonodesma Turner, 216
5/5/06 2:45:30 PM
516
INDEX
gonomela Lower, 46, 431, 432, 433 Gossypium, 305, 471 gracilistria Turner, 38, 288, 289, 290, 291, 292 gracilivalva sp. nov., 1, 248, 250, 251, 252 grandaevana Lienig, 307 Grapholita Treitschke, 4, 17, 18, 24, 47, 50, 54, 55, 66, 111, 117, 120, 183, 310, 371, 396, 397, 398, 405, 406, 407, 408, 426, 427, 443, 448–454, 455, 456, 461, 467 Grapholita-group, 4, 50, 396, 397, 398, 407, 426, 427, 461 Grapholitha Treitschke, 183, 288, 306, 448, 450, 452, 456 Grapholitini, 1, 4, 5, 9, 17, 26, 49, 50, 51, 54, 79, 92, 219, 221, 298, 299, 396, 397, 398, 399, 430, 461 gratulata Meyrick, 271 Gravitarmata Obraztsov, 306 Griselda Heinrich, 468 Gymnandrosoma Dyar, 46, 54, 59, 398, 426, 430–433, 435, 439 Gypsonoma Meyrick, 220, 300, 307 gypsopa Diakonoff, 199 hacobiani sp. nov., 1, 30, 83, 85, 86, 87, 88, 89 haematina Diakonoff, 199 halysideta Walsingham, 269, 271 hapalaspis Meyrick, 235 Hardenbergia, 450 harmonia Meyrick, 111, 113, 115, 117, 119 harmonica Meyrick, 129, 134 Harmosma Diakonoff, 111, 117 harpana Hübner, 229 hartigiana Saxesen, 134 Hearnia, 261, 269 hebesana Walker, 216 Hedulia Heinrich, 423 Hedya Hübner, 129, 133, 134, 136, 138, 139, 142, 144 Hedycarya, 71 Heleanna Clarke, 18, 41, 51, 57, 58, 317, 345–349, 394 helicana, 170, 171, 172, 173 Helictophanes Meyrick, 1, 18, 37, 60, 170, 220, 221, 250, 257–262, 288, 295, 298 Heligmocera Walsingham, 301 helota Meyrick, 68, 73 Hemerosia Stephens, 271 hemicapnodes Diakonoff, 199 hemicosma Lower, 266, 268, 269, 271 hemicyclica Diakonoff, 199 hemidoxa Meyrick, 36, 239, 240, 241, 242 Hemimene Hübner, 398, 403 hemiochra Turner, 235 hemitypa Diakonoff, 139 Hendecaneura Walsingham, 317 Hendecasticha Meyrick, 349, 360, 376 herbifera Meyrick, 98 Hermenias Meyrick, 1, 18, 42, 54, 59, 320, 330, 349, 360, 362, 367, 369, 370, 371–375, 378 Herminias Lower, 371, 375 Herpystis Meyrick 17, 18, 43, 51, 61, 216, 299, 300, 348, 380–383 hesperialis Walker, 134 Heterochorista Diakonoff, 108 Heteroschistis Diakonoff, 120, 228, 275 hians Meyrick, 32, 139, 141, 142 Hibbertia, 233 hibbertiana Meyrick, 229, 231, 232, 233, 235 Hibiscus, 269, 305 hilaris Turner, 239, 240, 241, 242
120503•Olethreutine Moths 4pp.in516 516
hilarocrossa Meyrick, 443 hilarograpta Meyrick, 102 himerodana Meyrick, 235 Hippocastaneaceae, 510 Hippocrateaceae, 430, 509 Hiroshiinoueana Kawabe, 177, 182, 183 historica Meyrick, 222, 223, 224, 225 Holocola Meyrick, 18, 24, 42, 51, 54, 57, 59, 61, 235, 299, 349, 350, 355, 356, 360, 362–371, 372, 374, 375, 376, 379 Homona Walker, 225 honesta Meyrick, 40, 367, 370 hortaria Meyrick, 184, 322, 323, 324, 325 Hulda Heinrich, 219, 469 hydrargyra Meyrick, 98 Hylotropha Turner, 334, 336, 338 Hyphantidium Scott, 464, 467 hypolepta Turner, 111, 117, 120 hypomolybda Turner, 365, 367, 371 Hyposarotis Diakonoff, 426 hyposcopa Lower, 464, 467 Hystrichophora Walsingham, 221 Icelita Bradley, 18, 43, 59, 60, 300, 383–388, 390 icelitodes Diakonoff, 388, 389, 390 idia Diakonoff, 182 idicopoda Diakonoff, 176 Idiomorpha Turner, 68, 73 Ilex, 324 imbrifera Meyrick, 371 implexa Meyrick, 378, 379 incondita Diakonoff, 108 indentata Bradley, 43, 357, 384, 385, 386, 387, 388 indentatus Dyar, 356 infectana Meyrick, 235 infensa Meyrick, 356, 357, 361, 362, 376 infimama Walker, 361 infimana Walker, 361 iniqua Meyrick, 134, 135, 137, 138, 139 injectiva Heinrich, 423 inquinatana Hübner, 422, 423 insignata Kuznetzov, 331 insinuans Meyrick, 380, 382, 383 insuetana Kuznetzov, 343 insulana Aurivillius, 301, 305, 306 intricata Turner, 170 Intsia, 387 iodryas Meyrick, 383 ionia Wileman & Stringer, 405, 406 Irianassa Meyrick, 17, 18, 37, 61, 65, 220, 262–266, 288 iridescens Meyrick, 451 iridorphna Meyrick, 443 iridosoma Meyrick, 439 iris Diakonoff, 306 irrorea Lower, 104 ischalea Meyrick, 371 isogramma Meyrick, 289, 290 Ixonympha gen. nov., 1, 48, 51, 54, 65, 397, 398, 426, 464–467 Jambosa, 132 janthinana Duponchel, 452 japonica Kawabe, 330 Jasminum, 269 jejuna Meyrick, 380, 382, 383 johngreeni sp. nov., 1, 95, 96, 97, 98, 99
5/5/06 2:45:31 PM
INDEX
Juncaceae, 50, 206, 209, 507 jungiana Guenée, 448 jungiella Clerck, 448 Karacaoglania Koçak, 426 Kenneliola Paclt, 423, 426 koenigiana Fabricius, 37, 49, 221, 266, 267, 268, 269, 270, 271 Kundrya Heinrich, 321, 322, 324 labidophora Diakonoff, 121, 124 Lagerstroemia, 219 Lamiaceae, 343, 508 Lamyrodes Meyrick, 229, 234, 235 lancealana Hübner, 207 lantana Busck, 301, 302, 303, 304, 305, 306 Lantana, 115, 182, 305 laricana Heinemann, 349, 353 Larix, 349, 353 Lasiognatha Diakonoff, 129, 133, 134 lasiophora Lower, 361 lasiosoma Meyrick, 98 Laspeyresia Hübner, 93, 219, 239, 242, 264, 266, 271, 396, 403, 406, 408, 412, 416, 422, 424, 425, 426, 450, 451, 452, 454, 456, 459, 461, 464 Laspeyria Germar, 396, 426 Lathronympha Meyrick, 406 latiferreana Walsingham, 422 Lauraceae, 50, 71, 76, 96, 430, 506, 509 Lavatera, 209 lavaterana Millière, 306 lechriaspis Meyrick, 355 Lecythidaceae, 120, 124, 348, 506, 509 lecythocera Meyrick, 123, 124 lediana Linnaeus, 452 Leguminivora Obraztsov, 1, 44, 55, 66, 397, 398, 406, 407– 411, 412, 426 Lepidunca Diakonoff, 349 lepta Horak & Sauter, 38, 272, 273, 274, 275 leptalea Diakonoff, 290 Leptarthra Lower, 271, 399, 402, 403 Leptia Guenée, 207 Leptocera Diakonoff, 124, 196 leptochlora Turner, 94, 98 Leptospermum, 353, 357, 367 leptotypa Turner, 334, 336, 337, 338 leucatma Turner, 170 leucognoma Clarke, 321 leuconephela Turner (Holocola), 370 leuconephela Turner (Statherotis), 196 leuconota Turner, 370 leucopleura Turner, 370 leucopyga Turner, 371 leucotreta Meyrick, 398, 426, 427, 428, 429, 430 leucotrigona Turner, 32, 148, 149, 150, 151 levatana Kuznetzov, 330 Licigena Diakonoff, 426 ligniferana Walker, 361 ligyropis Meyrick, 330 Liliaceae, 506 limnephilana Meyrick, 359, 362 liphaema Turner, 361 Lipoptycha Lederer, 403 Litchi, 115, 439, 468 litchivora Baixeras, 302
120503•Olethreutine Moths 4pp.in517 517
517
liturata Turner, 371 Lobesia Guenée, 17, 18, 23, 25, 26, 31, 49, 50, 59, 61, 62, 80, 84, 99, 100, 111–120, 133, 142, 144, 215 Lobophora Turner, 142, 144 Loboschiza Diakonoff, 17, 18, 37, 49, 51, 65, 79, 80, 220, 221, 222, 225, 242, 254, 266–271, 288 lomacula Lower, 412 Lomaschiza Lower, 111, 117, 120 Lomaschizodes Diakonoff11, 117 longestriata Durrant, 230, 231, 235 longigula sp. nov., 1, 408, 409, 410 longipalpana Möschler, 302, 305 Lophostemon, 357 Loranthaceae, 1, 51, 138, 182, 399, 402, 466 Loranthaceae, 1, 51, 138, 182, 399, 402, 466, 475, 509, 510 Loranthacydia Horak, Common & Komai, 44, 51, 54, 55, 66, 397, 398, 399–403, 461 Loranthacydia-group, 397, 398, 400 Loranthus, 182, 402 loricata Diakonoff, 29, 74, 75, 76, 77, 78 loxomochla Turner, 216 lucida Meyrick, 456 lucifera Meyrick, 199 lucifera Turner, 371 lugubris Felder & Rogenhofer, 68, 69 lundana Fabricius, 229 lunulana Denis & Schiffermüller, 448 luteiceps Kuznetzov, 416, 418 lutescens Meyrick, 234 lychnospila Turner, 170 lygistis Diakonoff, 456 Lythrum, 183, 197 Lytraceae, 507 Macadamia, 71, 138, 432, 439 macroma Turner, 349 macropetana Meyrick, 349, 357, 359, 360, 361 macrorhis Diakonoff, 331 macrosaris Meyrick, 330 Magnoliaceae, 96, 506 malesana Meyrick, 426 Mallotus, 340 Malus, 221 Malva, 305, 471 Malvaceae, 51, 124, 209, 269, 301, 305, 506, 508 Mangifera, 138 Maranta, 408 marcidella Walsingham, 305 marginana Haworth, 206 marginifasciatus Walsingham, 68 marmaropa Meyrick, 199, 201, 202 martia Meyrick, 268, 269, 271 Matsumuraeses Issiki, 397, 426, 427, 439, 440, 441, 442, 443, 447 maturicolor Diakonoff, 199 mediana Walker, 271 mediocunea Turner, 370 Megaherpystis Diakonoff, 320 Megalota Diakonoff, 18, 33, 64, 80, 100, 161, 165, 170–173 megalozona Diakonoff, 229 Mehteria Koçak, 239, 242 Meiligma Diakonoff, 176 Melaleuca, 115, 353, 367 melanancalis Meyrick, 330 melanochlaena Meyrick, 330
5/5/06 2:45:31 PM
518
INDEX
melanocosma Turner, 40, 314, 315, 316, 317 melanocycla Common, 134 Melanodaedala gen. nov., 1, 18, 40, 60, 317–321 melanographa Turner, 371 melanomochla Meyrick, 348, 349, 474 melanoneura Meyrick, 320, 321 Melia, 269 Melichrus, 367 Melissopus Riley, 422, 468 mellifera Diakonoff, 187 menthivora Oku, 216 Mesocallyntera Diakonoff, 73, 76, 78 mesoscia Meyrick, 235 mesosticha Turner, 362 metabola Turner, 362 metacycla Diakonoff, 170 metallocosma Lower, 399, 400, 401 402, 403 metallocosma Turner, 8, 262 metallota Lower, 139 Metaschistis Diakonoff, 196, 198, 199 Metaselena Diakonoff, 18, 25, 26, 38, 57, 58, 79, 220, 221, 222, 225, 228, 262, 270, 271–275 Metaspasma Diakonoff, 68, 71, 73 metaspra Diakonoff, 330 Metendothenia Diakonoff, 168, 170 Metrioglypha Diakonoff, 18, 34, 64, 80, 183–187, 188, 191, 203 Metriophlebia Diakonoff, 427 Metrosideros, 378 microchlamys Diakonoff, 124, 196 Microclita Diakonoff, 341, 343, 345 Microcorses Walsingham, 67, 68, 71, 73 Microcorsini, 1, 17, 27, 29, 49, 50, 54, 57, 67, 76, 79, 92, 177, 178 microgrammana Guenée, 423 Microsarotis Diakonoff, 1, 47, 55, 65, 397, 398, 426, 454–459, 461 miltographa Meyrick, 252, 253, 254, 255, 257 miltoxantha Meyrick, 257 Mimarsinania Koçak, 461 Mimosa, 138 Mimosaceae, 51, 138, 152, 154, 161, 173, 349, 367, 427, 435, 439, 506, 507, 509 minuta sp. nov., 1, 41, 327, 328, 331, 333 minutana Kearfott, 310 mirabilis Kuznetzov, 71 mixanthes Meyrick, 262 modificana Kuznetzov, 326 molesta Busck, 50, 398, 448, 453, 454 Molybdocrates Diakonoff, 156, 159 monela Clarke, 384, 385, 386, 387, 388 Monilia Walker, 355, 357, 361 Monimiaceae, 71, 506 monochroa Diakonoff, 36, 237, 238, 239 monosema Diakonoff, 216 Monosphragis Clemens, 307 Moraceae, 238, 383, 507, 509 mormopa Meyrick, 129, 130, 131, 132, 133, 134 morosa Meyrick, 371 mosaica Lower, 33, 161, 162, 163, 164, 165, 172 mosaica Meyrick, 257 multilinea Turner, 402, 403 munda Diakonoff, 330 Murraya, 250 Myrica, 357
120503•Olethreutine Moths 4pp.in518 518
myriolychna Turner, 250, 261, 262 Myrsinaceae, 50, 81, 84, 87, 92, 307, 313, 506, 508 Myrtaceae, 1, 51, 54, 71, 96, 115, 132, 138, 193, 299, 343, 349, 353, 357, 367, 374, 378, 430, 432, 454, 506, 507, 508, 509 mysterica Turner, 129, 130, 131, 132, 133, 134 naevana Hübner, 321 Nannobactra Diakonoff, 207, 208, 210 neaera Meyrick, 330 nebritana Treitschke, 448 negligens Diakonoff, 216 nemorivaga Tengström, 299 Nenomoshia Clarke, 252, 256 Neoanathamna Kawabe, 282 Neodasyphora Bae & Liu, 112 Neohermenias Diakonoff, 362, 369, 370, 374 Neolobesia Bae & Komai, 112 Neopotamia Diakonoff, 128, 151, 161, 162, 165, 169, 173, 176, 177, 190, 206 Neopotamia-group, 128, 161, 162, 173, 176, 177, 190, 206 neothela Turner, 41, 184, 341, 342, 343, 344, 345 Neothenia Diakonoff, 216 Nephelium, 132, 138, 183, 193 nephobola Turner, 93 neurosticha Turner, 212 nigrovenana Kuznetzov, 330, 474 nimbata Turner, 344, 345 niphada Diakonoff, 341, 343 niphostetha Turner, 103, 104 niphosticha Turner, 370 nitida Turner 365, 366, 371 niveimacula sp. nov., 1, 44, 411, 412, 414, 415 nobilis Diakonoff, 199 Noduliferola Kuznetzov, 18, 41, 58, 220, 299, 317, 341–345 Norma Heinrich, 321, 322, 324 Noteraula Meyrick, 207, 210 noteraula Walsingham, 207 Nothofagus, 246 notialis sp. nov., 1, 83, 84, 85, 86, 87, 88 Notocelia Hübner, 68, 306, 310, 313, 317 Notocydia gen. nov., 1, 44, 54, 55, 65, 397, 398, 406, 407, 408, 411–415, 426 notosphena Turner, 371 Nuntiella Kuznetzov, 313, 476 Nyctidea Diakonoff, 165, 168 obeliscana Meyrick, 367, 370 obscura Wollaston, 306 ocellana Denis & Schiffermüller, 349, 350, 351, 352, 353, 354, 355, 376 ochronephes Turner, 370 ochronota Turner, 355 ochropa Meyrick, 403, 405, 406 ochropepla Turner, 235 ochrophara Turner, 235 ochroplaca Turner, 93 ochroptila Turner, 30, 89, 90, 91, 92, 93 olbophora Turner, 38, 275, 276, 278 Oleaceae, 51, 269, 508 Olethreutes Hübner, 68, 79, 80, 86, 124, 128, 134, 139, 142, 144, 151, 161, 169, 176, 182, 190, 196, 205, 257 Olethreutes-group, 80, 128, 134, 139, 142, 144, 151, 161, 190 Olethreutini, 1, 5, 11, 17, 19, 25, 26, 28, 49, 50, 53, 54, 62, 79, 80, 81, 85, 92, 93, 99, 100, 107, 109, 119, 120, 121, 126,
5/5/06 2:45:31 PM
INDEX
128, 133, 142, 146, 148, 151, 152, 153, 159, 161, 162, 165, 170, 173, 177, 178, 184, 205, 206, 219, 220, 221, 294, 298, 299, 331, 383, 396, 397, 399 ombrodelta Lower, 25, 46, 57, 398, 436, 437, 438, 439 Omphalea, 261 Omphaletis, 261 Opadia Guenée, 452 ophiocosma Turner, 121, 122, 123, 124, 125 Ophiorrhabda Diakonoff, 18, 29, 32, 49, 62, 80, 119, 128– 134, 139, 142, 144, 162, 178, 196, 206 opsia Meyrick, 370 optanias Meyrick, 208, 209, 210 opulenta Diakonoff, 156, 157, 158, 159, 161 orientis Kuznetzov, 418 Oriodryas Turner, 17, 18, 38, 54, 61, 220, 221, 239, 242, 253, 257, 275–278, 280 osteomelesana Swezey, 378, 379 ostracitis Meyrick, 225, 226, 228, 229 otiosana Clemens, 307 Owenia, 269 Oxalidaceae, 152, 155, 506 Oxysemaphora Diakonoff, 1, 18, 26, 30, 50, 64, 79, 80, 81, 82–89, 92, 100, 120, 128, 168, 196, 220 Oxysemaphora-group, 50, 79, 80, 81, 82, 86, 100, 120, 128 pachnitis Meyrick, 364, 368 pachyneura Turner, 321 Paedisca Treitschke, 306, 307, 334, 336 pagerostoma Diakonoff, 403, 405, 406 Palaeobia Meyrick, 229, 234, 235, 392, 395 palamedes Meyrick, 454, 456, 458 pallidula Meyrick, 383 pallifimbriana Bradley, 439 palmicola Meyrick, 378, 379 palmodes Meyrick, 280, 282 Pammene Hübner, 397, 406, 426, 427, 447, 461 Pammenemima Diakonoff, 44, 50, 55, 65, 397, 398, 403–406 Pammenitis Diakonoff, 426 Pammenodes Danilevsky & Kuznetzov, 461 Pammenopsis Kuznetzov, 1, 26, 47, 55, 66, 288, 397, 398, 426, 443–448 Pandanaceae, 96, 506 panegyrica Diakonoff, 353 panolbia Turner, 355 Parabactra, 210, 383 parachlora Diakonoff, 97 paradisea Diakonoff, 117 Paralobesia Obraztsov, 119, 474 Parapammene Obraztsov, 48, 55, 66, 397, 398, 426, 461–464 parastrepta Meyrick, 408 Parasuleima Clarke, 301, 305 Parepisimia Diakonoff, 108, 109, 111 Parienia Berg, 349, 353 Parinari, 461 Parthenium, 307, 309, 469 parvisignana Meyrick, 450, 451 parvulana Walker, 120 passercula Turner, 211 patris Diakonoff, 210 pauliani Diakonoff, 456 pedias Meyrick, 212, 215 pellopis Turner, 363, 367, 370 Pelochrista Lederer, 300 peltophora Meyrick, 111, 112, 113, 114, 115, 116, 117, 119, 120
120503•Olethreutine Moths 4pp.in519 519
519
peltosema Lower, 43, 392, 393, 394, 395 pendulata Meyrick, 192, 193, 194, 195, 196 penechra Diakonoff, 355 Penthina Treitschke, 68, 156, 173, 306, 422 Penthostola Diakonoff, 93, 94, 97 peraea Meyrick, 370 percnobathra Meyrick, 285 percnophylla Meyrick, 73, 78 perdigna Kuznetzov, 242 peregrinana Möschler, 306 perfixa Turner, 370 perfracta Diakonoff, 436 pericyphana Meyrick, 367, 371 Peridaedala Meyrick, 317, 320, 321, 330, 331, 334, 345 Periphoeba Bradley, 18, 38, 60, 220, 221, 242, 243, 246, 278–282 periptycha Turner, 370 permixtana Hübner, 111 perplexa Turner, 313 persinuata sp. nov., 1, 417, 418, 419 perspectana Walker, 371 perversa Turner, 306 pervicax Meyrick, 45, 419, 421, 422 pessota Meyrick, 399, 400, 401, 402, 403 petrifraga Diakonoff, 121, 124 petulantana Kennel, 461 Phaecadophora Walsingham, 162, 163, 165 Phaecasiophora Grote, 100, 102 phaedropa Turner, 306 phaeoloma Turner, 355 phaeoscia Turner, 370 phaeosigma Turner, 32, 129, 130, 131, 132, 134 Phalaena Linnaeus, 422, 423, 426, 448, 452 Phanerophlebia Diakonoff, 436 phanerops Diakonoff, 199 Phaneta Stephens, 300 Phanetoprepa Obratsov, 423 Phaseolus, 450 Phaulacantha Diakonoff, 185 phaulomorpha Meyrick, 443 phaulopa Meyrick, 207 Philalcea Stephens, 229 phileris Lower, 235 phileris Meyrick, 229, 230, 235 philobrya Meyrick, 324 phloeotis Meyrick, 285, 286, 287, 288 phoenicoplaca Turner, 156 Phoxopteris Treitschke, 210, 229 Phoxopteryx Sodoffsky, 229, 234 Phthinolophus Dyar, 356 Phthoroblastis Lederer, 461 phycitipalpia sp. nov., 1, 326, 327, 328, 329, 331, 332, 341 phyllanthana Meyrick, 135, 136, 137, 138, 139 phyllodes Lower, 34, 184, 185, 186, 187 physalodes Meyrick, 345, 346, 348, 349 physophora Lower, 31, 111, 112, 113, 114, 115, 116, 117, 118, 119, 120 Pimelodendron, 348 Piper, 221, 241, 242 Piperaceae, 221, 241, 507 Pirireisia Koçak, 112, 118 Pittosporaceae, 84, 450, 506 Pittosporum, 84, 87 placida Meyrick, 101, 104 plagana Haworth, 206, 207
10/5/06 12:01:48 PM
520
INDEX
plana Meyrick, 226, 227, 228, 229 Planchonia, 124 Platypeplus Walsingham, 129, 133, 134, 139, 178, 439 plebeiana Meyrick, 306 plebejana Zeller, 22, 23, 40, 51, 57, 301, 302, 303, 304, 305, 306 pleurogramma Clarke, 341, 343 pleurosema Turner, 216 plinthinana Meyrick, 362, 363, 365, 367, 368, 369, 370 plumbosana Bradley, 406 plumboviridis Diakonoff, 98 Plutographa Diakonoff, 313, 317 Poaceae, 50, 206, 209, 221, 290, 305, 430, 507, 508, 509 Podognatha Diakonoff, 18, 33, 63, 100, 119, 146, 152, 156–161 poecilaspis Meyrick, 264, 266 poetica Meyrick, 252, 253, 254, 255, 256, 257 Pogonozada Hampson, 436 Poincianna, 439 Polemograptis Meyrick, 271, 476 poliophylla Turner, 370 Polyalthia, 159, 256 Polychrosis Ragonot, 82, 111, 112, 117, 118, 120, 129, 133, 170, 212, 215, 216 polymetalla Turner, 35, 218, 219 polymita Turner, 371 polyochtha Diakonoff, 34, 200, 201, 202, 203 polyphaea Turner, 302, 303, 304, 305, 306 polyplega Turner, 371 Polyscias, 324 Pomaderris, 233 pomana Fabricius, 426 Pomatophora Diakonoff, 190 pomonana Denis & Schiffermüller, 426 pomonella Linnaeus, 22, 45, 57, 398, 406, 422, 423, 424, 425, 426 Pongamia, 443 postvittana Walker, 22 pottsi Clarke, 133, 139 praeceps Meyrick, 196, 197, 198, 199 Proschistis Meyrick, 18, 26, 34, 64, 80, 120, 196, 199–203, 205 Proselena Meyrick, 235 prospera Meyrick, 37, 257, 258, 259, 260, 261, 262, 294 Protancylis Diakonoff, 396 Proteaceae, 71, 115, 138, 193, 221, 234, 430, 432, 439, 506, 507, 509, 510 Proteopteryx Walsingham, 306 Proteoteras Riley, 317, 336 Protithona Meyrick, 349, 374 Protobactra Diakonoff, 207, 210 protosema Diakonoff, 34, 48, 196, 197, 198, 199, 200 prunivora Walsh, 454 Prunus, 221 psammetalla Lower, 212, 213, 214, 215, 216 psammitis Turner, 210 Psegmatica Meyrick, 349 Pseudacroclita Oku, 235 Pseudancylis gen. nov., 1, 18, 38, 59, 282–285 Pseudoclita Bradley 348, 349 Pseudogalleria Ragonot, 426, 427, 430, 439 pseudonectis Meyrick, 418, 419 Pseudopammene Komai, 397, 426 Pseudophiaris Obraztsov, 242, 270 Pseudotomoides Obraztsov, 423
120503•Olethreutine Moths 4pp.in520 520
pseusta Diakonoff, 74, 76 pseustis Meyrick, 235 Psidium, 132, 138, 357, 378, 432 Psilacantha Diakonoff, 100, 102 Psydrax, 389 Pternidora Meyrick, 17, 18, 38, 66, 220, 285–288, 290 ptiladelpha Meyrick, 306 ptychora Meyrick, 408 pullatana Bradley, 69, 73 punctidiscanum Dyar, 430 purpurascens Diakonoff, 199 pycnochra Meyrick, 426 Pyralis Linnaeus, 229, 266, 271, 426, 448 pyrrhocrossa Meyrick, 34, 203, 204, 205 pyrrhopa Lower, 37, 252, 253, 254, 255, 256, 257 pythonias Meyrick, 330 quartaria Diakonoff, 129 Quercus, 71 querula Meyrick, 71 quietana Meyrick, 350, 351, 352, 353, 355 ramella Linnaeus, 229 Ranunculaceae, 183, 197, 507 Rapanea, 84, 87, 92, 307, 313 rebellis Meyrick, 158, 160, 161 reliquana Hübner, 111, 118 reticulata Turner, 68, 73 Retinia Guenée, 306 rhacodyta Meyrick, 245, 246 Rhadinoscolops Obraztsov, 266, 270 Rhamnaceae, 233, 234, 324, 507, 508 Rhectogonia Diakonoff, 18, 31, 59, 80, 97, 120, 124, 125–128 Rhizophoraceae, 439, 510 Rhodacra Diakonoff, 18, 34, 63, 80, 203–205 rhodantha Meyrick, 173, 176 Rhodonympha Diakonoff, 173, 175, 176 Rhodosphaera, 324 Rhopalovalva Kuznetzov, 331 Rhopaltriplasia Diakonoff, 331 Rhopobota Lederer, 18, 40, 51, 58, 184, 299, 300, 307, 317, 321–325, 330, 345, 388 rhothia Meyrick, 357 Rhus, 320, 330, 476 Rhyacionia Hübner, 300, 306 rhynchias Meyrick, 437, 438, 439 Ricinus, 161 rivulifera Turner, 373, 374, 375 roerigii Zacher, 427 Rosa, 138 Rosaceae, 138, 221, 233, 324, 425, 454, 506, 507, 508, 509, 510 rostrifera Meyrick, 284, 285 Rubiaceae, 389, 509 rubrisignis Turner, 235 ruficomana Meyrick, 351, 352, 355 rupifera Meyrick, 203, 205 rusticula Meyrick, 380, 383 Rutaceae, 50, 84, 87, 233, 250, 430, 432, 439, 506, 507, 509, 510 Ruthita Clarke, 257, 261 rutilescens Turner, 46, 433, 434, 435 Saccharum, 290 Salacia, 430 Salicaceae, 138, 233, 507
5/5/06 2:45:32 PM
INDEX
salicella Linnaeus, 136, 138 Saliciphaga Falkovitsh, 162, 163 saligneana Clemens, 307 saloris Diakonoff, 166 sanderyi sp. nov., 1, 456, 457, 458, 459 Sapindaceae, 115, 132, 138, 182, 183, 193, 406, 425, 439, 464, 506, 507, 509, 510 sapphiropa Meyrick, 262, 264 sappiroflua Diakonoff, 196, 198, 199 Sasa, 221 sauteri Horak, 17, 18, 478, 479, 480, 481, 482, 483, 484, 485, 486 saxana Walker, 361 scalopias Meyrick, 210 scambodes Meyrick, 258, 259, 260, 262, 298 scaristis Meyrick, 129, 130, 131, 132, 133, 134 schistaceana Snellen, 221, 288, 289, 290, 291, 472 Schlechtendahlia (Aphididae), 330, 476 Schoenotenes Meyrick, 403 Sciaphila Treitschke, 216, 313, 356, 361, 467 sciodelta Meyrick, 235 scioessa Turner, 430 scoliomelas Diakonoff, 362 scopulosana Meyrick, 40, 317, 318, 319, 320, 321 scythropa Meyrick, 210 seclusana Walker, 448 sedifera Meyrick, 212, 213, 214, 215, 216 seditiosana Meyrick, 371 segetana Meyrick, 232, 235 segregana Kuznetzov, 330, 333 Selania Stephens, 397, 426 selectana Christoph, 461, 463 semaeophora Diakonoff, 193 Semasia Stephens, 422, 423 semicanella Walker, 22, 42, 355, 356, 357, 358, 359, 360, 361 Semniotes Diakonoff, 190 semnodryas Meyrick, 196, 199 Semnostola Meyrick, 220 senior Diakonoff, 182 Senna, 154, 412 seriana Kennel, 423, 470 servilisana Walker, 361 servillana Duponchel, 422 Sesbania, 418, 439, 443 sicariana Meyrick, 370 siculana Hübner, 229 Sida, 305 siderea Turner, 138, 139 Sideria Guenée, 229 siderostola Diakonoff, 76, 78 silvana Kuznetzov, 423 Simaethis Leach, 475 simpsonii Busck, 426 sinapichroa Turner, 44, 402, 403 Sisona Snellen, 102 Socioplana Diakonoff, 176 sollicitana Meyrick, 370 solomonensis Bradley, 193 Sophora, 408, 418 sordida Turner, 29, 68, 69, 70, 71, 72, 73 Sorolopha Lower, 1, 18, 26, 30, 50, 63, 64, 80, 93–99, 128, 184 spanistis Meyrick, 371 Spanistoneura Diakonoff, 426 spermatophaga Diakonoff & Bradley, 229, 297, 298
120503•Olethreutine Moths 4pp.in521 521
521
sphaerocopa Meyrick, 93 sphaerocosmana Meyrick, 104 sphenophora Turner, 362 spiladorma Meyrick, 343, 345 Spilonota Stephens, 1, 18, 24, 26, 42, 49, 54, 59, 61, 80, 299, 300, 330, 331, 349, 350–355, 357, 360, 361, 362, 367, 369, 370, 371, 372, 376, 378, 379, 387, 388 Spilonota-group, 1, 54, 80, 299, 331, 349, 372, 376, 378 Spinobactra Diakonoff, 207, 210 spinosa Diakonoff, 207 spirographa Diakonoff, 212, 215, 216 spodostola Turner, 364, 367, 368, 371 sponditis Meyrick, 165 Sporocelis Meyrick, 199, 202 spumans Diakonoff, 166, 168 squamosa Diakonoff, 73, 74, 76, 77, 78 Statheromeris Diakonoff, 190, 193 Statherotis Meyrick, 18, 34, 50, 57, 63, 80, 148, 149, 151, 161, 168, 183, 187, 188, 190–196 Statherotis-group, 149, 161, 183, 188, 190 Steganoptycha Stephens, 306, 310, 375 stenampyx Diakonoff, 236, 238, 239 stenophylla Turner, 370 stenotes Clarke, 375, 378, 379 Sterculiaceae, 430, 509 Steriphotis Meyrick, 111, 117, 120 Stigmonota Guenée, 448, 450, 451, 452 stilpna Turner, 235 stilpnosticta Turner, 161 straminea Butler, 207 straminea Meyrick, 207 strenuana Walker, 40, 307, 308, 309, 310 Strepsiceros Meyrick, 356, 361, 362, 370, 371 Strepsicrates Meyrick, 18, 22, 42, 54, 57, 61, 99, 107, 108, 299, 300, 349, 355–362, 369 striphromita Turner, 371 Strobila Sodoffsky, 422 strobilana Hübner, 422 strobilella Linnaeus, 423 Strophedra Herrich-Schäffer, 397, 426, 427, 466, 467 Stylobasium, 154 Styphelia, 378 subpallida Turner, 362 subsequana Herrich-Schäffer, 335, 336 subversana Zeller, 310 Surianaceae, 152, 154, 506 Sycacantha Diakonoff, 17, 18, 26, 31, 63, 99, 100–104, 105, 107, 108, 111, 119, 120, 161 Sycacantha-group, 99, 100, 111, 119, 120, 161 symplecta Turner, 104 symploca Turner (Eucosma), 285, 311, 312, 313 symploca Turner (Lobesia), 118, 120 synneurota Meyrick, 306 synomotis Meyrick, 230, 231, 232, 234, 235 syntaractis Turner, 212 Syntozyga Lower, 17, 18, 35, 50, 64, 206, 210, 212–216 Syzygium, 96, 132, 378 Tamarindus, 456 tamias Diakonoff, 156, 157, 158, 161 Taniva Heinrich, 219, 469 taophanes Meyrick, 443 tapina Turner, 370 tarachodes Meyrick, 371 tatarana Bradley, 383, 386, 387, 388
5/5/06 2:45:32 PM
522
INDEX
Taxaceae, 508 telifera Chrétien, 331 temenopsis Meyrick, 97 Temnolopha Lower, 18, 33, 62, 100, 129, 139, 161, 162–165, 168, 172, 173, 178, 190, 206, 257 Templemania Busck, 17, 18 tenebrosa Turner, 180, 181, 182, 183 tenuitexta Diakonoff, 362 tephraea Meyrick, 412 Tephrosia, 418 testudinea Turner, 209, 210 tetanota Meyrick, 138 tetramita Turner, 44, 404, 405, 406 Tetramoera Diakonoff, 18, 38, 61, 221, 288–292, 298 tetrarcha Meyrick, 149, 150, 151, 195 tetrazancla Turner, 450, 452 thalassinana Meyrick, 42, 362, 365, 366, 367, 368, 369, 371 Thaumatotibia Zacher, 46, 55, 59, 396, 397, 398, 426, 427–430, 435, 439 Theaceae, 182, 430, 509 Theorica Diakonoff, 119, 156, 159, 160 Thiodia Hübner, 300 thoenarcha Meyrick, 270, 271 Thylacandra Diakonoff, 427 Thysanocrepis Diakonoff, 18, 39, 61, 151, 156, 161, 220, 278, 284, 288, 292–295, 298 thysanota Meyrick, 330 thystas Meyrick, 1, 184, 185, 186, 187 Tia Heinrich, 219 Tiliaceae, 464, 510 tinctoria Meyrick, 382, 383 Tinea Linnaeus, 330, 448, 471, 476 Titanotoca Diakonoff, 398, 403, 406 Tmetocera Lederer, 350 Tokuana Kawabe, 282 tonica Meyrick, 34, 183, 187, 188, 189, 190 tonsoria Meyrick, 122, 124 Toona, 51, 221, 297, 298, 470, 508 Toonavora gen. nov., 1, 18, 39, 51, 58, 60, 221, 229, 290, 295–298 tornocosma Turner, 306 tornocycla Turner, 306 tornosema Turner, 361 tornosticha Turner, 449, 452 Tortrix Linnaeus, 73, 111, 124, 156, 206, 207, 216, 229, 257, 321, 350, 403, 422, 423, 426, 448 Tospitis, 271 tothastis Meyrick, 300, 395, 396 toxosema Turner, 196 trachynota Meyrick, 337 Trachyschistis Meyrick, 18, 32, 54, 63, 129, 134, 136, 139–142, 144 tranquilla Meyrick, 370 transfixa Turner, 361 transtrifera Meyrick, 118, 120 transversa Diakonoff, 216 trepida Meyrick, 38, 279, 281, 282
120503•Olethreutine Moths 4pp.in522 522
triangulana Meyrick, 166, 349, 360, 363, 365, 366, 367, 368, 369, 370 tricentra Meyrick, 418, 419 trichograpta Meyrick, 440, 442, 443 tricia Clarke, 348 tricolorana Meyrick, 33, 165, 166, 167, 168, 169, 170 trigonana Walsingham, 71 Tritopterna Meyrick, 18, 41, 51, 57, 58, 299, 317, 331, 336, 338–341 truculenta Meyrick, 210 tumulata Meyrick, 321 turifera Meyrick, 271 uberana Meyrick, 1, 37, 257, 258, 259, 260, 261, 262, 295 Ukamenia Oku, 177, 178, 182, 183 uncimacula Turner, 33, 161, 170, 171, 172, 173 uptoni sp. nov., 1, 36, 243, 244, 245, 246, 247, 248 venosana Zeller, 35, 207, 208, 209, 210, 211 vera Diakonoff, 173 Verbenaceae, 115, 138, 182, 305, 506, 508 verutana Zeller, 206 Vigna, 408, 418 vinculata Meyrick, 33, 156, 157, 158, 159, 160, 161 vindemians Meyrick, 165, 166, 167, 168, 169, 170 Vitaceae, 115, 506 viteana Clemens, 119 Vitex, 343 vitiosa Meyrick, 370 Vitis, 115, 187, 507 volutana Meyrick, 235 vulgana Walker, 183, 187 vulnerata Walsingham, 271 wahlbergiana Zeller, 161, 190 Whittenella gen. nov., 1, 18, 43, 54, 58, 300, 392–395 Williella Horak, 17, 18 Xanthium, 309 xanthorhina Diakonoff, 108 Xenolepis Diakonoff, 119 Xenopotamia Diakonoff, 161 Xenosocia Diakonoff, 349, 353 xuthochyta Turner, 211 xylistis Lower, 111, 113, 116, 118, 120 zaleuta Meyrick, 199, 201, 202, 205 zapyrana Meyrick, 17, 18, 24, 47, 449, 450, 451, 452 Zeiraphera Treitschke, 300 Zingiberaceae, 163, 507 Zizyphus, 234 Zomaria Heinrich, 119, 151, 152, 156, 159, 160, 161 Zomaria-group, 119, 151, 152, 156, 159, 160, 161 Zomariana Diakonoff, 18, 33, 50, 54, 57, 62, 100, 151, 152–156, 159, 162, 178, 206 zopherana Meyrick, 24, 42, 367, 370 zophophanes Turner, 46, 428, 429, 430 zophotypa Turner, 362
5/5/06 2:45:33 PM
