Claudio Jose Barros de Carvalho is Full Professor at the Department of
Zoology of the Universidade Federal do Parana; fellow of the CNPq, with many papers published in Systematics, Bionomics and Biogeography of Diptera. A former student in the Museu ~acional,
Rio de Janeiro, he did his
Ph.D. studies in Curitiba, and in the ~arIy
1990's, he spent a year at the
~atural History Museum in London.
To
date. 20 graduate students have obtained their degrees under him. ~Iarda Souto Couri is Professor at the
.\Iuseu Nacional, Universidade Federal do Rio de Janeiro; C~Pq.
fellow of
the
with expressive scientific
contributions on Systematics, Bionomics and Phylogeny of Diptera. \Yith a Ph.D. degree in Parasitology trom the Universidade Federal Rural do Rio de Janeiro and a postdoctoral project developed at the Natural History Museum, London, she has under her leadership a team of researchers and students. She has also supelTised several M.Sc. and Ph.D. candidates in Zoology.
Muscidae (Diptera) of the Neotropical Region:
TAXONOMY
.. UFPR
Reitor Carlos Roberto Antunes dos Santos Vice-Reitor Romolo Sandrini Diretor da Editora da UFPR Luiz Carlos Ribeiro
Conselho Editorial Alberto Pio Fiori Andre de Macedo Duarte Fany Reicher Izaura Hiroko Kuwabara Jose Carlos Cifuentes Vasquez Leilah Santiago Bufrem Manoel Eduardo A. Camargo e Gomes Maria Benigna Martinelli de Oliveira Pedro Ronzelli Junior Sergio Herrero de Moraes Victor Manoel Pelaez Alvarez
Muscidae (Diptera) of the Neotropical Region:
TAXONOMY
Edited by Claudio Jose Barros de Carvalho
~ UFPR
© Claudio Jose Barros de Carvalho
Muscidae (Diptera) of the Neotropical Region: Taxonomy Coordena~ao
editorial: Marildes Rocio Artigas Santos Revisao de texto: Gabriel Augusto Rodrigues de Melo Revisao final: do Autar Editora~ao eletronica: Alquimia Esrudio de Arte Final Fotos: Philippe Wauters Capa: Se9ao de Programa9ao Visual da Universidade Federal do Rio de Janeiro Serie Pesquisa n. 68 Coardena9ao de Processos Tecnicos. Sistema de Bibliotecas, UFPR Muscidae (Diptera) of the Neotropical Region: taxonomy / edited by Claudio Jose Barros de Carvalho. - Curitiba : Ed. UFPR, 2002. 287 p. : il. - (Pesquisa ; 68) ISBN 85-7335-091-1 1. Muscidae. 2. Diptera. 3. Entomologia. I. Carvalho, Claudio Jose Barros de. II. Titulo. III. Serie.
CDD 595.770913 CDU 595.77(213)
ISBN 85-7335-091-1 Ref. 315 Apoio: Funda9ao Araucana Editora UFPR Centro Politecnico - Jardim das Americas Caixa Postal 19 .029 81531-980 - Curitiba - Parana - Brasil tel.Jfax: (41) 361-3380/361-3381/267-5973 e-mail:
[email protected]
2002
FOREWORD
iosystematic revisions are vitally essential in biological research as guide and reference work to the continually expanding knowledge of the fauna and flora. They bring together, assimilate and organize scattered information from around the world so that students and researches can quickly ascertain the current status of the taxonomic hierarchy and find reference to many kinds of information. The dipterous of the family Muscidae constitute an important group of human and animal pest, widely distributed in the world fauna. Larvae breed mainly in decaying plant and animal material or manure. Adults of many species bite or passively vector pathogens for disease such as typhoid fever, dysentery, anthrax, etc. Although numerous, ubiquitous, and economically important, muscids have not had the attention they deserve. More than 4000 described species are recorded worldwide, but only 84 genera and about 840 species of them are known from the Neotropical Region and the actual number that must occur in this region is inestimable. No comprehensive monograph has ever published to this area of the world and the present contribution of Claudio Jose Barros de Carvalho and Marcia Souto Couri is well warranted. Subfamilies, tribes and genera are revised, and keys for identification of all genera and species with Neotropical occurrence are proposed. The authors have undertaken a formidable task, and the present work will constitute a significant step in the systematics of N eotropical muscids providing a basic reference for further research.
B
Jose Henrique Guimaraes Associate Professor, rCB Universidade de Sao Paulo February 2001
PREFACE
ow great are God's riches! How deep are his wisdom and knowledge! How impossible to explain his decisions or to understand his methods! As Scripture says, "Who could ever known the mind of the Lord?" Who could ever be his counsellor? ... For all things were created by him and all things exist through him and for him. To him be the glory forever! Amen. [Romans, Ch. 11:33-36] apud McALPINE (1989). The scientific literature shows that there is no consensus about the definition of some words commonly used by a taxonomist or a systematist. Systematics and some related words as taxonomy, classification and biological system have received different definitions by different authors (MINELLI, 1993). Among the definitions still accepted, there are the ones of SIMPSON (1961) for Systematics, Taxonomy and zoological classification. According to this author, Systematics is the scientific study of the groups, of the diversity of the organisms and of all the relations among them. On the other side, Taxonomy is the theoretic study of the classification, including its bases, principles, procedures and rules. Zoological classification is the arrangement of the animals into groups having their similarities as a basis. In his point of view, Taxonomy is part of the Systematic and so, subordinated to it. However, about 30 years ago, NELSON (1970) introduced a new concept of Comparative Biology (WILEY, 1981;AX, 1987; BROOKS & McLENNAN, 1991), where characters of the diverse taxa are analysed comparatively, clearly seeking for similarities and differences among them. This is the study of the diversity patterns, looking for the comprehension of the origin of the patterns of similarities and differences among the groups (AMORIM, 1997). So, Comparative Biology is understood as the study of the biological diversity under a historical perspective (NELSON & PLATNICK, 1981). Its main scope is the reconstruction of the similarity and dissimilarity patterns that exist inside
H
the organic diversity, both concerning the diversity of forms or attributes as well as their geographical distribution. Systematics and Taxonomy are areas that integrate the Comparative Biology. According to some authors (MINELLI, 1993), Taxonomy can be understood as a synonym of Systematics and has the role to produce taxonomic information, including classification, nomenclature, description, keys to identification and to show the biological inter-relations among the analysed taxa. In other words, Systematics has, as one of its main roles, to present a general reference system on the biological diversity (HENNIG, 1966). The scarce know ledge of the dipterofauna of the neotropics has other reasons than the different academic interpretations of Systematics and Taxonomy described above. About 30,000 species ofDiptera in the neotropics are known nowadays (Nelson Papavero, personal communication), what represents a very timid number in relation to the real number of species that live in this region. In Brazil, two main points bring difficulties to the knowledge of the dipterofauna: the abundant richness associated with the reduced number of qualified persons to study it. As it was pointed by THOMPSON (1990), new technologies, research training and also new work conditions are necessary to the comprehensible know ledge of the Diptera of any region. The union of efforts of several levels is necessary to change this reality in a relatively short period of time. But for sure, one of the first steps is the publication of basic information for the identification of the species as well as information on the phylogenetic relations among them and also on the patterns of their distribution (CARVALHO, 1998). Using this know ledge for a more consolidated for a more consolidate study of the biodiversity, as well as, through the development of new theories in Systematics, and a better comprehension of the phylogeny, this scenery should change quickly. Unfortunately, in the threshold of the Third Millennium, the knowledge of the Diptera of the Neotropical Region is still incipient (GRAZIA et ai., 2000). The present publication is a great step to the comprehension of the muscid biodiversity. Before undertaking any study of a basic or practical nature, it is always necessary to know with whom we are dealing, or, in other words, to know who is whothat is the beginning of everything.
References AMORIM, D. de S. 1997. Elementos basicos de Sistematica Filogenetica; 2' edi'tao, revista e ampliada. Holos & Sociedade Brasileira de Entomologia, Ribeirao Preto, 276 p. AX, P. 1987. The Phylogenetic System. The systematization of organisms on the basis of their phylogenesis. John Wiley & Sons, Chichester, 340 p. BROOKS, D.R. & McLENNAN, D.A. 1991. Phylogeny, Ecology, and Behavior. The University of Chicago Press, Chicago, 434 p. CARVALHO, CJ.B. de. 1998. Taxonomista de insetos, uma especie em extin'tao no Brasil. Inf. Soc. Entomol. Bras. 23: 1. GRAZIA, J.; CJ.B. de CARVALHO; L.M. de ALMEIDA; M.M. CASAGRANDE & O.H.H. MIELKE. 2000. A biodiversidade de insetos no Brasil no Terceiro Milenio: dificuldades e perspectivas. Entomol. Vect. 7: 123-141. HENNIG, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, IlL, 263 p. McALPINE, J.E 1989. Phylogeny and classification of the Muscomorpha, p. 1397-1518. In: McAlpine, J.E & D.M. Wood (eds.), Manual ofNearctic Diptera, volume 3. Research Branch, Monograph No. 32, p. 1333-1581. MINELLI, A. 1993. Biological Systematics; the state of the art. Chapman & Hall, London, 387 p. NELSON, GJ. 1970. Outline ofa theory of comparative biology. Syst. Zool. 20: 471-472. NELSON, G.E & PLATNICK, N.L 1981. Systematics and Biogeography, Cladistics and Vicariance. Columbia University Press, New York, 567 p. SIMPSON, G.G. 1961. Principles of Animals Taxonomy. Columbia University Press, New York, 254 p. THOMPSON, EC. 1990. Biosystematic information dipterists ride the third wave, p. 179-201. In: M. Kosztarab & C.W. Schaefer. Systematics on the North American Insects and Arachnids: status and needs. Virginia Agricultural Experiment Station Information series 90-1. Blacksburg: Virginia Polytechnic Institute and State University. WILEY, E.O. 1981. Phylogenetics; the theory and practice of phylogenetic systematics. New York, Wiley Sons, 439 p.
Claudio Jose Barros de Carvalho April 2002
ACKNOWLEDGEMENTS This book is a result of about 25 years dedicated to the study of the muscids flies. During this period an intense scientific exchange and a great friendship grew among us since the beginning of our scientific life in the Museu Nacional, Rio de Janeiro. Along this whole path, many common colleagues and friends have given us valuable help and to all these people we would like to offer our special thanks - Adrian Charles Pont (Oxford University Museum), Oxford, Peter Skidmore, Swansea, and our many colleagues at The Natural History Museum (London), The National Museum ofNatural History (Washington), The Canadian National Collection (Ottawa), Universidade Federal do Parana (Curitiba) and the Museu N acional, Universidade Federal do Rio de Janeiro (Rio de Janeiro), for study facilities and much valuable discussion. We also are grateful to the Conselho Nacional de Desenvolvimento Cientifico e Tecnol6gico, an agency of the Brazilian Government, for scientific and technological development, for the support provided by a grant to CJBC (Proc. Nr. 300043/86-4) and to MSC (Proc. Nr. 300386/80-0). More specifically to the elaboration of this book, we would like to thank Luciane Marinoni and Sonia Maria Noemberg Lazzari (Universidade Federal do Parana, Curitiba), Jose Albertino Rafael (Instituto Nacional de Pesquisas daAmazonia, Manaus), all our present graduates and undergraduates students for their support and critical reviews of earlier versions of the manuscript. To Denise Pamplona (Museu N acional, Rio de Janeiro) and one anonymous referee for their careful and critical review of the final version of the manuscript. To Jose Henrique Guimaraes for the foreword. To Gabriel Augusto Rodrigues de Melo (Universidade Federal do Parana, Curitiba) for revising the English. We owe a special debt to Philippe Wauters, who, with his sensibility, enthusiasm and photography skills, knew how to capture the spirit of the flies. We also thank the Section ofPrograma9ao Visual of Un iversidade Federal do Rio de Janeiro for the cover design. Claudio Jose Barros de Carvalho Marcia Souto Couri
CON'!'ENTS PART I. BASAL GROUPS By Claudio Jose Barros de Carvalho & Marcia Souto Couri INTRODUCTION .................................................................................................... 17 KEY TO NEOTROPICAL GENERA OF MUSCIDAE ............................................. 21 DIAGNOSIS AND KEY TO SPECIES ..................................................................... 37 SUBFAMILY ATHERIGONINAE FAN, 1965 ........................................................... 38 Atherigona Rondani, 1856 ................................................................................. 38 SUBFAMILY MUSCINAE LATREILLE, 1802 ......................................................... 39 Tribe Muscini Latreille, 1802 ................................................................................ .40 Biopyrellia Townsend, 1932 ............................................................................... .40 Cyacyrtoneura Townsend, 1931 ........................................................................ .41 Dasymorellia Malloch, 1923 ............................................................................... .41 Morellia Robineau-Desvoidy, 1830 ..................................................................... .42 Musca Linnaeus, 1758 ....................................................................................... 48 Neomyia Walker, 1859 ....................................................................................... 49 Neorypellia Pont, 1972 ....................................................................................... 49 Parapyrellia Townsend, 1915 .............................................................................. 50 Polietina Schnabl & Dziedzicki, 1911.. .................................................................. 51 Sarcopromusca Townsend, 1927 .......................................................................... 55 Xenomorellia Malloch, 1923 .............................................................................. 57 Tribe Stomoxyini Meigen, 1824 .............................................................................. 58 Haematobia Le Peletier & Serville in Latreille et aI., 1828 .................................... 59 Neivamyia Pinto & Fonseca, 1930 ......................................................................... 59 Stomoxys Geoffroy, 1762 ..................................................................................... 60 SUBFAMILY AZELIINAE ROBINEAU-DESVOIDY, 1830 ...................................... 62 Tribe Azeliini Robineau-Desvoidy, 1830 ..................................................................... 62 Azelia Robineau-Desvoidy, 1830 ........................................................................... 62 Drymeia Meigen, 1826 ....................................................................................... 63 Hydrotaea Robineau-Desvoidy, 1830 ................................................................... 63 Micropotamia Carvalho, 1993 .............................................................................. 68 Ophyra Robineau-Desvoidy, 1830 ....................................................................... 70 Potamia Robineau-Desvoidy, 1830 ....................................................................... 74 Thricops Rondani, 1856 ....................................................................................... 74
Tribe Reinwardtiini Brauer & Bergenstamm, 1889 .................................................... 76 Brachygasterina Macquart, 1851 ........................................................ ,............... 76 Chaetagenia Malloch, 1928 ............................................................................... 78 Correntosia Malloch, 1934 ................................................................................. 79 Dalcyella Carvalho, 1989 ................................................................................. 79 Itatingamyia Albuquerque, 1979 ......................................................................... 80 Muscina Robineau-Desvoidy, 1830 ...................................................................... 80 Palpibracus Rondani, 1863 ................................................................................ 82 Philornis Meinert, 1890 ...................................................................................... 88 Psilochaeta Stein, 1911 ...................................................................................... 96 Reinwardtia Brauer & Bergenstamm, 1889 .......................................................... 97 Synthesiomyia Brauer & Bergenstamm, 1893 ....................................................... 98 SUBFAMILY PHAONIINAE MALLOCH, 1917 .......................................................... 98 Dolichophaonia Carvalho, 1993, stat. rev ........................................................... 99 Helina Robineau-Desvoidy, 1830 ......................................................................... 105 Phaonia Robineau-Desvoidy, 1830 ....................................................................... 119 Souzalopesmyia Albuquerque, 1951 .................................................................. 126
PART II. APICAL GROUPS By Marcia Souto Couri & Claudio Jose Barros de Carvalho INTRODUCTION ..................................................................................................... 133 DIAGNOSIS AND KEY TO SPECIES ..................................................................... 135 SUBFAMILY CYRTONEURININAE SNYDER, 1954 .............................................. 135 Arthurella Albuquerque, 1954 ............................................................................. 135 Cariocamyia Snyder, 1951 ................................................................................. 136 Charadrella Wulp, 1896 ..................................................................................... 137 Chortinus Aldrich, 1932 ..................................................................................... 139 Cyrtoneurina Giglio-Tos, 1893 ............................................................................. 139 Cyrtoneuropsis Malloch, 1925 ............................................................................. 141 Mulfordia Malloch, 1928 ..................................................................................... 149 Neomuscina Townsend, 1919 ................................................................................ 149 Neomuscittiopsis Albuquerque & Lopes, 1992....................................................... 155 Neurotrixa Shannon & Del Ponte, 1926 ................................................................. 155 Pseudoptilolepis Snyder, 1949 .............................................................................. 156 Xenothoracochaeta Malloch, 1921 ....................................................................... 158 SUBFAMILY MYDAEINAE VERRAL, 1888 ............................................................. 160 Brontaea Kowarz, 1873 ..................................................................................... 160
Graphomya Robineau-Desvoidy, 1830 ................................................................. 163 Hemichlora Wulp, 1893 ................................"..................................................... 167 Mydaea Robineau-Desvoidy, 1830 ......................................................................... 167 Myospila Rondani, 1856 ................................................................................... 170 Scenetes Malloch, 1936, stat.rev.................................................... ..................... 171 Scutellomusca Townsend, 1931 ............................................................................ 172 SUBFAMILY COENOSIINAE VERRAL, 1890 .......................................................... 173 Tribe Limnophorini Villeneuve, 1902 ........................................................................... 174 Agenamyia Albuquerque, 1953 ........................................................................... 174 Albertinella Couri & Carvalho, 1997 ..................................................................... 176 Drepanocnemis Stein, 1911 ................................................................................. 177 Limnophora Robineau-Desvoidy, 1830 .................................................................. 178 Lispe Latreille, 1797 ............................................................................................ 182 Lispoides Malloch, 1920 ........................................................................................ 184 Pachyceramyia Albuquerque, 1955 ........................................................................ 187 Rhabdotoptera Stein, 1919 ................................................................................... 189 SpUo gona Schnabl, 1911 ....................................................................................... 190 Syllimnophora Speiser, 1923 ................................................................................. 192 Tetralnerinx Berg, 1898 ........................................................................................ 200 Thaumasiochaeta Stein, 1911 ............................................................................... 20 1 Tribe Coenosiini Verral, 1890 ................................................................................... 205 Apsil Malloch, 1929 ............................................................................................ 205 Bithoracochaeta Stein, 1911 ................................................................................ 211 Coenosia Meigen, 1826 ........................................................................................ 214 Cordiluroides Albuquerque, 1954 .......................................................................... 218 lnsulamyia Couri, 1982 ..................................................................................... 219 Neodexiopsis Malloch, 1920 ................................................................................ 220 Notoschoenomyza Malloch, 1934 .......................................................................... 237 Oxytonocera Stein, 1911 ...................................................................................... 238 Pentacricia Stein, 1898 ...................................................................................... 239 Pilispina Albuquerque, 1954 ................................................................................ 240 Plumispina Albuquerque, 1954 .............................................................................. 241 Reynoldsia Malloch, 1934 ..................................................................................... 242 Schoenomyza Halliday, 1833 ................................................................................. 247 Schoenomyzina Malloch, 1934 .............................................................................. 251 Spathipheromyia Bigot, 1884 .............................................................................. .254 Stomopogon Malloch, 1930 ............................................................................... 257
REFERENCES ...................................................................................................... 263
PART I
BASAL GROUPS By Claudio Jose Barros de Carvalho* & Marcia Souto Couri**
INTRODUCTION The Muscidae occurs in all biogeographical regions and contains more than 4000 species (PONT, 1986a, 1989). In the Neotropical Region, about 800 species are recognised (CARVALHO et al., 1993). Muscidae species have extremely varied habitsthe larvae can be coprophagous, saprophagous or carnivorous (SKIDMORE, 1985) and the adults of many species live on decaying animal or vegetable matter. Muscidae is one of the most interesting families ofDiptera, due to the association of many species with the antropobiocenosis (GREENBERG, 1971). Some species have a great medical and veterinary importance, acting as vectors of many diseases and other species have a considerable ecological importance. The adults are very abundant in rural or semi-rural areas. At high altitudes, they form a very high proportion ofthe fauna, both in number of species as in individuals (PONT, 1986a). The larvae of some species are agriculturally important, being primarily pests of plantations. Some species attack stored products with agricultural and commercial importance. * **
Universidade Federal do Parana, Departamento de Zoologia, C.P. 19020, Curitiba, 81.531-980, Parana, Brazil; email:
[email protected] Museu Nacional do Rio de Janeiro, Departamento de Entomologia, Quinta da Boa Vista, Rio de Janeiro, 20.940040, Brazil; e-mail:
[email protected]
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
17
Part I - Basal Groups
Muscidae can be recognised by diverse characters among the Muscomorpha (AMORIM & CARVALHO, 1992) as, for example, the absence ofa vertical series of setae on meron and the presence of developed calypteres. These characters identify the Muscoidea (sensu McALPINE, 1989), but the Muscidae shows some exclusive characters, a condition that indicates the monophyly of the family. Some authors (ROBACK, 1951; HENNIG, 1965, 1973; SKIDMORE, 1985; McALPINE, 1989; MICHELSEN, 1991; PONT & CARVALHO, in preparation) consider the family monophyly based on the presence ofthree derived characters: (1) female with the abdomen showing less than 7 pairs of spiracles (spiracles 6 and 7 were lost, but some genera not related amongst themselves, have reacquired the spiracle 6). This reduction in the number of spiracles is a unique apomorphy among the Calyptratae, (2) male reproductive system without the accessory pair of glands. Among the Muscoidea (McALPINE, 1989), this condition is present in the three other families - Fanniidae, Scathophagidae and Anthomyiidae, (3) larval mouth-hooks closely appressed to one another. Although the Muscidae are considered to be a family satisfactorily known worldwide (THOMPSON, 1990), in the Neotropical Region its taxonomy is still in incipient levels (CARVALHO, 1989a). Many classifications of this family were proposed by the authors. Until the beginning of the 1960's, typological classifications were presented (e.g. ROBACK, 1951; HENNIG, 1956, 1957, 1958, 1959, 1960a, 1960b, 1962a, 1962b, 1962c, 1962d, 1963a, 1963b, 1964), but HENNIG (1965) proposed the first classification for Muscidae based on cladistic methodology. Starting about that time, some catalogues (PONT, 1972, 1977, 1980, 1986a, 1989) were published following the classification proposed by HENNIG (1965), with few modifications. In the beginning of the nineties, CARVALHO et al. (1993) published a catalogue for the Neotropical Region following the classification proposed by CARVALHO (1989d). The natural classification of the family, however, still needs improvements (HUCKETT & VOCKEROTH, 1987). Many taxa are known only from the original descriptions or only from the adults forms. In this way, important characters for the understanding of the family were not properly evaluated, considering the family as a whole (CARVALHO, 1989a). A more consistent natural classification will be possible only when the Neotropical genera are fully revised. A fundamental contribution to the studies on the biology of the muscid of the 18
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Co uri
world was presented by SKIDMORE (1985), where the immature stages of about 440 muscid species were treated. His classification of the family was proposed mainly based on characters of the immature stages. Four years latter, CARVALHO (1989d) proposed a classification for the muscids, on the basis of a cladistic analysis of characters of the adult morphology. Species representing 27 genera, 12 of them exclusively Neotropical were utilised. The resulting cladogram was used to propose a phylogenetic classification with seven subfamilies and eight tribes: Acanthipterinae, Atherigoninae, Muscinae (Stomoxyini and Muscini), Azeliinae (Azeliini and Reinwardtiini), Phaoniinae, Mydaeinae (Mydaeini and Graphomyini) and Coenosiinae (Coenosiini and Limnophorini). This classification is very similar to the one proposed by SKIDMORE (1985). From 1989 on, many revisions of Neotropical muscid genera were published (CARVALHO, 1989a, 1989b, 1989c, 1993a, 1999; COURI & CARVALHO, 1992, 1993, 1997b;COURI&PAMPLONA, 1992;CARVALHO&COURI, 1993;ARAlJJO & COURI, 1993; COURI, 1995, 1996a, b, 1998a, 1999a, 2000a, b, in press; COURI & LAMAS, 1993; PALKA-ROCHA & CARVALHO, 1994; CARVALHO & PONT, 1998; MOTTA & COURI, 1999; PAMPLONA, 1999; COELHO, 2000). Several papers with descriptions of new genera or partial generic revisions were also recently published (COURI & ALBUQUERQUE, 1979; COURI, 1982a, b, 1989; PAMPLONA, 1983, 1986 a, b, c, 1992; COURI & LOPES, 1985, 1986, 1988a; CARVALHO, 1985a, b, 198ge; PAMPLONA & COURI, 1989, 1995, 1998b; LOPES, 1992; CARVALHO & COELHO, 1993; COURI & CARVALHO, 1995; COELHO, 1998a, b). These complete or partial revisions represent 50% of genera recognised by CARVALHO et al. (1993) for the Neotropical Region. Some keys for the identification of the Neotropical muscid genera are available in the literature (COURI & LOPES, 1985, 1986, 1988b; LOPES & COURI (1989a), VOCKEROTH, 1996; COURI & PONT, 1999). The patterns of distribution of the South American muscid species are scarcely known. A small number of papers on the historical biogeography have been published for subgroups of Muscidae, a few of them dealing with Neotropical groups. HENNIG (1965) indicated that the first muscid invasion in South America occurred from the Northern Hemisphere, during the Upper Cretaceous or at the beginning of the Tertiary Period, under Muscidae (Diptera) of the Neotropical Region: TAXONOMY
19
Part I - Basal Groups
a clearlydispersionist view. Recently CARVALHO (1999) presented an alternative hypothesis for the origin of the muscids in the Neotropical Region, based on the ideas proposed by AMORIM & PIRES (1996) and PONT & CARVALHO (1997). The transoceanic relations of some genera were also investigated by COURI & CARVALHO (2000). For the southern area of the South America, CARVALHO & COURI (in press) presented a hypothesis for the explanation of the distribution pattern of two small muscid clades. The scope of this chapter is to revise the taxonomy of the basal groups of Neotropical Muscidae (see CARVALHO, 1989d). Diagnoses for the subfamilies, tribes and genera, discussions of the supra-generic relationships, and keys for identification of all genera and species of basal groups are presented. The classification followed here was the one proposed by CARVALHO et al. (1993). The type-species of each genera are cited before the diagnosis. Known synapomorphic and autapomorphic characters were included in the diagnosis, and the monophyly of the subfamilies, tribes and genera are discussed when possible. The terminology used was taken mainly from McALPINE (1981) and COELHO (1997), the last paper in particular for the morphology of the proboscis. Some Latin abbreviations commonly used in taxonomic papers (see PAPAVERO, 1994) were also used. The key to genera was based mainly in the following papers: COURI & LOPES (1985,1986, 1988b), LOPES & COURI (1989a), VOCKEROTH (1996) and COURI & PONT (1999). VOCKEROTH (1996) presented a key for the identification of the genera of Muscidae occurring in Central America. In this key, he included Fannia RobineauDesvoidy and Euryomma Stein, genera of Fanniidae. This family is one of the most basal groups within the Muscoidea, and perhaps among the Calyptratae (VOSSBRINK & FRIEDMAN, 1989; MICHELSEN, 1991; CARVALHO et al., 1993 and BERNASCONI et al., 2000). The keys to species were based on diagnostic characters available in the literature, the study of type-material for some species and adaptations on some published keys. As in the Catalogue of the Neotropical Region by CARVALHO et al. (1993), the geographical coverage is the continent of South America as far north as, and including, Mexico; the islands of the Caribbean and the Antilles, including the Bahamas but excluding Bermuda; the Falklands (Malvinas), Galapagos Islands, Easter Island, and Juan Fernandez Islands, in the southern Atlantic and Pacific Oceans respectively. The known geographical distribution in the Neotropical Region was included in the keys. It was not possible to 20
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
include all species in the keys, since some of them are insufficiently known in the literature and the type-material was not examined. Most of the illustrations have been already published, but a few modifications were introduced. Abbreviations for the Museums and Institutions from which material was examined are as follows: AMNH - American Museum ofNatural History, New York, USA BMNH - The Natural History Museum, London, UK CNC- Canadian National Collection, Ottawa, Canada DZUP - Departamento de Zoologia, Universidade Federal do Parana (Cole<;ao de Entomologia Pe. Jesus Santiago Moure), Curitiba, Brazil IML- Instituto Miguel Lillo, Tucuman, Argentina INHS - Illinois Natural History Survey, Illinois, USA INPA - Instituto Nacional de Pesquisas da Amazonia, Manaus, Brazil MNHNP - Museum National d'Histoire Naturelle, Paris, France MNRJ - Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil MZSP - Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, Brazil NMW - N aturhistorisches Museum, Vienna, Austria SMT - Staatliches Museum fur Tierkunde, Dresden, Germany UMO - Hope Entomological Collections, University Museum, Oxford, UK USNM - National Museum of Natural History, Washington D.C., USA UZMC - Universitetets Zoologisk Museum, Copenhagen, Denmark WSU - Washington State University, Pullman, USA ZMHU - Museum fUr N aturkunde, Humboldt - Universitat, Berlin, Germany
KEY TO THE NEOTROPICAL GENERA OF MUSCIDAE 1.
Proboscis elongated, strongly sclerotized, not retractile, labella reduced (Figs. 39, 42,44); arista with long cilia on the dorsal surface and bare on ventral or, at most, with 3-4 ventral cilia (Figs. 42, 44); prosternum setulose ........................................ .2 Proboscis retractile, faintly or moderately sclerotized, labella not reduced; arista bare (Fig. 1) or setulose (Fig. 4), not as described above; pro sternum bare or
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
21
Part I - Basal Groups
setulose ................................................................................................................ 4 2.
Palpus measuring about one-third of proboscis length; propleuron setulose on center; meron setulose (Mexico, Guatemala, EI Salvador, Nicaragua, Costa Rica, Panama, Puerto Rico, Cuba, Jamaica, Virgin Is., St. Vincent, Haiti, Dominica, Trinidad, Tobago, Venezuela, Guyana, Colombia, Peru, Ecuador, Galapagos Is., Bolivia, Brazil, Uruguay, Argentina, Chile, Easter Is.; cosmopolitan) ................. Stomoxys Geoffroy [page 60] Palpus longer than one-third of proboscis length; propleuron and meron bare ........... 3
3.
Arista bare on ventral surface; notopleuron with no covering setulae (Mexico, Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Puerto Rico, Cuba, Jamaica, Virgin Is., Haiti, S. Domingo, Colombia, Venezuela, Guyana, Brazil, Chile, Argentina; Europe, Asia, North Africa, Hawaii; Nearctic Region)
.................................................... ...... Haematobia Le Peletier & Serville [page 59] Arista with 3-4 ventral cilia (Figs. 37,39,42,44); notopleuron with covering setulae (Trinidad, Guyana, Surinam, Peru, Brazil) ..... .Neivamyia Pinto & Fonseca [page 59] 4.
Head angular in profile; antenna long; antennal insertion above mid-level of eye (Fig. 1); dorsocentral presutural setae very short and fine, almost indistinct from the covering setulae (Mexico, El Salvador, Panama, Trinidad, Virgin Is., Jamaica, Puerto Rico, Santo Domingo, Barbados, Cuba, St. Lucia, St. Kitts Is., Colombia, Guyana, Ecuador, Galapagos Is., Peru, Venezuela, Brazil, Paraguay, Argentina; pantropical)
.............................................................................. ..Atherigona Rondani [page 38] Head shape not as above; antennal insertion below mid-level of eye (Fig. 96); dorsocentral pre sutural setae developed or not differentiated from the covering setulae .................................................................................................................. 5 5.
Lower calypter enlarged, subtruncate posteriorly and with the anteromedian angle extending below base of scutellum; if calypter glossiform, then arista plumose (Fig. 4); general body coloration usually metallic blue ............................................................ 6 Lower calypter glossiform, at most a little enlarged, not extending below base of
22
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Co uri
scutellum; if so, then the antenna reaches epistoma; arista bare or plumose; general body coloration usually not metallic blue ........ :....................................................... 15 6.
Radial sector of the wing, in dorsal vIew, with a long seta near at the insertion of the humeral vein (Fig. 34) (Peru, Brazil) ....................... Xenomorellia Malloch [page 57] Radial sector of the wing, in dorsal view, without a long seta at the insertion of the humeral vein ........................................................................................................... 7
7.
Lower calypter glossiform ..................................................................................... 8 Lower calypter enlarged ........................................................................................ 9
8.
Dorsocentral setae 2:4; lower calypter without cilia ventrally; hind tibia with a strong posterodorsal seta (calcar) (Mexico, Guatemala, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Argentina) ..................................... .Dasymorellia Malloch [page 41] Dorsocentral setae 2:2; lower calypter with cilia ventrally, on basal half; hind tibia with a small posterodorsal seta (calcar), shorter than the diameter of tibia (Brazil)
........................................................................ ...............Neorypellia Pont [page 49] 9.
Supra-squamal ridge setulose ............................................................................... 10 Supra-squamal ridge bare ...................................................................................... 11
10. Katepistemal setae 1:3; intrapostalar seta strong; apex of abdominal tergite 5 gold yellow (Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Brazil, Argentina)
.......................................................................... Sarcopromusca Townsend [page 55] Katepistemal setae 1:2; intrapostalar seta absent; tergite 5 not differing from other abdominal tergites in colour (Mexico, Cuba, Costa Rica, Brazil; Nearctic and Palaearctic Regions, north-west Oriental) ...................... Neomyia Walker [page 49] 11. Katepisternal setae 0: 1 or 0:2 (the anterior one weak); male: anterointernal ommatidia very enlarged, with about the same diameter as anterior ocellus (Mexico, Costa Rica, Panama, Trinidad, Venezuela, Bolivia, Brazil, Paraguay, Argentina)
..................................................................... ......... Biopyrellia Townsend [page 40] Muscidae (Diptera) of the Neotropical Region: TAXONOMY
23
Part I - Basal Groups
Katepisternal setae 1: 1 or 1:2; male: anterointernal ommatidia not enlarged as described above .................................................................................................................. 12 12. Acrostichal setae 0: 1; metepimeron setulose above hind coxa; humeral callus with 3-4 setae (Mexico, EI Salvador, Costa Rica, Jamaica, Panama, Colombia, Venezuela, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Paraguay, Argentina; Nearctic, Palaearctic, Afrotropical and Australian Regions)............... .. Morellia Robineau-Desvoidy [page 42] Acrostichal setae forming a different pattern; metepimeron bare above hind coxa; humeral callus with 2-3 setae ................................................................................ 13 13. Dorsocentral setae 2:4, presuturals strong; general coloration of the body not metallic (Mexico, Guadeloupe Is., Belize, Guatemala, San Salvador, Nicaragua, Costa Rica, Panama, Nassau, Jamaica, Puerto Rico, Haiti, Dominica, Cuba, Barbados, Mona Is., St. Lucia, St. Vincent, Virgin Is., Grenada, Trinidad, Colombia, Venezuela, Guyana, Surinam, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is., Easter Is.; Nearctic and Palaearctic Regions; north-western Oriental; cosmopolitan) ...................................................Musca Linnaeus [page 48] Dorsocentral setae absent or reduced; when present, only weakly distinguishable from the covering setulae; general coloration of the body blue or green metallic .............. 14 14. Katepisternal setae 1:2; acrostichal setae 0:0 (Mexico, Cuba, Puerto Rico, Jamaica, Dominica, Guadeloupe Is., Trinidad, Colombia, Venezuela, Guyana, Peru, Bolivia, Brazil, Paraguay) ................................................. Parapyrellia Townsend [page 50] Katepisternal setae 1: 1 (Venezuela, Peru, Brazil)..... Cyacyrtoneura Townsend [page 41] 15. Anepimeron with a few setulae ............................................................................ 16 Anepimeron usually bare (except in Thaumasiochaeta Stein) .............................. 33 16. Postalar wall setulose .......................................................................................... 17 Postalar wall bare ................................................................................................. 20 17. Hind coxa with cilia on posterior side .................................................................... 18 24
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Hind coxa bare on posterior side ......................................................................... 19 18. Facial edge with no cilia; lower calypter on dorsal surface with fine cilia on disk; wing with vein CuAl bare on both surfaces (Costa Rica, Nicaragua, Panama, Venezuela, Brazil, Paraguay) ..................................... Pseudoptilolepis Snyder [Part II, page 156] Facial edge with short cilia (Fig. 96); lower calypter on dorsal surface with no cilia; wing with vein CuA 1 setulose on both surfaces (Brazil) ............................ .
........................................................................ ....... Chaetagenia Malloch [page 78] 19. Acrostichal setae 1: 1,2:2 or 2: 1; female: one proclinate fronto-orbital seta; male: cercal plate, on ventral surface with spiny processes (Figs. 26-32) (Mexico, Venezuela, Bolivia, Peru, Brazil, Argentina) ................ .Polietina Schnabl & Dziedzicki [page 51] Acrostichal setae 0: 1; female: proclinate fronto-orbital seta absent; male: cercal plate on ventral surface with no processes (Mexico, Costa Rica, Panama, Cuba, Dominican Republic, Trinidad, Puerto Rico, Venezuela, Ecuador, Peru, Brazil, Uruguay, Argentina, Chile; Unitcd States of America) ....................................Philomis Meinert [page 88] 20. Antenna long, reaching epistoma and placed inside a deep concavity; male dichoptic, surstylus long, about 3 times the length of the cercal plate (Part II, Figs. 7, 8) (Mexico, Panama, Trinidad, Tobago, Ecuador, Brazil) ...... Charadrella Wulp [Part II, page 137] Antenna not reaching epistoma, not placed in a concavity; male holoptic or dichoptic, sursty Ius shorter than described above .................................................................. 21 21. Supra-squamal ridge setulose .............................................................................. 22 Supra-squamal ridge bare .................................................................................... 23 22. Dorsocentral setae 2:3; pro sternum bare; katepisternal setae 1: 1 (Bolivia, Peru)
.................................................................... ...Mulfordia Malloch [Part II, page 149] Dorsocentral setae 2:4; prosternum setulose; katepisternal setae 1:2 (Brazil)
............................................................ SCutellomusca Townsend [Part II, page 172] 23. Parafacial setulose, with one or more series of setulae; palpus enlarged apically (Part Muscidae (Diptera) of the Neotropicai Region: TAXONOMY
25
Part I - Basal Groups
n, Figs. 83-96); male: dichoptic, width of frons about one third of head width (Mexico, Bahamas Is, Virgin Is., Cuba, Puerto Rico, Guadeloupe Is., Guatemala, Guyana, Peru, Bolivia, Brazil, Argentina, Paraguay, Chile; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions) ........................... .Lispe Latreille [Part n, page 182] Parafacial bare; palpus not enlarged or not enlarged as described above; male: holoptic or dichoptic ......................................................................................................... 24 24. Katepisternal setae 2:2 ........................................................................................ 25 Katepisternal setae forming a different pattern ...................................................... 26 25. Meron setulose; wing with vein R j with setae on basal third of dorsal surface; pro sternum with lateral setulae (Chile )..... Arthurella Albuquerque [Part n, page 135] Meron bare; wing with vein R j with cilia over its whole dorsal surface, except on the basal fifth; pro sternum bare (Brazil, Uruguay, Argentina) ............................ ..
................................................ .Neurotrixa Shannon & Del Ponte [Part II, page 155] 26. Arista with short cilia usually only on dorsal surface, some specimens with 1-2 ventral cilia (Mexico, Guatemala, Costa Rica, Brazil) ....Hemichlora Wulp [Part II, page 167] Cilia on arista forming a different pattern .............................................................. 27 27. Arista pubescent, plumes at most as long as basal width of arista; katepimeron and meron setulose .................................................................................................... 28 Arista ciliated, cilia longer than basal width of arista; katepimeron and meron usually bare .................................................................................................................... 29 28. General body coloration blue metallic; hind tibia with posterodorsal seta (calcar) developed; female: interfrontal seta present (Chile ).......Dalcyella Carvalho [page 79] General body coloration not metallic; hind tibia with a short posterodorsal seta (calcar), only slightly longer than the covering setulae; female: interfrontal seta absent (Guatemala, Costa Rica, Panama) ....................................... Chortinus Aldrich [Part II, page 139] 29. Dorsocentral pre sutural setae 1; veins R4+5 andM divergent apically; vein R4+5 with a 26
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Co uri
slight undulation subapically (Colombia, Venezuela, Peru, Bolivia, Brazil, Paraguay, Argentina) ............................... ......... .xenothoracochaeta Malloch [Part II, page 158] Dorsocentral presutural setae 2; vein R4+5 usually without a subapical undulation; vein M straight or curved apically................................................................................ 30 30. Antennal axis much longer than oral axis; male: dichoptic; female: ovipositor long (Figs. 97-98) (Brazil) .................................... .Itatingamyia Albuquerque [page 80] Antennal axis similar to oral axis in length; male: hoI optic; female: ovipositor long or short. ................................................................................................................... 31
31. Vein M strongly curved apically; vein R4+5 ending before wing apex (Part II, Fig. 15); apical portion of Rs with setulae on ventral surface; vein R] bare on both surfaces (Mexico, Guatemala, Costa Rica, Panama, Trinidad, Cuba, Venezuela, Colombia, Guyana, Peru, Brazil, Paraguay, Argentina; United States of America)
.......................................................... .... .Neomuscina Townsend [Part II, page 149] Apical section of vein M not so curved as in the Neomuscina species, only slightly bent forward [except in Cyrtoneuropsis brunnea (Hough, 1900)]; apical portion of Rs bare on ventral surface; vein Rl usually setulose .............................................. 32 32. Wing with vein R] setulose on ventral surface; female: ovipositor without microtrichia on the membrane between segments 6-7 and 7-8; ovipositor medium or short, and not delicate; cerci short and flat; tergites 6 and 7 scletotized (Mexico, Belize, Guatemala, Nicaragua, Costa Rica, Panama, Trinidad, EI Salvador, Cuba, Puerto Rico, Colombia, Venezuela, Guyana, Suriname, Ecuador, Bolivia, Brazil, Argentina)
.............................................................. Cyrtoneuropsis Malloch [Part II, page 141] Wing with vein R] bare on ventral surface; female: ovipositor with microtrichia on the membrane between segments 6-7 and 7-8; ovipositor long; cerci long and not flat; tergites 6 and 7 not sclerotized (Mexico, Belize, Guatemala, EI Salvador, Nicaragua, Costa Rica, Panama, Grenada, Trinidad, Venezuela, Colombia, Guyana, Peru, Brazil)
............................................................. Cyrtoneurina Giglio-Tos [Part II, page 139] Muscidae (Diptera) of the Neotropical Region: TAXONOMY
27
Part I - Basal Groups
33. Hind coxa, on posterior surface, with a row of weak setae ................................... 34 Hind coxa bare on posterior surface .................................................................... 35 34. Wing with vein Al ending abruptly, at most extending to half the distance between the apex of the cubital cell and the wing margin; male: enlarged ommatidia (Mexico, Argentina; Nearctic, Palaearctic, Afrotropical and Oriental Regions) ............................................................. ..........Azelia Robineau-Desvoidy [page 62]
Wing with vein Al extending more than half the distance between the apex of the cubital cell and the wing margin; male: ommatidia normal (Mexico; Nearctic, Palaearctic and Oriental Regions) ................................................... Thricops Rondani [page 74] 35. Gena, at least, usually with a distinct, upward curved seta (except in Ophyra); wing with subcostal vein meeting the costal vein along its basal third; female: proclinate orbital seta present. ............................................................................................. 36 Gena without the seta described above; wing with the subcostal vein meeting costal vein along its distal third or beyond; female: proclinate orbital seta absent... .......... .40 36. Gena with no differentiated setae; general coloration of the body black metallic; female: fronto-orbital seta weak, shorter than frontal setae; ocellar triangle long, almost reaching lunule (Mexico, Nicaragua, St. Vincent, Puerto Rico, Jamaica, Cuba, Trinidad, Tobago, Guyana, Venezuela, Ecuador, Peru, Bolivia, Brazil, Uruguay, Argentina, Chile, Galapagos Is.; Nearctic and Western Palaearctic Regions; Eastern Pacific) .................................................................... Ophyra Robineau-Desvoidy [page 70]
Gena with strong and upward curved seta; general body coloration not as above; female: fronto-orbital seta strong, longer than frontal setae; ocellar triangle short..... 37 37. Male: fore femur with a distinct concavity on its ventral surface, and usually with 1-2 processes near the concavity; female: interfrontal setae present (Mexico, Costa Rica, Ecuador, Peru, Colombia, Bolivia, Brazil, Argentina, Chile, Juan Fernandez Is.; Nearctic and Palaearctic Regions; North India; cosmopolitan) .................................................................. Hydrotaea Robineau-Desvoidy [page 63] 28
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Male: fore femur without a distinct concavity or processes on its ventral surface; female: interfrontal setae absent. ................................. '..................................................... 38 38. Arista plumose; sternite 1 setulose (Mexico, Peru; United States of America)
.............................................................. ..... Potamia Robineau-Desvoidy [page 74] Arista pubescent with hairs as long as or shorter than basal width of arista; sternite 1 bare .................................................................................................................... 39 39. Prealar weak, slightly longer than adjacent ground-setulae; hind tibia with a strong posterodorsal seta (calcar) at apical third; female: fronto-orbital plate with many setae (Mexico, Brazil, Argentina) .................................Micropotamia Carvalho [page 68] Pre alar strong; hind tibia usually with 3 or more strong posterodorsal setae more or less uniformly distributed along tibial length and as developed as anterodorsal setae; female: fronto-orbital plate different from above (Mexico; Nearctic, Palaearctic and Oriental Regions) ........................................................ .. .Drymeia Meigen [page 63] 40. Wing with Rs node or base of vein R4+5 usually with one cilia on dorsal surface (except some species of Phaonia) ................................................................................... .41 Wing with Rs node or base of vein R4+5 with no cilia on dorsal surface ................... 51 41. Pro sternum setulose laterally; prealar seta absent.. .............................................. .42 Pro sternum bare; prealar seta absent or present.. ................................................ 46 42. Vein R J with cilia on apical half of dorsal surface .................................................. 43 Vein R J bare on dorsal surface ............................................................................ 45 43. Gena strongly projected; fore tibia with no median setae (Mexico, Colombia, Ecuador, Peru, Bolivia, Brazil, Argentina, Chile, Juan Fernandez Is.)
................ ................................................ Syllimnophora Speiser [part II, page 192] Gena not projected; fore tibia with or without median setae ................................ ..44 44. Dorsocentral setae 0: 1 (Brazil) ..... Albertinella Couri & Carvalho [Part II, page 176] Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Dorsocentral setae forming a different pattern, never 0: 1 (Mexico, Bahamas, Puerto Rico, Cuba, Belize, Guatemala, Costa Rica, Panama, St. Vrncent, Jamaica, Venezuela, Colombia, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is.; Nearctic, Palaearctic, Afrotropical, Oriental and Australian
Regions).......................... .Limnophora Robineau-Desvoidy [in part] [Partil, page 178] 45. Dorsocentral postsutural setae 3; gena strongly projected; male: palpus long, enlarged and flat (Peru, Bolivia) ............................. Thaumasiochaeta Stein [Part il, page 201] Dorsocentral postsutural setae 3-4; gena not or only slightly projected; palpus different from described above (Mexico, Bahamas, Puerto Rico, Cuba, Belize, Guatemala, Costa Rica, Panama, St. Vincent Is., Jamaica, Venezuela, Colombia, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is.; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions)
.................................... Limnophora Robineau-Desvoidy [in part] [Part il, page 178] 46. Apical section of vein M not curved forward ...................................................... ..47 Apical section of vein M slightly to strongly curved forward .................................. .48 47. Head in profile almost twice as high as its greatest length, distinctly narrower at vibrissae than at antennal base; facial ridges covered up to slightly above middle of flagellomere with dense, short setulae (Cuba, Puerto Rico) .... .. Scenetes Malloch, 1936, statrev. [part il, page 171] Head in profile not as above; facial ridges bare (Mexico, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions ) ........Mydaea Robineau-Desvoidy [Part il, page 167] 48. Katepistemal posterior setae 2, the anterior seta absent; katepimeron and meron with black setulae (Mexico, Cuba, Puerto Rico, Peru, Brazil, Chile, Venezuela, Uruguay, Argentina; Nearctic, Palaearctic, Afrotropical and Oriental Regions. Essentially cosmopolitan in temperate and tropical areas, absent in New Zealand and Oceania, except Micronesia, more diverse in Afrotropical and Oriental Regions)
................................................. Graphomya Robineau-Desvoidy [Part II, page 163] 30
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Katepisternal setae 3, the anterior seta present; katepimeron and meron bare ...... .49 49. Parafacial with cilia on upper part; katepisternal setae 1: 1: 1 (Galapagos Is.; Nearctic Region) ............................................................. Pentacricia Stein [Part II, page 239] Parafacial with no cilia on upper part; katepistemal setae not 1: 1: 1.. ...................... 50 50. Hind tibia with a strong posterodorsal seta (calcar) along its apical three-quarters (Mexico, Guatemala, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, Uruguay, Argentina; cosmopolitan) ............. .Phaonia Robineau-Desvoidy [page 119] Hind tibia with no strong posterodorsal seta on apical three-quarters (Mexico, Costa Rica, Panama, Puerto Rico, Haiti, Cuba, S., Domingo, Jamaica, Guatemala, Venezuela, Guyana, Ecuador, Peru, Bolivia, Brazil, Argentina, Chile, Falkland Is.; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions)
....................................................................... .Myospila Rondani [Part II, page 170] 51. Labella not reduced (Figs. 109-118); prestomal teeth not developed .................... 52 Labella reduced (Part II, Fig. 146); prestomal teeth developed ............................ 64 52. Apical section of vein M strongly curved forward ................................................ 53 Apical section of vein M straight or only slightly curved forward ............... ,.......... 56 53. General body coloration metallic blue; eye setulose; sternite 1 bare (Venezuela,
Ecuador) ......................................... Reinwardtia Brauer & Bergenstamm [page 97] General body coloration not as above; eye bare; sternite 1 setulose ....................... 54 54. Arista pubescent, with the cilia shorter than the basal diameter of arista; facial border with no setulae; female: ovipositor with 5 spiracles (Mexico, Nicaragua, Costa Rica, Panama, Virgin Is., Jamaica, Cuba, St. Domingo, Puerto Rico, Trinidad, Guyana, Colombia, Venezuela, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is.; pantropical)
................................................... Synthesiomyia Brauer & Bergenstamm [page 98]
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Arista plumose, with length of plumae similar to the width of the flagellomere; facial border with setulae or bare; female: ovipositor with 5 or 6 spiracles .. :.................... 55 55. Facial border with developed setulae that continue above the vibrissae up to the basal quarter of the antenna; lower calypter glossiform; wing with cilia on both surfaces, from the node to vein R 2+3 ; female: ovipositor with the 6th spiracle present (Colombia, Brazil) ............................................... ........... Cariocamyia Snyder [Part II, page 136] Facial border bare, with very short setulae, not as described above; lower calypter enlarged, developed; wing with no cilia on vein R 2+3; female: ovipositor with 5 spiracles
(Brazil) .......................... .Neomusciniopsis Albuquerque & Lopes [Part II, page 155] 56. Hind tibia with one strong posterodorsal seta (calcar) on its apical third ................. 57 Hind tibia with no calcar...................................................................................... 62 57. Anterior intra-alar postsutural seta placed posteriorly to level of supra-alar seta; wing with apical section of vein M slightly curved forward (Mexico, Guatemala, Costa Rica, Venezuela, Peru, Brazil, Uruguay, Argentina, Chile; Nearctic and Palaearctic Regions) .................................................... Muscina Robineau-Desvoidy [page 80] Anterior intra-alar postsutural seta usually placed anteriorly to supra-alar seta; wing with apical section of vein M straight or slightly curved forward ............................. 58 58. Flagellomere not dilated; female: ovipositor with sternite 8 reduced to 2 posterior sclerites; clypeus, anteriorly, with a strong end hook-shaped in lateral view and with a prominent sclerotized area posteriorly (Figs. 109-118) (Mexico, Jamaica, Panama, Nicaragua, St. Vincent, Ecuador, Peru, Brazil, Argentina, Chile; southern of United States of America) ................ .Dolichophaonia Carvalho, 1993, stat. rev. [page 99] Flagellomere slightly to strongly dilated; female: ovipositor with sternite 8 developed along its whole extension; proboscis not as above ................................................. .59 59. Flagellomere strongly dilated (Figs. 94, 95); general body coloration metallic blue (Chile, Argentina) ........................................... .B rachygasterina Macquart [page 76]
32
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Flagellomere only slightly dilated; general body coloration blue, yellow or metallic green .................................................................................................................. 60 60. Eyes bare; male: dichoptic with interfrontal setae present (Argentina) .............................................................................. Correntosia Malloch [page 79]
Eyes ciliated; male: holoptic ................................................................................. 61 61. Posterior notopleural seta long, similar in length to anterior seta; notopleura sparsely setulose (Colombia, Ecuador, Peru, Bolivia, Brazil, Uruguay, Argentina, Chile, Juan Fernandez Is.) .................................................. ............ Psilochaeta Stein [page 96] Posterior notopleural seta short, shorter than anterior seta; notopleuron with no covering setulae (Chile, Argentina) .......................................... PaZpibracus Rondani [page 82] 62. Head quadrate, antenna long, reaching the epistoma; ocellar setae absent; general body coloration yellow (Peru, Bolivia, Brazil) .......... .. SouzaZopesmyia Albuquerque [page 126] Head with another shape; length of antennae variable, but not reaching epistoma; ocellar setae present; general body coloration yellow............................................ 63 63. Arista plumose, with the longest cilia longer than the width of the flagellomere; sternite 1 setulose or bare (Mexico, Jamaica, Guatemala, Costa Rica, Panama, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Argentina, Chile, Falkland Is.; Nearctic, Palaearctic, northern parts of Oriental and Afrotropical Regions) ...................................................... ................ HelinaRobineau-Desvoidy [page 105]
Arista bare; sternite 1 with strong setulae near the posterior margin (Part II, Fig. 20) (Mexico, Jamaica, Cuba, Puerto Rico, Haiti, St. Vincent Is., Peru, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions) ........................ .Brontaea Kowarz [Part II, page 160] 64. Wing with cilia in other veins besides vein c. ....................................................... 65 Wing veins with no cilia, except vein C ................................................................. 66 65. Sternite 1 setulose; katepisternal setae 2:2 (Guatemala, Venezuela, Colombia, Peru, Muscidae (Diptera) of the Neotropical Region: TAXONOMY
33
Part I - Basal Groups
Bolivia, Argentina, Chile, Juan Fernandez Is.; Nearctic Region) ................ .
.................. ................................. ................... Lispoides Malloch [Part II, page 184] Sternite 1 bare; katepisternal setae not 2:2 (Mexico, Colombia, Peru, Bolivia, Argentina; cosmopolitan with many species occurring in high altitudes) .................................. .
..... ............................ ....... ........... ....... ....... ...... .Spilogona Schnabl [Part II, page 190] 66. Dorsocentral setae 2:4; female: ovipositor with the hypoproct reduced and leaf-shaped, in lateral view, very setulose (Bahamas Is., Peru, Chile; southeastern United States of America) ............................................... ........ Tetramerinx Berg [Part II, page 200] Dorsocentral setae 1-2:2-3; female: ovipositor not as above ................................. 67 67. One strong dorsocentral presutural seta, sometimes preceded by a second short seta .................................................................................................................... 68 Two strong dorsocentral pre sutural setae, more or less similar in length ................. 80 68. Postsutural dorsocentral setae 2 ............................................................................ 69 Postsutural dorsocentral setae 3 .......................................................................... 70 69. Basal scutellar seta shorter the apical pair; frons longer than wide; hind tibia with long median setae on anterodorsal, anteroventral, posterodorsal and sometimes posteroventral surfaces (Mexico, Jamaica, Cuba, Puerto Rico, Costa Rica, Venezuela, Guyana, Surinam, Peru, Brazil, Paraguay, Uruguay, Argentina; United States of America) ................................................ .Bithoracochaeta Stein [Part II, page 211] Basal scutellar seta as long as or longer than the apical pair; frons wider than long; distribution of setae on hind tibia not as above (Peru, Argentina, Chile, Juan Fernandez Is.) .................................................. ...... Schoenomyzina Malloch [Part II, page 251] 70. Hind tibia with at least one anterodorsal supramedian seta in addition to the median setae .................................................................................................................... 71 Hind tibia without any anterodorsal supramedian seta in addition to the median setae .................................................................................................................. 74 34
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _'"F
Claudio Jose Barros de Carvalho & Marcia Souto Couri
71. Hind tibia without any posterodorsal supramedian seta; frons wider than longer (part II, Fig. 145) (Mexico, Ecuador, Peru, Argentina, Uruguay, Chile; Nearctic, Palaearctic, Afrotropical and Oriental Regions) ........................... Schoenomyza Halliday [in part] [part II, page 247] Hind tibia with at least one posterodorsal supramedian seta; frons variable ............ 72 72. Arista bare; lower calypter about twice as long as upper calypter (Peru, Bolivia) ........... '" ................................................. .. Stomopogon Malloch [Part II, page 257] Arista not as above; calypteres similar in size ....................................................... 73 73. Hind tibia with preapical setae on anterodorsal and dorsal surfaces (peru, Bolivia, Argentina, Uruguay, Chile, Juan Fernandez Is.)....... .. Notoschoenomyza Malloch [part II, page 237] Hind tibia with preapical setae on anterodorsal, dorsal and posterodorsal surfaces (Peru, Bolivia, Chile, Argentina) .......... ......Spathipheromyia Bigot [Part II, page 254] 74. Katepisternal seta 0: 1; disc of katepisternum with numerous setulae and hairs; lower proepimeral seta directed upwards, male: prealar seta present (Peru, Bolivia) ................................................................. Drepanocnemis Stein [Part II, page 177] Katepisternal setae 1: 1: 1; disc of katepisternum with few setulae or hairs; lower proepimeral seta directed downwards, male: prealar seta absent... ........................ 75 75. Calypteres similar in length (Mexico, Ecuador, Peru, Argentina, Uruguay, Chile; Nearctic, Palaearctic, Afrotropical and Oriental Regions) ................ . ...................................................... Schoenomyza Haliday [in part] [Part II, page 247] Lower calypter about twice as long as upper calypter.......................................... 76 76. Arista with plumes only on basal two-thirds; hind tibia with one supramedian posterodorsal seta (Brazil) ......................P lumispina Albuquerque [Part II, page 241] Arista with plumes along its entire length or only on basal two-thirds but usually with only very short hairs; hind tibia only rarely with a supramedian posterodorsal setae ................................................................................................................... 77 77. Hind femur with 3 pre apical setae, on anterodorsal, dorsal and posterodorsal to Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
posterior surfaces ................................................................................................ 78 Hind femur with 2 preapical setae, on anterodorsal and posterodorsal surfaces ..... 79 78. Antennal insertion well above mid-level of eye (Part IT, Fig. 118); one pair of postsutural intra-alar setae; lower calypter transverse (Mexico, St. Vincent Is., Colombia, Peru, Bolivia, Brazil).................................... Cordiluroides Albuquerque [Part II, page 218] Antennal insertion at mid-level of eye (Part II, Fig. 121); two pairs of postsutural intra-alar setae; lower calypter glossiform (Mexico, Guatemala, Costa Rica, Panama, Jamaica, Puerto Rico, St. Vincent Is., Venezuela, Colombia, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Paraguay, Uruguay, Argentina, Chile, Juan Fernandez Is.; Nearctic
Region) .................................................... .Neodexiopsis Malloch [Part IT, page 220] 79. Gena greatly reduced, almost linear (Part II, Figs. 4 3a, b); female: ovipositor of medium length; tergites and sternites broad (Part II, Figs. 47-48) (Panama, Brazil)
.............................................................Agenamyia Albuquerque [Part II, page 174] Gena of moderate width; female: ovipositor long; tergites and sternites long and narrow (Mexico, Venezuela, Guyana, Ecuador, Bolivia, Brazil, Uruguay, Argentina, Chile; cosmopolitan) ................................................... Coenosia Meigen [Part II, page 214] 80. Apical scutellar setae shorter than the basal setae (Puerto Rico, Brazil)
................................................................. .Pilispina Albuquerque [Part II, page 240] Apical scutellar setae as long as the basal setae ...................................................... 81 81. Fore tibia on posterior surface without setae ........................................................ 82 Fore tibia on middle of posterior surface with, at least, one seta ............................ 83 82. Arista with hairs restricted to basal half; lower calypter about 1.5 times as long as upper calypter (Brazil) ...................................... Insulamyia Couri [Part II, page 219] Arista with hairs along its entire length; both calypteres small, similar in size
(Peru) .......................................................... Rhabdotoptera Stein [Part II, page 189]
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Co uri
83. Palpus rather enlarged apically (Part II, Figs. 83-88); gena very deep; eyes reduced in size (Part II, Figs. 89-95), with few setae around vibrissae (Argentina, Chile)
............................................................................. Apsil Malloch [Part II, page 205] Palpus slender; gena variable; eyes not or hardly reduced, with few or many setae around vibrissae ................................................................................................... 84 84. Hind tibia without setae on posterodorsal surface; antennal flagellomere modified apically (part II, Fig. 135) (Peru, Bolivia) ........ Oxytonocera Stein [part II, page 238] Hind tibia with at least a median and sometimes also a supramedian seta on posterodorsal surface; antennal flagellomere not modified apically.......................... 85 85. Katepisternal setae 2:2; arista short and enlarged basally (Part II, Fig. 56); claws and pulvilli small; male: hypandrium not tubular; female: ovipositor of moderate length, with large tergites (Part II, Figs. 60-61) (Bahamas Is., Brazil; South America; North
America) ......................................... Pachyceramyia Albuquerque [Part II, page 187] Katepisternal setae 1: 1: 1; arista long and slender; claws and pulvilli long; male: hypandrium tubular, very elongated; female: ovipositor long, with slender tergites (Argentina, Chile )......................................... .Reynoldsia Malloch [Part II, page 242]
DIAGNOSIS AND KEY TO SPECIES Muscidae has eight subfamilies (HENNIG, 1965; PONT, 1977, 1980, 1986a, 1989; SKIDMORE, 1985; CARVALHO, 1989d), including Cyrtoneurininae (see Part II), 173 genera and more than 4000 species described to the world (CARVALHO et ai., 1993). The phylogeny of the family has been studied (HENNIG, 1965; SKIDMORE, 1985; CARVALHO, 1989d) but the phylogenetic relationships among the subgroups are still in debate (PONT & CARVALHO, in preparation). The basal groups considered are Atherigoninae, Muscinae, Azeliinae and Phaoniinae (CARVALHO, 1989d) and the classification adopted here is at in CARVALHO et al. (1993) catalogue, including the Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
taxonomic changes published afterwards to it. Table 1 (page 130) presents a summary genera occurring in Brazil, South America and the Neotropical Region.
Subfamily Atherigoninae Fan, 1965 Diagnosis. Eyes dichoptic in both sexes, with very weak sexual dimorphism; ovipositor with tergites 6 and 7 largely developed, tergite 8 complete. Monophyly. Subfamily with only one genus. See monophyly under the genus Atherigona. Comments. The only genus, Atherigona Rondani, has 195 described species (PONT, 1986b), only one, Atherigona orientalis Schiner, occurring in the neotropics.
Atherigona Rondani, 1856 (Figs. 1-3) Type-species. Anthomyia varia Meigen (orig. des.). Diagnosis. Head angular (Fig. 1), with a long and very sunken face; frontoorbital plate with one pair of reclinate orbital setae; antennal flagellomere very long; antenna long, almost reaching lower facial margin in frontal view; arista bare; palpus strongly differentiated between the two sexes; prementum of proboscis glossy, or thinly or partly dusted; thoracic setae very reduced in size; acrostichal setae 0: 1; prescutellar pair weak; dorsocentrals reduced, setulose and hardly distinct from ground setulae; one to two short postsutural intra-alars; prealar absent; subalar sclerite, anepimeron, meron and anatergite bare; katepisternals 1:2, lower one weaker, equidistant from the upper two; posterior spiracle small, bare on margins; katatergite pilose; hind tibia without calcar; wing veins, except costal, bare; veins R4+5 and M subparallel at wing margin (Fig. 2); Al +Cu~ reaching over halfway from crossvein Cu~ to wing margin; lower calypter of Phaonia-type; stemite 1 bare; ovipositor long (Fig. 3). Reference. PONT & MAGPAYO, 1995. Monophyly. According to PONT (1986b), there is doubt about the monophyly of the genus, with a great number of autapomorphies in relation to the ground plan of Muscidae. The genus is one of the most basal in the family (CARVALHO, 1989d).
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Geographical record. Mexico, EI Salvador, Jamaica, Cuba, Puerto Rico, Trinidad, Virgin Is., Santo Domingo, Barbados, St. Lucia, St. Kitts Is., Panama, Colombia, Venezuela, Guyana, Ecuador, Galapagos Is., Peru, Brazil, Paraguay, Argentina; cosmotropical. Comments. Only Atherigona orientalis Schiner, 1868 is known from the Neotropical Region.
__".....,....,......_-:::>
R4 +5
3 Figs. 1-3, characters of Atherigona (modified from PONT & MAGPAYO, 1995): 1. head, dorsolateral view; 2, wing, dorsal view; 3, ovipositor, dorsal view. Abbreviations: ar, arista; M, vein; R4+5 , vein; tg, tergite.
Subfamily Muscinae Latreille, 1802 Diagnosis. Arista plumose; anepimeron setulose; most of the genera with vein M strongly curved forward apically; Musca-type egg (according to HERTING, 1957); larvae coprophagous. Monophyly. HENNIG (1965) justified the monophy ly of the subfamily based mainly on characters of the ovipositor -long and tubular and sternite 8 well developed along its whole length. CARVALHO (1989d) justified the monophy ly of the subfamily based on 7 characters of the ovipositor, associated with the shape of the egg (Muscatype) and the presence of setulae in the anepimeron. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Comments. Traditionally, the Muscinae has been divided into two tribesMuscini and Stomoxyini, as presented by CARVALHO (1989d) and CARVALHO et al. (1993).
Tribe Muscini Latreille, 1802 Diagnosis. Most of the genera with lower calypter strongly enlarged, Muscatype, sub-truncate posteriorly, with its anteromedian angle extending at least slightly under base of scutellum; cereal plate, on its ventral surface, with spiny processes. Monophyly. HENNIG (1965) considered the tribe monophyletic, based on the setulose anepimeron, plumose arista and tortuous slits of posterior larval spiracles (see SKIDMORE, 1985). CARVALHO (1989d) indicated the monophyly ofthe tribe based on the enlarged Musca-type lowercalypter (see HENNIG, 1965). Comments. The tribe has 23 genera worldwide, 11 of them occurring in the neotropics, with about 58 species ascribed to this region.
Biopyrellia Townsend, 1932 Type-species. Musca bipuncta Wiedemann (orig. des.). Diagnosis. General body coloration dark metallic blue; male with anterointemal ommatidia very enlarged, with about the same diameter as anterior ocellus; vibrissa weak, inserted above oral margin; dorsocentral setae 0: 1 or 0:2 (the anterior one weak); acrostichal 0:0 or 0: 1 (weak); humerall; intra-alars 0: 1; supra-alars 0:2; intrapostalar seta absent; katepistemals 0: 1 or 0:2 (the postero-anterior one weak); lower calypter enlarged, about twice as long as upper calypter; apical portion of vein M strongly bent forward. Reference. PAMPLONA, 1986a. Monophyly. Not yet established. Geographical record. Mexico, Costa Rica, Panama, Venezuela, Trinidad, Bolivia, Brazil, Paraguay, Argentina. Comments. Monobasic genus: Biopyrellia bipuncta (Wiedemann, 1830), validated by PAMPLONA (1986a).
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Cyacyrtoneura Townsend, 1931 Type-species. Cyacyrtoneura cyanea Townsend (orig. des.). Diagnosis. General body coloration metallic blue with abdomen metallic green; katepistemal setae 1: 1. Reference. TOWNSEND, 1931. Geographical record. Venezuela, Peru, Brazil. Monophyly. Not yet established. Comments. Monobasic genus: Cyacyrtoneura cyanea Townsend, 1931. This species is known only from the type-material (ZMHU), which has not been examined after of the original description.
Dasymorellia Malloch, 1923 Type-species. Dasymorellia trichops Malloch (orig. des.). Diagnosis. Male holoptic; female with a pair of fronto-orbital proclinate setae; arista plumose; humeral 4; prealar strong; dorsocentrals 2:4; acrostichals 2:2 or 2:3; katepistemals 1:2; wings with cilia at node on both surfaces (except D. benoisti); apical portion of vein M bent forward; lower calypter glossiform, about twice as long as upper calypter; fore tibia on dorsal surface with 2 preapical setae; hind tibia with a strong posterodorsal seta (calcar). Reference. PAMPLONA, 1983. Monophyly. PAMPLONA (1983) did not present unique characters to support the monophyly of the genus. Geographical record. Mexico, Guatemala, Ecuador, Brazil, Colombia, Venezuela, Bolivia, Argentina, Peru. Comments. D.fulvipes (Bigot, 1887) (Mexico), D. saphirina (Seguy, 1935) (Ecuador) and D. spinifera (Wulp, 1883) (Argentina) were not included in the key due to lack of decisive information in the available descriptions (types not seen). PAMPLONA (1983) did not include D. saphirina in this genus, a lapsus corrected by CARVALHO et al. (1993). Nine species are known in the Neotropical Region.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Key to species (modified from PAMPLONA, 1983) 1.
Antenna and palpus yellow (Brazil) .......................... D.j1avipalpis Pamplona, 1983 Antenna and palpus brown ..................................................................................... 2
2.
Eye strongly haired in both sexes; parafacial, facial and gena shiny; gena with many hairs; acrostichals 2:3 .............................................................................................. 3 Eye haired in both sexes; hairs short and sparse in female (almost bare in D. callidimera); parafacial, facial and gena with grey pruinescence; gena with weak and
sparse hairs; acrostichals 2:2 .................................................................................. 4 3.
Wing bare on dorsal surface, mid femur with no anterodorsal setae (Ecuador) .................................................................................. .D. benoisti Pamplona, 1983
Wmg with about 5 cilia on dorsal surface, beginning at node, mid femur with 4 anterodorsal setae on middle third (Venezuela, Colombia) .................... .D. seguyi Pamplona, 1983 4.
Hind femur with 2 preapical anterodorsal setae, hind tibia with 5 anteroventral setae on apical half (Colombia, Ecuador, Peru, Bolivia, Brazil, Argentina) .........................................................................................D. trichops Malloch, 1923
Hind femur with a row of anterodorsal setae, hind tibia with less than 5 anteroventral setae on apical half. ................................................................................................. 5 5.
Legs yellow, hind femur with one median posteroventral seta (Mexico, Guatemala) ............................................................................... .... D. callidimera (Bigot, 1887)
Legs brown, hind femur with 2 median posteroventral setae (Colombia) .................................................................................. ... D. nigritibia Snyder, 1949
Morellia Robineau-Desvoidy, 1830 (Figs. 4-22, Plate 1) Type-species. Morellia agilis Robineau-Desvoidy (TOWNSEND, 1916) = hortorum (Fallen). 42
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Diagnosis. Male holoptic; anterointemal ommatidia enlarged; eyes bare or with very few cilia; palpus a little spatulate apically; arista plumose on basal two-thirds or threequarters and bare on apically (Fig. 4); acrostichals 0: 1; dorsocentrals 0:2, 0:3 or 1:2; intraalars 1: 1 or 1:2; supra-alars 0:3 or 1:3; katepisternals 1:2; anepimeron setulose; meron bare; metepimeron haired above hind coxa; M strongly curved forward apically (Fig. 5); stemite 1 setulose; ovipositor long with tergites very thin and possessing strong spines (Figs. 6a, 6b, 7a, 7b). Reference. PAMPLONA, 1986b. Monophyly. No revision has been publishing for this genus. Its species are found in all biogeographic regions of the world. Denise Pamplona (personal communication) is finishing a worldwide revision and a cladistic analysis for the species of Morellia. Geographical record. Mexico, Jamaica, El Salvador, Costa Rica, Panama, Venezuela, Colombia, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Paraguay, Argentina; Nearctic, Palaearctic, Afrotropical and Australian Regions. Comments. There are 19 species recorded to the Neotropical Region, 14 of them occurring in South America.
Key to species (modified from PAMPLONA, 1986b) 1.
Humeral callus yellow............................................................................................. 2 Humeral callus blue or dark metallic green ............................................................. 3
2.
Humeral callus with 4 setae; proepimeron setulose; wings without clouds (Mexico, EI Salvador, Costa Rica, Jamaica, Cuba, Haiti, Puerto Rico, Virgin Is., Venezuela; Nearctic Region) ..................................................... ......................M. basalis (Walker, 1853) Humeral callus with 3 setae; proepimeron bare; wings with clouds (Peru, Bolivia, Brazil) ...........................................................................M. humeralis (Stein, 1918)
3.
Wing without clouds .............................................................................................. 4 Wing with clouds (Figs. 12-22) ............................................................................. 7
4.
Antenna brown ..................................................................................................... 5 Antenna yellow...................................................................................................... 6
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
5.
Palpus yellow; lower half of fronto-orbital plate and upper half of gena grey pruinose; lower calypter brown (Brazil) ..................................... ..M. hirtitibia Pamplona, 1986 Palpus brown; fronto-orbital plate and gena entirely shiny; lowercalypteryellow (South America) ............................................... ................M. diversipalpis (Rondani, 1863)
6.
Intra-alar setae frequently 1:2; supra-alar setae frequently 0:3; mid femur on anteroventral surface without setae; hind femur with a row of dorsal setae; sursty Ius with a median internal seta (Fig. 8); paramere with setulae apically (Figs. 9,10); epiproct with spines on base, closely placed to each other (Fig. 11) (Brazil)
.................................................................................. ....M. dalcyi Pamplona, 1986 Intra-alar setae frequently 1: 1, supra-alar setae frequently 1:3; mid femur with a row of anteroventral setae; hind femur without dorsal setae; surstylus without a median internal seta; paramere with setulae apically; epiproct with spines frequently shorter than in M. dalcyi (Mexico, Costa Rica, Jamaica, Colombia, Ecuador, Venezuela, Peru, Brazil, Paraguay, Argentina) ............. M. violacea (Robineau-Desvoidy, 1830) 7.
Wing with no more than 2 faint, small spots (Figs. 12-16) ...................................... 8 Wing with at least 2 conspicuous spots (Figs. 17-22) .......................................... 13
8.
Flagellomere yellow.............................................................................................. 9 Flagellomere brown ............................................................................................ 10
9.
Wing with a discrete cloud on humeral vein; mid femur on posterodorsal surface with a group of strong and short apical setae, forming a rigid tuft; hind tibia on anteroventral surface with 5-6 setae on apical half (Brazil) .................. .
............................ .............................. M. paulistensis Pamplona & Mendes, 1995 Wing with a discrete brown cloud on humeral vein and crossvein r-m (Fig. 14); no tuft on mid femur; quetotaxy of hind tibia not as above (Mexico, Cuba, Puerto Rico, Galapagos Is., Brazil) ........................................ M. ochricornis (Wiedemann, 1830) 10. Pro sternum setulose ............................................................................................. 11 44
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Pro sternum bare ................................................................................................ 12 11. Hind tibia, on anterodorsal surface, with 4 setae evenly distributed from base to apex and with one seta on basal third of posterodorsal surface (Panama) ....................................................................................... .M. affinis Malloch, 1925
Hind tibia, on anterodorsal surface, with 2 setae on apical half and with one seta on apical third of posterodorsal surface (Brazil) ..............M. concacata Pamplona, 1986 12. Dorsocentral setae 1:2, hind tibia with 6 posteroventral setae (Brazil) .................................................................................. .... M. roppai Pamplona, 1986
Dorsocentral setae 0:2, hind tibia without posteroventral setae (Brazil) .............................................................................. .M. sinopensis Pamplona, 1986
13. Wing brown from base to apex along costa, this stripe connected to a dark cloud over crossvein r-m (Cuba, Peru, Bolivia) ......................M. semimarginata (Stein, 1918) Wing not as above ............................................................................................... 14 14. Crossvein dm-cu without spots (Fig. 19) ............................................................. 15 Crossvein dm-cu spotted (Fig. 17) ....................................................................... 16 15. Wing with a long cloud extending from apical portion of vein Sc to wing apex (Costa Rica, Guyana, Peru, Brazil, Paraguay) ............................M. nigricosta Hough, 1900 Wing cloud not as above (Fig. 22) (Brazil) ....................M. lopesae Pamplona, 1986 16. Cell r1 with 1 spot (Fig. 21) (Brazil) ...................... M. xanthoptera Pamplona, 1986 Cell r 1 with 2 spots (Fig. 22) ............................................................................... 17 17. Flagellomere brownish yellow with a basal yellow spot; hind femur with weak and short anteroventral setae only on apical third; male: anteroventral, ventral and posteroventral surfaces with few setae on apical half (Brazil) ........................................................................... ........ M. couriae Pamplona, 1986
Flagellomere brownish yellow but lacking a basal yellow spot; hind femur, on Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
anteroventral surface, with weak and short setae from base to apex; male: anteroventral, ventral and posteroventral surfaces with many setae on apical 'half (Brazil)
........................................................... .M. dendropanacis Pamplona e Couri, 1995
fl
st7
IO,3mm
6a
6b
-cer
7a
7b
Figs. 4-7, characters de Morellia (modified from PAMPLONA, 1986c; PAMPLONA & COURI, 1995): 4, Morellia sp., antenna, lateral view; 5, M. concacata, wing, dorsal view; 6a, M. concacata, ovipositor, dorsal view; 6b, M. concacata, ovipositor, ventral view; Fig. 7a, M. dendropanacis, ovipositor, dorsal view; 7b, M. dendropanacis, ovipositor, ventral view. Abbreviations: ar, arista; cer, cerci; f1, f1agellomere; M, vein; R I , vein; R2 +3 , vein; Sc, subcostal vein; st, sternite.
46
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
pal'
2mm
~
~2 2m01
201m
201m
2mm
Figs. 8-22, characters of Morellia (modified from PAMPLONA, 1986c; PAMPLONA & COURI, 1995): 8, M. dalcyi, left surstylus, dorsal view; 9, M. dalcyi, aedeagus, ventral view; 10, M. dalcyi, aedeagus, lateral view; 11, M. dalcyi, ovipositor, dorsal view; 12, M. affinis, wing, dorsal view; 13, M. concacata, wing, dorsal view; 14, M. ochricornis, wing, dorsal view; 15, M. roppai, wing, dorsal view; 16, M. sinopensis, wing, dorsal view; 17, M. couriae, wing, dorsal view; 18, M. humeralis, wing, dorsal view; 19, M. iopesae, wing, dorsal view; 20, M. nigricosta, wing. dorsal view; 21, M. xanthoptera, wing, dorsal view; 22, M. dendropanacis, wing, dorsal view. Abbreviations: dis, distiphallus; dm-cu, crossvein; epi, epiphallus; gon, gonopod; hyp, hypandrium; par, paramere; r-m, crossvein.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Musca Linnaeus, 1758 Type-species. Musca domestica Linnaeus (I. C. Z. N., 1925, Opinion 82). Diagnosis. Male holoptic; anterointernal ommatidia enlarged; eyes bare; arista enlarged on basal fourth, plumose; pre sutural acrostichals not developed; dorsocentral 2:4; katepisternals 1:2; anepimeron setulose; prosternum setulose; lower calypter broad, subtruncate posteriorly, with its anteromedian angle extending, at least, to slightly under base of scutellum; mid tibia without ventral setae; vein M strongly curved forward apically; sternite 1 setulose. References. HENNIG, 1963b, 1964. Monophyly. According to HENNIG (1965), the monophyly of this genus is not well established. Geographical record. Mexico, Guadeloupe Is., Guatemala, Belize, San Salvador, Nicaragua, Costa Rica, Panama, Nassau, Jamaica, Puerto Rico, Haiti, Dominica, Cuba, Barbados, Mona Is., St. Lucia, St. Vincent, Virgin Is., Grenada, Trinidad, Venezuela, Guyana, Surinam, Colombia, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is., Easter Is.; Nearctic and Palaearctic Regions; northwestern Oriental; cosmopolitan. Comments. Only M. domestica is recorded for the Neotropical Region. The following key also includes M. autumnalis De Geer, 1776, based on its economic importance and the possibility that it might invade the Neotropical Region. The larvae of both species can be easily distinguished by the posterior end ofthe third instar (STEYSKAL, 1973).
Key to species 1.
Frons totally black in male and grey in female, not shiny (Mexico, Nearctic and Palaearctic Regions, northwestern Oriental) ................M. autumnalis De Geer, 1776 Frons reddish brown around and under antenna in male and silver shiny in female (Mexico, Guadeloupe Is., Guatemala, Belize, San Salvador, Nicaragua, Costa Rica, Panama, Nassau, Jamaica, Puerto Rico, Haiti, Dominica, Cuba, Barbados, Mona Is., St. Lucia, St. Vincent, Virgin Is., Grenada, Trinidad, Venezuela, Guyana, Surinam,
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Colombia, Peru, Ecuador, Galapagos Is., Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is., Easter Is.; cosmopoIltan) .......M. domestica Linnaeus, 1758
Neomyia Walker, 1859 Type-species. Neomyia gavisa Walker (mon.). Diagnosis. General body coloration metallic green; male holoptic; eyes bare; arista plumose; presutural acrostichals not developed; dorsocentral2:3; katepisternals 1:2; anepimeron setulose; prosternum setulose; lower calypter broad, subtruncate posteriorly, with its anteromedian angle extending, at least, to slightly under base of scutellum; mid tibia with a strong, ventral seta apically; vein M strongly curved forward apically; sternite 1 setulose. Reference. HENNIG, 1963b (as caesarion). Monophyly. Not yet established. Geographical record. Mexico, Cuba, Costa Rica, Brazil; Nearctic and Palaearctic Regions, northwestern Oriental. Comments. Only Neomyia cornicina (Fabricius, 1781) is found in the Neotropical Region.
Neorypellia Pont, 1972 Type-species. Neopyrellia neglecta Townsend (orig. des.). Diagnosis. General body coloration metallic green; male holoptic; anterointernal facets elongated; eyes bare; interfrontal setae absent; arista plumose; prosternum bare; pre sutural acrostichals 1; posterior dorsocentral2 strong setae; intra-alar 1: 1; supra-alars 2; katepisternals 1:2; anepimeron setulose; calypteres glossiform, the upper one with basal half ciliated ventrally; base of veins R 2+3 and R 4+5 with cilia on dorsal surface; stem vein with cilia on both surfaces; M strongly curved forward apically; hind tibia with a small posterodorsal seta (calcar), shorter than the diameter of tibia; sternite 1 setulose. Reference. ALBUQUERQUE, 1955a. Monophyly. Not yet established. Geographical record. Brazil. Comments. Monobasic genus: Neorypellia neglecta (Townsend, 1939). Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Parapyrellia Townsend, 1915 (Figs. 23-25) Type-species. Musca violacea Fabricius of authors (orig. des.) =maculipennis Macquart Diagnosis. General coloration metallic green; male holoptic with anterointernal facets elongated; eyes bare; arista plumose; interfrontal setae absent; prosternum bare; dorsocentrals 1:2; supra-alars 2; katepisternals 1:2; anepimeron bare; calypteres glossiform, the lower one enlarged; vein M strongly curved forward apically (Figs. 23,24); sternite 1 setulose. Reference. ALBUQUERQUE & LOPES, 1979a. Monophyly. Not yet established. Geographical record. Mexico, Cuba, Puerto Rico, Jamaica, Dominica, Trinidad, Guadeloupe Is., Colombia, Venezuela, Guyana, Peru, Bolivia, Brazil, Paraguay. Comments. The two known species occur only in the Neotropical Region.
Key to species 1.
Male: hind tarsi without a strong spine; female: calypter brown; wing not as below (Fig. 23) (Mexico, Cuba, Puerto Rico, Jamaica, Dominica, Trinidad, Guadeloupe Is., Colombia, Venezuela, Guyana, Peru, Bolivia, Brazil, Paraguay)
.......................................................................... P. maculipennis (Macquart, 1846) Male: hind tarsi with a strong spine (Fig. 25); female: calypter yellow; wing as in Fig. 24 (Brazil) ............................................... P. oportuna Albuquerque & Lopes, 1979
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
~ 23
met
Figs. 23-25, characters of Parapyrellia (modified from ALBUQUERQUE & LOPES, 1979a): 23, P. lI1aclllipennis, wing, dorsal view; 24, P. oportllna, wing, ventral view; 25, P. oportllna, hind leg, lateral view. Abbreviation: met, metatarsus.
Polietina Schnabl & Dziedzicki, 1911 (Figs. 26-32, Plate 1) Type-species. Aricia pruinosa Macquart (orig. des.) = Mydaea concinna Wulp. Aricia pruinosa Macquart, 1846, preoc. Zetterstedt, 1845. Diagnosis. Eyes bare or ciliated; male holoptic; arista with long cilia on both surfaces, longer near base, measuring about twice the length of the pedicel; one proclinated orbital seta; palpus filiform or slightly spatulate apically; presutural acrostichals developed or not; dorsocentrals 2:3 or 2:4; prealar strong and long; pro sternum setulose laterally; membrane around prosternum bare; one long proepisternal setae and one long upward directed proepimeral seta; notopleuron with 3 setae, the median one short; postalar wall ciliated; margin of posterior spiracle ciliated; katepisternals 1:2; calypteres bare on dorsal surface, lower calypter about twice as long as upper one; vein R] on dorsal surface ciliated from base to apex; R 4+5 on dorsal surface with cilia from node to apex or almost so, on ventral surface with sparse cilia up to near crossvein r-m or near the apex; hind tibia, on posterodorsal surface, with a strong calcar and a strong pre apical seta; sternite 1 setulose; male: sternite 5 longer than wide; cercal plate on ventral surface with spiny processes (Figs. 26-32); female: ovipositor long, tergites elongated, wider basally. Reference. COURI & CARVALHO, 1997b. Monophyly. Genus monophyletic, based at least, on the presence of setae on Muscidae (Diptera) of the NeotropicaI Region: TAXONOMY
51
Part I - Basal Groups
postalar wall, a derived character among the Muscidae (COURI & CARVALHO, 1997b). Geographical record. Mexico, Venezuela, Bolivia, Peru, Brazil, Argentina. Comments. COURI & PAMPLONA (1997) transferred Cyrtoneurina ingrata Couri, 1982, Cyrtoneurina prima Couri & Machado, 1990 and Cyrtoneurina stellata Couri, 1992 to this genus. These species were included in the key below. Key to species (modified from COURI & CARVALHO, 1997b)
Male 1.
Postsutural dorsocentral setae 3 ............................................................................ 2 Postsutural dorsocentral setae 4 ............................................................................ 8
2.
Hind tibia, on anteroventral surface, with less than 4 setae on middle-third (usually 3) .......................................................................................................................... 3 Hind tibia, on anteroventral surface, with more than 4 setae on middle-third (usually 5) ......................................................................................................................... 7
3.
Presutural acrostichal setae not developed; spiny lower process of cercal plate with 3 spines similar in length (Fig. 27) (Mexico )............. .l~ wulpi Couri & Carvalho, 1997 Presutural acrostichal setae developed (one or two pairs); spiny lower process of cercal plate with 3 spines, one more developed than the others ............................ ..4
4.
Spines on lower process of cercal plate directed upward (Fig. 28) (Brazil)
.............................................................................. ..... P. minor Albuquerque, 1956 Spines on lower process of cercal plate directed downward (Fig. 29) (Brazil) ......... 5 5.
Lower process of cercal plate with 5 spines; hind tibia on anterodorsal surface with at most 5 setae on middle third (Brazil) ................ P. univittata Couri & Carvalho, 1996 Lower process of cercal plate with less than 5 spines; hind tibia on anterodorsal surface with more than 5 setae on middle third ....................................................... 6
6.
Lower process of cercal plate with 3 spines (Fig. 29) (Brazil) .......................... ..
52
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
.......................................................................... ....... .P. bicolor Albuquerque, 1956 Lower process of cercal plate with 4 spines (Brazil) ..... ..P. nigra Couri & Carvalho, 1996 7.
Wings not spotted; spiny lower process of cercal plate with 3 downwarddirected spines, 1 more developed than others (Fig. 26) (Peru, Brazil) .......... . ..................................................................................... P. jlavithorax (Stein, 1904)
Wings with brown spots on stigma, apical third of vein R 2+3 and on crossveins r-m and dm-cu (Brazil) .............................................................. ....... P. stellata (Couri, 1982) 8.
Spiny lower process of cercal plate with 4 well developed spines (Fig. 30) (Brazil) ....................................................................................... P. steini (Enderlein, 1927)
Spiny lower process of cercal plate with less than 4 spines ...................................... 9 9.
Spiny lower process of cercal plate with 2 developed spines, one almost vestigial (Fig. 39) (Mexico) .............................................. ............. P. concinna (Wulp, 1896) Spiny lower process of cercal plate with only one developed spine (Fig. 32) (Peru, Bolivia, Brazil, Argentina) ............................................ ..... P. orbitalis (Stein, 1904)
Female 1.
Postsutural dorsocentral setae 3 ............................................................................. 2 Postsutural dorsocentral setae 4 ............................................................................. 7
2.
Tergite 5 paler than remainder of abdomen ............................................................. 3 Tergite 5 concolours with remainder of abdomen .................................................. ..4
3.
Tergite 5 yellow on apical half; hind tibia with about 5 anteroventral setae on middle third (Venezuela, Peru, Bolivia, Brazil) ........................... P.jlavithorax (Stein, 1904) Tergite 5 entirely light brown; hind tibia with about 2 anteroventral setae on middle third (Brazil) .............................................. ......... P. prima (Couri & Machado, 1990)
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
4. Fore leg brown; hind tibia on anterodorsal surface with no more than 5 setae (Brazil)
................................................................................... P minor Albuquerque, 1956 Fore leg yellow or light brown; hind tibia on anterodorsal surface with more than 5 setae ..................................................................................................................... 5 5.
Fore leg light brown (Brazil) ........................... P. univittata Couri & Carvalho, 1996 Fore leg yellow................................................... ,......................................... '" ..... 6
6.
Hind tibia on anterodorsal surface with 7 setae on middle third (Brazil)
............................................................................... ? nigra Couri & Carvalho, 1996 Hind tibia on anterodorsal surface with 8-9 setae on middle third (Brazil)
................................................................................... ? bicolor Albuquerque, 1956 7.
Hind tibia on anteroventral surface with 5 or more setae (Brazil) .............. .
................................................................... .................. P. major Albuquerque, 1956 Hind tibia on anteroventral surface with no more than 4 setae .................................. 8 8.
Abdomen brown, with yellow, translucent areas on tergites 1+2 and 3, tergite 5 yellow; antennal flagellomere brown (Mexico, Venezuela) ............... P. rubella (Wulp, 1896) Abdomen not as above (see below); antennal flagellomere yellow.......................... 9
9.
Hind tibia on anteroventral surface with 4 setae on middle third; legs yellow, mid femur brown apically; hind femur brown on apical half (Brazil) ............ .
................................................................ ....................... P. steini (Enderlein, 1927) Hind tibia on anteroventra1 surface with 2 setae on middle third; leg colour not as below................................................................................................................... 10 10. Legs brown; fore femur, fore tibia and mid femur yellowish brown; abdomen silver pruinose and with yellow lateral spots on basal half of tergites 1-3 (Brazil)
.............................................................................. ? distincta Couri & Lopes, 1987 Legs and abdomen not as above (see below) ......................................................... 11 54
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
11. Abdomen yellowish with dark brown vittae on apex of tergites 1-4, tergite 5 with yellow apex (Mexico) ....................................................... P. concinna (Wulp, 1896) Abdomen not as above ........................................................................................ 12 12. Legs yellow (Peru, Bolivia, Brazil, Argentina) ......................P. orbitalis (Stein, 1904) Legs brown, with apical portion of fore and hind femora yellow (Brazil)
.......................................................................................... P. ingrata (Couri, 1982) Species of Polietina not included in the key and their respective geographical distribution Species
Geographical distribution
P. basicincta (Stein, 1904)
Peru, Bolivia
P. jlavidicincta (Stein, 1904)
Colombia, Venezuela, Bolivia
P. mellina (Stein, 1904)
Bolivia
Sarcopromusca Townsend, 1927 Type-species. Sarcopromusca arcuata Townsend (mon.) =pruna (Shannon & Del Ponte). Diagnosis. General coloration blackish with grey pruinescence; male holoptic; arista enlarged at basal fourth, plumose; acrostichals 0:2; dorsocentra13 :4; meron setulose;
katepisternals 1:3; anepimeron setulose; vein M strongly curved forward apically; lower calypter with cilia on internal border; mid tibia on ventral surface with a pre apical seta; apex of abdomen yellow gold; sternite 1 setulose. Reference. PAMPLONA, 1991. Monophyly. Not yet established. Geographical record. Guatemala, Nicaragua, Costa Rica, Panama, Colombia, Brazil, Argentina. Comments. Only two species are known, both restricted to the Neotropical Region.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
55
Part I - Basal Groups
Key to species 1.
Costal margin without an elongated brown cloud (Guatemala, Nicaragua, Costa Rica, Brazil, Argentina) ........................................ S. pruna (Shannon & Del Ponte, 1926) Costal margin with an elongated brown cloud (Costa Rica, Panama, Colombia, Brazil, Argentina) ................................................. ................ S. sarcophagina (Wulp, 1896)
1 mm
1mm
1mm
Figs. 26-32, characters of cereal plate, ventral view, of Polietina (modified from COURI & CARVALHO, 1997b): 26, P. flavithorax; 27, P. wUlpi; 28, P. minor; 29, P. bicolor; 30, P. steini; 31, P. concillfla; 32, P. orbitalis. Abbreviation: pr, process.
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Xenomorellia Malloch, 1923 (Figs. 33-36, Plate 1) Type-species. Xenamarellia haiti Malloch (orig. des.). Diagnosis. Male holoptic; arista plumose, bare on apical fourth; prealar seta long and strong; acrostichals 2:2; dorsocentrals 2:4; anepimeron setulose; M curved forward apically (Fig. 33); stem vein with cilia on dorsal surface and with a long seta near the insertion of the humeral crossvein (Fig. 34); radial node with cilia on both surfaces; hind tibia with 2 posterior setae; hind coxa with posterior cilia; sternite 1 setulose. Reference. ALBUQUERQUE, 1952. Monophyly. Not yet established. Geographical record. Peru, Brazil. Comments. A Neotropical genus with only two described species.
Key to species 1.
Antenna and palpus black; setae on basal half of anteroventral surface of hind femur at least twice as long as the width of the tibia, on apical half with about 5 setae of unequal lengths (Fig. 35) (Brazil) .................. ...... .x. mantanhesa Albuquerque, 1952 Antenna and palpus dark yellow; setae on basal half of anteroventral surface of hind femur less than 1,5 times as long as the width of the tibia, on apical half with a series of long setae (Fig. 36) (Peru, Brazil) ............................................
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
x. haiti Malloch, 1923
57
Part I - Basal Groups
hum
O,5mm
1 rnm
35
36
Figs. 33-36, characters of Xenomorellia (modified from ALBUQUERQUE, 1952): 33, X. montanhesa, wing, dorsal view; 34, X. montanhesa, wing, humeral vein, dorsal view; 35, X. montanhesa, hind leg, lateral view; 36, X. holti, hind leg, lateral view. Abbreviations: ad, anterodorsal surface; av, anteroventral surface; hum, humeral vein; sv, stem vein.
Tribe Stomoxyini Meigen, 1924
Diagnosis. Proboscis elongated, adapted to haematophagy, strongly sclerotized; labella reduced and not retractile; arista with long dorsal cilia and generally bare or at most with very few ventral cilia; male: epiphallus absent; female: 2 spermathecae. Monophyly. Tribe undoubtedly monophyletic (HENNIG, 1965, 1973), having Muscini as sister-group (CARVALHO, 1989d). Comments. This tribe have species distributed allover the world, with 10 described genera. Only three genera and six species are recorded to the Neotropical Region.
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
m
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Haematobia Le Peletier & Serville in LATREILLE et ai., 1828 Type-species. Conops irritans Linnaeus (WESTWOOD, 1840: 140 and I.C.Z.N., 1974, Opinion 1008). Diagnosis. General coloration dark greyish, with yellowish tinge and often yellow setae; head width 1.25 times its height; arista plumose only on dorsal surface; palpus spatulate, a little narrowed on basal quarter to half, grooved inside, reaching tip ofhaustellum when proboscis flexed; prostemum setulose; propleuron and postalar wall bare; prealar seta barely differentiated or vestigial; lower calypter about twice as long as upper one; vein AJlong. Reference. HENNIG, 1964. Monophyly. Not yet established. Geographical record. Mexico, Guatemala, EI Salvador, Nicaragua, Costa Rica, Panama, Puerto Rico, Cuba, Jamaica, Virgin Is., Haiti, S. Domingo, Colombia, Venezuela, Guyana, Brazil, Argentina, Chile; Nearctic Region; Europe, Asia, North Africa, Hawaii. Comments. This genus has species distributed in all biogeographical regions, but only Haematobia irritans (Linnaeus, 1758) occurs in the Neotropical Region. This species probably came from Venezuela to Brazil in the beginning of the eighties, and its widespread occurrence in this country was due to dispersion (VALERIO & GUIMARAES, 1983).
Neivamyia Pinto & Fonseca, 1930 (Figs. 37-44) Type-species. Neivamyia Zutzi Pinto & Fonseca (orig.des.)
=flavicornis
Malloch. Diagnosis. Head slightly wider than high; palpus stout subcylindric and reaching tip ofhaustellum when proboscis flexed; arista plumose on upper surface and with about 3 or 4 shorter cilia below on middle portion (Figs. 37,42,44); prostemum setulose; propleura and postalar wall bare, with at least 2 long postsutural dorsocentral setae; vein Al moderately long; lower calypter about twice as long as upper one; mid and hind tibiae with a submedian seta. Reference. LOPES & MANGABEIRA FILHO, 1938; LOPES, 1955. Monophyly. Not yet established. Adrian Charles Pont (personal communication)
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
understands that Neivamyia could be synonymized with Stomoxys, since there are no significant differences between them. Geographical record. Trinidad, Guyana, Surinam, Peru, Brazil. Comments. A Neotropical genus with only four species.
Key to species (males only) 1.
Male: orbital proclinate setae on frons .................................................................... 2 Male: no proclinate setae on frons ......................................................................... 3
2.
Frons about one-fifth of head width (Figs. 38, 39) (Colombia) ....................... .
...... ........ ...... ........ .... ..... ...... ........ ......... ..... ....................... .N. antenusi Lopes, 1955 Frons less than one-fifth of head width (Peru, Brazil) (Fig. 40) .......................... ..
.................. ......... ..... ......... ......... ....... .................... .........N. latifrons Malloch, 1932 3.
Frons less than one-tenth of head width (Fig. 41); proboscis measuring about the same length as the maximum height of the eye (Fig. 42) (Trinidad, Guyana, Surinam,
Brazil) ..................................................................... .N.Jlavicomis (Malloch, 1928) Frons more than one-tenth of head width (Fig.43); proboscis shorter than the maximum height the eye (Fig. 44) (Brazil) .. ..............N. travassoi Lopes & Mangabeira, 1938
Stomoxys Geoffroy, 1762 Type-species. Conops calcitrans Linnaeus (LC.Z.N., 1957, Opinion 441). Diagnosis. General coloration brownish grey to yellow brown; head slightly wider than high; vertex about one-fourth of head width in male and well over one-third in female; arista plumose; pal pus slender, subcylindric, scarcely one-third as long as haustellum; flagellomere about 2,5 as long as pedicel; prostemum and proepistemum setulose; prealar seta absent. Reference. HENNIG, 1964. Monophyly. Not yet established, but probably monophyletic (see discussion under Neivamyia). 60
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
O,5mm
1 mm
1 mm 1 mm
prb Figs. 37-44, characters of Neivamyia (modified from LOPES & MANGABEIRA, 1938; LOPES, 1955): 37, N. travassoi, antenna, lateral view; 38, N. antenusi, head, dorsal view; 39, N. antenusi, head, lateral view; 40, N. tatifrons, head, dorsal view; 41, N. ftavicornis, head, dorsal view; 42, N. ftavicornis, head, lateral view; 43, N. travassoi, head, dorsal view; 44, N. travassoi, head, lateral view. Abbreviations: ar, arista; pip, palpus; prb, proboscis.
Geographical record. Mexico, Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Puerto Rico, Cuba, Jamaica, Virgin Is., St. Vincent, Haiti, Dominica, Trinidad, Tobago, Venezuela, Guyana, Colombia, Peru, Ecuador, Galapagos Is., Bolivia, Brazil, Uruguay, Argentina, Chile, Easter Is.; cosmopolitan. Comments. The genus has species distributed in all biogeographical regions, but only Stomoxys calcitrans (Linnaeus, 1758) occurs in the Neotropical Region.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Subfamily Azeliinae Robineau-Desvoidy, 1830 Diagnosis. Arista not plumose, with the cilia shorter than the basal width of arista. Ovipositor with tergites 6 and 7 divided anteriorly; tergite 8 sc1erotized, anchor-type. Monophyly. Not established yet. Comments. SKIDMORE (1985) and CARVALHO (1989d) recognised the subfamily; the first author based on characters of the immature forms and the second, on derived characters of the adults listed in the diagnosis. The subfamily is divided in to two tribes, Azeliini and Reinwardtiini, tentatively indicated as sister-groups by CARVALHO (1989d).
Tribe Azeliini Robineau-Desvoidy, 1830 Diagnosis. Gena, in most species, with strong, upcurved setae; subcostal vein reaching apical margin of wing in a smooth curve; distiphallus with apical part of juxta spinulose (Figs. 53-56); female with a proc1inate fronto-orbital setae. Monophyly. Probably Azeliini is a monophyletic tribe (CARVALHO, 1993b). In most genera 2 character-states are found: a proc1inate fronto-orbital in female, and subcostal smooth. The former character-state is in the ground plan of the Muscidae (HENNIG, 1965). The second character is probably in the ground plan of Muscoidea (sensu McALPINE, 1989). However the most striking character in the tribe is the configuration of the male distiphallus, with the juxta spinulose (CARVALHO, 1993b). Comments. In the world there are nine genera, seven of them with species occurring in the Neotropical Region. 24 species are known to this region.
Azelia Robineau-Desvoidy, 1830 Type-species. Azeliajlorea Robineau-Desvoidy (RONDANI, 1866) =triquetra Wiedemann. Diagnosis. Wing with vein Al ending abruptly, extending at most halfway between the apex of the posterior cubital cell and the margin of the wing; hind coxa with a row of weak posterior setae; male with enlarged ommatidia; female with interfrontal setae. Reference. SNYDER, 1957b.
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Claudio Jose Barros de Carvalho & Marcia Souto Couri
Monophyly. Not yet established. Geographical record. Mexico, Argentina. Comments. Genus with species distributed all over the world, but after CARVALHO (1993b), only one species, Azelia neotropica Snyder, 1957, has been recognised in the neotropics. VOCKEROTH (1996) in his key to the Muscidae of Central America, mentioned this genus to Mexico and Panama, but unfortunately he did not provide specific identification of the material.
Drymeia Meigen, 1826 Type-species. Drymeia obscura Meigen (mon.) =hamata (Fallen). Diagnosis. General coloration blackish; male wing brownish; female wing light brown; arista pubescent; prealar strong; hind tibia usually with 3 or more strong posterodorsal setae which are more or less uniformly distributed along tibial length. References. HUCKETT, 1975; VOCKEROTH, 1996. Monophyly. Not yet established. Geographical record. Mexico; Nearctic, Palaearctic and Oriental regions. Comments: It seems that only Drymeia aterrima Wulp, 1896 occurs in Mexico (CARVALHO et al. 1993), although VOCKEROTH (1996) did not include this species in his key to the Muscidae of Central America. The syntypes males and females are in the BMNH and they are conspecifics.
Hydrotaea Robineau-Desvoidy, 1830 (Figs. 45-70) Type-species. Musca meteorica Linnaeus (des. CURTIS, 1839). Diagnosis. General coloration black or bluish-black, not metallic; gena with strong, upward-directed setae; thorax with no vittae or with 2 or 3 white, pruinose vittae; arista pubescent; dorsocentral setae 2:4; notopleuron covered with setulae, with the 2 setae of similar size; male: fore femur on ventral and posteroventral surfaces with 2 toothed processes; fore tibia flattened along basal half or two-thirds of ventral surface; female: inter-segmentar membrane of the ovipositor with granulations. Reference. PALKA-ROCHA & CARVALHO, 1994. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
63
Part I - Basal Groups
Monophyly. Probably monophyletic. After the works of SKIDMORE (1985) and PALKA-ROCHA (1992), it has been hypothesized, based on a large number of characters, that Ophyra Robineau-Desvoidy is the sister-group of Australophyra + Hydrotaea Robineau-Desvoidy. PONT (1986a, 1989) considered Ophyra a junior synonym of Hydrotaea, a classification not followed by PAMPLONA & COURI (1989) and CARVALHO et al. (1993). Geographical record. Mexico, Costa Rica, Colombia, Ecuador, Peru, Bolivia, Brazil, Argentina, Chile, Juan Fernandez Is.; Nearctic and Palaearctic Regions; northern India; cosmopolitan. Comments. This genus is found in all biogeographical regions; only eight species occur in the Neotropical Region. Hydrotaea Juscocalyptrata Macquart, 1855 was not included in the key because this species is known only from one male in very bad condition, deposited at UMO (PALKA-ROCHA & CARVALHO, 1994).
Key to species (modified from PALKA-ROCHA & CARVALHO, 1994) 1.
Male: unknown; female: general coloration shiny black; wing darker on apical half (Colombia) ............................................ ..................... H. nubilicosta Malloch, 1923 Male: eyes ciliated or bare; prealar setae 1 or 2; sternite 5 similar or wider in length than in width (Figs. 49-52); surstylus, in posterior view, rounded or elongated (Figs. 45-48); distiphallus as in Figs. 53-56; hypandrium, in dorsal view, triangular or rounded (Figs. 57-60); female: general coloration black to bluish black, not shiny; wing hyaline or uniformly brownish ............................................................................................. 2
2.
Halter with the knob light brown or yellow; male: eyes bare; mid tibia on ventral surface with long and fine setae on apical half; hind femur on ventral surface with 2 strong basal spines (HUCKETT, 1954; ALBUQUERQUE, 1957a); female: prealar absent; mesonotum with 2 white, pruinose vittae that reach the third and fourth pairs of dorsocentral postsutural setae; ovipositor without granulations on the intersegmentar membrane 6-8 (Figs. 61-66) (Chile; North America) .................H. acuta Stein, 1898 Halter with the knob black; male: eye ciliated; mid tibia, on ventral surface with no setae on apical half; hind femur on ventral surface with no spines; female: prealar
64
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
present; mesonotum with no vittae; ovipositor not as described above (Figs. 62-65; 67-70) ................................................................................................................... 3 3.
General coloration bluish black; calypter dark brown; male: cerci with anterior margin angulated (Fig. 45); female: ovipositor without granulations on membrane of segment 8 (Fig. 67) (Colombia, Bolivia, Argentina, Chile )... .H. cyaneiventris Macquart, 1851 General coloration black; calypter not as described above; male: cerci with the anterior margin not as described above (Figs. 46-48); female: ovipositor with granulations on membrane of segment 8 (Figs. 63-65; 68-70) ........................................................ ..4
4.
Eye with long and dense cilia; calypteres yellow; fore femur, on anterior surface, with setae; male: prealar setae 1; surstyIi on posterior view elongated (Fig. 48); hypandrium in dorsal view rounded (Fig. 60); female: prealar 1; mid tibia, on anterodorsal surface with no setae on limit of the apical third (Colombia) ..................H. villosa Stein, 1904 Eyes with short and sparse cilia; calypteres not yellow; male: prealar 1 or 2; fore femur, on anterior surface, with a row of setae on basal half; surstyli on posterior view rounded (Figs. 45-47); hypandrium in dorsal view, triangular (Figs. 57-59); female: prealar 2; fore femur, on anterior surface, with cilia or with a row of weak setae on basal half; mid tibia, on anterodorsal surface, with one seta on the limit of the apical third ..................................................................................................................... 5
5.
Outer vertical seta similar in length to the inner ones; male: prealar 2; fore tibia on anteroventral surface, with a row of short and strong setae on apical third; stemite 5 longer than wider (Fig. 50); cerci as in Fig. 46; distiphallus with apical spines developed (Fig. 54); hypandrium as in Fig. 58; female: mesonotum with no vittae (Venezuela, Ecuador; North America; Hawaii) .......................................H. houghiMalloch, 1916 Outer vertical seta shorter than the inner ones; male: prealar 1 or 2; fore tibia not as described above; stemite 5 with similar length and width (Fig. 51); cerci as in Fig. 47; distiphallus with apical spines not as described above (Fig. 55); hypandrium as in Fig. 59; female: mesonotum with three polinose vittae (Ecuador, Peru, Bolivia, Brazil, Argentina, Chile) ............................................. ................H. nicholsoni Curran, 1939
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
7
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Part I - Basal Groups
Q,5mm
53
54
55
56
Figs. 45-60, characters de Hydrotaea (modified from PALKA-ROCHA & CARVALHO, 1994): 45, H. cyaneiventris, cerci, epandrium and surstyli, posterior view; 46, H. houghi, cerci, epandrium and surstyli, posterior view; 47, H. nicholsoni, cerci, epandrium and surstyli, posterior view; 48, H. villosa, cerci, epandrium and surstyli, posterior view; 49, H. cyaneiventris, sternite 5, posterior view; 50, H. houghi, sternite 5, posterior view; 51, H. nicholsoni, sternite 5, posterior view; 52, H. villosa, sternite 5, posterior view; 53, H. cyaneiventris, aedeagus, lateral view; 54, H. houghi, aedeagus, lateral view; 55, H. nicholsoni, aedeagus, lateral view; 56, H. villosa, aedeagus, lateral view; 57, H. cyaneiventris, hypandrium, dorsal view; 58, H. houghi, hypandrium, dorsal view; 59, H. nicholsoni, hypandrium, dorsal view; 60, H. villosa, hypandrium, dorsal view. Abbreviations: aed apo, aedeagal apodeme; cer, cerci (cereal plate); dis, distiphallus; epd, epandrium; epi, epiphallus; gon, gonopod; sur, surstylus.
66
~~~~
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _EW"
Claudio Jose Barros de Carvalho & Marcia Souto Couri
66
67
68
69
70
Figs. 61-70, characters of Hydrotaea (modified from PALKA-ROCHA & CARVALHO, 1994): 61, H. acuta, ovipositor, dorsal view and spermathecae; 62, H. cyaneiventris, ovipositor, dorsal view and spermathecae; 63, H. houghi, ovipositor, dorsal view and spermathecae; 64, H. nicholsoni, ovipositor, dorsal view and spermathecae; 65, H. villosa, ovipositor, dorsal view and spermathecae; 66, H. acuta, ovipositor, ventral view; 67, H. cyaneiventris, ovipositor, ventral view; 68, ovipositor, ventral view, H. houghi; 69, ovipositor, ventral view, H. nicholsoni; 70, H. villosa, ovipositor, ventral view. Abbreviation: spr, spermathecae.
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Part I - Basal Groups
Micropotamia Carvalho, 1993 (Figs. 71-76) Type-species. Phaonia minuscula Albuquerque (orig. des.). Diagnosis. Parafacial very narrow in both sexes, strongly tapering downward; head of female with proclinate orbitals; interfrontal setae absent, frontal plate with several setae; arista pubescent, with hairs shorter than basal width of the arista; gena with a strong up curved seta; dorsocentral setae 2:3; pre sutural acrostichals developed; prealar short, slightly longer than adjacent ground-setulae; anepimeron bare; katepistemals 1:2, the lower one weak; posterior spiracle small in both sexes, smaller than the knob of halter, bare on margins; wing veins bare, except C; vein Sc smoothly curved on apical portion; mid femur with a preapical anterior seta; mid tibia with two median posterior setae; hind coxa bare on posterior apical margin; hind tibia with a strong calcar on apical third; stemite 1 bare; male: distiphallus with spiny and sclerotized processes; female: ovipositor long (Figs. 71,72). Reference. CARVALHO, 1993b. Monophyly. Probably monophyletic. According to CARVALHO (1993b), Micropotamia share an apomorphic character - absence of interfrontal setae in female with Megophyra Emden, 1965, which has only species occurring in the Oriental Region (PONT, 1977). Geographical record. Mexico, Brazil, Argentina. Comments. This genus was proposed for three South American species, but Phorbia fuscisquama Wulp, considered insertae sedis by CARVALHO et al. (1993), also belongs to this genus and is included in the key below. This species is known only from Mexico.
Key to species (modified from CARVALHO, 1993b) 1.
Antenna dark brown with the pedicel yellowish; thorax dark brown with humeral callus yellowish; halter yellow (Brazil) .... M. amazonica (Albuquerque & Lopes, 1982) Antenna entirely dark brown; thorax black, halter yellow, dark brown at base ......... 2
2.
Male: hind femur enlarged on apical third (AlBUQUERQUE, 1955a:Fig.1);cercalplate as in Fig. 75; stemite 5 as in Fig. 76 (Brazil) ............ M. minuscula (Albuquerque, 1955)
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Claudio Jose Barros de Carvalho & Marcia Souto Couri
Male: hind femur not enlarged on apical third; cereal plate not as above ................. 3 3.
Male: hind femur with short setae on anteroventral face; cereal plate as in Fig. 73; sternite 5 as in Fig. 74; female: ovipositor not as below (Brazil) ........................ .
..............................................................~ .............M. cilitibia (Albuquerque, 1955) Male: hind femur with only long setae on whole anteroventral surface; cereal plate not as above; female: ovipositor with narrow stemites and tergites (Mexico) ..................... .
.............................................................. .... M.fuscisquama (Wulp, 1896), comb.n.
1 mm
73 E E
1
mm/
1mm
Figs. 71-76, characters of Micropotamia (modified from CARVALHO, 1993b): 71, M. amazonica, ovipositor, dorsal view; 72, M. amazonica, ovipositor, ventral view; 73, M. cilitibia, cereal plate, posterior view; 74, M. cilitibia, sternite 5, dorsal view; 75, M. minuscula, cereal plate, posterior view; 76, M. minuscula, sternite 5, dorsal view.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Ophyra RobineaugDesvoidy, 1830 (Figs. 77-88) Type-species. Ophyra nitida Robineau-Desvoidy (des. RONDANI, 1866) = ignava Harris. Diagnosis. General coloration metallic black; male holoptic; eyes bare; ocellar triangle shiny, large and long, sometimes reaching lunule; frontal setae 6-11 pairs; proclinate seta and interfrontal setae present in female; arista bare or pubescent; dorsocentral setae 2:4; katepisternals 1: 1; anepimeron bare; vein M straight; ovipositor long with narrow tergites. Reference. PAMPLONA & COURI, 1989. Monophyly. See discussion under genus Hydrotaea. Geographical record. Mexico, Nicaragua, St. Vincent, Puerto Rico, Jamaica, Cuba, Trinidad, Tobago, Venezuela, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Argentina, Chile; Nearctic and Western Palaearctic Regions; Eastern Pacific). Comments. Besides the seven species recognised in the Neotropical Region, Ophyra ignava was recently identified in Chile (FIGUEIROA, 1999). VOCKEROTH (1996) followed PONT (1986a, 1989) and did not recognise Ophyra as a separate genus (see discussion under genus Hydrotaea).
Key to species (modified from PAMPLONA & COURI, 1989) 1.
Male ................................................................................................................... 2 Female ................................................................................................................ 8
2.
Palpus yellow or light brown; hind trochanter, on ventral surface, with a tuft of fine, hooked setae (Fig. 77) .......................................................................................... 3 Palpus dark brown or black; hind trochanter, on ventral surface, without a tuft of fine setae ..................................................................................................................... 4
3.
Mid femur with 6 short spinules on basal third; mid and hind femur brown to black; pedicel and flagellomere brown (Mexico, Nicaragua, St. Vincent; Puerto Rico, Jamaica, Cuba, Trinidad, Tobago, Guyana, Venezuela, Ecuador, Galapagos Is., Peru, Bolivia,
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Brazil, Uruguay, Argentina, Chile; Nearctic and Palaearctic Regions; Eastern Pacific)
.......................................................................... 0. aenescens (Wiedemann, 1830) Mid femur with 2-3 long spinules on basal third; mid and hind femur yellow; pedicel and flagellomere yellow (Cuba) ........................................... O. cubana Gregor, 1974 4.
Mid femur, on basal half of ventral surface, with a row of 10-12 strong setae (Figs. 78-79) .................................................................................................................. 5 Mid femur, on ventral surface, with a row of 2-6 setae (two rows in O. ignava, Fig. 80) ....................................................................................................................... 6
5.
Wing brown on upper half; calypteres yellow with brown margins (Brazil)
............................................................................... 0. solita ria Albuquerque, 1958 Wings uniformly light brown; calypteres infuscated with dark brown margins (Brazil) ................................................................................ O. albuquerquei Lopes, 1985 6.
Hind tibia strongly curved, its anteroventral, ventral and posteroventral surfaces with many fine and long setae on basal third (Fig. 81) (?Venezuela, Chile; Nearctic and Palaearctic Regions; North-west Oriental) ............................ 'O. ignava (Harris, 1780) Hind tibia not curved, setae on its anteroventral, ventral and posteroventral surfaces not as above ......................................................................................................... 7
7.
Fore tarsomeres yellow on ventral surface; mid femur on ventral surface with 4-6 fine cilia basally (Fig. 82); hind tibia on anteroventral, ventral and posteroventral without long setae on apical two-thirds (Brazil, Chile; Nearctic, Afrotropical, Oriental and Australasian regions ......................................... O. chalcogaster (Wiedemann, 1824) Fore tarsomeres uniformly dark brown to black; mid femur on ventral surface with 2 short and strong setae basally (Fig. 83); hind tibia on anteroventral, ventral and posteroventral with a series of long setae on apical two-thirds (Fig. 84) (Chile; Nearctic, Palaearctic and Afrotropical Regions; North India) ...... O. capensis (Wiedemann, 1818)
8.
Ocellar triangle long, reaching lunule (Fig. 85), strongly sclerotized, reddish-brown
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
(Chile; Nearctic, Palaearctic and Afrotropical Regions; North India) ............... . ................................................................................. 0. capensis (Wiedemann, 1818) Ocellar triangle short to long, not strongly sclerotized, blackish ................................ 9
9.
Ocellar triangle long, reaching lunule or ending very close to it (Figs. 86,88) ......... 10 Ocellar triangle shorter, not reaching lunule and not ending close to it (Fig. 87) ............. 13
10. Palpusyellow........................ " ................................................................................ 11 Palpus dark brown to black .................................................................................. 12
11. Pedicel and flagellomere brown, ocellar triangle broad with a round apex (Fig. 86); calcar long; anterior katepisternal seta hair-like (Mexico, Nicaragua, St. Vincent, Puerto Rico, Jamaica, Cuba, Trinidad, Tobago, Venezuela, Guyana, Ecuador, Galapagos Is., Peru, Bolivia, Brazil, Uruguay, Argentina, Chile; Nearctic and western Palaearctic Regions; eastern Pacific ) .................................... .0. aenescens (Wiedemann, 1830) Pedicel and flagellomere yellow, ocellar triangle narrow; calcar short; anterior katepisternal seta well developed (Cuba) ............................... O' cubana Gregor, 1974 12. Wing brown on upper half; calypteres yellow with light brown margins (Brazil) ................................................................................ 0. solitaria Albuquerque, 1958 Wing uniformly light brown; calypteres brown with dark brown margins (Brazil)
................................................................................. 0. albuquerquei Lopes, 1985 13. Upper calypter uniformly yellowish; fore tarsi usually with tarsomeres yellow on ventral surface (not so evident as in male); hind tibia on anteroventral surface with one seta (Brazil, Chile; Nearctic, Afrotropical, Oriental and Australasian Regions)
...................................................................... 0. chalcogaster (Wiedemann, 1824) Upper calypter slightly brownish with dark brown margins; fore tarsi usually with tarsomeres uniformly brown; hind tibia on anteroventral surface with 2 setae (?Venezuela, Chile; Nearctic and Palaearctic Regions, northwestern Oriental) ......................................................................................... 0. ignava (Harris, 1780) 72
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
C::::i'~;;f1Imtr ) 78
c
~
~';1~
81
82
c·····,::·· . ,;·. ? 83
oc tr
85
\rmmNJ 86
87
88
Figs. 77-88, characters of Ophyra (modified from PAMPLONA & CO URI, 1989): 77, O. aenescens, male, trochanter and mid femur; 78, O. salitaria, male, mid femur; 79. O. albuquerquei. male. mid femur; 80, O. ignava. male, mid femur; 81. O. ignava, male, hind tibia; 82, O. chalcagaster, mid femur; 83, O. capensis, male, mid femur; 84, O. capensis, male, hind tibia; 85, O. capensis, female, ocellar triangle; 86, O. aenescens, female, ocellar triangle; 87, O. chalcagaster, female. ocellar triangle; 88, O. albuquerquei, female, ocellar triangle. Abbreviation: oc tr, ocellar triangle.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Potamia Robineau-Desvoidy, 1830
(Figs. 89-93) Type-species. Potamia littoralis Robineau-Desvoidy (des. SEGUY, 1937). Diagnosis. Gena with 1-3 differentiated and upward-curved anterior setae; arista with cilia about twice as long as the basal diameter of the arista; katepistemum bare; male: distiphallus with spicules but not strongly sclerotized (Fig. 91); female: fronto-orbital proclinate setae present. References. SNYDER, 1955; CARVALHO, 1981. Monophyly. Probably monophyletic; this genus is one ofthe most basal of the tribe, judging from the presence of a primitive configuration of the male distiphallus (CARVALHO, 1993a), with spicules (Fig. 91), but without the strong sclerotization as found in Micropo tam ia, Ophyra, Australophyra Malloch (Australian genus), Hydrotaea (Figs. 53-56) and in the monophyletic group formed by the genera Azelia and Thrichops. Geographical record. Mexico, Peru, United States of America. Comments. Only two species occur in the Neotropical Region, both in Peru. Key to species
1. Male: cercal plate, in posterior view, without a cleft on posterior margin (Fig. 90); surstylus, in lateral view, as Fig. 89 (Peru) ..................... P plumata (Carvalho, 1981) Male: cercal plate, in posterior view, with a cleft on posterior margin (Fig. 92); surstylus, in lateral view, as Fig. 93 (Mexico, Peru; Nearctic Region) ... ... P scabra (Giglio-Tos, 1893)
Thricops Rondani, 1856
Type-species. Anthomyza hirtula Zetterstedt (orig. des.). Diagnosis. General coloration yellow; more than one inclined frontal setae present; vein Al extending more than halfway to wing margin; hind coxa with hairs on posterior apical margin; hind tibia with one posterodorsal setae. References. HUCKETT, 1975; VOCKEROTH, 1996.
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
apoede
89 90
92
93
Figs. 89-93, characters of Potamia (modified from CARVALHO, 1981; SNYDER, 1955): 89, P. plumata, epandrium, cereal plate and surstyli, lateral view; 90, P. plumata, epandrium, cereal plate and surstyli, posterior view; 91, P. plumata, aedeagus, lateral view; 92, P. scabra, epandrium, cerci and surstyli, posterior view; 93, P. scabra, epandrium, cereal plate and surstyli, lateral view. Abbreviations: apo ede, aedeagal apodeme; dis, distiphallus; epi, epiphallus; gon, gonopod.
Monophyly. Not proposed yet. Geographical record. Mexico; Nearctic, Palaearctic and Oriental regions. Comments. This genus, represented by Thricops diaphanus (Wiedemann, 1817) was reported for the first time in the Neotropical Region (Oaxaca, Mexico) by VOCKEROTH (1996). The male of this species can be recognised by a pronounced strong apical thorn on the ventral surfaces of hind tibia (HUCKETT, 1975).
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Tribe Reinwardtiini Brauer & Bergenstamm, 1889 Diagnosis. Hind femur, on posterodorsal surface, usually with a developed calcar; haustellum with a ciliated sclerotization; male: distiphallus membranous; female: frontoorbital usually absent; interfrontal setae absent. Monophyly. This tribe is probably not monophyletic (CARVALHO, 1989d), being the genera assembled on basis of primitive characters in relation to the ground plan of the Azeliinae. In the cladistic analysis of the family, CARVALHO (1989d) found only one synapomorphic character for this tribe - haustellum with a posterior ciliated sclerotization, but this character is homoplastic, occurring in many muscid genera. According to PONT (1986a, 1989), this tribe includes the most primitive genera of the family. Comments. A cosmopolitan tribe, with 16 genera, 11 of the them occurring in the Neotropical Region (CARVALHO etaZ., 1993). SKIDMORE (1985) called attention to the high number of Reinwardtiini genera occurring in the Neotropical Region. There are 85 species recorded from the neotropics.
Brachygasterina Macquart, 1851 (Figs. 94-95) Type-species. Brachygasterina vioZaceiventris Macquart (orig. des.). Diagnosis. Male holoptic, eyes with cilia; arista bare; flagellomere strongly dilated (Fig. 94-95); palpus dilated; interfrontal setae present or absent in female; presutural acrostichal developed; prealar seta present or absent; dorsocentrals 2:4; katepimeron bare; katepisternals 1: 1; R node with or without cilia; calcar present and strong; veins R4+5 and M parallel apically; ovipositor long, tergite 6 anchor-shaped, tergite 7 a little larger, sternites 6 and 7 divided posteriorly. Reference. CARVALHO, 1989b. Monophyly. Genus monophyletic, having the strongly dilated antenna as an autapomorphy (CARVALHO, 1989b). Geographical record. Chile, Argentina. Comments. The genus has only three species endemic to southern South America.
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Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Key to species (modified from CARVALHO, 1989b) 1.
General coloration metallic violet-blue; humeral callus yellowish; calypteres dark brown; interfrontal setae present in female (Chile) ......... .B.fulvohumeralis (Malloch, 1922) General coloration violet-blue; calypteres brownish with dark brown margins; interfrontal setae absent in female .......................................................................... .2
2.
Wing with Rs node bare on both surfaces; hind tibia on anterodorsal and anteroventral surfaces with at most 3 setae (frequently 2); prealar seta similar in length or much longer than notopleural anterior seta; sternite 1 bare; male: hind femur on posteroventral surface with long setae on apical half, shorter on basal half; female: fore tibia on dorsal surface without setae (Chile, Argentina) .............. .B. violaceiventris Macquart, 1851 Wing with Rs node with cilia on both surfaces; hind tibia on anterodorsal and anteroventral surfaces with at least 4 and 3 setae respectively; prealar seta absent in both sexes; sternite 1 setulose; male: hind femur on posteroventral surface bare on apical half and with longer setae on basal half; female: fore tibia on dorsal surface with 4 short setae on apical half (Chile, Argentina) ..................... .B. major Malloch, 1934
94
95
Figs. 94-95, characters of Brachygasterina (modified from CARVALHO, 1989b) 94, B. major, antenna, lateral view; 95, antenna, lateral view, B. violaceiventris; Abbreviations: ar, arista; ped, pedicel.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Chaetagenia Malloch, 1928 (Fig. 96) Type-species. Chaetagenia stigmatica Malloch (orig. des.). Diagnosis. Male dichoptic, eyes bare; facial ridges with short cilic" arista plumose, with long cilia on upper surface and short on lower; female with; directed setae on head; prealar seta present, moderately developed; scutellur;: on its lateral margins; prostemum with few setulae laterally; katepistemal posterior upper one, about twice the length of the others; metathoracic spiraclt: on its lower margin; anepimeron setulose; katepimeron with short and postalar wall setulose; lower calypter about twice as long as the upper one; on apical portion of stem vein and on veins R4+5 and CuA!; hind coxa with Cil::~lC)r surface; calcar absent; stemite 1 setulose. Reference. COURI & LAMAS, 1993. Monophyly. The bifid configuration ofthe distiphallus is very characl:(;":-l:siic and considered here as an apomorphy. This kind of aedeagus is also found in Pseudopri;o2epis Snyder, 1949 (LOPES & CARVALHO, 1985). Geographical record. Brazil. Comments. Monobasic genus: Chaetagenia stigmatica Malloch, 1928. COURI & LAMAS (1993) synonymized the other four described species with the type-species.
Fig. 96, Chaetagenia stigmata, head, lateral view
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Correntosia Malloch, 1934 Type-species. Correntosia bicolor Malloch (orig. des.). Diagnosis. Male dichoptic, eyes bare; arista pubescent, with cilia shorter than the basal width of the arista; flagellomere weakly dilated apically; interfrontal setae present in male; presutural acrostichals developed; prealar seta strong; wing veins bare; calcar present; abdomen elongated, longer than thorax length; ovipositor long, tergite 6 anchortype; tergite 7 a little larger than 6; stemite 8 entire; microtrichia absent. References. CARVALHO, 1989b; PONT, 2001. Geographical record. Argentina. Monophyly. Correntosia is close related to Palpibracus Rondani, but the presence of interfrontal setae in males is a rare character among the Muscidae, and it is not found in Palpibracus. Comments. This genus was for a long time monobasic (Correntosia bicolor Malloch, 1934), but recently PONT (2001) transferred Psilochaeta camifex Stein, 1911 to this genus. Differently from the type-species, C. camifex has orange antennal scape and pedicel, and yellow palpus (PONT, 2001).
Dalcyella Carvalho, 1989 Type-species. Helina veniseta Dodge (orig. des.). Diagnosis. Male holoptic, eyes with cilia; arista pubescent, with cilia shorter than basal width of the arista; flagellomere weakly dilated apically; interfrontal setae present in female; pre sutural acrostichals developed; prealar seta strong; anepimeron, katepimeron and meron setulose; vein R 4+5 with 2-3 cilia on both surfaces; calcar strong; abdomen oval, shorter than thorax length; ovipositor long, tergite 6 anchor-type; tergite 7 narrow; stemites 6 and 7 divided posteriorly; stemite 8 developed over its whole extension. Reference. CARVALHO, 1989c. Monophyly. DODGE (1967) described the type-species in Helina RobineauDesvoidy, because he could not relate it to any other muscid genera. Many species which would not conveniently fit Helina, have been assigned to it, and consequently this genus, as well other muscid genera (e.g. Phaonia), has been a depository of poorly studied species (SKIDMORE, 1985). Nowadays Helina is seen as an aggregate of distinct genera
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
(HUCKETT & VOCKEROTH, 1987). CARVALHO (1989c) erected Dalcyella for H. veniseta but no autapomorphic character was pointed out. Geographical record. Chile. Comments. Monobasic genus: Dalcyella veniseta (Dodge, 1967).
Itatingamyia Albuquerque, 1979 (Figs. 97-98) Type-species. Itatingamyia bivitatta Albuquerque (orig. des.). Diagnosis. Male dichoptic; antennal axis longer than oral axis; arista with short cilia, longest cilia shorter than twice of the basal width of pedicel; prealar and anterior intra-alar setae absent; dorsocentral setae 2: 3; acrostichals 0: 1, ciliform; pro sternum bare; anepimeron with 2-3 setae; katepisternals 1:2; posterior spiracle with cilia on margin; lower calypter glossiform, about as long as upper one; wing veins bare; veins R 4+s and M divergent at apex prosternum bare; sternite 1 setulose; female: ovipositor with cerci surpassing the hypoproct; tergite 6 anchor-type (Fig. 97), sternite 8 developed all its extension (Fig. 98). References. ALBUQUERQUE, 1979; CARVALHO & COURI, 1993. Monophyly. ALBUQUERQUE (1979) included the genus among the Mydaeinae, based mainly on the ovipositor shape. CARVALHO & COURI (1993), based on the shape of the female cerci (very long, surpassing the hypoproct), the development of sternite 8 and the shape on sternite 6, transferred the genus to Reinwardtiini, in the Azeliinae. Geographical record. Brazil. Comment. Monobasic genus: Itatingamyia bivitatta Albuquerque, 1979.
Muscina Robineau-Desvoidy, 1830 Type-species. Musca stabulans Fallen (des. COQUILLET, 1910). Diagnosis. Male holoptic, eyes bare; arista plumose with cilia on basal threefourths and bare on the apical fourth; interfrontal setae present in female; anterior intra-alar postsutural seta placed posteriorly to level of supra-alar seta; presutural acrostichals developed; prealar seta short, but distinct; anepimeron bare; meron setulose; wing with
80
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Claudio Jose Barros de Carvalho & Marcia Souto Couri
apical section of vein M slightly curved forward, veins bare; calcar strong, submedian; sternite 1 setulose. Reference. SNYDER, 1955. Monophyly. HENNIG (1965) had indicated the affinity of this genus with the Hydrotaeini based on the morphology of the ovipositor, while SKIDMORE (1985) placed this genus close to Synthesiomyia andPasseromyia based on characters of the immature stages. Geographical record. Mexico, Costa Rica, Guatemala, Peru, Venezuela, Brazil, Uruguay, Argentina, Chile; Nearctic and Palaearctic Regions. Comments. In the Neotropical Region, induding Mexico, there are four species, two of them occurring in South America.
Key to species 1.
Flagellomere dark brown ....................................................................................... 2 Flagellomere entirely or partly yellow...................................................................... 3
2.
Pedicel reddish-brown (Mexico; Nearctic and Palaearctic Regions) ........................ .
....................... .................. ................ ......... ................ ..... .M. levida (Harris, 1780) Apex of pedicel rather reddish; flagellomere lighter at base of pedicel (cosmopolitan)
.............................................................................. .....M. stabulans (Fallen, 1817) 3.
Pedicel and the longitudinal posterior half of flagellomere yellow; abdomen with greyish pruinescence (Mexico; Nearctic Region) ........................ M. dorsilinea (Wulp, 1896) Pedicel reddish and flagellomere rather reddish; abdomen reddish yellow translucent, last segment darker (South America) ........................ .M. principalis (Schiner, 1868)
Muscidae (Diptera) of the Neotropicai Region: TAXONOMY
81
Part I - Basal Groups
0,4mm 0,4 mm
97
98
Figs. 97-98, characters of ltatingamyia (modified from CARVALHO & COURI, 1993): 97, l. bivitatta, ovipositor, dorsal view; 98, I. bivitatta, ovipositor, ventral view.
Palpibracus Rondani, 1863 (Plate 1) Type-species. BrachypaZpus pilosus Macquart (aut.). Diagnosis. Male holoptic, eyes with cilia; arista pubescent, with cilia shorter than basal width of the arista; flagellomere weakly dilated apically or not; interfrontal setae present or absent in female; pre sutural acrostichals developed; prealar seta strong; katepimeron setulose or bare; wing veins bare; calcar present; abdomen oval, shorter than 82
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
thorax length; ovipositor long, tergite 6 anchor-type; tergite 7 thin (except in P. veneris); sternites 6 and 7 divided posteriorly (except in P. lancifer); sternite 8 developed over its whole extension. Reference. CARVALHO, 1989a. Monophyly. Probably monophyletic. CARVALHO (1989a) did not indicate putative synapomorphic characters for this genus, but pointed out its uniqueness among the muscids. Geographical record. Chile, Argentina. Comments. The species of Palpibracus have a very restricted distribution, occurring only in Chile and Argentina, in the Andes, southern of 30° S (CARVALHO, 1989a). LOPES & KHOURI (1996) described Palpibracus carvalhoi based on material from Chile. The type-material was not examined but, based on the original description, this species seems to be very close morphologically to P. univittatus (Bigot, 1857). PAMPLONA & COURI (2000) described P. valdiviensis from Valdivia, Chile, based only in female specimens. Based on their description it seems that it is close to P. apicalis (Malloch, 1934). Both species, P. carvalhoi and P. valdiviensis were not included in the key below.
Key to species (modified from CARVALHO, 1989a) 1.
Katepimeron setulose on posterior margin, with black cilia (sometimes yellow) in front of posterior spiracle ....................................................................................... 2 Katepimeron bare ................................................................................................ 8
2.
Antenna and palpus dark brown; mid tibia on posteroventral surface bare; male: head width similar to its height; eyes with anterointemal facets very enlarged; frontal setae limited to the anterior half offrons (Chile, Argentina) ............ P. veneris (Bigot, 1888) Antenna and palpus at least partly yellow; mid tibia on posteroventral surface with one or more setae; male: head wider than high; eyes with anterointernal facets not very enlarged; frontal setae over entire frons, those on upper frons weaker.. ............. 3
3.
Male: fore coxa without a strong apical spur; fore tibia on anteroventral surface with 1
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seta on apical third; hind tibia on posteroventral surface with a series of long setae; female: interfrontal setae absent; mid tibia on posteroventral surface with only 1 submedian seta (Chile ).................................................. F separatus (Malloch, 1934) Male: fore coxa with a strong apical spur; fore tibia on anteroventral surface with 2-3 setae on apical third; hind tibia on posteroventral surface bare, with small cilia on basal or apical half, never over its whole extension; female: interfrontal setae present; mid tibia on posteroventral surface with 1-3 submedian setae ................................ .4 4.
Male: parafaciallarge, its width, at the level of pedicel, similar to the width of gena at the lower margin of eyes; mid femur with no depression; female: frontal vitta totally reddish; fore tibia on anteroventral surface with 1-2 submedian seta (Chile, Argentina)
...................................................................... .............. F reynoldsi (Malloch, 1934) Male: parafacial, at the level of pedicel, about two-thirds as wide as gena at the lower margin of eyes; mid femur concave, anterior surface with a strong or faint median depression; female: frontal vitta dark brown sometimes with the anterior half reddish; fore tibia on anteroventral surface with only one submedian seta ...................... '" ...... 5 5.
Fore coxa entirely black; male: 10-11 pairs of frontal setae; mid femur on anterior surface without short spines; hind tibia on posterodorsal surface with 5 setae, the calcar submedian; female: frontal vitta dark brown (Chile, Argentina) ....................... .
....................................................................................... F similis (Malloch, 1934) Fore coxa yellow or at most dark brown on apical half; male: at most, 9 pairs of frontal setae (maximum); mid femur on anterior surface with 1-2 series of short spines apically; hind tibia on posterodorsal surface with 3-4 setae, the calcar inserted on the apical third; female: frontal vitta partly reddish ........................................................ 6 6.
Male: mid and hind femora yellowish; mid femur, on the median depression, without ground setulae; female: hind femur yellow, except for a black preapical ring; hind tibia on anteroventral surface with a series of 4-5 setae on apical half; posterodorsal surface with 2 setae on middle third (Chile) ................................... F chilensis (Bigot, 1885)
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Male: mid and hind femora partly dark brown; mid femur, on the median depression, with the usual ground setulae; female: hind femur yellow on the basal third and black on the apical two-thirds; hind tibia on anteroventral surface with a series of 5-6 setae on apical half; posterodorsal surface with 2-3 setae on middle third ......................... 7 7.
Male: fore tibia on anteroventral surface with 2 setae on apical half; ventral surface with 3-4 weak setae on apical third; posterodorsal surface with 0 or 2 setae on apical fifth; hind femur yellow, black on apical two-thirds; hind tibia black, anteroventral surface with a series of weak setae over its whole extension, longer toward the apex; posteroventral surface with cilia only on apical third, posterodorsal surface with 3-4 short setae evenly spaced along tibial length; mid tibia on posteroventral surface with 1-3 setae on apical half (Chile) ...................................... .l~ confusus (Malloch, 1928) Male: fore tibia on anteroventral surface with a series of very weak setae on apical half; posterodorsal surface with 1 median seta; hind femur yellow with a black ring on middle third; hind tibia yellowish; anteroventral surface with a series of weak setae along its whole length; posteroventral and ventral surfaces with cilia on apical half, longer apically; posterodorsal surface with 3 setae on middle third; female: unknown (Chile, Argentina) ............................................ ............P. pilosus (Macquart, 1851)
8.
General coloration metallic blue; abdomen, at most with tergite 5 partly orange yellow................................................................................................................... 9 General coloration yellow to black, thorax, at least, partly yellow; abdomen never metallic blue ........................................................................................................... 10
9.
Eyes with long and conspicuous cilia; legs black with tarsi partly yellowish white (more evident on fore leg of male); abdomen with the apical half of tergite 5 orange yellow; male: 12-16 pairs of frontal setae; fore coxa without an apical spur; fore tibia on anterior and anteroventral surfaces with fine, long cilia along its whole length; mid tibia on posterior surface with 4-5 setae on basal two-thirds; female: fore tibia on posterodorsal surface bare; mid femur on posteroventral surface with a series of setae
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on basal half (Chile, Argentina) ......................................... P. apicalis (Malloch, 1934) Eyes with short and sparse cilia; legs yellow, except for coxa, trochanter and tarsi black; abdomen entirely metallic violet blue; male: 8 pairs of frontal setae; fore coxa with a strong apical spur; fore tibia on posterior surface with long and strong setae along its length; female: fore tibia on posterodorsal surface with 2 short setae inserted at the base of the apical third; mid femur on posteroventral surface with one strong seta on basal third (Chile, Argentina) ............................... P. lancifer (Malloch, 1934) 10. Male: mesonotum yellow with a black median vitta; fore coxa with a strong apical spur; female: interfrontal setae present or absent; hind tibia on anteroventral surface with 3-5 setae on apical half. ................................................................................ 11 Male: mesonotum yellow with a black median vitta or black with grey pruinose vittae; fore coxa bare; female: interfrontal setae present; hind tibia on anteroventral surface with 2-3 setae on apical half.. ................................................................................ 13 11. Antenna dark brown, with scape, pedicel and base of flagellomere yellow; palpus yellow; male: hind tibia with a median curvature; anteroventral surface with a series of short setae, shorter than the width of tibia, over its whole length; posteroventral surface with a row of cilia on its whole extension, longer on apical fifth; female: frontal vitta yellow, a little darker posteriorly (Chile ) ........................... P. univittatus (Bigot, 1857) Antenna dark brown, pedicel a little lighter; palpus dark brown; male: hind tibia with or without a curvature; anteroventral surface with a series of long setae, longer than the width of tibia, over its whole length; posteroventral surface with a series of long and fine setae on apical two-thirds; female: frontal vitta dark brown, reddish in middle ornot. .................................................................................................................. 12 12. Gena orange yellow; mesonotum yellow with one narrow median vitta; male: mid tibia on posterodorsal surface with 2-4 setae evenly spaced along tibial length; hind tibia with no median curvature; female: interfrontal setae absent (Chile, Argentina)
.................................................................................... P. spicatus (Malloch, 1934) 86
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Gena dark brown; mesonotum yellow with one wide median vitta; male: mid tibia on posterodorsal surface with 4-6 setae evenly spaced along tibial length; hind tibia with a median curvature; female: interfrontal setae present (Chile)........... P. albuquerquei Carvalho, 1989 13. Palpus yellow with a brown base; mesonotum black with 3 grey pruinose vittae and small yellow areas; male: antenna inserted at level above the middle of eyes; female: abdomen dark yellow (Chile, Argentina) ..................... P. peruvianus (Malloch, 1929) Palpus dark brown; mesonotum yellow with 1-3 vittae, sometimes fused; male: antenna inserted at a level below the middle of eyes; female: abdomen dark brown .............. 14 14. Male: eyes with anterointernal facets strongly enlarged; fore tibia on anteroventral surface with 1 seta on middle third; ventral surface with no cilia on apical third; posteroventral surface with 1-2 setae on middle third; female: frontal vitta dark brown with only the anterior half reddish (Chile, Argentina) ..... P. nigriventris (Malloch, 1928) Male: eyes with anterointernal facets enlarged; fore tibia on anteroventral surface with 2-3 setae on apical third; ventral surface with 1-4 long cilia on apical third; posteroventral surface with 2-3 setae on middle third; female: frontal vitta reddish on almost all extension ................................................................................................ 15 15. Mesonotum yellow with a large black vitta (sometimes narrow in female); male: fore tibia on ventral surface with 4 long cilia on apical third; hind tibia with a median curvature, posteroventral surface with a series of fine cilia on middle third; female: mid tibia on ventral surface with one submedian seta; hind femur strongly curved and slightly enlarged preapically (Chile, Argentina) ......................................P.fasciculatus (Malloch, 1934) Mesonotum yellow with 3 black vittae (sometimes fused); male: fore tibia on ventral surface with 1-2 long median cilia; hind tibia without a median curvature, posteroventral surface with a series of strong cilia, absent on the basal two-thirds; female: mid tibia on ventral surface with no setae; hind femur only weakly curvature and not enlarged preapically (Chile )...................................................... P. trivittatus (Malloch, 1934)
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Philornis Meinert, 1890 (Figs. 99-108, Plate 2) Type-species. Philornis molesta Meinert (mon.). Diagnosis. Male holoptic (Fig. 100) or intermediate (Fig. 102); palpus a little enlarged apically; arista with long plumae; mesonotum usually with four brown vittae; anepimeron hairy; postalar wall with a tuft of cilia; posterior spiracle with cilia; dorsocentral setae 2:4; calcar absent; anal vein long; veins R 4+5 and M divergent, the latter a weakly bent forward apically (Fig. 99). References. COURI, 1989, 1999b. Monophyly. Genus certainly monophyletic having as sister-group Passeromyia with occurrence on the Afrotropical, Oriental and Australian Regions (COURI & CARVALHO, 2000). Several authors (e.g. PONT, 1974; SKIDMORE, 1985) had already called attention to the similarities among the two genera. Geographical record. Mexico, Costa Rica, Panama, Cuba, Dominican Republic, Trinidad, Puerto Rico, Venezuela, Ecuador, Peru, Brazil, Chile, Uruguay, Argentina; United States of America. Comments. P. cinnamomina (Stein), P. convexa (Stein), P. mima (Townsend), P. nielseni Dodge, P. obscura (Wulp), P. umanani Garcia, P. steini (Pont) and P. molesta Meinert were not included in the key, since we had access only to the data from the literature, which did not provide decisive species characters. Also, the last mentioned species is known only from one larval phase. Philornis species are largely distributed in the Neotropical Region, with three species known from the U.S.A., P. angustifrons (Loew, 1861), P. mimicola Dodge, 1968, and P. porteri Dodge, 1955, the latter known only from Florida (COURI, 1999b).
Key to species (modified from CO URI, 1999b) 1.
Vein R4+5 and/or CuA 1 with cilia on ventral surface and/or dorsal surface (Fig. 99); male: eyes approximated (Fig. 100); female: inner margin eye parallel (Fig. 10 1).......... 2 Veins bare; male: distance between eye greater than diameter of anterior ocellus (Fig. 102); female: inner eye margin converging above (Fig. 103) ....................................... 10
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2.
Gena yellow-haired ............................................ ,................................................. 3 Gena black-haired ................................................................................................ 4
3.
Wing slightly brown, calypteres uniforrnlybrown (Costa Rica) ........P schildiDodge, 1963 Wing hyaline, calypteres light yellow, except for brown margin of upper calypter (Brazil)
.................................................................................. ...... P setinervis Dodge, 1963 4.
Intra-alar setae 1:3 (Panama) ...... ................................ P rettenmeyeri Dodge, 1963 Intra-alar setae 1: 1 or 1:2 ........................................................................................ 5
5.
Mid femur on posterior surface with three preapical setae (Brazil) ........................ ..
...................................................................................... P amazonensis Couri, 1983 Mid femur on posterior surface with two preapical setae ......................................... 6 6.
Vein R4+5 with cilia only on ventral surface ................................................................ 7 Vein R 4+5 with cilia both on dorsal and ventral surfaces ........................................... 9
7.
Supra-alar setae 1:5; intrapostalar seta as long as posterior intra-alar one (Brazil)
..................................... .......................................................... P lopesi Couri, 1983 Supra-alar setae 1:3; intrapostalar seta shorter than posterior intra-alar one .............. 8 8.
Vein CuA! bare; vein R 4+5 on ventral surface with 2-3 cilia (Trinidad, Peru, Brazil)
.............................................................................................P aitkeni Dodge, 1963 Vein CuA! on ventral surface with 2 cilia; vein R 4+5 on ventral surface with 7 cilia (Brazil) .................................................. ..................... P rufoscutellaris Couri, 1983 9.
Vein R4+5 with 2 cilia on dorsal surface and 5 on ventral surface; vein CuA! on ventral surface bare (Mexico) ................................................. Pfasciventris (Wulp, 1896) Vein R 4+5 with 1-3 cilia on dorsal surface and 3-4 on ventral surface; vein CuA l on ventral surface with at least one cilia (Panama) ...................... .P zeketi Dodge, 1963
10. Gena black-haired ................................................................................................. 11 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Gena yellow-haired .............................................................................................. 15 11. Mid femur on posterior surface with 3 preapical setae (Trinidad, Brazil)
......................................................................... .... P downsi Dodge & Aitken, 1968 Mid femur on posterior surface with 2 preapical setae ......................................... 12 12. Hind tibia on anterodorsal surface with 2 setae on middle third; flagellomere dark brown (Argentina) ....................................................... ... P blanchardi Garcia, 1952 Hind tibia on anterodorsal surface with 1 setae on middle third; flagellomere yellow.... 13 13. Legs brown; tergite 5 reddish (United States of America.) ......P porteriDodge, 1955 Legs yellow with brown tarsi; tergite 5 brown with gray pruinescence....................... 14 14. Intrapostalar seta long, approximately 0.80 as long as posterior intra-alar one (Fig. 104); basicosta and epaulet brown (Guatemala, Panama, Peru, Brazil; United States of America) .................................................................... P mimicola Dodge, 1968 Intrapostalar seta ciliform; basicosta and epaulet yellow (Panama, Trinidad)
............................................................................... P nigra Dodge & Aitken, 1968 15. Hind tibia on anterodorsal surface no median setae or at most one seta.................. .16 Hind tibia on anterodorsal surface with more than one seta .................................. .19 16. Costal margin of wing with a brown elongated spot extending from middle of wing to apex (Fig. 105) (Peru, Brazil) ........................................... Pfomicosta Dodge, 1968 Wing hyaline ........................................................................................................ 17 17. Thorax light brown with one median brown vitae (Brazil) .......P univittata Dodge, 1968 Thorax brown with gray pruinescence and with four brown vittae .......................... .18 18. Mid tibia on posterior surface with 3 setae; intrapostalar seta ciliform; calypter light brown (Costa Rica, Panama, Trinidad, Ecuador, Brazil) .......P. falsifica Dodge & Aitken, 1968 Mid tibia on posterior surface with 2 setae; intrapostalar seta developed; calypter 90
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white (Brazil) .................................................... ................. P. petersoni Couri, 1984 19. Wing with dark brown spots on crossveins r-m and dm-cu (well marked in P. masoni, Fig. 106) .............................................................................................................. 20 Wing without brown spots on crossveins r-m and dm-cu ..................................... .21 20. Scutellum dark brown on sides, with extreme apex yellow; upper calypter with brown margin; fore tibia on dorsal surface with setae only at apex (Uruguay) ................ ..
...............................................................................................P. lnasoni Couri, 1986 Scutellum entirely brown; upper calypter yellowish; fore tibia on dorsal surface with 3-4 short setae on middle third (Argentina) .............................. P. seguyi Garcia, 1952 21. Wing light brown with brown clouds around longitudinal veins (Fig. 107) ............. 22 Wing without clouds, sometimes crossvein r-m surrounded by a brown spot.. ........ 25 22. Mid femur on posterior surface with 3 preapical setae; mid tibia on posterior surface with more than 3 setae on middle third (Mexico, Ecuador, Brazil) ............................. ..
............ ...............................................................................P. grandis Couri, 1984 Mid femur on posterior surface with 2 preapical setae; mid tibia on posterior surface with 1-3 setae on middle third .............................................................................. 23 23. Distance between eyes, at level of anterior ocellus, 0.20-0.22 of head width in male and 0.32-0.34 in female (Brazil) ................................ P. sabroskyi Albuquerque, 1957 Distance between eyes, at level of anterior ocellus, 0.06-0.09 of head width in male and 0.19-0.23 in female ....................................................................................... 24 24. Hind tibia on anterodorsal surface with 3 setae; calypteres white, except for brown margin of upper calypter (Brazil) ................................... P. obscurinervis Couri, 1984 Hind tibia on anterodorsal surface with 4 setae; calypteres light brown, margin of upper calypter dark brown and of lower one yellow (Costa Rica)
........................................................................................P. carinata Dodge, 1968 25. Hind tibia on anterodorsal surface with more than 5 setae .................................... .26 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Hind tibia on anterodorsal surface with at most 4 setae ............................................ 27 26. Legs yellow with brown tarsi in both sexes; calypteres light brown, margin of upper calypter brown; male: hind tibia not bowed (Mexico) .... ............... P bella Couri, 1984 Legs yellow in male and brown in female; calypteres white, margin of upper calypter brown; male: hind tibia distinctly bowed (Fig. 108) (Brazil, Argentina) .................. .
....... .............. ........... ........ ..... ...... ........ .............. ............... P torquans Nielsen, 1913 27. Distance between eyes, at level of anterior ocellus, of 0.03-0.05 of head width in males and of 0.09-0.14 in female ......................................................................... 28 Distance between eyes, at level of anterior ocellus, more than 0.05 if head width ........ 30 28. Basicosta brown; tergites 4 and 5 paler than thorax (Trinidad, Panama) ................ ..
...................................................... .............. ......... P querula Dodge & Aitken, 1968 Basicosta yellow; tergites 4 and 5 concolour with thorax ....................................... 29 29. Legs yellow in both sexes; mid femur on posterior surface with 3 preapical setae (Cuba, Costa Rica, Trinidad, Guyana, Brazil; United States of America) ..........
.................................................................................... P angustifrons Loew, 1861 Legs yellow in male and brown in female; mid femur on posterior surface with 2 pre apical setae (Trinidad, Brazil) ............................ .P. glaucinis Dodge & Aitken, 1968 30. Basicosta brown .................................................................................................... 31 Basicosta yellow.................................................................................................. 35 31. Wing without spots ............................................................................................. 32 Wing with brown spot at crossvein r-m ............................................................... 33 32. Abdomen dark brown with grey pruinescence; frons red at lunule; male: distance between eyes, at level of anterior ocellus, 0.10-0.12 of head width (Puerto Rico, Trinidad, Venezuela, Brazil) .............................. .P. deceptiva Dodge & Aitken, 1968 Abdomen brown with grey pruinescence, entire tergite 1+2 and basal half oftergite 3 92
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yellow; frons brown; male: distance between eyes, at level of anterior ocellus, 0.060.07 of head width (Costa Rica, Panama, Cuba, Dominican Republic, Puerto Rico, Trinidad, Venezuela, Brazil, Argentina) ............................... .P pici (Macquart, 1854) 33. Thorax light brown with yellow pruinescence; mid tibia on posterior surface with 3 setae (Trinidad, Brazil)....................................... P sanguinis Dodge & Aitken, 1968 Thorax brown with grey pruinescence; mid tibia on posterior surface with 2 setae ...... 34 34. Katepistemum with black cilia; hind tibia on anterodorsal surface with 2 setae (Brazil)
........................... ................................................................. P vulgaris Couri, 1984 Katepistemum with yellow cilia; hind tibia on anterodorsal surface with 3 setae (Brazil) ........................................................................................ p mediana Couri, 1984
35. Wing with black spot at crossvein r-m ................................................................... 36 Wing without a spot at crossvein r-m .................................................................. 38 36. Supra-alar setae 1:5; frons of both sexes with margins strongly diverging toward lunule (Brazil) .............................................. ................................. Pfrontalis Couri, 1984 Supra-alar setae 1:3; frons of both sexes with margins slightly diverging toward lunule .................................................................................................................. 37 37. Calypteres light brown, margin of upper one dark brown (Peru) .................. .
.................................................................................. .. P vespidicola Dodge, 1968 Calypteres white, margin of upper one brown (Peru) ...... .P albuquerquei Couri, 1983 38. Mid tibia on posterior surface with 3 setae; legs of both sexes yellow...................... 39 Mid tibia on posterior surface with 2 setae; legs of male yellow and of female brown (male of P spermophila unknown) ...................................................................... .40 39. Calypteres white, margin of upper one brown, greater ampulla brown, darker in male (Brazil) .............................................. ................................. P diminuta Couri, 1984 Calypteres white, greater ampulla yellow (Brazil) ...................P insularis Couri, 1983 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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40. Flagellomere brown (Trinidad, Brazil) .................P. trinitensis Dodge & Aitken, 1968 Flagellomere yellow............................................................................................. 41 41. Hind femur on anteroventral surface with 3 strong setae on apical third; hind tibia on anterodorsal surface with 3-4 setae (Trinidad) ......................... P. gagnei Couri, 1983 Hind femur on anteroventral surface with 4 strong setae on apical third; hind tibia on anterodorsal surface with 2 setae (Jamaica, Trinidad) .................. . ..........................................................................P. spermophila (Townsend, 1895)
1 mrn
1mm
Figs. 99-103. characters of Philornis (modified from co URI. 1999b): 99. P. aitkeni. wing. dorsal view; 100. P. aitkeni. male. head. frontal view; 101. P. aitkeni. female. head. frontal view; 102. P. downsi. male. head. frontal view; 103. P. dOlVnsi. female, head, frontal view.
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1 mm
1 mm
1 mm
r-m
1 mm
Figs. 104-108, characters of Phi/omis (modified from COURI, 1999b): 104, P. mimicola, mesonotum, dorsal view; 105, P. fumicosta, wing, dorsal view; 106, P. masoni, wing, dorsal view; 107, P. grandis, wing, dorsal view; 108, P. torquans, hind leg, lateral view. Abbreviations: ipal, intrapostalar seta; r-m, crossvein; dm-cu, crossvein.
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Psilochaeta Stein, 1911 (Plate 2) Type-species. Limnophora chlorogaster Bigot (orig. des.) = chlorogaster Wiedemann. Diagnosis. General coloration metallic violet blue, sometimes with greyish abdomen (P. chlorogaster); male holoptic; eyes with cilia; arista pubescent, with cilia smaller than the basal width of the arista; flagellomere slightly dilated apically; pre sutural acrostichal developed; prealar seta strong; katepimeron with few setulae; mesonotum with 3 dark vittae on dorsal surface; dorsocentral setae 2:3-4; acrostichals 2-3:3-6 but not forming lines; humerals 3; katepisternals 1-2:3-4; wing veins bare; calcar present; sternite 1 bare; female: ovipositor long; tergite 6 anchor-type; tergite 7 narrow; sternites 6 and 7 divided posteriorly. Reference. CARVALHO, 1989b. Monophyly. Psilochaeta was always associated to Brachygasterina Macquart, probably because of the same metallic coloration and, in part for their partly overlapping geographical distribution. CARVALHO (1989c) did not find autapomorphic characters for the genus. Geographical record. Colombia, Peru, Ecuador, Bolivia, Brazil, Uruguay, Argentina, Chile, Juan Fernandez Is. Comments. Among the endemic genera of Reinwardtiini in the southern South America, Psilochaeta has the largest known geographical distribution, with some important species for the antropobiocenosis (CARVALHO, 1989c). Psilochaetafulvolateralis Stein was not included in the key, since we could not exam its type. The holotype is probably deposited atSMT (PAPAVERO, 1971; CARVALHOetal., 1993). Psilochaeta carnifex Stein, 1911 (Chile) was recently transferred to Correntosia by PONT (2001).
Key to species (modified from CARVALHO, 1989c) 1.
Thorax dark brown with greyish pruinescence, with 4 brown vittae; abdomen metallic greenish, prealar seta shorter than the anterior notopleural seta; female: ovipositor longer than abdomen length (peru, Ecuador, Bolivia, Brazil, Uruguay, Chile, Argentina)
.........................................................................P. chlorogaster (Wiedemann, 1830) 96
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Thorax metallic violet blue, with 2 or 4 dark brown vitta:e; abdomen metallic blue to greenish blue; prealar seta similar to the anterior notoplemal in length; female: ovipositor shorter than abdomen length ................................................................................... 2 2.
Thorax with 4 dark brown vittae; notopleural setulae sparse; veins R4+5 and M parallel apically; male: distance between eyes shorter than flagellomere diameter; female: parafacial narrow, about half as wide as height of gena at lower margin of eyes (Brazil, Chile, Argentina) ................................... P. pampiana (Shannon & Del Ponte, 1926) Thorax with 2-4 dark brown vittae, more conspicuous presuturally; notoplemal setulae concentrated around posterior notoplemal seta; veins R4+5 and M divergent apically; male: distance between eyes similar to flagellomere diameter; female: parafacial wide, as wide as height of gena at lower margin of eyes .................................................. 3
3.
General coloration uniform; male: antennal axis longer than oral axis; antennal insertion below mid-level of eye; cercal plate with a large triangular, membranous area anteriorly and lower margin convex; paramere acute apically (Colombia, Brazil, Uruguay, . Argentina, Chile, Juan Fernandez Is.) .........................P. chalybea (Wiedemann, 1830) General coloration not uniform; greenish blue abdomen contrasting with mesonotum coloration; male: antennal axis shorter than oral axis; antennal insertion above midlevel of eye; cercal plate with a narrow triangular membranous area and lower margin almost linear; paramere obtuse apically (Chile) ................ P. violescens (Dodge, 1967)
Reinwardtia Brauer & Bergenstamm, 1889 Type-species. Reinwardtia tachinina Brauer & Bergenstamm (mon.) . Diagnosis. General coloration metallic blue; eye setulose; gena high; arista with very short cilia; prealar strong; katepimeron and anepimeron bare; katepisternals 1:2; calcar strong; wing veins bare; vein M strongly cmved forward to R4+5; lower calypter not very enlarged, about 2.5 times as long as upper calypter; abdominal setae strong; sternite 1 bare.
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References. BRAUER&BERGENSTAMM, 1889; HENNIG, 1965. Monophyly. The monophyly of the genus is not defined yet. HENNIG (1965) related this genus to Calliphoroides Malloch, which has only one described species from New Zealand. Geographical record. Venezuela, Ecuador. Comments. Monobasic genus: Reinwardtia tachinina Brauer & Bergenstamm, 1889.
Synthesiomyia Brauer & Bergenstamm, 1893 Type-species. Synthesiomyia brasiliana Brauer & Bergenstamm (orig. des.) =nudiseta Wulp. Diagnosis. Eye bare; arista with very short cilia; acrostichal presutural not differentiated; prealar strong; katepimeron and anepimeron bare; dorsocentrals 2:4; katepistemals 1:2; wing veins bare; vein M strongly curved forward apically; lower calypter very enlarged; stemite 1 setulose. Reference. HENNIG 1963a. Monophyly. The monophyly of the genus is not defined yet. S. nudiseta is closely related to Muscina based both on characters of the larvae and adults (SKIDMORE, 1985). Geographical record. Mexico, Nicaragua, Costa Rica, Panama, Virgin Is., Jamaica, Cuba, St. Domingo, Puerto Rico, Trinidad, Guyana, Venezuela, Colombia, Peru, Ecuador, Galapagos Is., Bolivia, Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is.; pantropical (SKIDMORE, 1985). Comments. Monobasic genus: Synthesiomyia nudiseta (Wulp, 1883). This species has been amply distributed throughout the world by human transport (HENNIG, 1965).
Subfamily Phaoniinae Malloch, 1917 Diagnosis. Calcar present or absent; anepimeron bare; female: ovipositor elongated; cerci free, surpassing the hypoproct; sternite 8 reduced. Monophy ly. HENNIG (1965) pointed out that the Phaoniinae did not constitute 98
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a monophyletic group. In CARVALHO's (1989d) cladistic analysis for the Muscidae, the subfamily showed no derived character. Helina Robineau-Desvoidy came out in a trichotomy at the base of the subfamily cladogram. Comments. The subfamily has 27 genera in the world, four of them in the Neotropical Region; 158 species are ascribed to this region.
Dolichophaonia Carvalho, 1993, stat. rev. (Figs. 109-120, Plate 2) Type-species. Phaonia brasiliensis Albuquerque (orig. des.). Diagnosis. Male holoptic; eye with short cilia; arista plumose; pre sutural acrostichals often differentiated; dorsocentral setae 2:3-4; prealar present, except in D. vockerothi, shorter than notopleural anterior seta; katepisternals 1:2; meron haired or not; wing veins bare; vein M parallel or very slightly forward-curved apically; calcar present, about twice as long as the basal width of hind tibia; female: clypeus, in lateral view, with a strong, hook-shaped anterior tip, posteriorly with a prominent sclerotization (Figs. 109118); ovipositor with large tergites and sternites (119-120). Reference. CARVALHO, 1993a. Monophyly. The monophyly of the genus was indicated by CARVALHO (1993a) based on the cladistic analysis of the family (CARVALHO, 1989d), where 2 characters of the female proboscis supported its monophyly. CARVALHO (1993a) also indicated the restrict geographical distribution, with only 2 species occurring in the United States of America, as an evidence for the monophyly of the genus. Geographical record. Mexico, Jamaica, Panama, Nicaragua, St. Vincent, Brazil, Argentina, Chile, Ecuador, Peru; south of United States of America. Comments. The synonymy between Dolichophaonia with Phaonia proposed by VOCKEROTH (1996) was not considered here, as he probably did not analyse the characters supporting the monophyly of Dolichophaonia indicated by CARVALHO (1993a). This is essentially a Neotropical genus with 27 species, two of them occurring in the southern of United States of America.
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Key to species (modified from CARVALHO, 1993a) 1.
Two proepimeral setae developed; mid tibia with three median posterior setae; male and female palpus filiform ...................................................................................... 2 One proepimeral setae developed; mid tibia often with 2 median posterior setae; female palpus more dilated than in male; if female palpus filiform, then tergite 5 bright yellow.................................................................................................................. 5
2.
Wing with or without a brown cloud on erossvein dm-eu; meron bare ..................... 3 Wing with two dark brown clouds on erossvein dm-eu; meron haired .................... ..4
3.
Dorsoeentral setae 2:4; mid and hind femora yellow apically; wing without clouds on erossvein dm-eu; male: eye densely haired (Chile )............ .D.femorata (Stein, 1911) Dorsoeentral setae 2: 3; mid and hind femora dark brown apically; wing with a cloud on erossvein dm-eu; male: eye with sparse hairs (Chile) ...... .D. regina Carvalho, 1993
4.
Palpus yellow; Dagellomere dark brown, except for yellow base (Mexico; United States of Ameriea) ........................................................ D. texensis (Malloch, 1923) Palpus dark brown apically; flagellomere entirely dark brown (United States of America) .......................................................... ........... D. limbinervis (Stein, 1918)
5. Sternite 1 setulose ................................................................................................ 6 Stemite 1 bare ..................................................................................................... 11 6.
Palpus yellow; hind femur with 2 preapical dorsal setae (Ecuador, Peru)
............................................................................ D. compressipalpis (Stein, 1911) Palpus light to dark brown; hind femur with 1 preapieal dorsal seta.......................... 7 7.
Dorsoeentral setae 2:3; both calypteres entirely whitish .......................................... 8 Dorsocentral setae 2:4; both calypteres or only the upper one with dark brown margins .................................................................................................................. 9
8. 100
Prealar strong, a little shorter than the anterior notopleural seta; wing yellowish, more Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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evident on costal margin (Peru) ............................... .D. sensitarsis (Carvalho, 1983) Prealar weak:, shorter than the anterior notopleural and similar to posterior notopleural; wing hyaline, with costal margin, from R j to apex, strongly dark brown (Brazil)
.......................................................................... ..... D. paranaensis Carvalho, 1993 9.
Upper calypter entirely yellowish; wing hyaline; cercal plate with a deep posterior incision (Brazil) ......................................... .D. santoamarensis (Albuquerque, 1958) Upper calypter yellowish with dark brown margins; wing hyaline with the costal margin yellowish; cercal plate with only a shallow posterior incision .................................... 10
10. Pa1pus yellow basally; tergite 5 dark brown, yellow apically; one prescutelar seta (Brazil) ............................................................. .D. catharinensis (Carvalho, 1983) Palpus entirely dark brown; tergite 5 light brown; 2 prescutelar setae (Brazil)
................................................................... ...... D. machadoi (Albuquerque, 1958) 11. Prealar absent (St. Vincent) ................................... .D. vockerothi (Carvalho, 1983) Prealar present, shorter than notopleural anterior seta .......................................... 12 12. One intra-alar postsutural seta; tergite 5 bright yellow (Jamaica) ....................... .
...................................................... ..................... D.jamaicensis (Carvalho, 1983) Two intra-alars postsutural setae; tergite 5 not as above ........................................ 13 13. Wing with inconspicuous clouds on crossveins; calypteres whitish; dorsocentral setae 2:3 ........................................................................................................................ 14 Wing with conspicuous clouds on crossveins; calypteres whitish to yellowish with dark margins or not; dorsocentral setae 2:3-4 ....................................................... 15 14. Palpus yellowish; flagellomere partly yellow, especially its base; lateral thoracic vittae interrupted at suture; calcar inserted on apical fifth of hind tibia (Nicaragua)
......................................................................... ............. D. trigonata (Wulp, 1896) Palpus dark brown; flagellomere dark brown; lateral thoracic vittae extending up to Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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middle of mesonotum; calcar inserted on apical fourth of hind tibia (Mexico)
......................................................................................... .D. unica Carvalho, 1993 15. Wing with a cloud on basal crossvein dm-cu and another on apex of vein R 1••••••••• 16 Wing without clouds on basal crossvein dm-cu or vein R1 ...................................... 17 16. Palpus light brown; calypteres whitish; dorsocentral setae 2:4 (Brazil) ............... .
............................................................................... D. anoctiluca (Carvalho, 1983) Palpus dark brown, except for yellow base; calypteres yellowish with dark brown margins; dorsocentral setae 2:3 (Brazil) ................. D. noctiluca (Albuquerque, 1958) 17. Wing with two clouds on crossvein dm-cu ........................................................... 18 Wing with one cloud on crossvein dm-cu ............................................................. 20 18. Upper calypter yellowish with dark brown margin; wing with costal margin yellowish (Brazil, Uruguay, Argentina) ...................... .D. trigona (Shannon & Del Ponte, 1926) Upper calypter whitish; wing without yellowish costal margin ................................. 19 19. Mid and hind femora yellowish; prealar seta a little shorter than anterior notopleural seta (Argentina) ......................................................... ... D. cacheuta (Snyder, 1957) Mid and hind femora dark brown; prealar seta strongly shorter than anterior notopleural seta (Argentina) ....................................................... .. D. catamacla (Snyder, 1957) 20. Meron haired below posterior spiracle; male: tergite 3 with discal setae (Brazil)
...................................................... .................. D. tachnoides (Albuquerque, 1958) Meron bare; male: tergite 3 without discal setae ..................................................... 21 21. Frontal vitta light brown (Brazil) ............................ D. gallicola (Albuquerque, 1958) Frontal vitta dark brown ...................................................................................... 22 22. Thorax light brown (Brazil) ................................ D. brasiliensis (Albuquerque, 1958) Thorax dark brown .............................................................................................. 23 102
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23. DorsocentraI setae 2:3; wing with only a inconspicuous cloud on crossvein dmcu ....................................................................................................................... 24 Dorsocentral setae 2:4; wing with a conspicuous cloud on crossvein dm-cu ............. .25 24. Palpus yellow; wing yellowish; acrostichal setae not differentiated; tergite 5 yellow at apex (Panama, Ecuador, Peru, Argentina) .................. .D. spontanea Carvalho, 1993 Palpus dark brown; wing hyaline with dorsal margin yellowish; presutural acrostichal setae differentiated; tergite 5 concolour with other abdominal tergites (Brazil) ........................................................................... .D. simplex (Albuquerque, 1958) 25. Wing hyaline without yellowish or dark brown costal margin; upper calypter whitish; post-humeral seta as long as anterior notopleura (Brazil) ................... . ....................................................... .......................... D. giacomeli (Carvalho, 1981) Wing hyaline with costal margin yellowish to dark brown; upper calypter yellowish, sometimes with dark brown margin; post-humeral seta shorter than anterior notopleural. .......................................................................................................... 26 26. Upper calypter uniformly yellowish; one prescutelar seta; mid femur yellowish (Brazil) ..............................................................................D. elongata (Albuquerque, 1958) Upper calypter yellowish with dark brown costal margin; two prescutelar setae; mid femur dark brown (Brazil) ...................................... .D. plaumanni (Carvalho, 1983)
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.>. :' ..... ):~:+
~ ;,
:,0';'
"
109
O,5mm
stn6
O,5mm
1,Omm
stn8
hyp
Figs. 109-120, female proboscis of Dolichophaonia, lateral view (modified from CARVALHO, 1993a): 109, D. anoctiluca; 110, D. brasiliensis; III, D. cacheuta; 112, D. catharinensis; 113, D. jemorata; 114, plaumanni; 115, D. santoamarensis; 116, D. simplex; 117, D. spontanea; 118, D. trigona; Figs. 119-120, D. regina, ovipositor; 119, ventral view; 120, dorsal view. Abbreviations: c1p, clypeus; hyp, hypoproct; pre, prementum; stn, sternite; spt, spermatheca.
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Helina Robineau-Desvoidy, 1830 (Plate 2) Type-species. Helina euphemioidea Robineau-Desvoidy (COQUILLET, 1901: 137) =pertusa Meigen. Diagnosis. Male hoi optic or dichoptic; eyes slightly pubescent; arista plumose; presutural acrostichal setae differentiated or not; dorsocentral setae 2:3; fore tibia without median setae; veins R 4+5 and M conspicuously divergent apically; pro sternum and anepimeron bare; wing veins bare; prealarpresent in male, fIliform; calcar absent; mesonotum with four dark vittae; abdomen usually without dark dorsal spots on tergites. Geographical record. Mexico, Jamaica, Guatemala, Costa Rica, Panama, Venezuela, Colombia, Peru, Ecuador, Bolivia, Brazil, Argentina, Chile, Falkland Is.; Nearctic, Palaearctic, northern parts of Oriental and Afrotropical Regions. References. MALLOCH, 1934; SNYDER, 1941. Monophyly. The genus is probably not monophyletic. Two subgenera are considered-Helina s.s. and Quadrularia Huckett, 1965 (see SKIDMORE, 1985). The genus is considered a repository of poorly studied species. See Monophy ly item under Phaonia. Geographical record. Mexico, Panama, Guatemala, Costa Rica, Jamaica, Colombia, Venezuela, Peru, Ecuador, Bolivia, Brazil, Argentina, Chile, Falklands Is.; Nearctic, Palaearctic, Afrotropical, northern Oriental and Australian Regions. Comments. Helina needs a thorough taxonomic revision. The keys for identification available in the literature (STEIN, 1911; MALLOCH, 1934; SNYDER, 1941) were all included here, with modifications (some species appear in more than one key). At the end of this section, there is a list of species not included in the keys, with their known geographical distribution. Helina cruciata Snyder, 1941 was included by SNYDER (1941) in his key (see below) because he thought this species might occur also in the Neotropical Region.
Key to species (modified from STEIN, 1911) 1.
Postsutural dorsocentral setae 3 ............................................................................ 2 Postsutural dorsocentral setae 4 ........................................................................... 10
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2.
Fore tibia with setae .............................................................................................. 3 Fore tibia without setae .......................................................................................... 5
3.
Wing with anterior margin dark brown (Bolivia) ..........H. nigromarginata (Stein, 1911) Wing with anterior margin not dark brown ............................................................. .4
4.
Pre sutural acrostichal setae differentiated (Peru) ...................H. biseta (Stein, 1904) Presutural acrostichal setae not differentiated (Peru) ..... H. echinogaster (Stein, 1911)
5.
General polinosity yellowish; antennae yellowish-white; legs light yellow (Panama, Peru, Bolivia, Brazil) ............................................... ..H. angustipennis (Stein, 1911) General polinosity grey; antennae dark brown or yellowish; legs never light yellow ................................................................................................................... 6
6.
Wing with anterior margin light brown (peru; Brazil) .....H. pedella (Wiedemann, 1830) Wing with anterior margin hyaline or, if margin pigmented, then prealar seta absent... .. 7
7.
Male dichoptic (Peru) .............................................. ....H. eurycephaZa (Stein, 1911) Male holoptic ....................................................................................................... 8
8.
Prealar seta present (Peru) .....................................H. consanguinea (Stein, 1911) Prealar seta absent. ............................................................................................... 9
9.
Dorsal dark vittae on mesonotum not well marked; palpus black (Peru, Brazil) .................................................................................H. nigrina (Wiedemann, 1830)
Dorsal dark vittae on mesonotum well marked; palpus yellow (Bolivia) ........................................................................................H. piliceps (Stein, 1911)
10. Fore tibia with setae (Peru, Brazil) ....................................H. tesselata (Stein, 1904) Fore tibia without setae (Peru) ............................................... H.fallax (Stein, 1911)
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Key to species (modified from MALLOCH, 1934) 1.
Metallic violet-blue species; calypteres and halter black; eyes conspicuously longhaired; arista almost bare; palpus black, strap-like; prealar seta over half as long as the one behind it; legs entirely black (Argentina) ....................H. viola Malloch, 1934 Black species, with more or less distinct grey or brownish-yellow dusting; calypteres and halter pale ........................................................................................................ 2
2.
Mid and hind femora entirely orange-yellow............................................................. 3 Mid and hind femora partly or entirely black ............................................................ 5
3.
All femora and tibiae entirely orange-yellow; male: hind femur with about 6 strong setae on apical half or more of anteroventral surface, unarmed on basal half of posteroventral surface (Chile)....... ...................... H. australis Carvalho & Pont, 1993 Fore femur almost entirely black; fore tibia usually with darkened base; mid and hind femora and tibiae orange yellow; male: hind femur with a greater portion of the anteroventral surface bristled ................................................................................. 4
4.
Hind femur in both sexes with long curved setae on apical half of anteroventral surface that decreases in length and strength toward the middle of femur; posteroventral surface without long setae; arista with its longest hairs about three times as long as its basal diameter (Chile, Argentina) ..............................................H. connexa Malloch, 1934 Hind femur in male with a complete series of anteroventral setae, weaker near base, and a series of long, slender setae on basal half of posteroventral surface; in female with a less extensive anteroventral series and fewer and shorter posteroventral setae; arista with its longest hairs hardly longer than its basal diameter (Chile, Argentina)
......................................................... ..................... H. nigrimana (Macquart, 1851) 5.
Postsutural dorsocentral setae 3 ............................................................................ 6 Postsutural dorsocentral setae 4 .............................................................................. 8
6.
Mid and hind femora broadly orange yellow at bases, blackened at apices; wing with
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crossveins distinctly with brown clouds, the inner one the more conspicuous, almost spot-like; abdomen densely grey dusted with a pair of small dark brown spots on tergite 3 and tergite 4 (Chile, Argentina) ............................... H. xena Malloch, 1934 Mid and hind femora blackened at bases; wing with crossveins not noticeably clouded; abdomen without evident dark paired spots on dorsum ......................................... 7 7.
Frons quite densely pale grey dusted, at narrowest point about 1.5 times as wide as flagellomere; orbits not bristled immediately in front of anterior ocellus; arista short pubescent; mid femur with apical third and hind femur with apical three-fourths brownish-yellow (Peru, Argentina) ....................................H. discolor (Stein, 1911) Frons entirely black, at narrowest point about three times as wide as flagellomere; orbits with setae on their entire extent, upper seta on each outcurved; arista short-haired, the longest hairs above, fully twice as long as basal diameter of arista; mid and hind femur with apical third reddish-yellow (Argentina) .............. ... .H. rufoapicata Malloch, 1934
8.
Mid tibia with a strong submedian seta on posteroventral surface; frons at narrowest point wider than flagellomere; prealar seta much less than one-third as long as the one behind it. ............................................................................................................... 9 Mid tibia with posterior but no posterodorsal submedian setae; other characters not as above in toto .................................................................................................. 11
9.
Arista with its longest hairs not longer than its basal diameter; mid tibia with 1-2 submedian anterodorsal setae (Peru, Argentina) .............. ....H. discolor (Stein, 1911) Arista with much longer hairs ............................................................................... 10
10. Mid tibia without submedian anterodorsal setae (Chile) ......H. chilensis Malloch, 1934 Mid tibia with 1-2 submedian anterodorsal setae (Chile, Argentina) ................ ..
......................... .............................................................. .H. simplex Malloch, 1934 11. Prealar seta less than one-third as long as the one behind it; hind femur without long setae on basal half of posteroventral surface; mid and hind femora orange-yellow to 108
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reddish yellow on at least the apical third ............................................................. 12 Prealar seta at least one-third as long as the one behind it, frequently much longer; hind femur entirely black or with fine long setae on at least basal half of posteroventral surface ................................................................................................................ 14 12. Mid and hind femora with less than their basal half blackened; male: frons at narrowest point about twice as wide as flagellomere (Chile, Argentina) ... .H. simplex Malloch, 1934 Mid and hind femora with more than their basal half blackened............................. 13 13. Male: frons at narrowest point hardly wider than flagellomere; orbits unarmed on upper third in front of ocelli (Chile, Argentina) ......H.juscocalyptrata Malloch, 1934 Male: frons at narrowest point about three times as wide as flagellomere; orbits bristled on entire extent (Argentina) ........................................H. ruJoapicata Malloch, 1934 14. Arista short haired, its longest hairs about five times as long as its basal diameter; hind femur in male black, with a faint yellowish-brown tinge at extreme apex, in female much more broadly fulvous-yellow, posteroventral surface in both sexes with no well developed erect setae on basal half; prealar seta over half as long as the one behind it; fore tibia rarely with a submedian posterior seta; calypter and bases of wing-veins fulvous-yellow (Chile )........................................................ H. bigoti Malloch, 1934 Arista pubescent, its longest hairs not or very little longer than its basal diameter; hind femur distinctly fulvous or reddish-yellow at apex and in male at least with a series of long fine erect setae at least on basal half of posteroventral surface; prealar seta less than half as long as the one behind it; calypter fulvous-yellow, but bases of wing veins fuscous (Chile, Argentina) ....................................... H. nigrimana (Macquart, 1851)
Key to species (modified from SNYDER, 1941)
Male 1. Strong postsutural dorsocentral setae 3............................................................ .2 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Strong postsutural dorsocentral setae 4 ................................................................ 12 2.
Costal margin, crossveins and apex of veins broadly infuscated; stem of halter at base of knob darkened; hind femur with slender, almost hair-like setae on at least basal half of posteroventral surface ........................................................................................ 3 Wings not so marked or exactly not as above ......................................................... .4
3.
Frontal vitta at narrowest part at most 1.5 times as wide as the diameter of anterior ocellus; when viewed in profile, antenna inserted opposite middle of eyes; flagellomere narrow not more than two and 2.5 times as long as pedicel (Colombia) ............ ..
............. ................................................................. .......... .. H. vierecki Snyder, 1941 Frontal vitta at narrowest part almost 2.5 times as wide as the distance across posterior ocelli, inclusive; when viewed in profile, antenna inserted opposite dorsal third of eye; flagellomere broad almost 4 times as long as pedicel (Colombia, Peru, Bolivia)
...................................................................... .......... H. marginipennis (Stein, 1904) 4.
One or two pairs of strongly developed pre sutural acrostichal setae ........................... 5 With at most a pair of well developed acrostichal setae which are never more than half as long and strong as the adjacent dorsocentral setae, or eyes not separated by a distance greater than diameter ocelli, inclusive ......................................................... 6
5.
Deep blackish pruinescent species with only the apices of mid and hind femora orange; hind tibia with three anterodorsal setae and three or four shorter posterior ones on middle (Argentina) ..................................................... H. ruJoapicata Malloch, 1934 Light yellowish-gray pruinescent species with at most only the fore femur indistinctly darkened; hind tibia with only one anterodorsal and no posterior setae (Ecuador)
............................................................... ...................... H. acrosticalis Snyder, 1941 6.
Longest hairs on arista shorter than or only as long as greatest diameter of arista........ 7 Longest hairs on arista at least half as long as the width of flagellomere ...................... 9
7. 110
Palpus, at least on basal two-thirds, pedicel and fore femur yellowish to fulvous Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Claudio Jose Barros de Carvalho & Marcia Souto Couri
(Ecuador, Peru) .......................................................... ;..... ..H. acrinis Snyder, 1941 Palpus, pedicel and fore femur black ..................................................................... 8 8.
Hind tibia with only one anteroventral seta which is situated on the apical fourth (Chile, Argentina) ................................................................. .........H. xena Malloch, 1934 Hind tibia with two or three anteroventral setae which are situated on the apical two thirds (?Mexico, Colombia, Peru) ...................................H. adelpha (Schiner, 1868)
9.
Hind tibia with one or more posterodorsal setae on at least basal half; crossveins very narrowly infuscated; abdominal stemites with a marginal fringe oflong setae (Mexico, North America) .............................................. ...............H. lasiosterna Snyder, 1941 Hind tibia without a basal posterodorsal seta; crossveins very broadly infuscated; abdominal stemites without long marginal setae ................................................... 10
10. Hind femur with at most only the anteroventral row of setae (Mexico, Costa Rica)
..................................................................................... H. refusa (Giglio-Tos, 1893) Hind femur with a complete row of anteroventral and posteroventral setae or long bristly hairs ............................................................................................................ 11 11. Presence of a conspicuous dark spot at base of discal cell below juncture of veins R 2+3 and R 4+5; crossveins clear (Brazil) ....................H. poeciloptera (Schiner, 1968) Absence of a conspicuous dark spot at base of discal cell below juncture of veins R4+5 and M; crossveins infuscated (Mexico ).....................H. signatipennis (Wulp, 1896) 12. All legs black or very dark brown ........................................................................ 13 At least some parts of the legs much lighter coloured............................................. 15 13. Margins of upper calypter darkened; hind femur without posteroventral hair-like setulae or setae (Ecuador) ............................................. .....................H. browni Snyder, 1941 Margins of upper calypter not darkened; hind femur with an almost complete row of posteroventral hair-like setae or setulae................................................................... 14
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14. Hind tibia with a series of very long slender posteroventral and ventral setae on basal three-quarters which are strongest at the base and become shorter toward apex (Peru) ................................................. ...............................H. longipila (Stein, 1918) Hind tibia without long hair-like posteroventral and ventral setulae on basal threequarters (Ecuador) .................................................... ........H. equator Snyder, 1941 15. With only one pre sutural acrostichal seta (Mexico )...............H. copiosa (Wulp, 1896) With one or more pairs of distinct presutural acrostichal setae which are always stronger than the adj acent acrostichal setae ....................................................................... 16 16. Vein M strongly bent posteriorly at apex; crossvein dm-cu strongly curved; both crossveins broadly infuscated (Peru) .....................................H. inepta (Stein, 1911) Vein M not strongly bent posteriorly at apex; other characters not exactly as above .................................................................................................................. 17 17. Mid tibia with a strong median posteroventral seta; frons distinctly wider than width of flagellomere; prealar less than one third as long as the seta behind it... ..... '" ............ 18 Mid tibia with no posteroventral setae or if present the other characters not exactly as above.,. .. " ......................................................................................................... 20 18. Longest hairs on arista not longer than its greatest diameter; mid tibia with one or two submedian anterodorsal setae (Peru, Argentina) ................... H. discolor (Stein, 1911) Longest hairs on arista distinctly longer than its greatest diameter............................ 19 19. Mid tibia with no anterodorsal setae; scutellar setulae descending below level of marginals (Chile) ........................... ...................................H. chilensis Malloch, 1934 Mid tibia with one or two anterodorsal setae; scutellar setulae not descending below level of marginals (Chile or Argentina) ................................H. simplex Malloch, 1934 20. All femora and tibia entirely fulvous (Chile )..........H. australis Carvalho & Pont, 1993 At least saIne parts offemora or tibia infuscated .................................................... 21
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21. Prealar less than one-third as long as first postsutural dorsocentral seta ................... 22 Prealar as long and strong as the first postsutural dorsocentral seta ....................... .25 22. Hind femur and tibia with long, slender posteroventral setae at base (Peru)
...................................................... ................................. H.longipila (Stein, 1918) Hind femur and tibia not so armed ........................................................................ 23 23. Frons at narrowest part equal to length of flagellomere; parafacial setae extending all the way to vertex (Argentina) .............................. .......H. ruJoapicata Malloch, 1934 Frons at narrowest part less wide as length of flagellomere; at most only slightly wider than distance across posterior ocelli, inclusive ........................................................ 24 24. Mid tibia without median anterodorsal setae; hind femur and tibia darkened only on basal fourth (Peru) ................................................. ...........H. townsendi Snyder, 1941 Mid tibia with one or two median anterodorsal setae; hind femur darkened on basal half and the hind tibiae darkened on basal two-thirds (Argentina) ............. .
................ ............................... ....................... ...... H. Julvocalyptrata Malloch, 1934 25. Longest hairs on arista fully one-half as long as the width of flagellomere; hind femur with anteroventral hair-like setae not as long as the diameter offemur at the portion in which they are situated; legs mostly blackish (Chile ) ............H. bigoti Malloch, 1934 Longest hairs on arista not longer than its greatest diameter; hind femur with anteroventral hair-like seta at least 1.5 as long as diameter offemur at the portion in which they are situated; legs mostly yellow........................................................... 26 26. Mid tibia with one or more short but distinct submedian anterodorsal setae; hind femur with a number oflong, hair-like posteroventral setae which are distinctly longer than diameter of femur at the portion in which they are situated (Chile, Argentina)
..................................................................................... .H. connexa Malloch, 1934 Mid tibia with no anterodorsal setae; hind femur without long, hair-like posteroventral setae (Chile, Argentina) ...........................................H. nigrimana (Macquart, 1851) Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Female 1. Strong postsutural dorsocentral setae 3................................................................. 2 Strong postsutural dorsocentral setae 4 ................................................................ 13 2.
Crossveins, costal margins and apex of vein R4+S and M broadly infuscated; no prescutellar acrostichal seta; prealar absent; halter black (Colombia, Peru, Bolivia) ...............................................................................H. marginipennis (Stein, 1904)
At most only the crossveins infuscated or not exactly as above ................................. 3 3. Hind tibia with one or more distinct posterodorsal setae on basal half or less .......... ..4 Hind tibia with no posterodorsal setae .....................................................................7 4. Frontal vitta with a pair of interfrontal setae; gena almost half as high as eyes; fore tibia without median posterior and anterodorsal setae; prealar almost as long and strong as posterior notopleural (North America; ?Neotropical Region) ....H. cruciata Snyder, 1941 Frontal vitta without interfrontal setae; gena less than one-fourth of eye height; fore tibia with median posterior or anterodorsal setae or with both .................................. 5 5. Pedicel mostly or entirely light yellow to orange (Mexico) ..... .H. parvula (Wulp, 1986) Pedicel entirely black or dark brown ...................................................................... 6 6. Hind tibiae with two or three posterodorsal setae; longest hairs on arista not quite as long as greatest width of flagellomere (Mexico )..................... .H. copiosa (Wulp, 1896) Hind tibia with only one posterodorsal seta on basal third or less; longest hairs on arista not over half as long as greatest width of flagellomere (Mexico) .............. .. ............................................................................................H. caneD Snyder, 1941
7.
Arista with hairs at most slightly longer than the greatest diameter of arista............... 8 Arista with hairs distinctly longer than greatest diameter of arista, usually at least half as long as greatest width of flagellomere ................................................................ 12
8. Thorax, abdomen and palpus largely or entirely yellow in ground colour; fore tibia 114
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with one median anterodorsal seta (Ecuador, Peru) ................H. acrinis Snyder, 1941 Thorax entirely and abdomen and palpus usually black; fore tibia without median anterodorsal seta ........................................................... 9.
0 .......................................
9
Mid tibia with two median anterodorsal setae (Ecuador) ...H. brevivena Snyder, 1941 Mid tibia without median anterodorsal setae ......................................................... 10
10. Without presutural acrostichal setae; crossveins at most only moderately infuscated (Mexico) ..................................................................... .......... H. socia (Wulp, 1896) With one or more pairs of short but distinct pre sutural acrostichal setae; crossveins very broadly infuscated ........................................................................................... 11 11. Mid and hind femora infuscated on apical third, the basal two-thirds orange-yellow; hind tibia with only one anteroventral seta which is situated on apical fourth (Chile, Argentina) ......................................................... ..................... H. xena Malloch, 1934 All femora almost entirely infuscated; hind tibia with one anteroventral seta situated on its middle (?Mexico, Colombia, Peru) .............................H. adelpha (Schiner, 1868) 12. Veins R J and R 2, crossveins and M at base of discal cell broadly infuscated; crossveins most distinctly so infuscated; katepisternals 1:2 (Brazil) ... H. poeciloptera (Schiner, 1968) Only the crossveins infuscated; katepisternals 2:2 (Mexico; Costa Rica)
.................. H. refusa (Giglio-Tos, 1893) and (Mexico) H. signatipennis (Wulp, 1896) 13. Metallic violet-blue non pruinescent species with entirely black calypter and halter (Argentina) ..................................................... .....................H. viola Malloch, 1934 Species not so coloured ....................................................................................... 14 14. Legs entirely black to dark brown ......................................................................... 15 Legs partly or entirely yellow to reddish .............................................................. 17 15. Margins of both calypteres and entire halter black; scutellar setulae descending well below level of marginals, especially at base of scutellum; abdomen distinctly bluish, Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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with sparse greyish pruinescence (Ecuador) ............................ H. notha Snyder, 1941 Margins of calypteres and entire halter not black; at most with one or two setulae descending below level of marginals on middle of scutellum. .................................. 16 16. Crossvein dm-cu joining the vein M at about the middle of the portion beyond the crossvein r-m; costal cell basad the vein R1 light brownish hyaline, scarcely darker than the lightest portion of wing membrane (Ecuador) ........ ....H. anubes Snyder, 1941 Crossvein dm-cu joining vein M well beyond the middle of the portion beyond cross vein r-m; costal cell before juncture basad vein R1 almost opaque, black, very much darker than the lightest portion of wing membrane (Ecuador) ........ H. equator Snyder, 1941 17. Mid tibia with a median posteroventral seta (Chile or Argentina) ........................... .
...................................................... ................................ H. simplex Malloch, 1934 Mid tibia without a submedian posteroventral seta............................................... 18 18. Mid tibia with two or more strong median anterodorsal setae ............................... 19 Mid tibia bare or with a single very weak median anterodorsal seta ........................ 21 19. Prealar as long as or longer than posterior notopleural seta; mid and hind femora entirely yellow (Chile, Argentina) ...................................H. connexa Malloch, 1934 Prealar at most half as long as the posterior notopleural seta ................................. 20 20. Thorax with dense, golden yellow pruinescence; all tibiae fulvous; without a strong pair of prescutellar acrostichal seta (Ecuador) ................. H. brevivena Snyder, 1941 Thorax with greyish pruinescence; only the hind tibia fulvous, others of legs black; with a pair of strong prescutellar acrostichal seta (Argentina) ............... ..
........................................................................... H.Julvocalyptrata Malloch, 1934 21. Vein M very strongly bent posteriorly at apex; crossvein broadly infuscated, dm-cu most broadly so, at its juncture with veins M and CuA 1; eye with short but distinct hairs (Peru) ........................................................................... H. inepta (Stein, 1911)
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Vein M subparallel with R4+5 at apex; cross veins not broadly infuscated ................ 22 22. Mid and hind femora entirely yellow (Chile, Argentina) ... .H. nigrimana (Macquart, 1851) Mid and hind femora mostly infuscated ................................................................. 23 23. Prealar distinctly longer than the posterior notopleural seta; mid tibia with four or five median posterior setae; fore and mid tibiae entirely fulvous (Chile) ............ ..
.......................................... ................................................. .H. bigoti Malloch, 1934 Prealar at most half as long as the posterior notopleural seta; mid tibia with two median posterior setae; fore and mid tibiae entirely infuscated (Ecuador) .............. .
.......................................................................................... .. H. anubes Snyder, 1941
Species of Helina not included in the key and their respective geographical distribution Species
Geographical distribution
H. albuquerquei Pont, 1972
Brazil
H. argentina Snyder, 1957
Argentina
H. auricolis Albuquerque, 1980
Brazil
H. circulatrix (Walker, 1861)
Mexico, Guatemala
H. crepedoseta Snyder, 1940
Mexico, Guatemala, Costa Rica, Panama
H. crocea Snyder, 1940
Brazil, Argentina
H. cyanicolor (Stein, 1911)
Bolivia
H. dasyophthalma Malloch, 1928
Peru, Bolivia
H. dasyops (Macquart, 1843)
Falkland Is.
H. dichroma (Wiedemann, 1830)
Brazil
H. discreta (Wulp, 1896)
Mexico, Jamaica
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H. etesia (Giglio-Tos, 1893)
Mexico
H. evecta (Harris, 1780)
Mexico, Venezuela; Nearctic, Palaearctic, northern parts of Oriental and Afrotropical Regions
lIS
H. Julvapoda Snyder, 1940
Mexico, Guatemala
H. gigantea Albuquerque, 1956
Brazil
H. grandis (Schiner, 1868)
South America
H. latipennis (Stein, 1904)
Colombia, Bolivia
H.leucocephala (Wulp, 1896)
Mexico, Panama
H. limbata (Wiedemann, 1830)
Brazil
H. luteola Albuquerque, 1956
Brazil
H. lyncii (Wulp, 1883)
Argentina
H. maculipes (Stein, 1918)
Mexico
H. meraca (Wulp, 1896)
Mexico; U. S. A.
H. monacha (Schiner, 1868)
South America
H. mulcata (Giglio-Tos, 1893)
Mexico
H. neosimplex Snyder, 1957
Argentina
H. neotropica Snyder, 1951
Peru
H. nivaloides Albuquerque, 1956
Brazil
H. nobilis Albuquerque, 1956
Brazil
H. parsura (Giglio-Tos, 1893)
Mexico
H. praecipua (Walker, 1853)
Brazil
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
H. procedens (Walker, 1861)
Mexico,U.S.A.
H. prolatifrons Snyder, 1940
Panama
H. regobarrosi Albuquerque, 1958
Brazil
H. rubripalpis (Wulp, 1896)
Mexico, Costa Rica, U.S.A.
H. sera (Giglio-Tos, 1893)
Mexico
H. tarsalis (Stein, 1918)
Mexico
H. trichops (Stein, 1918)
Peru
H. umbrosa (Wulp, 1896)
Costa Rica, Venezuela
H. walkeri Carvalho & Pont, 1993
Brazil, Paraguay
Phaonia Robineau-Desvoidy, 1830 Type-species. P haonia viarum Robineau -Desvoidy (des. COQUILLET, 1901 a: 140) = valida Harris. Diagnosis. Male dichoptic; eyes ciliated; arista plumose; dorsocentral setae 12:3-4; notopleuron with covering setulae and with 2 setae, the posterior one weaker; prealar seta present (absent in P. lentiginosa Snyder); lower calypter glossiform, Phaoniatype; Rs node bare or ciliated; vein M usually curved forward apically; hind tibia on posterodorsal surface with the calcar about as long as the width of the tibia at calcar insertion; female: ovipositor elongated, tubular, tergites narrow; stemite 8 reduced to 2 sclerites; microtrichia usually well developed only on the membrane; cerci free. Reference. COELHO, 2000. Monophy ly. Phaonia is a very heterogeneous genus with more than 300 species described in the world and it is certainly not monophyletic (HENNIG, 1965; ZINOVJEY, 1981; SKIDMORE, 1985; HUCKETT & VOCKEROTH, 1987; CARVALHO, 1989d). Except for the Neotropical Region, the species from other biogeographical regions have been segregated into several groups (seeZINOVJEV, 1981), howeverthe monophyly of none of these "groups" has been established. SKIDMORE (1985) considered Phaonia
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s.s. more closely related to subgenus Quadrularia of Helina than to the other species placed in Phaonia. Geographical record. Mexico, Guatemala, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Paraguay, Uruguay, Argentina; cosmopolitan. Comments: The species P costipennis (Snyder, 1957); P marginata Stein, 1918; P nervicincta Stein, 1918; P pteropsila Stein, 1918 andP tetragona Gaminara, 1930 were not included in this key. According to COELHO (2000), their original descriptions are not sufficient to recognise them. Besides this, their types have not been located and are probably destroyed.
Key to species (modified from COELHO, 2000) 1.
Wing on radial node with cilia on dorsal and/or ventral surface; dorsocentral setae 12:3-4 .................................................................................................................... 2 Wing on radial node with no cilia; dorsocentral setae 2:4 .......................................... 31
2.
Presutural dorsocentral seta 1............................................................................... 3 Presutural dorsocentral setae 2 .............................................................................. 5
3.
Prescutellar acrostichal setulae present; posterior spiracle with cilia on posteroventral margin; wing with dark clouds on apical portion of veins Sc and R}, apical half of~+3 and on crossveins r-m and dm-cu; female: faint cloud on apical portions ofR4+5 and M (Brazil, Peru, Bolivia) ............................................. P praesuturalis (Stein, 1904) Prescutellar acrostichal setulae absent; posterior spiracle bare on posteroventral margin; wing not as above ..................................................................................... 4
4.
General coloration brown to dark brown; fore tibia bare on anterior surface; male: tarsi with claws and pulvilli short, shorter than the length of tarsomere 5 (Brazil)
........................................................................... ....... P bigoti (Albuquerque, 1957) General coloration dark brown with tergite 1 and base oftergite 2 light brown; fore tibia with an anterior median seta; male: tarsi with claws and pulvilli similar to tarsomere 5 in length (Argentina) ................................................ P monochaeta Snyder, 1957 120
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5.
Postsutural dorsocentral setae 3 ............................................................................ 6 Postsutural dorsocentral setae 4 ........................................................................... .28
6.
Prealar seta absent (Brazil) ........................................ ...P. lengitinosa Snyder, 1957 Prealar seta present, similar to notopleural anterior seta in length ............................. 7
7.
General coloration light brown, at most, abdomen darker...................................... 8 General coloration dark brown to black, at most the scutellum yellowish brown; abdomen entirely dark brown or with tergites 1-3 light brown ............................... 14
8.
Thorax and abdomen light brown (Brazil) ..P. aurantica (Albuquerque & Medeiros, 1980) Abdomen dark brown, in contrast to thorax, or at most tergites 1+2 light brown ..... 9
9. Abdomen dark brown, with tergites 1+2 light brown (Brazil, Argentina) ...........................................................................P. shannoni Carvalho & Pont, 1993
Abdomen entirely dark brown ............................................................................. 10 10. Prescutelar acrostichal setulae present. .................................................................. 11 Prescutelar acrostichal setulae absent.. ................................................................. 13 11. Posterior spiracle with cilia on posteroventral margin (Brazil, Argentina) ....................................................................... P. nigriventris (Albuquerque, 1954)
Posterior spiracle bare on posteroventral margin .................................................. 12 12. Hind tibia with 4 median setae (Brazil) ...P. benjamini (Albuquerque & Medeiros, 1980) Hind tibia with 1-2- median setae (Brazil) ................ P. similata (Albuquerque, 1957) 13. Dark brown vitta on mesonotum reaching the row of intra-alar setae; male: ocellar setae strong (Brazil) ............................................... P. annulata (Albuquerque, 1957) Dark brown vitta on mesonotum not reaching the row of intra-alar setae; male: ocellar setae weak (Brazil) ........................................... P. grajauensis (Albuquerque, 1957) 14. Scutellum yellowish brown apically (Venezuela, Brazil, Argentina, Uruguay) Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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......................................................................................... P. trispila (Bigot, 1885) Scutellum concolour with thorax .......................................................................... 15 15. Posterior spiracle with cilia on posteroventral margin ............................................. 16 Posterior spiracle bare on posteroventral margin .................................................. 18 16. Male: tarsi with claws and pulvilli short, shorter than the length of tarsomere 5; female: ovipositor with microtrichia, except on tergite 6 (Peru, Brazil) .................................. ..
.................. ..... ............ ...... .............. ..... ................. ........ .... P. quercus Coelho, 1998 Male: tarsi with claws and pulvilli similar to tarsomere 5 in length; female: ovipositor not as above ........................................................................................................ 17 17. Wing with dark brown clouds only on crossveins r-m and dm-cu; male: paramere with a concavity on ventral surface; gonopod with the anterior region short and shorter than the width of paramere (peru, Bolivia, Brazil, Argentina) ................ P.latinervis (Stein, 1904) Wing with dark clouds on apices of veins R4+5 and M, another cloud extending from apices of Sc and R1 to apex of R2+3, and one cloud on crossveins r-m and dm-cu; male: paramere and gonopod not as above (Ecuador, Peru, Bolivia) ........................ .
.................................. ........................... ........................... .... P. maculosa Stein, 1911 18. Abdomen dark brown with tergites 1+2 and base oftergite 3light brown; male: tarsi with claws and pulvilli short, shorter than length of tars omere 5; female: ovipositor with microtrichia, except on tergite 6 (Panama, Brazil, Ecuador, Peru) .................. .
....... .......... ..... ........... ................. ....... ............... .............. P. hugonis Carvalho, 1989 Abdomen entirely dark brown; male: tarsi with claws and pulvilli long, longer than or similar in length to tarsomere 5; female: ovipositor not as above .............................. 19 19. Prescutelar acrostichal setulae present.. ................................................................ 20 Prescutelar acrostichal setulae absent. .................................................................. 27 20. Fore tibia with 1-2 anterior median setae ............................................................. 21
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Fore tibia bare on anterior surface ........................................................................ 24 21. Mid tibia bare on anterior surface ........................................................................ 22 Mid tibia with 1-2 anterior median setae ............................................................... 23 22. Wing with dark brown clouds only on crossveins r-m and dm-cu; male: distance between eyes, at level of anterior ocelli, about 0.26 of head width (Venezuela, Ecuador) ............................................................ ....... P. bipunctata (Schiner, 1868) Wing with dark brown clouds on apices of veins Sc, R] and M and on crossveins r-m and dm-cu; male: distance between eyes, at level of anterior ocellus, about 0.24 of head width (Colombia, Ecuador, Peru, Bolivia) ...................P. curvata (Stein, 1904) Hind tibia with two anterior and one posterodorsal sub-basal setae; dorsal surface with one strong apical seta (Peru, Boli via, Argentina) .......P. punctinervis Stein, 1911 23 Hind tibia with no anterior or posterodorsal sub-basal setae; dorsal surface without apical seta (Peru) .......................................................... .P. punoensis Coelho, 1998 24. Hind tibia with 1-2 sub-basal anterior setae ............................................................ 25 Hind tibia with no sub-basal anterior setae ........................................................... 26 25. Hind tibia with one submedian posterodorsal seta (Bolivia) .....P. soratiensis Coelho, 1998 Hind tibia with no submedian posterodorsal seta (Bolivia) .....P. nigripuncta Stein, 1911 26. Female: ovipositor with microtrichia only on membrane (Brazil, Ecuador, Peru)
..........................................................................................P. triseta (Curran, 1931) Female: ovipositor with microtrichia on membrane, tergites and sternites (Brazil)
......................................................................................... P. grandis (Couri, 1982) 27. Fore tibia with a median anterior seta (Mexico, Panama, Colombia, Ecuador, Bolivia)
...................................................................................P. abdita (Giglio-Tos, 1893) Fore tibia bare on anterior surface (Brazil; Bolivia; Argentina) ...P. advena Snyder, 1957 28. Fore tibia with 1-2 anterior median setae .............................................................. 29 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Fore tibia without anterior median seta .................................................................. 30 29. Hind femur on anterior surface with a preapical seta; hind tibia without an anterior sub-basal seta (Colombia, Ecuador) ............................. P. equatoralis Coelho, 1998 Hind femur on anterior surface bare; hind tibia with one anterior sub-basal seta (Bolivia)
......................................................................................P. boliviana Coelho, 1998 30. Apical portion of vein M strongly bent forward; male: gonopod with a long and curved anterior region, surpassing the paramere basally (Mexico, Ecuador, Peru, Bolivia) ............................................... ............................. P. venicurva (Stein, 1904) Apical portion of vein M slightly bent forward; male: gonopod not as above (Mexico) .................................................................. ...... P. mexala (Snyder, 1957) 31. Stemite 1 setulose ................................................................................................ 32 Stemite 1 bare ...................................................................................................... 34 32. Flagellomere entirely dark brown; calypteres yellow with dark brown margins; hind femur on anteroventral surface with setae only on apical half; meron with many setulae; male: paramere on ventral surface with a weak concavity; gonopod with the anterior region long, straight and not surpassing the width of the paramere (Mexico)
.....................................................................................P. nigerrima Carvalho, 1984 Flagellomere dark brown with a yellow base; calypteres whitish; hind femur on anteroventral surface with a complete series of strong setae; meron with few setulae ................................................................................................................ 33 33. Pedicel dark brown, yellow apically; wing hyaline; hind femur dark brown (Mexico) ............................................. ........................... P. robusta Carvalho, 1984 Pedicel entirely dark brown; wing yellowish; hind femur yellowish; male: paramere with the ventral surface almost straight, with a small cavity near its apex; gonopod with the anterior region long, straight and not surpassing the width of the paramere (Mexico) ................................................................ ....P. mexicana Carvalho, 1984 34. Arista with long plumae; cerci with a deep posterior incision .................................. 35 124
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Arista with few plumae; cerci not as above .......................................................... 36 35. Palpus dark brown, with a yellow base; humerals 3; fore tibia with 1-2 anteroventral submedian setae; male: paramere small; gonopod with the anterior region short (Mexico) .......................................................................... P. major Carvalho, 1984 Palpus entirely dark brown; humerals 2; fore tibia without an anteroventral submedian seta; male: paramere large; gonopod with the anterior region curved (Mexico)
........................................................................................ P. minuta Carvalho, 1984 36. Lower-posterior katepistemal seta weak; fore tibia without one anterior median seta; hind tibia on posterodorsal surface with a strong median seta, in addition the calcar (Ecuador) ..............................................................................P. ponti Coelho, 1998 Lower-posterior katepistemal seta strong; fore tibia with one anterior median seta; hind tibia on posterodorsal surface only with the calcar.. ....................................... 37 37. Hind tibia with one anterior submedian seta; male: paramere with a concavity on ventral surface (Ecuador) ...............................................P. carvalhoi Coelho, 1998 Mid tibia without an anterior submedian seta; male: paramere not as above (Ecuador) ......................................................................... P. hucketti Coelho, 1998
Species of Phaonia not included in the key and their respective geographical distribution Species
Geographical distribution
P. costipennis (Snyder, 1957)
Brazil
P. marginata Stein, 1918
Peru
P. nervicincta Stein, 1918
Paraguay
P. pteropsila Stein, 1918
Ecuador
P. tetragona Gaminara, 1930
Uruguay
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Souzalopesmyia Albuquerque, 1951 (Figs. 121-132) Type-species. Souzalopesmyia carioca Albuquerque (orig. des.). Diagnosis. Male dichoptic, frons narrower than in female, with reclinate orbital seta, and without interfrontal setae; ocellar setae absent; antenna long, reaching epistoma (Fig. 121); arista plumose, longest hairs as long as maximum antennal diameter; parafacial setulose on upper half; dorsocentral setae 1-2: 3; acrostichal setulae not distinct from ground setulae, except in female of S. amazonica; humerals 2; intra-alars 1:2; prealar absent; anepisternum with a short seta on upper anterior comer, meron bare; katepisternals 1:2 (not 1: 1: 1 as stated by ALBUQUERQUE, 1951); posterior spiracle small, triangular, with yellow setulae on its margin; prostemum bare; fore tibia with one posterodorsal submedian seta; fore tarsomere 1 with one ventral seta; mid femur with 1 anterodorsal, 1 dorsal, 1 posterodorsal and 1 posterior preapical seta; mid tibia with 2-4 posterior median setae, 1 strong ventral seta apically; hind tibia without calcar and with 1 anterodorsal median seta, and dorsal, anterodorsal and anteroventral surfaces each with one apical seta (Fig. 122); veins bare, except costa; vein M slightly curved forward apically (Figs. 123, 124); lower calypter of Phaonia-type; sternite 1 bare; ovipositor with long tergites; sternites and membranes covered with rnicrotrichia (Figs. 125, 126). Reference. CARVALHO, 1999. Geographical record. Peru, Bolivia, Brazil. Monophyly. Souzalopesmyia is a monophyletic genus based on the following synapomorphies: head elongated in lateral view; ocellar setae absent and general coloration yellow (CARVALHO, 1999). Comments. Souzalopesmyia species are rare and very similar to each other. The species, based on the known collected material, are found in forests and are probably nocturnal, since they have been very rarely collected during the day (CARVALHO, 1999).
Key to species (modified from CARVALHO, 1999) 1.
Inner vertical setae divergent (Fig. 121); crossvein dm-cu oblique, weakly curved (Fig. 124); female: 2 frontals (Brazil) ............................S. carioca Albuquerque, 1951 Inner vertical setae parallel; crossvein dm-cu oblique, almost straight (Fig. 123); female:
126
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3 frontals ................................................................................................................ 2 2.
Dorsocentrals setae 2:3; male: postocular row of setulae incomplete, not reaching epistoma; setulae black; female: some presutural acrostichal setulae stronger than the ground setulae (Brazil) ......................................... S. amazonica Albuquerque, 1951 Dorsocentrals setae 1: 3; male: postocular row of setae complete, setulae black and yellow, the latter beginning beyond basal half of eye; female: pre sutural acrostichal setulae undifferentiated from the ground setulae ........................................................ 3
3.
One proepistemal seta weak, about three-quarters as long as upper anepistemal setae; male: stemite 5 without sharp depression on posterior margin (Fig. 127); cercal plate heart-shaped (Fig. 128) (Brazil) ................................... S. paraensis Carvalho, 1999 One proepistemal seta strong, similar to the upper anepistemal setae; male: stemite 5 with sharp depression on posterior margin (Figs. 129, 130); cercal plate rounded (Figs. 131, 132) ................................................................................................... 4
4.
Species ranging from 8.0-9.0 mm in length; posterior katepistemal seta strong, about twice the length of the anterior one; male: both frontal setae cruciate; stemite 5 as in Fig. 129; cercal plate as in Fig. 131 (Peru, Bolivia) ........... S. singularis (Stein, 1919) Species ranging from 6.5 to 8.0 mm in length; posterior katepistemal seta very strong, about three times as long as the anterior one; male: lower frontal setae cruciate, upper reclinate; stemite 5 as in Fig. 130; cereal plate as in Fig. 132 (Brazil) .............. ..
................. ........... ..... ................ ................. .................... .S. sulina Carvalho, 1999
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
t
127
Part I - Basal Groups
~3 1 mm
1 mm
't', ••':.:::."
.•••
O,5mm
126
Figs. 121-126, characters of Souzalopesmyia (modified from ALBUQUERQUE. 1951): 121. S. carioca. head. frontal view; 122. S. carioca. hind leg. anterior view; 123. S. amazonica. male wing; 124. S. carioca. female; 125. S. carioca, ovipositor; 126, S. amazonica, ovipositor.
128
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Claudio Jose Barros de Carvalho & Marcia Souto Couri
129 130
Figs. 127-132, characters of Souzalopesmyia (modified from CARVALHO, 1999): 127, S. paraensis, sternite 5; 128, S. paraensis, cereal plate; 129, S. singularis, sternite 5; 130, S. sulina, sternite 5; 131, S. singularis, cereal plate; 132, S. sulina, cereal plate.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part I - Basal Groups
Table 1. Subfamilies, tribes and genera of basal Muscidae, including the number of species in Brazil, South America and the Neotropical Region. Subfamily
Genus
Tribe
Brazil
South
Neotropical
America Atherigoninae Muscinae
Muscini
Stomoxyini
Azeliinae
Azeliini
Reinwardtiini
Phaoniinae
Atherigona Biopyrellia Cyacyrtoneura Dasymorellia Morellia Musca Neomyia Neorypellia Parapyrellia Polietina Sarcopromusca Xenomorellia Haematobia Neivamyia Stomoxys Azelia Drymeia Hydrotaea Micropotamia Ophyra Potamia Thricops Brachygasterina Chaetagenia Correniosia Dalcyella ltatingamyia Muscina Palpibracus
I
I
I
I
I
I
I
2
7
9
14
17
1
I
19 2
I
I
I
I
I
I
2 13
2 15
2 18
2 2
2 2
2 2
I
I
I
3
4
4
1
1
I
0
1
I
0 3 4
0 7 3 6
8 4 7
0
2
2
0
0
I
0 I
3 I
3 I
I
I
2
I
I
I
I
I
I
I 4
17
I
I
0
2 17
Phi/om is
29
38
49
Psilochaeta Reinwardtia Synthesiomyia Dolichophaonia stat.rev. Helina
3
5
5
I
I
I
I
I
I
15
22
27
17
61
84
16
36
42
Phaonia Souwiopesmyia Total
I
1
37 genera
3
5
5
144
271
332
References (see end of Part II) 130
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
,
Claudio Jose Barros de Carvalho & Marcia Souto Couri ,,
Plate 1
•
•
•
•
Morellia humeralis (Stein, 1918)
Polietina nigra Couri & Carvalho, 1996
(6mm)
(7 mm)
•
Xenomorellia montanhesa Albuquerque, 1952
Palpibracus spicatus (Malloch, 1934)
(7 mm)
(5 mm) •
Muscidae (Diptera) ofthe Neotropical Region: TAXONOMY
131
•
Claudio Jose Barros de Carvalho & Marcia Souto Couri
Plate 2
•
•
I
..
..
Philornis trinitensis Dodge & Aitken, 1968 . (7 mm)
L Psilochaeta chlorogaster (Wiedemann, 1830) (6mm)
•
I I
Dolichophaonia gallicola (Albuquerque, 1958) (8mm)
Helina longipila (Stein, 1918) (8 mm) •
132
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
•
PART II
APICAL GROUPS By Marcia Souto Couri* & Claudio Jose Barros de Carvalho**
INTRODUCTION Part II of this book deals with the remaining three subfamilies of Muscidae Cyrtoneurininae, Mydacinae and Coenosiinae - considered as apical groups within Muscidae, in particular the two last subfamilies (CARVALHO, 1989d). As in Part I, the main scope of this section is to provide means of identifying the species included in these subfamilies. However, one of the difficulties to reach this goal is that some species could not be included in the identification keys, especially in the genera for which a thorough revision is needed. On the other hand, an indirect goal of this chapter, and for extension of this book as a whole, is to identify weak points in our knowledge of muscid taxonomic and diversity for the neotropics (e.g. taxonomic problems, need of generic revisions, need of more collecting in some groups with few known species, among others). CARVALHO et al. (1993) considered Cyrtoneurininae as a Muscidae subfamily, although recognising that' 'this is not a natural group, and several genera have already been transferred
* **
Museu Nacional do Rio de Janeiro, Departamento de Entomologia, Quinta da Boa Vista, Rio de Janeiro, 20.940040, Brazil; e-mail:
[email protected] Universidade Federal do Parana, Departamento de Zoologia, C.P. 19020, Curitiba, 81.531-980, Parana, Brazil; email:
[email protected]
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
to other subfamilies". Following this and considering the taxonomic changes and the new taxa described, from the last catalogue until now, the subfamily includes 11 genera and about one hundred species in the Neotropical Region. Cyrtoneuropsis Malloch and Neomuscina Townsend are, by far, the largest genera of Cyrtoneurininae, housing together more than 60% of the species of the subfamily. Three genera remain monobasic - Arthurella Albuquerque, Chortinus Aldrich and Neurotricha Shannon & Del Ponte. Some Cyrtoneurininae show a very interesting biology - e.g., Charadrella malacophaga Lopes breeds in dead gastropods (CARVALHO, 1980, 1985a). Viviparity has been recorded in some genera. The other two subfamilies seem to be close related, although this has not been clearly established. Mydaeinae has been traditionally treated as a separate subfamily, especially based on adult characters (CARVALHO, 1989d). SKIDMORE (1985), however, did not find characters from the larva morphology to "readily separate" them from the Reinwardtinae. Mydaeinae comprises seven genera, including Mydaea, a very speciose and poorly studied genus in South America. The Coenosiinae is the most speciose subfamily of Muscidae, housing over than 40% of all known muscid species in the Neotropical region. Two tribes are included in this subfamily - Limnophorini and Coenosiini. CARVALHO (1989d) provided evidence for the monophyly of the Coenosiini, although only two genera were included - Bithoracochaeta andNeodexiopsis. COURI & PONT (2000) proposed a worldwide classification for the genera of Coenosiini, using cladistic methodology. In their analysis, the monophyly of Coenosiinae was supported mainly by absence of a prealar seta, while the monophyly of Coenosiini was supported by the number and arrangement of the katepistemal setae (three setae forming an equilateral triangle), the dichoptic male frons, the presence of only one pair of reclinate orbital seta in both sexes, the presence of usually only one anterodorsal setae on hind tibia, and the elongated, tube-like hypandrium (COURI & PONT, 2(00). As a result of their analysis, some taxonomic changes were proposed, including the removal of six genera to tribe Lirnnophorini, four of them with occurrence in the neotropics - Agenamyia Albuquerque, Drepanocnemis Stein, Pachyceramyia Albuquerque and Rhabdotoptera Stein. Talking these changes into consideration, Coenosiini now comprises 16 genera, being Neodexiopsis Malloch the largest one and Limnophorini, 12 genera. About 43% of 28 genera of Coenosiinae found in South America are not represented in Brazil. The material and methods follow the Part I of this book, including the terminology and geographical distribution. 134
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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DIAGNOSIS AND KEY TO SPECIES The classification adopted here is the one proposed by CARVALHO et al. (1993), including the taxonomic updates and changes published posteriorly, especially the ones recently proposed by COURI & PONT (2000) on Coenosiini. Table 1 (page 259) summarizes the genera and species of the subfamilies Cyrtoneurininae, Mydaeinae and Coenosiinae occurring in Brazil, South America and the Neotropical Region.
Subfamily Cyrtoneurininae Snyder, 1954 Diagnosis. Male: dichoptic or holoptic; anepimeron setulose in most genera; hind tibia usually with no calcar; female: fronto-orbital seta absent. Monophyly. This is a polyphyletic subfamily (COURI & CARVALHO, 1997b). HENNIG (1965) stated that the Cyrtoneurininae could constitute a monophy letic group and several authors in the Neotropical Region followed his concept. However CARVALHO et al. (1993), despite including this subfamily in their catalogue for the Neotropical Region, did not recognised it as a natural group. Comments. After HENNIG's (1965) consideration about the monophyly of the Cyrtoneurininae, PONT (1972) included 15 genera in this subfamily but posteriorly some of them were excluded. Currently, there are 12 genera and 108 species in the Neotropical Region.
ArthurellaAlbuquerque, 1954 Type-species. Arthurella nudiseta Albuquerque (orig. des.). Diagnosis. Male holoptic; arista bare; pro sternum, anepimeron and me ron setulose; dorsocentrals 2:4; presutural acrostichals 2; intra-alars 2; katepisternals 2:2; vein R j with dorsal setulae on middle third; base of vein R 4+5 setulose on both surfaces; veins R 2+3 and R4+5 divergent apically; mid tibia with 4 setae on posterior surface; hind tibia with calcar; sternite 1 bare. References. ALBUQUERQUE, 1954a; LOPES, 1985a. Monophyly. Not yet established. Geographical record. Chile. Comments. Monobasic genus: Arthurella nudiseta Albuquerque, 1954. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
135
.~
Part II - Apical Groups
ARAUJO & COURI (1993) redescribed this species, including the description ofthe puparium and suggested that the species might belong in the Azeliinae, Reinwardtiini.
Cariocamyia Snyder, 1951 (Plate 1) Type-species. Cariocamyia maculosa Snyder (orig. des.). Diagnosis. Facial border with developed setulae that continue above the vibrissae up to the basal quarter of the antenna; male: strong pair of reclinate setae opposite to the anterior ocellus; female: interfrontal setae present; prosternum with numerous setulae over a large portion of the lateral area; proepisternum, anepimeron and suprasquamal ridge bare; meron with setulae only on the subtriangular portion above hind coxa; posterior thoracic spiracle with fine black hairs; vein R 2+3 setulose on both surfaces from node to midway to crossvein r-m; vein R4+5 with cilia on both surfaces from node to half of distance to crossvein r-m, strongly curved forward; lower calypter not truncate apically; mid femur with a short, subapical, anterodorsal seta; hind tibia without calcar and hind coxa bare above; female: sixth abdominal spiracle present Reference. SNYDER, 1951. Monophyly. One of the synapomorphies for Muscidae is the abdomen offemale showing less than 7 pairs of spiracles. This reduction in the number of spiracles is a unique apomorphy among the Calyptratae. Spiracles 6 and 7 were lost, but some genera, not related amongst themselves have reacquired the spiracle 6. Cariocamyia (see VOCKEROTH, 1972) has six spiracles, a condition acquired independently and considered apomorphic for the genus (see MICHELSEN, 1991). Only Achanthiptera Rondani, a genus apparently not phylogenetically related to Cariocamyia, has also reacquired the spiracle 6. This is a monobasic Palaearctic genus. Geographical record. Colombia, Brazil. Comments. Only two species occur in the region, both in South America.
Key to species 1.
Hind tibia on anteroventral surface with one seta (Colombia, Brazil) ........... .
...................................................................................... C. maculosaSnyder, 1951 136
Muscidae (Diptera) of the Neotropica1 Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
Hind tibia on anteroventral surface with two setae (Brazil) .............. .
...................................................... .............................. C. vibrissata (Stein, 1918)
Charadrella Wulp, 1896 (Figs. 1-9, Plate 1) Type-species. Charadrella macrossoma Wulp (mon.). Diagnosis. Male dichoptic; eyes with sparse, short and fme yellowish cilia; antenna long, reaching epistoma; arista plumose; the 2 upper pairs of frontal setae backwardly directed; dorsocentrals 3:3; acrostichals not differentiated; prealars 1 or 2; anepimeron setulose; proepimeron setulose; lower calypter enlarged reaching the area below scutellum; veins M and R4+5 divergent apically; calcar almost undifferentiated; sternite 1 setulose. References. CARVALHO, 1980, 1985a. Monophyly. Although the monophyly of C haradrella has not yet been established, a series of characters seems to support its monophyly, like the configuration of the head in both sexes and the shape of the lower calypter. Geographical record. Mexico, Panama, Trinidad, Tobago, Ecuador, Brazil. Comments. A very characteristic genus with three species in the Neotropical Region. Charadrella macrossoma, whose type-locality is in Yucatan, Mexico, has the largest geographical distribution within the genus, occurring from southern Mexico to Ecuador.
Key to species 1.
Male: femora yellow; fore and mid tibia light brown on apical third; sternite 5 triangular (Fig. 4); female: hypoproct rectangular (Fig. 5) (Brazil) ................ .
...................................................... ....................... C. albuquerquei Carvalho, 1985 Male: femora dark brown on basal third; fore and mid tibia dark brown on apical third; sternite 5 rectangular (Figs. 1,2); female: hypoproct triangular (Fig. 3) ............ .2 2.
Wmgs with a spot on apex of vein M (Fig. 6); male: cercal plate without median incision and with a prolongation on posterior margin (Fig. 7) (Mexico, Guatemala, Panama, Ecuador
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
Trinidad, Tobago, Venezuela, Guyana)................................ C. macrossoma Wulp, 1896 Wings without a spot on apex of vein M (Fig. 9); male: cereal plate with a deep median incision on posterior margin (Fig. 8) (Brazil) .... C. malacophaga Lopes, 1938
0,1 mm
0,1 mm
M
sur
0,1 mm
1 mm Figs. 1-9, characters of Charadrella (modified from CARVALHO, 1981, 1985a): I, C. malacophaga, male, sternite 5, dorsal view; 2. C. macrossoma, male, sternite 5, dorsal view; 3, C. malacophaga, female, hypoproct and cerci, dorsal view; 4, C. albuquerquei, male, sternite 5; 5, C. albuquerquei, female, hypoproct and cerci; 6, C. macrossoma, wing, dorsal view; 7, C. macrossoma, male, cerci and surstyli; 8, C. malacophaga, male, cerci and surstyli; 9, C. malacophaga, wing, dorsal view. Abbreviations: c pI, cereal plate (cerci); M, vein; sur, surstylus.
138
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
Chortinus Aldrich, 1932 Type-species. Chortinus bequaerti Aldrich (orig. des.). Diagnosis. Eyes with short and sparse cilia; male with enlarged anterointemal ommatidia; male holoptic; arista with short setulae; acrostichals 0: 1; dorsocentrals 2:4; intra-alars 0:2; prealar strong; scutellum bare on lateroventral margin; prostemum bare; propleuron bare; anepistemun, katepistemum and anepimeron with long cilia; veins R j and R4+5 with dorsal and ventral setulae; veins R4+5 and M almost parallel apically; stem vein on dorsal surface with short setulae on basal third; hind tibia on anteroventral, anterodorsal and posterodorsal surfaces with one short seta on apical third; female: ovipositor short. Reference. PAMPLONA & COURI, 1995a. Monophyly. PAMPLONA & COURI (199Sa) presented a key to segregate Chortinus from Cyrtoneurina, but did not indicate the monophyly of the genus. Geographical record. Guatemala, Costa Rica, Panama. Comment: Monobasic genus: Chortinus bequaerti Aldrich, 1932 (validated by PAMPLONA & COURI, 1995a).
Cyrtoneurina Giglio-Tos, 1893 (Fig. 10) Type-species. Cyrtoneurina uber Giglio-Tos (des. COQUILLET, 1910). Diagnosis. Male eyes with enlarged anterointemal ommatidia; arista with long cilia (12 to 19 cilia); dorsal secondary cilia short (Fig. 10), absent in C. arleriopsis; acrostichals 0: 1; dorsocentrals 2:4; intra-alars 0: 1; intrapostalar seta present; scutellum with setulae on lateroventral margin; prostemum bare or with a very fine lateral pubescence; propleuron bare; katepistemals 1:2; anepimeron setulose; vein R1 on ventral surface bare; vein R 4+5 with cilia on dorsal (except in C. costalis) and ventral surfaces (except in C. costalis and C. crispaseta); ovipositor long, epiproct with 2 setae, tergites slender. Reference. PAMPLONA, 1999. Monophyly. According to PAMPLONA (1999), Cyrtoneurina is monophyletic based on the following synapomorphies: presence of microtrichia on the membrane between the segments 6-7 and 7 -S. Geographical record. Mexico, Belize, Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Grenada, Trinidad, Colombia, Venezuela, Guyana, Peru, Brazil. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
Comments. COURI & PAMPLONA (2000) reevaluated the position of C alifusca Couri, 1982 and kept it in this genus.
Key to species (modified from PAMPLONA, 1999) 1.
Vein R] with dorsal cilia; entire palpus yellow......................................................... 2 Vein R] bare on dorsal surface; palpus brown, at least on basal half.. ....................... 3
2.
Hind femur on anterodorsal surface with a series of setae; anteroventral surface with 2 setae on apical third; posterodorsal surface with one preapical seta (Brazil) .......................................................................................... C. alifusca Couri, 1982 Setae on hind femur distributed not as above (peru, Brazil) .... C crispaseta Snyder, 1954
3. Vein R 4+5 bare on ventral surface; hind tibia on middle third of posterodorsal surface without setae (Brazil) ....................................................... C costalis (Walker, 1853) Vein R 4+5 with cilia on ventral surface; hind tibia on middle third of posterodorsal surface with one seta ............................................................................................. 4 4.
Prealar seta absent; hind femur on basal third of ventral surface with one strong seta with round apex (?Guatemala or Panama - La Pita) ........ C geminata (Stein, 1904) Prealar seta present; hind femur on basal third of ventral surface without one strong seta with round apex ............................................................................................... 5
5.
Arista with up to 12 cilia; secondary dorsal cilia absent; palpus with apical half yellow and basal half brown (Brazil) ........................................... C arleriopsis Couri, 1982 Arista with more than 12 cilia; secondary dorsal cilia present; palpus entirely brown ....... 6
6.
Body length about 4.0 mm; mid tibia on basal half of posterior surface with a complete series of short setae (Brazil) .................................................. C biseta Snyder, 1954 Body length about 5.0 to 6.0 mm; mid tibia on basal half of posterior surface without a series of short setae ............................................................................................ 7
140
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Marcia Souto Couri & Claudio Jose Barros de Carvalho
7.
Scutellum on basal half of lateroventral margin with up to 5 cilia; prostemum bare; arista with about 15 cilia (Costa Rica, Panama, Brazil) ......... c. confusa Snyder, 1954 Scutellum on basal half oflateroventral margin with more than 5 cilia; pro sternum with a very fine and inconspicuous pubescence; arista with about 19 cilia (Mexico, Guatemala, El Salvador, Belize, Nicaragua, Panama, Grenada, Trinidad, Colombia, Peru, Venezuela, Guyana) .................................................... c. uber Giglio-Tos, 1893
Species of Cyrtoneurina not included in the key and its respective geographical distribution Species
Geographical distribution
C. cylindrica (Macquart, 1846)
Brazil
O,5mm
10 Fig. 10, Cyrtoneurina uber (modified from PAMPLONA, 1999), antenna.
Cyrtoneuropsis Malloch, 1925 (Figs. 11-13) Type-species. Spilogasterveniseta Stein (orig. des.). Diagnosis. Male eye with enlarged anterointernal ommatidia; arista with long cilia (18 to 29 cilia, except C. seriata with 9, C. ocasionalis with 10 and C. spiloptera with 14); dorsal secondary cilia short or long (Fig. 11); acrostichals 0: 1; dorsocentrals 2:4 (2:3 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
in C. gluta and C. spiloptera); intra-alars 0: 1; intrapostalar seta absent or present; scutellum with many cilia on latera-ventral margin (few cilia in C. gluta, C. maculipennis and C. walkeri); prosternum setulose or bare, with fine lateral pubescence (c. neotrita and C. ocasionalis); propleuron bare or setulose in C. beebei, C. multomaculata and C. neotrita; katepisternals 1:2; anepimeron setulose; veins R] (except in C. brunnea and C. paUipes) and R4+5 with cilia on dorsal and ventral surfaces; ovipositor short or medium sized, tergites slender or large; sternite 8 with or without strong setae with round apex. References. PAMPLONA, 1999; PONT & PAMPLONA, 2000. Monophyly. According to PAMPLONA (1999), the genus is monophyletic and supported by five synapomorphies: presence of ventral cilia on vein R]; medium to short length of the non-delicate ovipositor; cerci short and flattened; tergites 6 and 7 very sclerotized and hypoproct with a lateral incision. Geographical record. Mexico, Belize, EI Salvador, Guatemala, Nicaragua, Costa Rica, Panama, Trinidad, Cuba, Puerto Rico, Colombia, Venezuela, Guyana, Suriname, Ecuador, Bolivia, Brazil, Argentina. Comments. PONT & PAMPLONA (2000) synonymized Paracyrtoneuropsis Pamplona under Cyrtoneuropsis. COURI & PAMPLONA (2000) transferred Cyrtoneurina brunnea and Cyrtoneurina pallipes to this genus and they are included in the key below.
Key to species (modified from PAMPLONA, 1999) 1.
Male: prealar seta absent or, if present, only slightly longer than ground setulae (Fig. 12); female: interfrontal seta absent... ..................................................................... 2 Male: prealar seta present and strong; female: interfrontal seta present... .................. 17
2.
Fore tibia with one seta on middle third; dorsocentrals 2:3 (Mexico, Guatemala, Guyana, Costa Rica, Panama, Canal Zone, Venezuela) ....... c. gluta (Giglio-Tos, 1893) Fore tibia without one seta on middle third; dorsocentrals 2:4 (the posterior one very short in C.fuscicosta) ............................................................................................ 3
3. 142
Pro sternum setulose ................................................................................................ 4
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
Prosternum bare .................................................................................................. 15 4.
Male ................................................................................................................... 5 Female .............................................................................................................. 11
5.
Hind tibia on ventral surface, at least, with one seta ................................................ 6 Hind tibia on ventral surface without a long submedian seta .................................... 7
6.
Hind tibia on ventral surface with one long submedian seta (Panama, Trinidad, British Guyana, Brazil) ................................................................ c. armipes (Stein, 1911) Hind tibia on ventral surface with three setae on middle third (Paraguay) ........................................................................................ C. paUipes (Stein, 1918)
7.
Wings without a brown cloud between apex of veins Sc and R J ; hind tibia on posterior to posteroventral surface with at most one seta ...................................................... 8 Wings with a brown cloud between apex of vein Sc and R J (sometimes faint); hind tibia on posterior to posteroventral surface with long setae or cilia ........................... 9
8.
Katepisternum with short and sparse ground setulae (Fig. 13); hind tibia on posterior surface with a series of setae (Mexico, Guatemala, Belize, El Salvador, Costa Rica, Panama, Colombia, Peru, Venezuela, Brazil, Paraguay) ............... c. gem ina (Wiedemann, 1830) Katepisternum bare; hind tibia on posterior surface without a series of setae (Guatemala, Belize, Nicaragua, El Salvador, Costa Rica, Panama, Grenada, Trinidad, Colombia, Peru, Venezuela, Guyana, Brazil?) ................................... C. immunda (Stein, 1911)
9.
Katepimeron with one seta and one setula; pal pus yellow; arista with about 9 cilia (Colombia, Peru, Brazil) ..................................................... C. seriata (Stein, 1911) Katepimeron bare; palpus brown, at least basally; arista with more than 20 cilia.......... 10
10. Hind tibia on anterodorsal surface with 2 setae on middle third; dm-cu straight; pro sternum bare (Peru, Guyana, Brazil) ............................. C. conspersa (Stein, 1911) Hind tibia on anterodorsal surface with 1 seta on middle third; dm-cu sinuose; Muscidae (Diptera) of the Neotropical Region: TAXONOMY
143
Part II - Apical Groups
prostemum setulose (Brazil) ................................... C.jlaviantennata (Couri, 1982) 11. Wings without a brown cloud between apex of veins Sc and R J ................................ 12 Wings with a brown cloud between apex of veins Sc and R J ( sometimes faint) ............ 13 12. Vein R J with cilia (Mexico, Guatemala, Belize, Salvador, Costa Rica, Panama, Colombia, Peru, Venezuela, Brazil, Paraguay) ..............................
c. gemina (Wiedemann, 1830)
Vein R J without cilia (Paraguay, Brazil) ................................... C. paZZipes (Stein, 1918) 13. Mesonotum, when viewed posterolaterally, with an conspicuous golden pruinescence, and without vittae (Peru, Guyana, Brazil) .......................... C. conspersa (Stein, 1911) Mesonotum, when viewed posterolaterally, with grey pruinescence, and with vittae .... 14 14. Arista with 8.bout 9 cilia; prealar seta absent; fore coxa and tibia brown (Colombia, Peru, Brazil) ..........................................................................
c. seriata (Stein, 1911)
Arista with about 20 cilia; prealar seta present; fore coxa yellow and fore tibia yellowish on basal half (Panama, Trinidad, Guyana, Brazil) ................
c. armipes (Stein, 1911)
15. Wings with brown clouds from base to apex of costal vein, on crossveins r-m and dm-cu (Guyana, Brazil) .................................................. C.juscicosta (Curran, 1934) Wings without brown clouds on costal vein, cross veins i·-m and dm-cu .................... 16 16. Presutural dorsocentral setae with similar length; male: hind tibia on ventral to posteroventral surface with one long submedian seta and with a series of shorter setae (Mexico, Guatemala, El Salvador, Belize, Panama, Colombia) ....................................
c. steini (Snyder, 1954)
Anterior presutural dorsocentral seta longer than the second; male: hind tibia on ventral to posteroventral surface without a long submedian seta and without a series of shorter setae (Mexico, Guatemala, Belize, El Salvador, Costa Rica, Panama, Colombia, Venezuela, Peru, Brazil, Paraguay) .............................
c. gemina (Wiedemann, 1830)
17. Prosternum setulose ............................................................................................ 18 Prostemum bare or with a very fine and inconspicuous pubescence .......................... 23 144
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Marcia Souto Couri & Claudio Jose Barros de Carvalho
18. Postsutural dorsocentrals 3 (Belize, Panama, Suriname, Ecuador, Peru, Bolivia, Brazil)
........................................................................... c. spiloptera (Wiedemann, 1830) Postsutural dorsocentrals 4 ................................................................................. 19 19. Propleuron bare .................................................................................................. 20 Propleuron ciliated ................................................................................................ 22 20. Stem vein bare; anepimeron ciliated only posteriorly (Panama) ...................... .
...................................................... ................................. C. mimica (Snyder, 1954) Stem vein ciliated; anepimeron entirely ciliated...................................................... .21 21. Vein R J with cilia after the humeral crossvein (Guatemala, Belize, Costa Rica, Panama, Peru, Guyana, Brazil) ............................................... ...........
c. dubia (Snyder, 1954)
Vein R J without cilia after the humeral crossvein (Brazil; Argentina) ............... . ..................................................................................... C. brunnea (Hough, 1900) 22. Lateroventral margin of scutellum with many short cilia reaching its apex; stem vein ciliated on ventral surface at level of humeral vein (?Costa Rica, Colombia, Peru, Guyana, Brazil, Bolivia) ............................................ c. multomaculata (Stein, 1904) Lateroventral margin of scutellum with many short cilia restricted to its basal half; stem vein bare on ventral surface at level of humeral vein (Brazil) .................. .
.................................................................................. C. protosetosa (Snyder, 1954) 23. Propleuron ciliated ............................................................................................. 24 Propleuron bare .................................................................................................. 25 24. Hind tibia on posterodorsal surface with one seta (Peru) ..... c. neotrita (Snyder, 1954) Hind tibia on posterodorsal surface without setae (Trinidad, Guyana) .......... .
.................................................................... ..................... C. beebei (Curran, 1934) 25. Postsutural dorsocentrals 3 (Brazil) ..................................... C. walkeri (Pont, 1972) Postsutural dorsocentrals 4 ................................................................................ 26 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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26. Hind coxa ciliated on posterior surface (Mexico, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Peru, Bolivia, Brazil, Argentina)
...................................................... .................................. C. veniseta (Stein, 1904) Hind coxa bare on posterior surface ....................................................................... 27 27. Prealar setae 2; arista with about 10 cilia (Brazil) ............. C ocasionalis (Couri, 1982) Prealar setae 1; arista with more than 16 cilia ...................................................... 28 28. Stem vein with cilia on ventral surface (Costa Rica, Panama, Guyana, Bolivia, Brazil)
...................................................... ..................... C. maculipennis (Macquart, 1843) Stem vein bare on ventral surface ........................................................................... 29 29. Stem vein without cilia on dorsal surface ................................................................ 30 Stem vein with cilia on dorsal surface ................................................................... 37 30. Wings without clouds on apex of vein R 2+3 ; hind femur on posteroventral to ventral surface with, at least, 2 strong basal setae ............................................................. 31 Wings with a brown cloud on apex of vein R 2+3; hind femur on posteroventral to ventral surface without strong basal setae ............................................................. 32 31. Pedicel brown; legs dark brown (Brazil) ....................... C pararescita (Couri,1995) Pedicel yellow; legs light brown (Mexico, Guatemala, Belize, Nicaragua Costa Rica, Panama, Cuba, Puerto Rico, Trinidad, Colombia, Ecuador, Venezuela, Guyana, Peru, Bolivia, Brazil, Paraguay) ............................................. ...... C rescita (Walker, 1861) 32. Halter yellow at least at knob .............................................................................. 33 Halter brown, with knob darker........................................................................... 35 33. Calypteres entirely whitish; arista with about 18 cilia; hind femur, on ventral surface without setae (Brazil) .............................................. .......... C similata (Couri, 1982) Calypteres brown or at least lower one with brown margins; arista with more than 21 146
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cilia; hind femur, on ventral surface with 3-4 setae ................................................ 34 34. Hind tibia on anterodorsal surface with one seta on middle third; arista with dorsal secondary cilia short; eyes bare (Brazil) .......................... C incognita (Snyder, 1954) Hind tibia on anterodorsal surface with 2 setae on middle third; arista with long dorsal secondary cilia; eyes with short and sparse cilia (difficult to observe) (Mexico, Guatemala, El Salvador, Costa Rica, Panama, Brazil, Paraguay) ................... .
...................................................... .............................. C. polystigma (Wulp, 1896) 35. Wing without brown clouds on apex of costal vein and on crossvein dm-cu (Panama, Grenada, Trinidad, Peru, Brazil) ................................... C praenubila (Snyder, 1954) Wing with a brown cloud on apex of costal vein and on crossvein dm-cu .............. 36 36. Humeral callus yellowish; calypteres white or yellowish; costal vein with an elongated brown cloud along its entire extension (Mexico, Nicaragua, Costa Rica, Panama, Trinidad) ............................................. .............................. C wulpi (Snyder, 1954) Humeral callus brownish; calypteres light brown with darker margins; costal vein with a brown cloud only on its apical half (Brazil) ............. Cjitscisquama (Snyder, 1954) 37. Pleura and fore leg yellow..................................................................................... 38 Pleura and fore leg light brown .............................................................................. 39 38. Proepimeral cilia yellow; wing with a brown cloud only on apex of vein R) (Brazil, Paraguay) ............................................. .............................. C mellina (Stein, 1918) Proepimeral cilia brown; wing with a brown cloud on upper margin from vein Sc up to wing apex (Brazil) ........................................................... C varicolor(Hough, 1900) 39. Anepimeron with long cilia; vein R 2+3 with a brown cloud; arista with long dorsal secondary cilia (Mexico, El Salvador, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Peru, Bolivia, Brazil, Argentina) ............ .
.......................................................................................... C. veniseta (Stein, 1904)
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Anepimeron with short cilia; vein R 2+3 without a brown cloud; arista with short dorsal secondary cilia (Mexico, Guatemala, El Salvador, Costa Rica, Panama, Guyana, Argentina) ..................................................... ............
c. inuber(Giglio-Tos, 1983)
Species of Cyrtoneuropsis not included in the key and its respective geographical distribution Species
Geographical distribution
C. monstrata (Wulp, 1896)
Mexico
1 mm
c Figs, 11-13, characters of Cyrtoneuropsis (modified from PAMPLONA, 1999): II, C. dubia, antenna; 12, C. dubia, thorax, lateral view; 13, C. gemina, katepisternum. Abbreviation: c, lower calypter.
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Mulfordia Malloch, 1928 Type-species. Mulfordiajerruginea Malloch (orig. des.). Diagnosis. Female: inner vertical setae long and strong, outer pair very short; dorsocentraI2:3; prealar absent; suprasquamal ridge setulose; katepisternal setae 1: 1; pro sternum and center of anepimeron bare; posterior spiracle with a few black hairs on hind margin; meron with fine hairs on center; sternite 1 bare; hind coxa setulose above on posterior surface; hind tibia without a posterodorsal seta; vein R4+5 with few short black setulae on its base, on ventral surface. Reference. MALLOCH, 1928. Monophyly. Not yet established. Geographical record. Bolivia, Peru. Comments. CARVALHO et aI. (1993b) placed M. longipes (Stein, 1911) tentatively in Mulfordia. This species was included in the key.
Key to species
1.
Veins R4+5 and M sub-parallel apically (Peru) .......................M. secunda Snyder, 1951 Vein M slightly curved forward .............................................................................. 2
2.
Pedicel and palpus fuscous; wings yellowish hyaline with yellow veins (Bolivia)
.................................................................................. .M. jerruginea Malloch, 1928 Entire antenna greyish black; palpus black; wings infuscated (Bolivia)
......................................................................................... .M. longipes (Stein, 1911)
Neomuscina Townsend, 1919 (Figs. 14-15) Type-species. Musca scutellaris Fabricius (orig. des.) =tripunctata Wulp. Diagnosis. Eyes bare; arista plumose; anepimeron with discal cilia; prosternum, proepimeron and proepisternum bare; postalar wall bare; supra and infra squamal ridges bare; anterior intra-alar seta absent; female with 2 posterior intra-alar; acrostichals 0: 1 or 1: 1; calcar absent; hind coxa on posterior surface bare; vein R 2+3 bare or with few cilia at Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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node, on dorsal and ventral surfaces; apical portion of stem vein setulose (Fig. 14) on ventral surface and usually with one or 2 setulae on dorsal surface; vein M curved forward and ending at or close to wing apex; vein R4+5 ending well before wing apex (Fig. 15). Reference. SNYDER, 1954a. Monophyly. Not yet established. Geographical record. Mexico, Guatemala, Costa Rica, Panama, Trinidad, Cuba, Guyana, Colombia, Venezuela, Peru, Brazil, Paraguay, Argentina; United States of America. Comments. Neomuscina needs an extensive revision. Except for SNYDER (1954a), there is no revisional paper and a few new species have been described after his work. At the end of the following key, there is a list of species that were not included in it. COURI & PAMPLONA (2000) transferred Cyrtoneurina scutellata Johnson to this genus.
Key to species (modified from SNYDER, 1954a) 1.
Vein R 4+5entirely bare ............................................................................................. 2 Vein R4+5 with one or more hairs on ventral surface beyond the node .................... 28
2.
With one or more pairs of distinct presutural acrostichal setae .................................. 3 Without distinct presutural acrostichal setae .......................................................... 13
3.
Meron with hairs or setulae below the spiracle (Mexico, Puerto Rico, Venezuela; southern United States of America.) .............................. .N. tripunctata (Wulp, 1896) Meron bare below spiracle .................................................................................... 4
4.
Mid femur with two strong preapical, dorsal to posterior setae ............................ . ...... ...... .......... .. (Brazil, Argentina) N. zosteris (Shannon & Del Ponte, 1926) [in part] ................................................................. (Brazil) N. ponti Lopes & Khouri, 1995 Mid femur with three strong preapical, dorsal to posterior setae ............................. 5
5.
150
Hind tibia with two median anterodorsal setae ....................................................... 6
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Hind tibia with one median anterodorsal seta.......................................................... 8 6.
Mid and hind femora fulvous on basal half or more (Venezuela) ................ .
.......................................... .......................................... .N. nudistigma Snyder, 1949 Mid and hind femora fulvous, with the possible exception of extreme base ............... 7 7.
Male: margin of upper calypter broadly infuscated; eyes facets not conspicuously enlarged above and in front; diameter of anterior ocellus subequal to diameter of one of the posterior ocelli; female: palpus conspicuously enlarged near apex, this apical portion 1.7 times the basal diameter (Mexico) .............. .N. vecta (Giglio-Tos, 1893) Male: margin of upper calypter not broadly infuscated; eyes facets conspicuously enlarged above and in front; diameter of anterior ocellus approximately two times that of one ofthe posterior ocelli; female: palpus not conspicuously enlarged apically (Mexico ) ..................................................... .............. .N. paralis (Giglio-Tos, 1893)
8.
Base of femora infuscated, usually one or more pairs of coxae with dark streaks ...... 9 Base offemora and all coxae entirely yellow to fulvous ......................................... 11
9.
Crossveins not surrounded by dark clouds ........................................................... 10 Crossveins surrounded by extensive, infuscated c1ouds ....................................... .23
10. Male: intra-alars 2; eye facets not conspicuously enlarged above and in front (Mexico, Costa Rica, Panama, Venezuela, Brazil) ....................... ..N. dorsipuncta (Stein, 1918) Male: intra-alar 1; eyes with an area of conspicuously enlarged facets above and in front (Venezuela) ........................................... ..................... N. macrops Snyder, 1949 11. Without dark accessory setulae adjacent to either the prothoracic or stigmatal seta (Brazil) .............................................. ............................. N. sanespra Snyder, 1949 With dark accessory setulae adjacent to both the prothoracic and the stigmatal seta..... 12 12. Dorsocentrals 2:3 (Mexico, Guatemala) ................................ .N. triseta Snyder, 1949 Dorsocentrals 2:4 (Mexico, Guatemala) ....................... N. mexicana (Macquart, 1843) Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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13. Dorsocentrals 2:3 ................................................................................................ 14 Dorsocentrals 2:4 ................................................................................................ 16 14. Mid femur with one posterodorsal and one posterior preapical setae (Barro Colorado Is., Trinidad) ..................................................................... .N. capalta Snyder, 1949 Mid femur with one dorsal, one posterodorsal and one posterior preapical setae ........ 15 15. Calypteres deep brown, the margins infuscated; hind femur with a complete row of anteroventral setae; vein R4+5 bare on ventral surface (Brazil) ... .N. inflexa (Stein, 1918) Calypteres white to pale yellow, the margins not infuscated; hind femur with only two to three long anteroventral setae at apex; vein R4+5 with a single ventral setulae beyond node (Barro Colorado Is.) .....................................................N. currani Snyder, 1949 16. Mid femur with 2 preapical setae ......................................................................... 17 Mid femur with 3 preapical setae .......................................................................... 23 17. Hind tibia with 1 submedian anterodorsal seta; thoracic pleura mostly fulvous ............ 18 Hind tibia with 2 submedian anterodorsal setae; thoracic pleura mostly infuscated ............................................................................................................................ 21 18. Mid and hind femora infuscated on at least basal half (Brazil) ..................... . .................... ..... .................. ..... ........ .......... ..... ........ ...... .. N. mediana Snyder, 1949 .................................... .................................. N. goianensis Lopes & Khoury, 1995
Mid and hind femora entirely fulvous to light brown ............................................... 19 19. Calypteres and their margins darkened (Brazil, Argentina) ............................ .. .................................................. .N. zosteris (Shannon and Del Ponte, 1926) [in part]
Calypteres and their margins not darkened .......................................................... .20
20. Wings with an uninterrupted dark cloud from apex of vein Sc to apex of vein R 2+3 ; fore tibiae with a short submedian anterodorsal seta (Trinidad, Guyana) ................................................................................... .N. nigricosta Snyder, 1949 152
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Wing with a distinct dark spot at apex of vein Sc and a faint spot near apex ofR2+3 , the interspace not darkened; fore tibia without a submedian anterodorsal seta
...................................................................... ..N. transporta Snyder, 1949 [in part] 21. Wings without a trace of a dark cloud along costal margin or surrounding crossveins; hind femur with 2-4 posteroventral setae on basal half only (Costa Rica, Panama)
........................................................................................... .N. atincta Snyder, 1949 Wings with a faint to distinct dark cloud along costal margin and surrounding cross veins r-m and dm-cu; hind femur with a complete row of strong posteroventral setae ...... .22 22. Thorax and femora entirely fulvous (South America) ................................ .. .................... ........ ........................ ...... ............N. transporta Snyder, 1949 [in part]
Thorax and at least the mid and hind femora mostly infuscated (Venezuela) .....................................................................N. nudistigma Snyder, 1949 [in part]
23. Posterior thoracic spiracle with several black setulae in at least the posteroventral comer in addition to the flap-like covering of hairs (Mexico, Guatemala, EI Salvador, Costa Rica, Panama, Venezuela, Guyana, Peru, Brazil, Paraguay) [N. pictipennis (Bigot, 1878) sensu lato] .......................................................................................... 24 Posterior thoracic spiracle without any black setulae in addition to the flap-like covering of hairs ................................................................................................................ 25 24. Thoracic pleurae and femora mostly infuscated (Mexico, Guatemala, EI Salvador, Costa Rica, Panama, Venezuela, Guyana, Peru) .........N. pictipennis nudinervis (Stein, 1918) Thoracic pleura and femora fulvous (Venezuela, Brazil, Paraguay) ................. .. .......... ...................... ...... ..... ............ ..........N. pictipennis pictipennis (Bigot, 1878)
25. Wings without infuscated areas ............................................................................ 26 Wings with infuscated areas ................................................................................ 27 26. The basal three-quarters or more of at least hind femur infuscated; male: intra-alar 1 (Panama, Trinidad, Brazil) ................................................N. instabilis Snyder, 1949 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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All femora entirely fulvous; male: intra-alars 2 (Paraguay) .. .N. schadei Snyder, 1949 ................................................................................ (Peru) N. stabilis (Stein, 1911) .......................................................... (Brazil) N. vittoriae Lopes & Khoury, 1995 27. Posteriorfourth offemora infuscated (Trinidad, Venezuela, Guyana) .................... .
....................................................................................... .N. similata Snyder, 1949 Posterior fourth of femora mostly fulvous (Brazil, Paraguay) ........................ .
................................................................................. ... .N. neosimilis Snyder, 1949 ............................. (United States of America, Florida) N. rufoscutella Dodge, 1955 28. Dorsocentrals 2:3 (Panama) .................................. N. currani Snyder, 1949 [in part] Dorsocentrals 2:4 ............................................................................................... 29 29. Costal margin darkened ...................................................................................... 30 Costal margin not darkened (Brazil) .............................. N. atincticosta Snyder, 1949 ................................................................. (Jamaica) N. scutellata (Johnson, 1919) 30. Margins of calypteres infuscated (Brazil) ......................N. tinctinervis (Stein, 1918) Margins of calypteres not infuscated (Colombia, Peru) ........... N. apicata (Stein, 1904)
Species of Neomuscina not included in the key and their respective geographical distribution
154
Species
Geographical Record
N. arcuata (Wiedemann, 1830)
Brazil
N. neotropica Curran, 1934
Guyana
N. sparsiplumata (Stein, 1918)
Brazil
N. tauota Lopes, 1984
Cuba Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
15
Figs. 14-15, characters of Neomuscina (modified from SNYDER, 1954a): 14, N. tripullctata, wing base, ventral view; IS, N. tripunctata, wing tip. Abbreviations: sv, stem vein; R4+5 , vein; M, vein.
Neomusciniopsis Albuquerque & Lopes, 1982 Type-species. Neomusciniopsis Iatifrontata Albuquerque & Lopes (orig. des.). Diagnosis. Male with wide frons, eyes separated by no more than 1,2 - 1,3 times the width of antenna; arista with moderately long plumae; dorsocentral setae 2:4; anepimeron and katepimeron bare; lower calypter enlarged, reaching scutellum; katepistemals 1:2, wing veins bare; vein M slightly curved forward apically; hind tibia without calcar; stemite 1 setulose. References. ALBUQUERQUE & LOPES, 1982; COURI et aI. (in press). Monophyly. Not yet established. Geographical record. Brazil. Comments. Monobasic genus: N. latifrontata Albuquerque & Lopes, 1982. The two previously described species were recently synonymized by COURI et al. (in press).
Neurotrixa Shannon & Del Ponte, 1926 Type-species. Neurotrixa mirata Shannon & Del Ponte (orig. des.) = felsina Walker. Diagnosis. Eyes with short and sparse cilia; arista pubescent; anepimeron partly ciliated; proepimeron and meron bare; intra-alar setae developed; vein R J with dorsal cilia; vein R4+5 with cilia from node to apex; dorsocentrals 2:4; katepistemals 2:2; hind tibia Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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with one median fine and short seta on anterodorsal, posterodorsal and anteroventral surfaces. Reference. LOPES, 1985b. Monophyly. Not yet established. Geographical record. Brazil, Uruguay, Argentina. Comments. Monobasic genus: Neurotrixafelsina (Walker, 1849).
Pseudoptilolepis Snyder, 1949 (Figs. 16-19) Type-species. Pseudoptilolepisfulvapoda Snyder (orig. des.). Diagnosis. Anepimeron, postalar wall, meron above posterior spiracle and proepimeron setulose; posterior thoracic spiracle with many setae on its superior margin; wings with vein M strongly curved forward apically; vein R 4+5 with weak cilia along its whole extension on ventral surface, on dorsal surface with strong and sparse cilia on its basal half; lower calypter with fine discal setae on dorsal surface; hind coxa with a series of small setae on posterior surface apically; calcar absent; male: aedeagus with the distiphallus bifid. Reference. LOPES & CARVALHO, 1985. Monophyly. The presence of cilia on the dorsal surface of the lower calypter, as well as the bifid aedeagus, suggest that the genus is monophyletic. These characters are rarely seen within the Muscidae, although a bifid aedeagus is also found in Chaetagenia Malloch, perhaps a synapomorphy uniting these two genera (SCHUHLr & CARVALHO, in press). Geographical record. Costa Rica, Nicaragua, Panama, Venezuela, Brazil, Paraguay. Comments. All the six described species occur in the Neotropical Region. The type material of P. latipalpis (Stein, 1918) was not examined, but it was included in the key based on characters mentioned in the original description.
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Key to species (modified from LOPES & CARVALHO, 1985) 1.
Postsutural dorsocentral setae 4 (Brazil) ................ ............ P. latipalpis (Stein, 1918) Postsutural dorsoeentral setae 3 ............................................................................ 2
2.
Proepistemum with diseal yellow hairs .................................................................... 3 Proepisternum bare ............................................................................................... 4
3.
Male with stemite 5 longer than wide (Fig 16); cercal plate with a upper incision that ends before the middle of the plate (Fig. 17) (Brazil) ........................ . ....................................................................... P. jluminensis Albuquerque, 1954
Male with stemite 5 wider than long (Fig. 18); cereal plate with a long upper incision that ends beyond the middle of the plate (Fig. 19) (Nicaragua, Venezuela, Brazil) .................................................................. ...................... P.fulvapoda Snyder, 1949
4.
Posterior femur yellowish brown; lowercalypteryellowish brown with darker margins (Nicaragua, Costa Rica, Panama, Venezuela, Brazil) .......... P. nigripoda Snyder, 1949 Femora yellow; lower calypter yellowish ............................................................... 5
5.
Fore tibia with an apical anterodorsal and dorsal seta subequal or equal to tibial width (Brazil, Paraguay) ...................................................... .P. nudapleura Snyder, 1949 Fore tibia without an apical anterodorsal seta or, if present, short, shorter than the dorsal apical and tibial width (Costa Riea, Venezuela, Brazil).............. P. confusa Snyder, 1949
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17
O,2mm
O,2mm
O,2mm O,2mm Figs. 16-19, characters of Pseudoptilolepis (modified from LOPES & CARVALHO, 1985): 16, P. jluminensis. sternite 5. male; 17, P. jluminensis, cereal plate and surstyIi; 18, P. julvapoda, sternite 5, male; 19, P. julvapoda, cereal plate and surstyli.
Xenothoracochaeta Malloch, 1921
(Plate 1) Type-species. Xenothoracochaeta prima Malloch (orig. des.). Diagnosis. Male holoptic; arista plumose; fIrst pair of frontal setae strong; scutellum with discal setulae extending up to the margins of the ventral surface; prealar seta present in both sexes (sometimes vestigial in male); meron setulose; anepimeron with cilia below calypter; katepistemals 1:2; metapleuron with a group of cilia; posterior spiracle with setae on posterior margin; veins R4+5 and M divergent apically; stemite 1 bare. 158
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References. SNYDER, 1949a; ALBUQUERQUE & LOPES 1979b. Monophyly. Not yet established. Geographical record. Venezuela, Colombia, Peru, Bolivia, Brazil, Paraguay, Argentina. Comments. There are six species in the Neotropical region, all them occurring in South America.
Key to species (modified from SNYDER, 1949a) 1.
Scutellar setulae descending onto ventral surface of scutellum; with two equally long and strong stigmatal setae ....................................................................................... 2 Scutellar setulae not invading ventral surface of scutellum; with a single stigmatal seta on each side or with a strong and a very much weaker one ...................................... 5
2.
Antenna and palpus mostly yellow.......................................................................... 3 Antenna and pal pus not yellow.............................................................................. .4
3.
Pro sternum setulose (Peru, Bolivia, Brazil, Paraguay) ......................................... ..
.................................................................... .......... X. ruJiguttata (Macquart, 1851) Prosternum bare (Brazil) ....................... .x. subreptilia Albuquerque & Lopes, 1979 4.
Antenna, palpus and entire scutellum infuscated (Venezuela) .......................... .
...... ................................... ... ..................... ............ .......x. caerulea (Snyder, 1949) Antenna mostly black, brownish at apex of pedicel and base of flagellomere; palpus black; scutellum greyish (Argentina) .................................. .x. golbachi Snyder, 1957 5.
Dorsocentrals 2:3 (Colombia) ................................ X.foscomarginata (Snyder, 1949) Dorsocentrals 1:3 (Colombia, Peru) ....................................X. prima Malloch, 1921
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Subfamily Mydaeinae Verral, 1888 Diagnosis. Prealar seta usually present; calcar absent; wings with the veins usually ciliated; female: ovipositor reduced; cerci reduced, sclerotized and with cilia only on outer surface; sternite 7 divided; egg Mydaea-type (according to HERTING, 1957). Monophyly. Not well established yet. SKIDMORE (1985) did not find unique characters for the subfamily. CARVALHO (1989d) found the shape of the stemite 7 (divided transversely in two pieces) as a synapomorphy for the genus-type of the subfamily. Comments. The subfamily has 22 genera in the world, eight of them occurring in the Neotropical Region. There are 46 species occurring in this region.
Brontaea Kowarz, 1873 (Figs. 20-32) Type-species. Anthomyia polystigma Meigen (des. COQUILLETT, 1910: 516). Diagnosis. Male holoptic; female: frons without proclinate orbitals and without crossed setae on frontal vitta; arista bare (longest hairs shorter than its basal diameter); intra-alars 2; dorsocentrals 2:4; prealar absent; intrapostalar 1; proepistemal 1; disk of notopleuron bare; postalar declivity, suprasquamal ridge, pro sternum and proepisternal depression bare; anepisternum with a short seta in upper anterior corner; anepimeron, greater ampulla and meron bare; katepistemals 1:2; methatoracic spiracle small, triangular, bare on margins; mid tibia with 1 submedian posterior seta and 1 strong ventral apical seta; hind tibia without calcar, with 1 short submedian anterodorsal seta, dorsal apical seta strong; veins bare; vein M slightly curved towards R 4+5 on apical portion; lower calypter "Phaonia-type", bare; sternite 1 with stiffsetulae laterally (Fig. 20); female ovipositor (Figs. 21, 22) and egg "Mydaea-type". Reference. CARVALHO & PONT, 1998. Monophyly. The genus Brontaea is probably polyphyletic. Some of its species have aPhaonia-type ovipositor (according to HERTING, 1957), while others, have a Mydaea-type ovipositor. All species herein considered have a Mydaea-type ovipositor. This genus is here considered as a Mydaeinae as previously suggested by SKIDMORE (1985) and then by CARVALHO (1989d) and CARVALHO & PONT (1998). 160
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Geographical record. Mexico, Cuba, St. VicentIs., Jamaica, Puerto Rico, Haiti, Peru, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. The genus occurs in all biogeographical regions but it is absent from New Zealand and Oceania and was accidentally introduced in Hawaii (CARVALHO & PONT, 1998).
Key to species (modified from CARVALHO & PONT, 1998) 1.
Mesonotum with dark brown vittae, very conspicuous and well-defined (Figs. 25); male: outer rows of presutural acrostichal setulae equidistant between the dorsocentral setae and each other; mesonotum with the dark central vittae continuing onto scutellum; abdomen with the dark markings on tergite 3 L-shaped (Fig. 29); cercal plate quadrate; female: sternite 2 wider than long (Mexico, Venezuela, Colombia, Peru, Bolivia, Brazil, Paraguay) ............................................................. .......... .B. nonnata (Bigot, 1885) Mesonotum with dark vittae absent or, when present, not well marked and not sharply defined; male: outer row of presutural acrostichal setulae usually closer to the dorsocentral setae than to each other; scutum with the dark central vitta discontinued at scutellum or continuing as an ill-defined and suffused mark; abdomen with the dark markings on tergite 3 L-shaped or quadrate; cercal plate not as described above .......... .2
2.
Palpus yellow, darkened at tip; larger species, body length 4.6-6.7 mm but only rarely less than 5.0 mm (a few female specimens); trochanters broadly yellow, especially on fore leg; anterior postsutural intra-alar seta reduced or absent, hardly distinct from ground setulae; male: abdomen extensively yellow on basal segments, and the dark markings quadrate especially on tergite 3 (Fig. 27); cercal plate heart-shaped; epiphallus abruptly enlarged apically, in dorsal view expanded mushroom-shaped; female: frontal triangle yellowish-grey dusted, very well marked; sternite 2 longer than wide (Mexico, St. Vincent, Jamaica, Puerto Rico, Haiti, Venezuela, Brazil, Argentina; Southern United States of America) .............................................................. B.debilis (Williston, 1896)
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Palpus black; smaller specimens, body-length 2.6-5.3 mm; trochanters only narrowly or not at all yellow; male: abdomen wholly black, the dark markings on tergites 3 and 4 as in figures 29 and 30; cereal plate and epiphallus not as described above; female: frontal triangle dull grey to grey, less well marked .................................................... 3 3. Crossvein dm-cu more or less straight; tergite 1+2 narrowly grey dusted along hind margin; sternite 5 wider than long, with 4-6 setulae placed on apical margin; anterior postsutural intra-alar seta very short, at most twice as long as ground setulae; male: ocellar setae very short, hardly longer than length of ocellar tubercle; 3-4 frontal setae; parafacial sunken, in lateral view partly obscured by eye; hind femur with 3-4 strong anteroventral setae preceded by a series of much shorter but distinct setae along basal part of this surface (Fig. 31); cercal plate expanded at base and narrowing suddenly in apieal part; distiphallus without spinules on basal part; female: proepimeral seta with no or at most one adjacent setula; fronto-orbital plate with 1-3 setulae outside lower frontal setae; tergites 4 and 5 with fewer, smaller and sparser brown seta-dots; sternite 7 present as short rod (Mexico, Cuba, Puerto Rico, Peru, Bolivia, Ecuador, Brazil, Chile; United States of America, mainly southern states)...... ........ B. quadristigma (Thomson, 1869) Crossvein dm-cu distinctly curved; sternite 5 longer than wide, with numerous setulae (usually 15-20) on lateral and apical margins; anterior postsutural intra-alar seta longer, at least twice as long as ground setulae, often longer; male: ocellar setae long and strong, at least equal to lowest frontal seta; 5-6 frontal setae; parafacial not sunken, visible in lateral view; hind femur with only 3-4 strong anteroventral setae in apical third (Fig. 32); cercal plate narrowing uniformly from base to apex; distiphallus with spinules on basal part; female: proepimeral seta with 3-6 adjacent setulae; frontoorbital plate with 4-6 setulae outside lower frontal setae; tergites 4 and 5 with numerous large brown seta-dots; sternite 7 present only as 2 seta-dots (Mexico, West Indies, Brazil; United States of America, Canada; Hawaii) ................B. delecta (Wulp, 1896)
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~20
epi
21
0,5 mm 0,25 mm
0,25 mm
1 mm
1 mm
Figs. 20-32, characters of Brontaea (modified from CARVALHO & PONT, 1998): 20, B. debilis, sternite 1; 21, B. debilis, ovipositor, lateral view; 22, B. normata, ovipositor, lateral view; 23, B. debilis, mesonotum, dorsal view; 24, B. quadristigma, mesonotum, dorsal view; 25, B. normata, mesonotum, dorsal view; 26, B. delecta, mesonotum, dorsal view; 27, B. debilis, abdomen, dorsal view; 28, B. quadristigma, abdomen, dorsal view; 29, B. llormata, abdomen, dorsal view; 30, B. delecta, abdomen, dorsal view; 31, B. quadristigma, hind femur and hind tibia, lateral view; 32, B. delecta, hind femur and hind tibia, lateral view. Abbreviations: epi, epiproct; hyp, hypoproct.
Graphomya Robineau-Desvoidy, 1830 (Figs. 33-37, Plate 1) Type-species. Musca maculata Scopoli (des. WESTWOOD, 1840: 141). Diagnosis. Characteristic marks on mesonotum and abdomen (Figs. 33-34); male holoptic; female without interfrontal setae; arista enlarged on basal third, plumose; dorsocentrals 2:4; prealar present; postalar declivity, suprasquamal ridge and pro sternum bare; anepimeron, center of proepimeron and prosternum bare; katepisternals 1: 1, both Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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on upper posterior angle; meron setulose on the upper margin, both in front of and below the spiracle; methatoracic spiracle bare on margins; mid tibia with 1 submedian posterior seta and 1 strong ventral apical seta; hind tibia without calcar; vein M curved towards R4+5 on apical portion (Fig. 35); vein R4+5 setulose basally base; lower calypter about twice the length of the upper one; stemite 1 setulose. References. MALLOCH, 1934; ARNTFIELD, 1975. Monophyly. The monophyly and affinities of Graphomya have been long regarded as a problem (SKIDMORE, 1985). HENNIG (1965) included it among the Mydaeinae, a concept followed by VOCKEROTH (1972). SKIDMORE (1985) also included the genus among the Mydaeinae, although having the same view as LOBANOV (1977), that Graphomya has more affinities with the Coenosiinae than with the Mydaeinae. Geographical record. Mexico, Cuba, Puerto Rico, Peru, Brazil, Chile, Venezuela, Uruguay, Argentina; Afrotropical, Palaearctic, Nearctic, Oriental. Essentially cosmopolitan in temperate and tropical areas, absent in New Zealand and Oceania, except Micronesia, better developed in Afrotropical and Oriental Regions (VOCKEROTH, 1972). Comments. Some South American records, especially those related to the typespecies, are probably misidentifications.
Key to species (modified from MALLOCH, 1934) 1.
Male ................................................................................................................... 2 Female ................................................................................................................ 4
2.
Frons at narrowest point as wide as length of flagellomere; parafacial with anterior part of frontal orbits, and most of gena, densely golden yellow dusted; abdomen black, with grey dust and the usual irregular tripunctate dorsum; tergite 5 conspicuously differentiated by dense golden yellow dusting; mid tibia with at least one anterodorsal seta beyond middle; 2 or more posterodorsal and 1 or 2 posteroventral setae (Peru, Brazil, Chile, Argentina) .................................. G. analis (Macquart, 1851) [in part] Frons at narrowest point not, or very little, over half as wide as length of flagellomere; parafacial dusting not as above; tergite 5 not conspicuously differentiated by goldenyellow dusting from the others; mid tibia without anterodorsal setae beyond
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middle .................................................................................................................. 3 3.
Pre sutural area between dorsocentrals with a broad black vitta that is but inconspicuously if at all divided by a central grey vitta (Puerto Rico, Venezuela, Uruguay, Argentina; Afrotropical, Palaearctic and Oriental Regions; Melanesia) .................. . .................................................................................. G. maculata (Scopoli, 1763) Pre sutural area between dorsocentrals with 2 narrow black vittae that are separated by a central dusted vitta as conspicuous as those on each side of the dorsocentrals (Peru, Argentina, Chile) ...................................................... G. chilensis Bigot, 1888
4.
Mid tibia with an anterodorsal seta beyond middle; tergite 5 with dense golden-yellow dust that contrasts sharply with the grey dusting of the other tergites; orbits, parafacial and gena with the same dense golden-yellow dusting (Peru, Brazil, Chile, Argentina) ......................................................................... G. analis(Macquart, 1851) [in part] Mid tibia without an anterodorsal seta beyond middle; tergite 5 not, or very slightly, more noticeably yellowish dusted than the other tergites; head usually with much less distinct yellow dusting ............................................................................................ 5
5.
Frontal orbits, parafacial and the greater part of gena densely golden-yellow dusted (Argentina) ...................................................................... G. auriceps Malloch, 1934 Frontal orbits, parafacial and most of gena grey dusted (Puerto Rico, Venezuela, Uruguay, Argentina; Afrotropical; Palaearctic and Oriental Regions; Melanesia) .................................................................................... G. maculata (Scopoli, 1763)
Species of Graphomya not included in the key and their respective geographical distribution Species
Geographical distribution
G. mendozana (Enderlein, 1935)
Argentina
G. meridionalis Townsend, 1892
Brazil
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G. mexicana Giglio-Tos, 1893
Mexico, Panama, Bolivia, Argentina
G. occidentalis Arntfield, 1975
Mexico, California, Hawaii (*)
G. panamensis Dodge, 1965
Panama
G. podexaurea (Enderlein, 1935)
Chile
G. stipata (Walker, 1853)
Puerto Rico, Venezuela, Guyana
(*) Keyed in ARNTFIELD (1975) with other North American Graphomya
Figs. 33-35, characters of Graphomya sp. (modified from ARNTFIELD, 1975): 33, mesonotum, dorsal view; 34, abdomen. dorsal view; 35. wing. dorsal view. Abbreviation: ntp ant. notopleural anterior seta.
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Hemichlora Wulp, 1893 Type-species. Curtonevra vittigera Bigot (mon.) =scordalus Walker. Diagnosis. Male hoI optic; female frons without interfrontal setae; arista with short cilia only on dorsal surface; intra-alars 2; dorsocentrals 2:4; prealar present; postalar declivity, suprasquamal ridge, pro sternum and proepisternal depression bare; anepimeron setulose; propleuron and prosternum bare; meron setulose below spiracle; katepisternals 1: 1, both setae on upper posterior angle; methatoracic spiracle bare on margins; hind tibia without calcar; veins bare; M slightly curved towards R4+5 on apical portion; lower calypter about 1.5 times the length of the upper one; sternite 1 setulose. Reference. LOPES, 1978. Monophyly. Not yet established. Geographical record. Mexico, Guatemala, Costa Rica, Brazil. Comments. Monobasic genus: Hemichlora scordalus (Walker, 1861).
Mydaea Robineau-Desvoidy, 1830 (Plate 2) Type-species. Mydaea scutellaris Robineau-Desvoidy (des. COQUILLET, 1901: 139) = corni Scopoli. Diagnosis. Male hoI optic; female frons without interfrontal setae; arista plumose, with plumae moderately long on basal half, becoming shorter on apical half; intra-alars 2; dorsocentrals 2:4; prealar present; anepimeron and meron bare; katepisternals 1:2; hind tibia without calcar; veins bare; vein M straight; lower calypter about twice the length of the upper one; sternite 1 bare; female ovipositor and egg "Mydaea-type". Reference. SNYDER, 1949a. Monophyly. Not yet established, but according to VOCKEROTII (1972) Mydaea and Myospila are closely related, especially based on the morphology of the ovipositor. Geographical record. Mexico, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. Mydaea is poorly studied in the Neotropical Region and a great number of new species to be described remain. VOCKEROTH (1996) synonymized Scenetes Malloch, 1936 with Mydaea, specially based on the structure of the ovipositor. However, the Neotropical Mydaea were not yet studied phylogenetic ally together with Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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the Mydaea of the other biogeographical regions. The genus has monophyletic "groups" that could be raised to a higher taxonomic level. Based on these considerations, we will not recognise this synonymy until a generic revision of Mydaea is done.
Key to species (modified from SNYDER, 1949a) 1. Dorsocentral setae 2:3 ........................................................................................... 2 Dorsocentral setae 2:4 .......................................................................................... 4 2.
Fore tibia without a median posterior seta; margins of calypteres darkened (Venezuela, Ecuador, Bolivia) .......... .................................................. M. nubivena Snyder, 1941 Fore tibia with a median posterior seta; margins of calypteres not darkened ............. 3
3.
Palpus and at least the second antennal segment yellow (Mexico) .......................... .
..... .................. .................. ............. .............. ....... ....... ..... ...M. meridia Snyder, 1941 Palpus and antenna entirely black (Mexico ) ........................... M. cresa Snyder, 1949 4.
Wing with a dark costal cloud; fore tibia with a median posterior seta (Ecuador) ...................................................................................... M. Jumicosta Snyder, 1941 Wing without a dark costal cloud; fore tibia without a median posterior seta ............ ..
S.
s
Margins of upper calypteres infuscated; crossveins r-m and dm-cu with narrow infuscated clouds (Mexico, Paraguay, Brazil) ...... ..............M. plaumanni Snyder, 1941 Margins of upper calypteres and crossveins nn and dm-cu not infuscated .................. 6
6.
Frontal vitta, at narrowest part of front, three to four times as wide as fronto-orbital plate (Venezuela) ........................................... .......................M. caras Snyder, 1949 Frontal vitta, at narrowest part of front, only as wide as width of fronto-orbital plate (Colombia) ....................................................... ..M. compressicornis Snyder, 1949
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Species of Mydaea not included in the key and their respective geographical distribution Species
Geographical distribution
M. castanea Albuquerque, 1955
Brazil
M. centralis Stein, 1911
Peru; Chile
M.fumipennis (Wulp, 1896)
Mexico
M. insons (Giglio-Tos, 1896)
Mexico
M. latomensis Snyder, 1957
Argentina
M. longiseta (Wulp, 1896)
Mexico
M. orba Stein, 1911
Bolivia
M. pansa (Giglio-Tos, 1893)
Mexico
M. pauciseta Stein, 1911
Peru
M. sexpunctata (Wulp, 1883)
Argentina
Unplaced species (CARVALH 0 etal. 1993) Species
Geographical distribution
M. inermis (Stein, 1904)
Bolivia
M. latipennis (Hough, 1900) *
Brazil
M. marginalis (Stein, 1904)**
Bolivia
* probably Cyrtoneurininae; ** probably Helina.
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Myospila Rondani, 1856 (Figs. 36-37) Type-species. Musca meditabunda Fabricius (orig. des.). Diagnosis. Male holoptic; female with a pair of interfrontal setae; arista enlarged on basal third, plumose; intra-alars 2; dorsocentrals 2:4; prealar present; postalar declivity, suprasquamal ridge, pro sternum, anepimeron and meron bare; katepisternals 2:2; methatoracic spiracle bare on margins; vein M slightly curved forward apically; vein R4+5 with few cilia basally; hind tibia without calcar; sternite 1 bare; female: ovipositor and egg Mydaea-type (Figs. 36-37). References: SNYDER, 1940; COURI & LOPES, 1988a, b. Monophyly. See discussion under Mydaea. Geographical record. Mexico, Costa Rica, Panama, Jamaica, Cuba, Puerto Rico, Haiti, S., Domingo, Guatemala, Venezuela, Guyana, Ecuador, Bolivia, Peru, Brazil, Argentina, Chile, Falkland Is.; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. Only six species are know in the Neotropical Region.
Key to species (modified from SNYDER, 1940) 1.
Thorax and abdomen entirely shiny blue; halter black (Argentina, Chile, Falkland Is.)
...................................................... ..............................M. cyanea (Macquart, 1843) Thorax and abdomen not shiny blue; halter yellow to orange ................................ '" .. 2 2.
Wing with a distinct darkened area on apices of the veins Sc and R J ; at most the margins of both calypteres distinctly infuscated (Peru, Brazil, Argentina)
............................................................... .M. obscura (Shannon & Del Ponte, 1926) 3.
Wings without a distinct darkened area on apices of the veins Sc and R J ; at most the margins of the upper calypter slightly brownish; the lower margined with yellow............ 3 Pedicel, flagellomere and the fore and mid tibiae black; intra-alars 2 (Mexico, Guatemala, Ecuador, Peru, Bolivia; Nearctic and Palaearctic Regions)
...................................................... .................... M. meditabunda (Fabricius, 1781) 170
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Pedicel and flagellomere light brown to orange or yellow; all tibiae light brown to orange; intra-alar 1................................................................................................ 4 4.
Palpus and flagellomere light brown or yellow; wing with a brown spot on cross vein rm (Mexico, Costa Rica, Panama, Puerto Rico, Haiti, Cuba, S. Domingo, Jamaica, Venezuela, Guyana, Peru, Bolivia, Brazil) ......................M. pallidicomis (Bigot, 1887) Palpus and flagellomere light brown; wings hyaline .................................................. 5
5.
Calypteres white with brown margins in female; hind tibia on anterodorsal surface with 2 setae on middle third (Brazil) ............M. matogrossensis Couri & Lopes, 1988 Calypteres entirely white in female; hind tibia on anterodorsal surface with 4 setae on middle third (Brazil) .........................................M.jluminensis Couri & Lopes, 1988
Figs. 36-37, characters of Myospi/a (modified from COURI & LOPES, 1988a): 36, M. matogrossensis, ovipositor, lateral view; 37, egg, dorsal view.
Scenetes Malloch, 1936, stat.rev. Type-species. Scenetes cardini Malloch (orig. des.). Diagnosis. Head in profile almost twice as high as its greatest length, distinctly narrower at vibrissae than at antennal bases, gena about as high as width of flagellomere and twice as high as width of parafacial on middle; facial ridges rather closely short setulose to a little above middle of flagellomere, the latter from 6 to 8 times as long as pedicel; arista plumose on basal half or more, bare apically; eyes almost bare, frons in both sexes about one-fifth of the head width at vertex, widened toward anterior margin; orbits narrow, each Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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with fourto six incurved anterior and two recurved posterior setae; interfrontal setae absent; ocellar and inner vertical setae long; outer vertical setae undeveloped; prostemum, center of anepimeron and meron bare; lower calypter narrowly rounded at apex and much larger than the upper calypter; vein M not curved forward apically; R 4+5setulose on both surfaces from base to about midway to the crossvein r-m; calcar absent. Reference. CARVALHO, 1985b. Monophyly. Not yet established. See discussion under Mydaea. Geographical record. Cuba and Puerto Rico. Comments. Monobasic genus: Scenetes cardini Malloch, 1936.
Scutellomusca Townsend, 1931 (Figs. 38-42) Type-species. Musca scutellaris Fabricius (orig. des.). Diagnosis. Eyes with short and sparse cilia; male holoptic; arista plumose; 2 orbital setae, the upper ones backward inclinated; prealar seta short; scutellum with lateral cilia on basal half; proepimeron setulose; katepistemals 1:2; lower calypter about twice as long as the upper one; suprasquamal ridge haired; katepimeron with few setulae; vein Rl with dorsal cilia; vein R4+5 with dorsal and ventral cilia; vein M strongly curved forward apically; mid tibia with a posteroventral seta inserted on the limit of basal third and a posterior seta on the limit of apical third; hind tibia with no setae on posterodorsal surface; stemite 1 setulose; male: cercal plate with short lower projections; female: ovipositor and egg "Mydaea-type" (Figs. 38-42). References. ALBUQUERQUE, 1954c; COURI & CARVALHO, 1992. Monophyly. Not yet established. Geographical record. Brazil. Comments. Monobasic genus: Scutellomusca scutellaris (Fabricius, 1805).
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38
0,3 mm
41
Figs. 38-42, characters of Scutellomusca scutellaris (modified from COURI & CARVALHO, 1992): 38, ovipositor, dorsal view; 39, ovipositor, ventral view; 40, ovipositor, lateral view; 41, egg, dorsal view; 42, egg, lateral view.
Subfamily Coenosiinae Verral, 1890 Diagnosis. Proboscis adapted to predation, prestomal teeth developed, prementum shiny; prealar seta absent; mid femur on anterior surface with a preapical seta; anepimeron bare; mesonotum with 3 or 5 dorsal vittae; calcar absent; abdomen with lateral spots on tergites. Monophyly. The subfamily is probably monophyletic, the strongest evidence being the absence of the prealar seta (COURI & PONT, 2000). Comments. Two tribes are traditionally included in the subfamily: Limnophorini and Coenosiini, as presented by SKIDMORE (1985), CARVALHO (1989d) and CARVALHO et al. (1993). The cosmopolitan distribution of several genera has been taken as an indication of the antiquity of the group (HENNIG, 1959). Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Tribe Limnophorini Villeneuve, 1902 Diagnosis. Prostemum setulose or bare; apical portion of vein M bent forward apically or not; female: stemite 8 usually absent; membrane on segment 8 usually with spicules; hypoproct with modifications. Monophyly. Not yet established, but the tribe is probably paraphyletic. Several Coenosiini genera were recently transferred to Limnophorini (COURI & PONT, 2000). Comments. There are 12 genera and 129 species recorded from the Neotropical Region. Agenamyia Albuquerque, 1953
(Figs. 43-50) Type-species. Agenamyiafumipennis Albuquerque (orig. des.) Diagnosis. Male dichoptic; eyes bare, at level of anterior ocellus separated by 3.43.6 of head width; gena and peristome linear (Fig. 43a, 43b); prestomal teeth developed; arista bare; one orbital reclinate seta; prealar seta absent; lower proepimeral seta downcurved; dorsocentral setae 1:3; anepimeron bare; scutellum bare on lower and lateral margins; prostemum bare; katepistemals 1: 1: 1, not equidistant from each other, the anterior one weaker than the others; lower calypter about twice the length of upper one; wing veins bare; M and R4+5 parallel apically (Fig. 44); calcar absent; stemite 1bare; cercal plate with lateral projections (Fig. 45); ovipositor moderately long with large tergites (Figs. 46-48). References. CARVALHO & COURI, 1992; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. Probably monophyletic, based on the presence of unique muscid characters as the shape of the gena and the linear peristome. Geographical record. Panama, Brazil. Comments. Agenamyia has been recently transferred to Limnophorini (COURI & PONT, 2000).
Key to species (modified from CARVALHO & COURI, 1992) 1. Male: hind femur with three strong anteroventral setae on middle third; cerci, in lateral 174
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view, with the apical third enlarged; stemite 5 as in figure 99; female: legs usually dark, sometimes femora and tibiae paler (Panama) ....... A. exotica Carvalho & Couri, 1992 Male: hind femur with two strong anteroventral setae on middle third; cerci, in lateral view with the apical third uniform; stemite 5 as in Fig. 50; female: unknown (Brazil) ............................................................................A.fumipennis Albuquerque, 1953
1 mm
~
U a
1 mm
"~o,'mm gn_~r.;:"5'fI:J'
44
43
O,Smm
Figs. 43-50, characters of Agenamyia (modified from ALBUQUERQUE, 1953; CARVALHO & COURI, 1992): 43, A. jumipennis, male head, (a) view frontal, (b) lateral view; 44, A. jumipennis, wing; 45, A. jumipennis, cereal plate; 46, A. exotica, ovipositor, ventral view; 47, A. exotica, ovipositor, dorsal view; 48, A. exotica, ovipositor, lateral view; 49, A. exotica, sternite 5, dorsal view; 50, A. jumipennis, sternite 5, dorsal view. Abbreviations: gn, gena; tg, tergite.
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Albertinella Couri & Carvalho, 1997 (Figs. 51-55) Type-species. Hypsomyia nebulicola Couri & Carvalho (orig. des.). Diagnosis. Male holoptic; eyes with cilia, more conspicuously in male; arista pubescent with plumes shorter than basal width of arista (some exemplars with 1-2 cilia more or less long on basal half of arista and easily differentiated from the others); gena slightly prominent (Fig. 51); prestomal teeth developed; labella reduced (Fig. 52); prosternum setulose laterally; mesonotum with two silver pruinose vittae coincident with dorsocentral row, in female the vittae reach the apex of scutellum, in male they are discontinuous and interrupted below suture up to posterior dorsocentral pair of setae and do not reach the apex of scutellum; acrostichal setae ciliform, not differentiated from ground setulae; prealar seta absent; lower proepimeral seta upcurved; dorsocentral setae 0: 1; katepistemals 1:2, the upper posterior seta strong and long, about three times the length of the others; anepimeron bare; lower calypter about twice the length of the upper one; vein R j with cilia on apical half of dorsal surface; vein R4+5 with cilia along its entire extension on both surfaces; hind tibia with a long and fine median anteroventral seta and another shorter submedian seta; no posterodorsal preapical setae; stemite 1 bare; ovipositor (Fig. 53-55) moderately long; segment 8 directed upwards with spicules (Fig. 55); hypoproct elongated, upward curved and with strong setae and spines; cerci short; egg Phaonia-type. References. COURI & CARVALHO, 1995, 1997a. Monophyly. Albertinella is the only Limnophorini genus with dorsocentral seta 0: 1. Geographical record. Brazil. Comments. Monobasic genus: Albertinella nebulicola (Couri & Carvalho, 1995).
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... _-------- ........
O,5mm
53 O,3mm
O,3mm
,
I
,
I I
I I
I I I
I
Figs. 51-55, characters of Albertinella nebulicola (modified from COURI & CARVALHO, 1995): 51, male head, lateral view; 52, female proboscis, lateral view; 53, ovipositor, dorsal view; 54, ovipositor, ventral view; 55, ovipositor, lateral view. Abbreviation: hyp, hypoproct.
Drepanocnemis Stein, 1911 Type-species. Drepanocnemis dorae Stein (des. MALLOCH, 1928: 471). Diagnosis. Male dichoptic; arista bare; 1 pair of reclinate orbital setae; prestomal teeth developed; labella reduced; prealar seta absent; lower proepimeral seta directed upward; dorsocentral setae 1:3; katepistemum with many ground setulae; lower calypter about twice the upper one; male fore tibia very broad, with a long, posterodorsal subbasal seta; stemite 1 bare. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
References. STEIN, 1911; MALLOCH, 1928; COURI & PONT 1999; COURI & PONT, 2000. Monophyly. COURI & PONT (2000) indicated five characters for the genus, three of them reversals: labella not developed; prealar seta present and disk of katepistemum with many setulae. The modification of the male fore tibia can be interpreted as an autapomorphy for the genus. Geographical record. Peru, Bolivia. Comments. Drepanocnemis was recently transferred from Coenosiini to Limnophorini (COURI & PONT, 2000). Only three species are known in the Neotropical Region: D. dorae Stein, 1911 (Peru); D. griseovirens Malloch, 1928 (Peru) and D. hirticeps Stein, 1911 (Peru, Bolivia). The characters available in literature are insufficient to identify and key out the species.
Limnophora Robineau-Desvoidy, 1830 (Plate 2) Type-species. Limnophora palustris Robineau-Desvoidy (des. COQUILLETT, 1910: 561) = maculosa (Meigen). Diagnosis. Male eyes separated by no more than one-fifth ofthe head width (at least in the South American species, see MALLOCH, 1934); eyes almost invariably bare; prestomal teeth developed; prealar seta absent; lower proepimeral seta upcurved; postsutural dorsocentral setae 3 or 4; prostemum setulose; apical portion of vein M slightly bent forward; wing with cilia on both sides of base of radial sector; stemite 1 bare; female: ovipositor with segment 8 directed upwards and with spicules; hypoproct elongated and with spines. References. STEIN, 1911; MALLOCH, 1934. Monophyly. Although Limnophora belong to a monophy letic group that includes many genera occurring in all biogeographical regions, HENNIG (1965) understood that the genus is paraphyletic. The Limnophora group is considered the sister-group of Spilogona (HENNIG, 1965; SKIDMORE, 1985). Geographical record. Mexico, Bahamas, Puerto Rico, Cuba, Belize, Guatemala, Costa Rica, Panama, St. Vincent Is., Jamaica, Venezuela, Colombia, Guyana, Bolivia, Peru, Ecuador, Galapagos Is., Brazil, Uruguay, Paraguay, Argentina, Chile, Juan Fernandez Is.; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. 178
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Comments. The genus needs an extensive taxonomic revision. The keys of STEIN (1911) and MALLOCH (1934) are presented below with modifications. The species not included in the key are listed at the end.
Key to species (modified from STEIN, 1911) 1.
Both crossveins r-m and dm-cu with a brown cloud (Mexico, Guyana) .................. . . .......................... ........ .................... ............. ........... ...... L. elegans Macquart, 1843
Crossveins r-m and dm-cu without brown clouds .................................................... 2 2.
Mesonotum almost entirely dark brown, vittae, if present almost imperceptible (Peru) ................................................................................. .L. marginipennis Stein, 1911
Mesonotum grey with 3 brown very distinct vittae ................................................... 3 3.
Wing completely hyaline or at least yellowish (Mexico, Peru, Brazil) .......................... .. . ............. .............................. ............. ......... ............ ......... ..... L. deleta (Wulp, 1896)
Wing distinctly blackish, specially on anterior margin ................................................ .4 4.
Orbit, face and gena with yellowish pruinescence (female) (Peru, Bolivia, Argentina) ........................................................................... ............... L. aurifacies Stein, 1911
Orbit, face and gena with grey or blackish pruinescence .......................................... 5 5.
Prescutum entirely black (Mexico, Guatemala, Costa Rica, Panama, Venezuela, Guyana, Ecuador, Peru, Bolivia, Brazil, Argentina) .......... .L. saeva (Wiedemann, 1830) Prescutum with spots or vittae ................................................................................ 6
6.
Arista with long plumae, longer than width of flagellomere (Peru, Guyana, Brazil, Argentina) ................................... ......................................... .L. piliseta Stein, 1919 Arista with plumae shorter than width of flagellomere ............................................. 7
7.
Mesonotal vittae forming a large vittae after suture; vein R] bare (Mexico, Bahamas, Puerto Rico, Cuba, Venezuela, Guyana, Peru, Brazil, Paraguay, Argentina, Chile;
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Nearctic region) ................................................................. .L. narona (Walker, 1849) Mesonotal vittae not forming a unique large vittae after suture; vein R J with short setae near apex ............................................................................................................. 8 8.
Mesothorax with a transversal brown vitta after suture (Mexico, Guatemala, Belize, Costa Rica, Puerto Rico, Venezuela, Peru, Brazil, Chile; southern United States of America) .............................................. ................... .L. corvina (Giglio-Tos, 1893) Mesothorax without a transversal brown vitta after suture ....................................... 9
9.
Arista slightly pubescent; mesonotal vittae almost confluent after suture (Chile)
...................................................... ............................................... L.laeta Stein, 1911 Arista distinctly pubescent or with short plumae; mesonotal vittae not confluent after suture (Bolivia) ....................................................................... L. alacris Stein, 1911
Key to species (modified from MALLOCH, 1934) 1.
Longest hairs on arista longer than width of flagellomere; eyes of male bare, separated at narrowest part of frons by a mere line, at which point the orbits obliterate the interfrontalia; hind femur with 2-3 preapical anteroventral setae, and a series of widely spaced weak setae on basal half or more of posteroventral surface (Peru, Guyana, Brazil, Argentina) .................... '" ..................................... .... .L. piliseta Stein, 1919 Longest hairs on arista not as long as the width of the flagellomere; other characters different from above .................................................................... '" ....................... 2
2.
Postsutural dorsocentral setae 3, all strong; longest cilia on arista about half as long as width of flagellomere (Peru, Argentina) ........................... .L. brevihirta Malloch, 1934 Postsutural dorsocentral setae 4; the two anterior pairs behind suture much shorter and weaker than the two posterior pairs ................................................................ 3
3. 180
Arista pubescent, longest shorter not longer than its basal diameter; both sexes with Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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some well-developed setae on basal half of posteroventral surface of hind femur (Argentina, Chile, Juan Fernandez Is.) ................ ......... .L. patagonica Malloch, 1934 Arista with longest hairs about twice as long as its basal diameter; female without well-developed setae on basal half of posteroventral surface of hind femur (Argentina, Chile) ........................................ species not named according to MALLOCH, 1934
Species of Limnophora not included in the key and their respective geographical distribution Species
Geographical distribution
L. aczeli (Snyder, 1957)
Argentina
L. albuquerquei (Lopes & Couri, 1987)
Brazil
L. altaneira (Albuquerque, 1954)
Belize, Venezuela, Brazil
L. cubana Johnson, 1919
Cuba
L. exulWilliston, 1896
Cuba,Jamaica,St. Vincent Is.
L. garrula (Giglio-Tos, 1893)
Mexico, Brazil, Argentina
L. iniqua (Stein, 1911)
Peru, Chile
L. leucotelus (Walker, 1853)
Brazil
L. limbata Bigot, 1885
Mexico, Chile
L. longiseta (Lopes & Khouri, 1989)
Brazil
L. marginata Stein, 1904
Colombia, Ecuador, Brazil, Argentina
L. minuscula (Wulp, 1896)
Mexico
L. nigrargentata (Albuquerque, 1954)
Brazil
L. ovativentris (Macquart, 1851)
Uruguay
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L. paranaensis (Albuquerque, 1954)
Brazil
L. paulistana (Lopes & Khouri, 1991)
Brazil
L. pica (Macquart, 1851)
Costa Rica, Jamaica, Brazil, Galapagos Is.
L. platystoma (Thomson, 1869)
Ecuador
L. spreta Malloch, 1921
Brazil
L. strigata (Giglio-Tos, 1893)
Mexico
L. vicaria (Walker, 1853)
Brazil
L. vicina Robineau-Desvoidy, 1830
Brazil
L. virgata (Wiedemann, 1830)
Brazil
L vittata (Macquart, 1851)
Brazil, Argentina.
Unplaced species of Limnophora s.lat. (CARVALHO etal., 1993) Species
Geographical distribution
* L. porteri (Brethes, 1920)
Chile
* Probably not Limnophorini Lispe Latreille, 1797 Type-species. Musca tentaculata De Geer (mon. LATREILLE, 1802: 462). Diagnosis. Male dichoptic; palpus enlarged apically, spatula-like; prestomal teeth developed; without interfrontal setae and proclinate supraorbital, parafacialia setulose; dorsocentral setae 2:3; prealar seta absent; anepimeron setulose in the center; lower proepimeral seta upcurved; katepistemals 1:2; male: gonopod absent; female: ovipositor with segment 8 directed upwards and with spicules; hypoproct elongated and with spines. References. SNYDER, 1949b, 1954b. 182
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Monophyly. A monophyletic group with many apomorphic characters (HENNIG, 1965; SKIDMORE, 1985). See sister-group relationships under Lispoides. Geographical record. Mexico, Bahamas Is., Virgin Is., Cuba, Puerto Rico, Guadalupe Is., Guatemala, Guyana, Bolivia, Peru, Brazil, Argentina, Chile, Paraguay; Nearctic, Palaearctic, Afrotropical, Oriental and Australian Regions. Comments. The genus needs an extensive taxonomic revision on the Neotropical Region. A key of SNYDER's (1949b) is here presented with modifications and the species not included in the key are listed at the end.
Key to species (modified from SNYDER, 1949b) 1.
Mid tibia with only one posterior median seta ........................................................ 2 Mid tibia with one submedian anterior or anterodorsal seta in addition to the posterior median seta ........................................................................................................... 4
2.
Hind femur with a row of short, stout, posteroventral setae on basal half (Argentina, Chile) ............................................................................ .L. setuligera (Stein, 1911) Hind femur with no posteroventral seta .................................................................. 3
3.
Male with a prolongation offore metatarsus (Guadalupe Is., Guatemala, Peru, North America, Palaearctic Region) ................................. .L. tentaculata (De Geer, 1776) Male with a simple fore metatarsus (Bolivia) ........................... .L. vilis (Stein, 1911)
4.
Pre sutural dorsocentral seta weak, the anterior one, if present, never so strong as the outter vertical, and the posterior one smaller then the anterior ocellar seta (Brazil)
......................................................................................... .. L. latana Snyder, 1949 Presutural dorsocentral seta strong; the anterior one as strong as the outer vertical and the posterior one as long as the anterior ocellar seta ................................................ 5 5.
Postsutural dorsocentral setae 4, the two anterior pairs weak (Mexico, Puerto Rico, Guyana, Brazil, Paraguay) .................................................. .L. seratina Wulp, 1896 Postsutural dorsocentral setae 3, all strong ............................................................. 6
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6.
The anterior postsutural dorsocentral seta strong; pedicel with only a small fulvous spot at apex; with well-defined dark abdominal markings (Mexico, Chile; United States of America) ...................................................... .......... .L. levis (Stein, 1911) The anterior postsutural dorsocentral seta very small, scarcely developed; pedicel entirely fulvous; posterior margins of abdominal tergites broadly fulvous and with illdefined markings (Brazil) .................................................... .L. lisarba Snyder, 1949
Species of Lispe not included in the key and their respective geographical distribution Species
Geographical distribution
L. albitarsis (Stein, 1898)
Bahamas Is.; Nearctic Region
L. bahama Snyder, 1958
Bahamas Is.
L. cotidiana Snyder, 1954
Mexico; Nearctic Region; Central and East Asia
L. nasoni (Stein, 1898)
Bahamas Is., Virgin Is., Cuba, Puerto Rico; Nearctic Region
L. rufitibialis Macquart, 1843
Puerto Rico, Brazil or Chile
Lispoides Malloch, 1920 Type-species. Limnophora aequifrons Stein (orig. des.). Diagnosis. Frons of male moderately wide, sometimes about as wide as that of the female; prestomal teeth developed; pro sternum bare; prealar seta absent; lower proepimeral seta upcurved; vein R4+5 bare; sternite 1 setulose; ovipositor with segment 8 directed upwards and with spicules; hypoproct elongated and with spines. References. STEIN, 1911; MALLOCH, 1934. Monophyly. HENNIG (1965) suggested that Lispoides and Lispe could be sister-groups. SKIDMORE (1985), based on characters of the larvae, indicated that Lispoides is not closely related to Limnophora and that the common characters found in both genera are due to adaptative convergence. 184
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Geographical record. Guatemala, Venezuela, Colombia, Bolivia, Peru, Argentina, Chile, Juan Fernandez Is.; Nearctic Region. Comments. Recently, PONT (2001) transferred Limnophora pubiceps Stein, 1911 to this genus.
Key to species (modified from STEIN, 1911) 1.
Eye distinctly haired; crossveins r-m and dm-cu surrounded by a brown cloud (Bolivia)
...................................................................................... .. L. elegantula Pont, 1972 Eye bare; crossveins r-m and dm-cu not surrounded by a brown c1oud ................... 2 2.
Crossvein r-m greyish (Bolivia, Peru) ........................L. abnonninervis (Stein, 1911) Crossvein r-m not greyish ..................................................................................... 3
3.
Halter whitish or greyish with daker margin .......................................................... ..4 Halter whitish without daker margin ....................................................................... 6
4.
Mesonotum bare beyond the strongest seta (Venezuela, Peru) ... .L. laevis (Stein, 1911) Mesonotum setulose beyond the strongest setae .................................................... 5
5.
Mid femur with strong and long setae on posteroventral surface; mid tibia with many setae; hind femur with strong setae on ventral surface, a little before distal apex (Peru)
.....................................................................................L. atrisquama (Stein, 1904) Mid femur with fine and short setae on posteroventral surface; mid tibia with only one median seta; hind femur bare on ventral surface (Peru) .. ....L. propinqua (Stein, 1911) 6.
Median vitta on frons thin in both sexes, or at most as wide as orbits; general coloration grey-whitish (Chile) ................................................................ L. diluta (Stein, 1911) Median vitta on frons wider than the orbits; general coloration dark .......................... 7
7.
Mesonotum almost without vittae; abdomen much longer than thorax (Bolivia, Chile)
............................................................................................ L. gracilis (Stein, 1911) Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Mesonotum with vittae faint or distinctly marked; abdomen as long as thorax ............ 8 8.
Mesonotum with faint dorsal vittae, all having the same width (Peru, Bolivia) ...................................................... ................................ L. nigribasis (Stein, 1911)
Mesonotum with distinct dorsal vittae, the median one thinner than the lateral ones (Peru, Bolivia, Chile) ................................................ ........... .L. triplex (Stein, 1911)
Key to species (modified from MALLOCH, 1934) 1.
Thorax densely pale, grey dusted; mesonotum with three conspicuous brownish-black vittae, the central one narrowest, of uniform width and sharply margined on its entire extent, occupying one-third of the intra-dorsocentral area; pleurae with a dark upper marginal vitta; frons of male about one fourth of the head width and of female about one third ................................................................................................................ 2 Thorax much less conspicuously vittate, central vitta neither uniformly wide nor sharply margined; pleurae without a dark upper marginal vitta; frons of male and of female about equally wide, at most one-fourth of the head width ....................................... 3
2.
Meron with some setulose hairs on upper margin in front of spiracle (Argentina, Chile) .............................................................................. .L. inaequifrons Malloch, 1934
Meron bare (Argentina, Chile?) .................................. .L. argenticeps Malloch, 1934 3.
With some hairs on fronto-orbital plate below level of lower frontal seta; abdomen less densely greyish dusted, the dark markings not sharply defined but more extensive, specially in male (Colombia, North America) .................. .L. aequifrons (Stein, 1898) No hairs on fronto-orbital plate below level oflower frontal setae; abdomen densely brownish-grey dusted, with the dark central line and subtriangular paired spots well defined and conspicuous in both sexes .................................................................. .4
4.
Mid tibia with one posterior seta; hind tibia with one long submedian anterodorsal seta (Peru) ................................................. ....................... .L. uniseta Malloch, 1934
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Mid tibia with several posterior setae; hind tibia with two or more unequal anterodorsal setae (Argentina) .......................................................... ,L. argentina Malloch, 1934 Species of Lispoides not included in the keys and their respective geographical distribution [including L. puhiceps (Stein, 1911) recently transferred to this genus by PONT,2001] Species
Geographical Distribution
L. guatemala Snyder, 1951
Guatemala
L. insularis Hennig, 1957
Juan Fernandez Is.
L. latifrons Snyder, 1957
Argentina
L. lopesi Albuquerque, 1955
Brazil
L. puhiceps (Stein, 1911)
Peru,Bolivia
Pachyceramyia Albuquerque, 1955 (Figs. 56-61) Type-species. Pachyceramyia enigmatica Albuquerque (orig. des.). Diagnosis. Head with the antennal axis longer than the oral axis (Figs. 56-57); male dichoptic; upper pair of frontal setae directed outward; arista short, wider on basal two-thirds, pubescent and a little longer than flagellomere; gena and facialia large; frontoorbital plate with a series of strong short cilia; prealar seta absent; acrostichal prescutellar seta ciliform; dorsocentral setae 2:4; katepisternals 2:2; wing veins bare; veins R4+5 and M parallel apically (Fig. 58); lower calypter about 1.5 as long as upper one; hind tibia with two anterodorsal (one median to submedian and another supramedian) , one posterodorsal on apical third and one posterodorsal preapical setae (Fig. 59); female: ovipositor moderately long with tergites 6 and 7 large (Figs. 60, 61). References. HUCKETT, 1936; ALBUQUERQUE, 1955b. Monophyly. Not yet established. Geographical record. Bahamas Is., South America, Brazil; United States of America. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Comments. P. pantherina (Wiedemann, 1824) was not recognised after the original description and is not herein included in the key. Key to species 1.
Postsutural dorsocentral setae 4 (Brazil) ............... P. enigmatica Albuquerque, 1955 Postsutural dorsocentral setae 3............................................................................ 2
2.
Arista plumose; hind tibia with the distal posterodorsal seta much longer (about 1.5 times) than proximal posterodorsal seta (United States of America)
................................................................................. P. longispina (Malloch, 1923)
1 mm
O.5mm
61 Figs. 56-61, characters of Pachyceramyia (modified from ALBUQUERQUE, 1955b; COURI & PONT, 2000): 56, P. enigmatica, head, lateral view; 57, P. enigmatica, % view; 58, P. enigmatica, wing; 59, P. enigmatica, hind tibia; 60, P. cordyluroides, ovipositor, dorsal view; 61, P. cordyluroides, ovipositor, ventral view. Abbreviations: tg, tergite; st, sternite.
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Arista bare or with very short setulae; hind tibia with the distal posterodorsal seta as long as or shorter than proximal posterodorsal seta (Bahamas Is.; United States of America) ............................................... ................... P. cordyluroides (Stein, 1898)
Species of Pachyceramyia not included in the key and its respective geographical distribution Species
Geographical Distribution
P. pantherina (Wiedemann, 1824)
South America
Rhabdotoptera Stein, 1919 (Figs. 62··67) Type-species. Shoenomyza striatipennis Stein (mon.). Diagnosis. Eyes with small and sparse cilia; male dichoptic; 1 pair of reclinate orbital setae; epistoma not produced; antenna short, apex of flagellomere rounded; arista bare; palpus slender; scutellum bare below and laterally; acrostichals present before and after suture; dorsocentrals 2:3, the anterior one about half as long as the posterior setae; prealar absent; prosternum bare; proepisternals 2; propleurals 1, all directed upwards; katepisternals 1: 1: 1, forming an equilateral triangle and with an extra seta laterally to anterior one (Fig. 62); lower calypter short, hardly projecting beyond the upper one; wings with bare veins; veins R 4+s and M parallel apically (Fig. 67); legs with claws and pulvilli very small; hind tibia on anterodorsal surface with one long median seta, anteroventral surface with 2 short setae on middle third, dorsal surface with a preapical seta; sternite 1 with few short setulae; male: sternite 5 sub-triangular (Fig. 63); cercal plate sub-quadrate (Figs. 64, 65); hypandrium curved (Fig. 66). Reference. COURI, 1996a. Monophyly. Not yet established. Geographical record. Peru. Comments. Monobasic genus: Rhabdotoptera striatipennis (Stein, 1898).
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ads
\ ~
~_622 0,2mm
63
hyp 0,05 mm
Figs. 62-67, characters of Rhabdotoptera striatipennis (modified from COURI, 1996a): 62, katepisternum; 63, sternite 5, male, dorsal view; 64, cereal plate and surstyli, dorsal view; 65, cereal plate and surstyli, lateral view; 66, aedeagus, lateral view; 67, wing, dorsal view. Abbreviations: ad s, additional katepisternum setae; hyp, hypandrium.
Spilogona Schnabl, 1911 Type-species. Aricia carbone lla Zettersdedt (mon). Diagnosis. Male frons intermediate; prestomal teeth developed; labellareduced; dorsocentral setae 2:3-4; katepisternals 1: 1-3; sternite 1 usually bare, when setulose, then eyes of male sub contiguous or separated by less than twice the distance across posterior ocellus inclusive; ovipositor moderately long, tergites 6 and 7 large. Reference. SNYDER, 1957b. 190
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Monophyly. Probably paraphyletic. HENNIG (1965) divided the genus into 3 species groups: species from New Zealand; species with the ovipositor with spines on genital plate; species without spines on genital plate. HENNIG (1965) called attention to the morphological similarity of the ovipositor, concerning the presence of spines, with those of Limnophora. Geographical record. Mexico, Colombia, Peru, Bolivia, Argentina. Genus cosmopolitan with many species occurring at high altitudes; in tropical regions, the species are mainly found in high altitudes and in highland areas (SKIDMORE, 1985). Comments. There are eight species in the Neotropical region, six of them in South America.
Key to species (modified from STEIN, 1911) 1.
Fore tibia with setae (Peru, Bolivia) ...................................S. hirticeps (Stein, 1911) Fore tibia with no setae .......................................................................................... 2
2.
Eye haired (Peru, Bolivia) ..................................................... S. trichops (Stein, 1911) Eye bare ............................................................................................................... 3
3. Eyes distinctly separated; halter black (Peru, Bolivia) .... .S. semicinerea (Stein, 1911) Eyes very close to each other; halter yellow (Colombia, Bolivia) .................. ..
......... ............. ......... .............. .............. .......... ............... .... S. aterrima (Stein, 1904)
Species of Spilogona not included in the key and their respective geographical distribution Species
Geographical Distribution
S. anthrax (Bigot, 1885)
Mexico; United States of America
S. argentiJrontata Snyder, 1957
Argentina
S. golbachi Snyder, 1957
Colombia, Brazil, Argentina
S. immaculiventris (Malloch, 1923)
Mexico; San Jose Is., San Francisco Is.
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Syllimnophora Speiser, 1923 Type-species. Limnophora aliena Stein (aut.). Diagnosis. Male: frons moderately wide; ocellar triangle white dusted usually extended to, or nearly to, the anterior margin; produced vibrissal angles; pro sternum setulose laterally; dorsocentral setae 2:3-4; presence of setulae at the base of vein R 4+5, on both surfaces and on upper surface at least to the apical third or more of vein Rl. References. MALLOCH, 1934; SNYDER, 1957b. Monophyly. Not yet established. Adrian Charles Pont (personal communication) understands that Syllimnophora is probably a monophyletic group but only an apical clade of the Limnophora branch. Geographical record. Mexico, Colombia, Peru, Bolivia, Brazil, Ecuador, Argentina, Chile, Juan Fernandez Is. Comments. There are 33 species recording to the Neotropical region, all of them also occurring in South America.
Key to species (modified from STEIN, 1911) 1.
Male: anterior tarsi with setae over whole extension of the ventral surface (Peru, Bolivia)
.................................................................................... .S. barbitarsis (Stein, 1911) Male: anterior tarsi without these setae ........................................................................ 2 2.
Postsutural dorsocentral setae 3; vein R4+5 forward curved apically.......................... 3 Postsutural dorsocentral setae 4; vein R 4+5straight apically...................................... .4
3.
Wing with no marginal spine; mid tibia with no anterodorsal setae; Male: frons wider at ocellar triangle than at the eye; halter black; fore tarsi compressed (Peru, Bolivia)
............................................................................. ... S. compressitarsis (Stein, 1911) Wing with marginal spine; mid tibia with one anterodorsal seta; male: frons narrower at ocellar triangle than at eyes; halter yellow; fore tarsi not compressed (Peru, Bolivia)
........................................................................... ........ S. spinuligera (Stein, 1911) 4. 192
Halter black ........................................................................................................... 5 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Halter yellow........................................................: ................................................ 7 5.
Mid tibia club-like form at end, enlarged and with dense setae over apical half, longer apicad (Peru, Bolivia) ........................................................ .S. clavitibia (Stein, 1911) Mid tibia not club-like and not bristled as described above ...................................... 6
6.
Mesonotum uniformly greyish, no visible vittae; abdomen with light grey-whitish clouds, never yellow (Peru, Bolivia) .......................................... ...S. candidifrons (Stein, 1911) Mesonotum with 3 large spots before suture; abdominal marks yellowish (Peru, Bolivia, Chile) ................................................. ................................... S. aliena (Stein, 1911)
7.
Mid tibia with 8 strong posterior setae, distributed from base to middle of tibia (Peru, Chile) ................................................. ............................... S. setitibia (Stein, 1911) Mid tibia at most with 2 posterior setae ................................................................. 8
8.
Mesonotum almost completely black, without visible vittae (Mexico, Peru, Bolivia)
...................................................................................... .S. gracilitarsis (Stein, 1911) Mesonotum greyish with 3 black, well-marked vittae .............................................. 9 9.
Eyes very close to each other................................................................................. 10 Eyes separated by a very visible and large vitta....................................................... 11
10. Anterior margin of wing very blackish (Colombia, Ecuador, Peru, Bolivia, Chile)
..................................................................................... .S. atrovittata (Stein, 1904) Anterior margin of wing slightly greyish (Peru) ........................... S. pura (Stein, 1911) 11. Anterior margin of wing blackish (Ecuador, Peru) .....................S. suavis (Stein, 1911) Wings more or less translucid ............................................................................... 12 12. Hind femur with 4-5 strong ventral setae, distributed from base to middle of tibia (Argentina, Chile) .............................................................. .. S. Jemorata (Stein, 1911) Hind femur with no setae on ventral surface ........................................................... 13 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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13. Abdomen as wide as thorax, the light marks not reaching lateral area; wing yellow at base (Peru, Bolivia) ............................................... .. S. compressijrons (Stein, 1911) Abdomen wider than thorax; abdominal marks reaching lateral area; wing blackish at base (Chile) ............................................... ........................ S. zebrina (Bigot, 1885)
Key to species (modified from MALLOCH, 1934) Male 1.
Halter with yellow knob ......................................................................................... 2 Halter with fuscous or black knob .......................................................................... 7
2.
Hind femur with about 6 moderately long and strong setae on apical half of posteroventral surface, and similarly bristled on apical half of anteroventral surface; dorsum of abdomen with central linear and lateral rounded and white dusted spots, only central part of lateral pair of tergite 5 yellowish (Argentina, Chile)
...................................................... .................................... S.femorata (Stein, 1911) Hind femur with no well developed setae on basal half of posteroventral surface, but with setae on apical half of anteroventral surface ...................................................... 3 3.
Wing attenuated at apex; crossvein dm-cu oblique, distance from its anterior extremity on vein M to margin of wing, directly below it, hardly more than equal to half the length of crossvein; all three pairs of lateral abdominal spots partly yellowish dusted, those on tergite 5 most distinctly so (Argentina) .............S. tenuipennis Malloch, 1934 Wing normal in form; crossvein dm-cu more or less oblique but distance from its junction with vein M to margin of wing, directly below it, normally over half as long as length of crossvein; abdomen usually with four pairs of white or yellowish dusted lateral spots or coloured otherwise ....................................................................... .4
4.
Lateral paired pale dusted spots large, tergite 1+2 with very distinct yellowish-grey dust at apex on the curve on that part on which the tergite is furnished with setae, the
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dust distinctly yellowish on tergites 3-5 (Chile ) ......................
s. zebrina (Bigot, 1885)
Lateral paired pale dusted spots small, tergite 1+2 faintly greyish dusted on each side below curve basal to the area covered with setae but not grey dusted at apex .......... .5 5.
Frons at narrowest point less than one-third as wide as an eye and not wider than length of flagellomere; lateral spots on tergites 3-4 silvery-white and situated on the curves, the pair on tergite 5 large and entirely golden-brown dusted (Chile, Argentina) ............................................................................... S. angustifrons Malloch, 1934 Frons at narrowest point about half as wide as an eye and distinctly wider than length of flagellomere; lateral spots on tergites 3 and 4 sometimes indistinct, when distinct, not on the curve, and the pair on tergite 5 largely grey, not entirely golden brown ....... 6
6.
Fore tibia with a submedian posterior seta (Chile) ............. S. dubitalis Malloch, 1934 Fore tibia without a submedian posterior seta (Argentina) ... S. latifrons Malloch, 1934
7.
Hind femur with the anteroventral setae confined to the apical half or slightly more of that surface ............................................................................................................ 8 Hind femur with the anteroventral setae extending to base but becoming shorter and finer basally......................................................................................................... 10
8.
Frons entirely silver-grey dusted between orbits, each orbit with a long, fine, outwardlycurved seta near upper extremity; frontal width at narrowest point fully half that of one eye; sternite 5 entirely dull, yellowish-grey dusted; setae on apical half on anteroventral surface of hind femur and basal half of posteroventral surface of mid femur very fine though long (Peru, Bolivia, Chile) ...............................................S. aliena (Stein, 1911) Frons with the usual conspicuously silvery-white dusted triangle extending to anterior margin and well defmed; frontal width distinctly less than that of one eye; stemite 5 either undusted or entirely pale grey dusted; setae on apical half of anteroventral surface of hind femur and basal half of posteroventral surface of mid femur long and strong ................................. 9
9.
The silvery-white dusted frontal triangle very much reduced, in front of ocelli hardly
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wider than anterior ocellus and reduced to a mere line about middle of frons where it becomes obsolete; tergite 1+2 not distinctly grey dusted above the lateral curve; stemite 5 entirely grey dusted (Chile, Argentina) .................... S angustifrons Malloch, 1934 The silvery-white dusted frontal triangle about twice as wide as anterior ocellus at upper extremity, gradually narrowed in front and almost reaching anterior margin of frons; tergite 1+2 distinctly dusted above lateral curve; stemite 5 not entirely dusted, shining black (Chile) .................................................... .S. nigriventris Malloch, 1934 10. Fourth and fifth segments of the fore tarsus much dilated, fourth wider than long, fifth about as wide as long; stemite 5 pale grey dusted, the pair oflarge spots on tergite 5 silvery-white dusted, yellowish only slightly and below the lateral curves (Argentina)
..................................................................................... .S. latimana Malloch, 1934 Fourth and fifth segments of fore tarsi not at all dilated, distinctly longer than wide; sternite 5 either partly or entirely yellow dusted, the pair of large lateral spots on tergite 5 golden-brown or yellow dusted (Chile )........... S inconspicua Malloch, 1934 Female 1.
Halter with knob yellow......................................................................................... 2 Halter with knobs dark brown to black ................................................................... 8
2.
Hind femur with some setae on basal half of posteroventral surface; each frontal orbit with two short, outwardly-curved upper setae (Argentina, Chile) .................... .
...................................................... ................................... S.femorata (Stein, 1911) Hind femur without any distinct posteroventral setae on basal half............................. 3 3.
Fore tibia with a strong submedian posterior seta (Chile )....S dubitalis Malloch, 1934 Fore tibia without a submedian posterior seta........................................................ .4
4.
Abdomen densely pale grey dusted, with two broad continuous brownish black vittae on dorsum and one below each lateral curve; mesonotum with a central dark chocolate brown vitta, separated from a broader deep black vitta on each side by a clear-cut
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whitish-grey dusted vitta that continues on scutellum; hind tibia with one submedian anterodorsal seta (Peru) ............................................. S. vittiventris Malloch, 1934 Abdomen not marked as above, with pale dusted markings usually definitely spotlike; mesonotal markings less sharp; hind tibia with at least two submedian anterodorsal setae .................................................................................................................... 5 5.
Tergite 5 with lateral dusted spots large, golden-brown; the other pairs small and largely whitish-grey; fore tibia sometimes with a seta on posterior surface placed less than one-third from apex; parafacial silvery (Chile, Argentina) ...................... .
................................................................................ .S. angustifrons Malloch, 1934 Tergite 5 with lateral dusted spots not entirely golden-brown, merging into grey on margins, the other pairs largely or entirely yellow dusted; fore tibia without posterior setae .................................................................................................................... 6 6.
Frontal triangle with a yellowish tinge, not entirely silver, ceasing abruptly at anterior ocellus, represented on vertex by two small white dusted spots close to bases of postvertical setae; the dusted lateral vitta on margin of mesonotum in front of wing bases yellow except in front; vittae over scutellum yellow apically (Argentina)
................................................................................ .S. tenuipennis Malloch, 1934 Frontal triangle silvery-white, usually distinct, or at least distinguishable along sides of ocellar triangle to vertex ......................................................................................... 7 7.
Mid tibia with one or more well-developed anterodorsal setae (Argentina)
...................................................................................... S. [atifrons Malloch, 1934
s.
Mid tibia without any anterodorsal setae (Chile )....................... zehrina (Bigot, 1885) 8.
Frons, when viewed from side against the light, entirely whitish-grey dusted and triangle a little more heavily white dusted; parafacial densely and uniformly pale grey dusted, with a hardly perceptible yellowish tinge below; gena with a more noticeably yellowish tinge; thorax with the black parts brownish or greyish-black, upper callus in front of wing base conspicuously greyish-white dusted (Peru, Bolivia, Chile) ............... .
.. .................. ..... .......... ........ ............................ ............... ........ S. aliena (Stein, 1911) Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Frons, when viewed from side against the light, not or very slightly white dusted and triangle densely white dusted and very conspicuously differentiated; parafacial with upper extremity black, lower part grey dusted and with a marked yellow or brown tinge; gena partly or entirely brownish-yellow; thorax with the dark parts intensely black ................................................................................................................... 9 9.
Gena broadly silvery-white behind, golden-brown in front; parafacial narrow ly yellow in front, broadly silvery-white behind; colour of thorax and abdomen intensely shining black; the two white dusted submedian vittae evanescent on anterior margin of mesonotum, widened before reaching suture; abdomen with dusted marks white except some yellowish dust at curves on tergites 4-5, each with a small well-defined white dusted spot close to extreme lateral margins (Argentina).................... ................ s. variceps Malloch, 1934 Gena entirely brownish-yellow dusted on disc; parafacial hardly showing a trace of grey dust behind; colour of thorax and abdomen brownish or greyish-black and not too shiny; submedian gray vittae entire; abdomen with dusted markings largely yellow and extreme lateral edges of tergites less distinctly spotted (Chile) .............. .. ....... .......................................................................... .S. inconspicua Malloch, 1934
Key to species (modified from SNYDER, 1957b)
1. Palpus fulvous (Argentina) ..............................................S.jlavipalpis Snyder, 1957 Palpus fuscous ..................................................................................................... 2 2. Halter fuscous to yellow........................................................................................ 3 Halter black .......................................................................................................... 4 3.
Oral margin projecting anteriorly well beyond juncture of parafacial and fronto-orbital plate; hind tibia with 3 anterodorsal setae; mid tibia with 2 median posterior setae; fifth fore tarsal segment wider than long; fourth segment about twice as long as wide (Chile) ................................................................................. .S. inconspicua Malloch, 1934
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Oral margin not projecting anteriorly beyond juncture of parafacial; hind tibia with one median anterodorsal seta; mid tibia with one median posterior seta; fifth fore tarsal segment almost square and the fourth only slightly longer than fifth (Argentina)
.............................................................................. .. S. latimanoides Snyder, 1957 4.
Mid tibia distinctly enlarged preapically and with a row oflong anterior and posterior setulae on apical half (Pem, Bolivia) ................................ s. clavitibia (Stein, 1911) Mid tibia with slender proportions throughout, without long anterior and posterior setulae, but with two or more median posterior setae .............................................. 5
5.
Mid tibia with 3 posterior setae; hind tibia with a median anteroventral seta; sternite 5 densely golden yellow pminescent (Ecuador) .........................S. browni Snyder, 1957 Mid tibia with one long and one short posterior seta; hind tibia without median anteroventral seta; sternite 5 greyish pminescent (Argentina) ........................ .
...................................................................................... .S. latimana Malloch, 1934
Species of Syllimnophora not included in the key and their respective geographical distribution Species
Geographical distribution
S. breviseta (Stein, 1911)
Bolivia, Peru
S. integra (Stein, 1911)
Bolivia
S. lispomina Hennig, 1955
Juan Fernandez Is.
S. nigra (Lopes & Couri, 1987)
Brazil
S. pauciseta (Stein, 1904)
Peru, Bolivia
S. steini Carvalho & Pont, 1993
Colombia
S. stigmatica (Lopes & Couri, 1987) Brazil
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Tetramerinx Berg, 1898 (Fig. 68) Type-species. Tetrachaeta unica Stein (aut.). Diagnosis. Male dichoptic; upper pair of frontal setae backward directed; antenna long; arista short and plumose, wider on basal two-thirds (Fig. 68); prealar seta absent; dorsocentral setae 2:4; anepimeron and meron bare; katepistemals 2:2; hind tibia with 14 anterodorsal setae; 1-2 anteroventral; 1-2 short posterodorsal and 1 preapical dorsal; sternite 1 bare; male: aedeagus long, gonopod short and dilated on center; female: ovipositor with reduced hypoproct, leaf shaped in lateral view, very setulose. References. LOPES & COURI, 1989b. Monophyly. Based on characters of the puparium, SKIDMORE (1985) considered Tetramerinx among the Coenosiini, and close to Coenosia. LOPES & COURI (1989b), based on the morphology ofthe ovipositor considered the genus as a Limnophorini, a position followed here. The monophyly of the genus is not yet established. Geographical record. Bahamas Is., Peru, Chile; southeast United States of America. Comments. Only two species described to the Neotropical Region - T. nigripes Stein, 1911 (Chile), T. rufitibia Stein, 1911 (from southeast of United States to Peru and Chile). After the original description, T. nigripes was not recognised and the characters available in literature are not sufficient to include it in an identification key. In North America only T. unica (Stein, 1898) is known. The two species described in Tetramerinx are not included in a key.
0,05
mm
Fig. 68, Tetramerinx unica (modified from LOPES & COURI, 1989b), male head, lateral view.
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Thaumasiochaeta Stein, 1911 (Figs. 69-82) Type-species: Thaumasiochaeta pilitarsis Stein (mon.). Diagnosis. Male dichoptic; frons very wide, about one-half of head width; epistomal projection variable: weakly projecting in T. variegata (Fig. 69), T. incaica and T. pilitarsis (Fig. 91), and strongly projecting in T. nigriceps (Figs. 72, 75) and T. longipalpis (Fig. 74); ocellar triangle with a pair oflong setae near anterior ocelli and about 6 other finer and shorter setae (only one pair in T. haustellata and T. longipaZpis); antenna with flagellomere wide, arista bare, enlarged at base; palpus long, broad and flattened (Fig. 76); dorsocentral setae 2:3; prealar seta absent; scutellum with setulae on lateral margin; prosternum setulose; proepisternals 2; proepimerals 2; humeral callus with 2 long setae; notopleuron with 2 long setae, the anterior a little longer than the posterior; with setulae in addition to the setae; katepisternum with long and fine cilia on disc; katepisternals 1: 1 (l:2 in T. ZongipaZpis) , the upper posterior one about twice as long as the other; lower calypter about 1.5 the length of upper one; anepimeron setulose, with long and fine setulae; meron with setulae below posterior spiracle; wing (Fig. 77 a) with strong spines along costal margin (Fig. 77b); stem vein with ventral surface bare (setulose in T. longipalpis); vein R] on dorsal surface with cilia on apical fourth (along entire length in T. longipalpis); base of veins R 2+3 and R4+5 with setulae on both surfaces; sternite 1 setulose; proboscis with haustellum totally sclerotized, shining (Fig. 78); hind tibia with a preapical seta on each of the posterodorsal, anterodorsal and dorsal surfaces; ovipositor with microtrichia on membrane of segment 7 and spicules on membrane of segment 8 (Figs. 79-80). Reference. COURI, 1999a. Monophyly. Not yet established. Geographical records. Peru, Bolivia. Comments. Seven species are described in the Neotropical region. The males are easy to identify since they vary in characters such as leg chaetotaxy and mid femur curvature. On the other hand, females are very similar and difficult to identify.
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Part II - Apical Groups
Key to adults (modified from COURI, 1999a) 1.
Katepistemals 1: 1; vein R J on dorsal surface setulose apically; vein R4+5 with setulae only basally............................................................................................................ 2 Katepistemals 1:2; vein R J on dorsal surface setulose along entire length; vein R 4+5 with cilia up to crossvein r-m (Peru) ............................... .T. longipalpis (Stein, 1911)
2.
Ocellar triangle with six or more setae in addition to the ocellar pair; male with or without a depression on basal third of hind femur................................................... 3 Ocellar triangle with no setae, apart from the ocellar pair; male without a depression on basal third of hind femur (Peru).................... ............ .T. haustellata (Stein, 1911)
3. Epistoma strongly projecting (Figs. 72, 75); lower calypter with margin brown (Peru)
........................................................................................ T. nigriceps (Stein, 1911) Epistoma not strongly projecting (Figs. 69-71); lower calypter with margin whitish or brown ......................................................................................................................... 4 4.
Lower calypter with brown margin; male: arista triangular apically (Peru, Bolivia)
.............................................................................................. T. pilitarsis Stein, 1911 Lower calypter with whitish margin; male: arista not triangular at apex ....................... 5 5.
Wing hyaline, without clouds; male: fore tibia without a very long preapical seta with triangular apex on dorsal surface (Peru) ................... .T. compressitarsis (Stein, 1911) Wing clouded; male: fore tibia with a very long preapical seta with triangular apex on dorsal surface ......................................... ;............................................................... 6
6.
Wing clouded as in Fig. 81 (Peru, Bolivia) ........................ T. variegata(Stein, 1911) Wing clouded as in Fig. 82 (Bolivia) .................................... ... .T. incaica (Stein, 1911)
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
1 mm
1
mm
74 1 mm
Figs. 69-75, characters view; 70, T. variegata, head, lateral view; 73, nigriceps, female head,
1 mm
of Thaumasiochaeta (modified from COURI, 1999a): 69, T. variegata, male head, lateral female head, lateral view; 71, T. pilitarsis, male head, lateral view; 72, T. nigriceps, male T. compressitarsis, male head, lateral view; 74, T. longipalpis, male head, lateral; 75, T. lateral view.
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Part II - Apical Groups
O,5mm
~'?fiIf 76
1 mm
b
~ O,5mm
O,2mm
1 mm Figs. 76-82, characters of Thaumasiochaeta (modified from COURI, 1999a): 76, T. pilitarsis, male palpus; 77a, T. pilitarsis, wing, 77b, costal margin of wing; 78, T. pilitarsis, female proboscis; 79, T. pilitarsis, ovipositor, dorsal view; 80, T. pilitarsis, ovipositor, ventral view; 81, T. variegata, wing; 82, T. incaica. wing. Abbreviations: hf, hypopharynx; Ibl. labella; Ie, labrum-epipharynx; pIp, palpus.
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Tribe Coenosiini Verral, 1888 Diagnosis. Eyes usually dichoptic in both sexes; usually one orbital rec1inate seta; lower proepimeral seta directed downward; katepistemal setae 1: 1: 1, forming an equilateral triangle; hind tibia usually with one anterodorsal seta; male: hypandrium tubular, elongated. Monophyly. The tribe is monophyletic; the number and disposition of the katepistemal setae - 1: 1:1 and fonning an equilateral triangle - is a synapomorphic character for the Coenosiini (COURI & PONT, 2000). Comments. Coenosiini is the largest tribe in the subfamily, with 29 genera in the world. In the Neotropical region, there are 16 genera and 221 species (COURI & PONT, 2000).
Apsil Malloch, 1929 (Figs. 83-102) Type-species. Apsil maculiventris Malloch (orig. des.). Diagnosis. Head flat, male dichoptic; 1 pair of rec1inate orbital setae; frons about as wide as half of head width at vertex; 3-4 inc1inate pairs offrontal setae and 1 rec1inate orbital seta; parafacial setulose; fronto-orbital plate bare; antenna short, inserted at midlevel of eye, apex of flagellomere round; arista with short plumes along its entire length; palpus somewhat dilated apically (Figs. 83-88); eye reduced in size; gena very high (Figs. 89-95); dorsocentral setae 2:3; prealar seta absent; lateral margin of scutellum bare; subbasal and apical pair of scutellar setae long; pro sternum bare; lower proepimeral seta directed downwards; proepistemals 2, directed upwards; intra-alars 1:2; supra-alars 1:2; katepistemum with few setulae on disc, the setae 1: 1: 1 and forming an equilateral triangle; 2 notopleural setae; wing veins bare; vein A,+CuA 2 10ng; sternite 1 bare; male: cercal plate longitudinally divided (Fig. 96), hypandrium tubular (Fig. 97); female: ovipositor long, with microtrichia, tergites 6 and 7 slender (Figs. 98-99); spennathecae elongate. References. COURI, 2000a; COURI & PONT 1999; COURI & PONT, 2000. Monophyly. Probably monophyletic (CARVALHO & COURI, in press), based on four synapomorphies: shape of the head (flat), presence of2 presutural dorsocentral setae, gena high and 4 frontal setae. Geographical record. Chile, Argentina. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
Comments. Among the Coenosiini, Apsil is one of the few endemic genera in southern South America (CARVALHO & COURI, in press). Recently, COURI (2002) described a new species from Chile which was included in the key below.
Key to species (modified from MALLOCH, 1934) 1.
Anepimeron setulose (Argentina) ................................ A.jlavipalpis Malloch, 1934 Anepimeron bare .................................................................................................. 2
2. Ma]e ................................................................................................................... 3 Female .............................................................................................................. 11 3.
Fore tibia thickened apically and with several series of erect lanceolate setae that are about twice as long as tibial diameter (Fig. 100); legs black, fore tarsi testaceousyellow, basal segment compressed, its greatest width exceeding that of apex of tibia, and its upper edge with a fringe of black setae that are decumbent and directed apically (Chile, Argentina) ........................................................... A. pennata Malloch, 1934 Fore tibia without lanceolate setae; fore metatarsus not compressed ...................... .4
4.
Tibiae and tarsi testaceous to fulvous yellow.......................................................... 5 Tibiae black, tarsi rarely partly yellowish ................................................................ 8
5.
Stigma and both crossveins of wing infuscated, and a small dark mark on cell r4+5 almost below apex of vein ~+3 which usually has a darker longitudinal central streak in it resembling a rudimentary vein; palpus not noticeably dilated at apex; abdomen fuscous, intermediate tergites darkest, all with a complete pale grey apical fascia; hind femur without a clump of erect hairs on posterior surface near base .......................... 6 Only crossveins infuscated and no dark mark on cell r4+5' sometimes with a very faint brown spot at the end of vein Sc ........................................................................... 7
6.
Palpus conspicuously dilated at apex; abdomen grey dusted, tergites 1+2-4 each with a pair of large subtriangular black marks and a pair of sub linear marks on disc of
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tergite S (Fig. 101); hind femur with a clump of fine erect hairs on posterior surface near base (Chile) .................................................. A. maculiventris Malloch, 1929 Pal pus not conspicuously dilated at apex; abdomen with brown marks as in Fig. 102; hind femur without a clump of fine erect hairs on posterior surface near base (Chile) ........................................................................................ A. diminuta Couri, 2002 7. Mid tibia with only one seta on middle; hind tarsus over three-fourths as long as its tibia and with the usual short hairs on posterior surface (Chile, Argentina)
................................................................. ..............A. maculipennis Malloch, 1934 Mid tibia with two setae on middle, one on anterodorsal and the other on posterior surface; hind tarsus less than two-thirds as long as its tibia, basal segment with longer, denser and finer hairs than usual on posterior surface (Chile, Argentina)
.......................................................................................... A. biseta Malloch, 1934 8.
Legs entirely black; hind tarsus much over two-thirds as long as its tibia; stemite S not glossy, and without any outstanding setae or process (Chile) ................ .
....................................................................................... .A. atripes Malloch, 1934 Legs never entirely black, apices of all tarsi, or at least of the fore pair, yellowish; hind tarsus less than.two-thirds as long as hind tibia; stemite Slargely glossy and of abnormal structure ...................................... '" ...................................................................... 9 9.
Hind trochanter with a strong spine on inner side that is abruptly bent near base; palpus slightly thickened apically; disc of scutellum and a similarly broad patch on hind margin of mesonotum conspicuously greyish-white dusted (Chile, Argentina)
................................................................................. ......... A. apicata Malloch, 1934 Hind trochanter unarmed but each hind coxa with two strong straight apical spines on inner side; palpus broadly and rather abruptly dilated on almost apical half; scutellum and center of hind margin of mesonotum not too noticeably greyish or brownish dusted ................................................................................................................ 10
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Part II - Apical Groups
10. Processes of sternite 5 with short stiff black apical setae; palpus spoon-shaped (Argentina) .................................................................. ...... .A. dilata Malloch, 1934 Processes of sternite 5 with short dense yellow hairs at apex, some black hairs on inner sides; palpus paddle-shaped (Chile, Argentina) ...... A . spatulata Malloch, 1934 11. Wing with a small dark spot on R 4+5 cell (Chile, Argentina) ...................... ..
........................................................................................ A . biseta Malloch, 1934 Wing without a small dark spot on R4+5 cell.. ......................................................... 12 12. Fore femur with posteroventral series of setae not stouter than those of posterodorsal series; fore coxa with two series of quite long and moderately strong black setae on entire anterior side; dark brown central vitta on mesonotum extending to posterior margin (Chile) .................................................................... A. atripes Malloch, 1934 Fore femur with posteroventral series of setae much stouter than those of posterodorsal series, curved, fitted for grasping prey; fore coxa with a few similarly curved setae on apical third ........................................................................................................... 13 13. Palpus very slightly thickened at apex; thorax with a conspicuous lavender-grey dusted mark on disc of scutellum that extends forward on center of mesonotum as far as length of scutellum (Chile, Argentina) ............................... A . apicata Malloch, 1934 Pal pus very distinctly dilated at apex, usually on apical half; thorax without such conspicuous pale dusted central mark, dust yellowish-grey or brownish-grey, scutellum sometimes entirely dark brown .............................................................................. 14 14. Scutellum grey dusted on disc, black or dark brown on sides (Chile, Argentina)
....................................................................................... A. spatulata Malloch, 1934 Scutellum entirely dark brown (Argentina) ......................... A. dilata Malloch, 1934
208
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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!83
~84
0,3mm
~85 ~86
&87
~
.) 88
0,1 mm
Figs. 83-96, characters of Apsil (modified from CO URI, 2000a): 83, A. apicata, female palpus; 84, A. atripes, female palpus; 85, A. dilata, female palpus; 86, A. maculiventris, female palpus; 87, A. pennata, female palpus; 88, A. spatulata, female palpus; 89, A. apicata, female head, frontal view; 90, A. atripes, female head, view frontal; 91, A. macltlipennis, female head, view frontal; 92, A. dilata, female head, view frontal; 93, A. macltlipennis, female head, view frontal; 94, A. pennata, female head, view frontal; 95, A. spatulata, female head, view frontal; 96, A. apicata, cereal plate, posterior view.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
209
Part II - Apical Groups
99 98
i:'I1:-,~:' "
,:,
':
Figs. 97-102, characters of Apsil (modified from COURI, 2000a, COURI, 2002; MALLOCH, 1934): 97, A. atripes, aedeagus, lateral view; 98, A. apicata, ovipositor, dorsal view; 99, A. apicata, ovipositor, ventral view; 100, A. pennata, male foreleg; 101, A. maculiventris, abdomen, dorsal view; 102, A. diminuta, abdomen, dorsal view. Abbreviations: hyp, hypandrium; tg, tergite.
210
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Bithoracochaeta Stein, 1911 (Figs. 103-117) Type-species. Anthomyia despecta Walker (orig. des.) = leucoprocta Wiedemann. Diagnosis. Male: holoptic, eyes bare; at the level of anterior ocellus separated by 0.35-0.36 of head width; margin of eyes convergent toward lunule; arista with basal cilia measuring about twice the basal width of the arista; flagellomere with pointed apex; 1 pair of reclinate orbital setae; dorsocentral setae 1:2; pre sutural acrostichal setae developed; prealar seta absent; katepisternal setae as in Figs. 103, 107, 109, 110, 111, 113, 114; proepimerals 2, the lower one downcurved; wing veins bare; hind tibia with one long anteroventral, anterodorsal and posterodorsal setae on middle third, some species also with a long posteroventral seta; sternite 1 bare; female: vertical internal seta long, backward directed and external vertical short seta, outward directed; interfrontal setae absent; proboscis with prestomal teeth developed, palpus with strong setae; sternite 1 bare; male: hypandrium tubular (Figs. 106, 108); female: ovipositor long; microtrichia present over entire ovipositor; cerci longer than epiproct and similar in length to hypoproct (Fig. 104). References. MOTTA & COURI, 1999; COURI & PONT 1999; COURI & PONT 2000. Monophyly. COURI & PONT (2000) in their cladistic analysis to the tribe, pointed out that the loss of the postsutural intra-alar seta is a synapomorphy for Bithoracochaeta and Insulamyia. In their analysis no autapomorphy was recognised for Bithoracochaeta. Geographical record. Mexico, Costa Rica, Cuba, Jamaica, Puerto Rico, Venezuela, Guyana, Suriname, Peru, Brazil, Paraguay, Uruguay, Argentina; United States of America. Comments. Bithoracochaeta pad/era Giglio-Tos, 1893 were not included in the key because the characters in the original descriptions were not sufficient. COURI & MARQUES (2001) recently described B. equatorialis from Ecuador, a species near B. flavicoxa Malloch, but it also was not included in the key.
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Key to species (modified from MOTTA & COURI, 1999) 1.
Hind tibia with 3 long setae on middle third, parafacial not visible in lateral view........ 2 Hind tibia with 4 long setae on middle third; parafacial visible in lateral view............ 3
2.
Fore coxa and femur dark brown; katepistemals as in Fig. 115; ovipositor with tergites 6-8 as in Fig. 116a (Costa Rica) .................................. .B. nigricomis Malloch, 1934 Fore coxa yellow and fore femur yellow except for slightly brown apex; katepistemals as in Fig. 105; tergites 6-8 thinner as in Fig. 117 (Costa Rica, El Salvador, Guyana, Brazil) ........................................................... .................B.flavicoxa Malloch, 1934
3.
Mid and hind femora yellow with a light brown band; katepistemals as in Fig. 107; male: tubular hypandrium short (Fig. 106) (Brazil, Argentina) .......................... .
.................................................................................... .B. atricomis Malloch, 1934 Mid and hind femora withoutthe light brown band; male: tubular hypandrium long ........4 4. Hypandrium without spines (Fig. 108); fore femur yellow; katepistemals as in Fig. 109 (Mexico, Cuba, Costa Rica, Panama, Brazil, Paraguay, Uruguay, Argentina; United States of America) ............................................... B. Zeucoprocta (Wiedemann, 1830) Hypandrium with spines; katepisternals different from those in Fig. 109 ...................... 5 5
Mid and hind femora brown .................................................................................. 6 Mid and hind femora differently coloured ............................................................... 7
6.
Mesonotum with a well-marked median vitta, reaching apex of scutellum; antenna with scape and pedicel and the basal extremity of flagellomere dark brown, rest of antenna yellow; ocellar triangle with golden pruinescence; katepistemals as in Fig. 110; cercal plate with the posterior margin without an incision (Puerto Rico, Brazil)
................................................................................ B. varicomis (Coquillett, 1900) Mesonotum with the median vitta hardly visible; antenna brown with silver pruinescence; ocellar triangle with silver pruinescence; katepisternals as in Fig. 111; cercal plate with 212
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
3 incisions on posterior margin, the median one slightly little deeper than the others (Fig. 112) (Brazil) ........................................... .B. maricaensis Couri & Motta, 1994 7.
Male: flagellomere brown with silver pruinescence; fore coxa yellow with the posterior surface brown; arista with cilia over its entire extension; katepistemals as in Fig. 113 (Brazil) .................................. ..................................B. plumata Albuquerque, 1955 Male: flagellomere yellow; fore coxa brown with yellow apex; arista with cilia only on basal two-thirds; katepistemals as in Fig. 114 (Guyana, Brazil, Peru)
......................................................................................... .B. annulata Stein, 1911
';j103
~ 105
~09 ~111 ~4 112
116 108
104
J;j15
Figs. 103-117. characters of Bithoracochaeta (modified from MOTTA & COURI, 1999): 103, B. nigricornis, katepisternum; 104, B. varicornis, ovipositor, dorsal view; 105, B. flavicoxa, katepisternum; 106, B. atricornis, aedeagus; 107, B. atricomis, katepisternum; 108, B. leucoprocta, aedeagus; 109, B. leucoprocta, katepisternum; 110, B. varicornis, katepisternum; 111, B. maricaensis, katepisternum; 112, B. maricaensis, cereal plate and surstyli; 113, B. plumata, katepisternum; 114, B. annulata, katepisternum; 115, B. nigricornis, katepisternum; 116, B. nigricomis, ovipositor, a, dorsal view, b, ventral view; 117, B. flavicoxa, katepisternum.
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Coenosia Meigen, 1826 Type-species. Musca tigrina Fabricius (WESTWOOD, 1840: 143). Diagnosis. Male dichoptic; frons paralell-sided, longer than wide; 1 pair of reclinate orbital setae; prestomal teeth developed; labella reduced; prealar seta absent; lower proepimeral seta downward directed; dorsocentral setae 1: 3; katepistemals 1: 1: 1, forming an equilateral triangle; lower calypter about 1.8 times the upper one; hind femur with anterodorsal and dorsal apical setae (posterodorsal setae lacking); hind tibia with one anterodorsal submedian seta and usually one much shorter anteroventral seta nearer the apex; stemite 1 bare; male: hypandrium tubular; female: ovipositor long with microtrichia, tergites and stemites slender; 3 spermathecae. References. STEIN, 1911; MALLOCH, 1934; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. COURI & PONT (2000) did not find derived characters for Coenosia. Geographical record. Mexico, Venezuela, Guyana, Ecuador, Bolivia, Brazil, Uruguay, Chile, Argentina; cosmopolitan. Comments. Coenosia needs a taxonomic revision in the Neotropical Region. Many of its species were no longer recognised after their original descriptions, and were not included in the keys. At the end, a list with all species not included in the key is presented.
Key to species (modified from STEIN, 1911) 1.
Legs black, at most the trochanter and very rarely the mid tibia slightly reddish ........ .2 At least the posterior tibia or part ofthe femora yellow........................................... 8
2.
Mid tibia with 1-2 anteroventral setae (Peru, Bolivia) ........ c. crassicauda Stein, 1911 Mid tibia without anteroventral setae ........................................................................ 3
3.
Crossveins r-m and dm-cu slightly fuscous ............................................................ .4 Crossveins not infuscated ........................................................................................ 5
4.
Wing with whitish longitudinal marks; abdomen with large clouds; claws and pulvilli very long (Peru) ...................................................................... c. picta Stein, 1911
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Wings with no longitudinal whitish marks; abdomen with very small clouds; claws and puvilli not enlarged (Peru) .................................................. C vittithorax Stein, 1911 5.
Antenna long, reaching or almost reaching epistoma; flagellomere about three times the length of pedicel. .............................................................................................. 6 Antenna shorter than described above, not reaching epistoma; flagellomere at most, 2.5 times the length ofpedicel. .................................................................................. 7
6.
Frons velvet black; orbits and median vitta not very visible Chile) .................. . ........................................................................... .................. C. atrifrons Stein, 1911 Frons grey; orbits and median vitta distinct (Chile) .................. C procera Stein, 1911
7.
Male: terminalia on dorsal view so wide as the last abdominal segment and surpassing it by about half of its length; species with about 5 mm long (Peru, Chile) .............................................................................................. C. tumida Stein, 1911 Male: terminalia on dorsal view thinner than the last abdominal segment; species with about 4 mm long (Peru, Bolivia) ....................................... C setiventris Stein, 1911
8.
Mid femur entirely or partly yellow........................................................................ 9 Mid femur entirely black ....................................................................................... 12
9.
Mid tibia from middle to apex of posterior surface with 4 very long setae (Peru) ....................................................................... ..................... C. pilitibia Stein, 1911 Mid tibia without these 4 setae ............................................................................ 10
10. All coxae yellow (Venezuela, Guyana, Peru, Bolivia, Brazil) .. C plumiseta Stein, 1911 All coxae grey..................................................................................................... 11 11. Claws and pulvilli very short (Argentina, Chile) ............. C rotundiventris Stein, 1911 Claws and pulvilli distinctly long ............................................................................. 12 12. All tibiae yellow; marks on abdomen very faint (Chile) .............. C iniqua Stein, 1911 Fore tibia black, except on base; marks on abdomen conspicuous (Chile) ............................................................................................. C. strenua Stein, 1911 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Key to species (modified from MALLOCH, 1934) 1.
Mesonotum with a pair of setae placed a short distance in front of the usual presutural pair that are over one-third as long as the latter and as long as the strong pair of acrostichals in line between them, dusting brownish grey, dense and uniform, with hardly a trace of darker vittae along dorsocentral series of setae; abdomen dull, densely brownish-grey dusted, with a pair of small brown, widely separated spots beyond middle on each tergite from third to fifth, and long strong setae on the last two; fore tarsus slender, about 1.25 as long as fore tibia, the basal segment half as long as the tibia (male) (Chile ).........................................................
c. chaetosa Malloch, 1934
Mesonotum with at most a pair of setulae in front of the usual pre sutural pair of dorsocentrals that are not one-fourth as long as the latter; dusting grey, vittae along dorsocentral series of setae quite distinct; abdomen slightly shining, and with larger fuscous paired spots; fore tarsus stouter, shorter, and with the basal segment not nearly half as long as its tibia ................................................................................... 2 2.
Mesonotum with a central brown line and two broad shining brown vittae that extend from the dorsocentrals laterally to or over the bases of the intra-alars; posterior margin of mesopleuron browned (Chile )....................................
c. inaequalis Malloch, 1934
Mesonotum without a central brown line, brown vittae along series of dorsocentral setae narrow, not extending laterally over midway to the intra-alars; posterior margin of mesopleuron grey (Argentina, Chile?) ......................... C. unifonnis Malloch, 1934
Species of Coenosia not included in the key and their respective geographical distribution
216
Species
Geographical Distribution
C. argentifrons (Malloch, 1934) (*)
Chile, Argentina
C. aurifera (Malloch, 1934) (*)
Argentina Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
C. benoisti Seguy, 1932
Ecuador
C. camorinensis Albuquerque, 1956
Brazil
C. cothurnata (Bigot, 1885)
Chile
C. curviventris Albuquerque, 1959
Brazil
C. enormis Albuquerque, 1956
Brazil
C. gaminarai Carvalho & Pont, 1993
Uruguay
C. inusitata (Malloch, 1934) (*)
Chile
C. latitibia Albuquerque, 1957
Brazil
C. leucotrichia Albuquerque, 1956
Brazil
C. longipede Albuquerque, 1956
Brazil
C. minuscularis Albuquerque, 1956
Brazil
C. neotropica Carvalho & Pont, 1993
Peru, Bolivia
C. noniesmaculata Gaminara, 1930
Uruguay
C. plumbea Albuquerque, 1954
Brazil
C. projecta (Malloch, 1934) (*)
Argentina
C. tarsata (Snyder, 1957)
Argentina
C. tinctipenis Albuquerque, 1956
Brazil
C. trichophthalma Albuquerque, 1959
Brazil
C. wulpi Pont, 1972
Mexico
(*) appearing in Austrocoenosia in MALLOCH (1934)
Unplaced species of Coenosia sensu CARVALHO et al. (1993) Species
Geographical Distribution
C. tenuior (Walker, 1853)
Brazil
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Cordiluroides Albuquerque, 1954 (Figs. 118-120, Plate 2) Type-species. Cordiluroides listrata Albuquerque (orig. des.). Diagnosis. Male dichoptic; eyes with cilia; 1 pair of reclinate orbital setae; antennal insertion placed well above mid-level of eye (Fig. 118); arista with short cilia; prestomal teeth developed; prealar seta absent; lower proepimeral seta downward directed; mesonotum with two brown vittae between the dorsocentral and the intra-alar row of setae; dorsocentral setae 1:3; katepistemals 1: 1: 1, forming an equilateral triangle; anepimeron bare; lower calypter about twice the lower one; wing veins bare; veins R4+5 and M parallel apically (Fig. 119); hind femur with 3 preapical setae, on anteroventral, dorsal and posterodorsal surfaces; hind tibia with one anterodorsal and one posterodorsal median to submedian, as well as one preapical dorsal seta (Fig. 120); stemite 1 bare; male: hypandrium tubular; female: ovipositor long with microtrichia. References. COURl & PAMPLONA, 1992; COURI & PONT 1999; COURI & PONT 2000. Monophyly. The systematic position of this genus among the Coenosiini was discussed by COURI & PAMPLONA (1992). More recently, COURl & PONT (2000), in a cladistic analysis of the tribe, indicated one synapomorphy for Cordiluroides + Neodexiopsis + HaroZdopsis (hind femur with three preapical dorsal setae). Geographical record. Mexico, St. Vincent Is., Colombia, Peru, Bolivia, Brazil. Comments. Five species are ascribed in this genus, three of them occurring in Brazil.
Key to species (modified from COURI & PAMPLONA, 1992) 1.
General coloration reddish; palpus light brown; coxa with blackish cloud near apex (Mexico) ............................................................... ............
c. bicolor (Bigot, 1885)
General coloration brown; palpus dark brown to black; coxa without blackish cloud near apex .............................................................................................................. 2 2.
Abdomen with a median brown vitta and a brown spot on lower half of the each segment (Mexico, St. Vincent Is., Colombia, Peru, Bolivia) ............. c. insularz's (Williston, 1896)
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Abdomen different coloured ................................."................................................. 3 3.
Arista, on basal half, with short cilia (cilia longer than the basal diameter of arista) and bare on apical half (Brazil) .................................... C geniculata (Macquart, 1851) Arista with cilia not as above ................................................................................. 4
4.
Ocellar setae absent; legs yellow, with apical half of femora and tarsi light brown; male: terminalia prominent (Brazil) ..................... C megalopyga Albuquerque, 1954 Ocellar setae present; legs dark brown with the femora-tibial junctions yellow; male: terminalia not prominent (Brazil) ................................. C listrata Albuquerque, 1954
118
~9
~o
2mm
1 mm
Figs" 118-120, characters of Cordiluroides (modified from ALBUQUERQUE, 1954d; COURI & PAMPLONA, 1992): 118, C. geniculata, head, lateral view; 119, C. listrata, wing; 120, C. listrata, hind femur and hind tibia.
Insulamyia Conri, 1982 Type-species. Insulamyia inusitata Couri (mon.). Diagnosis. Male dichoptic; 1 pair of reclinate orbital setae; prestomal teeth developed; labella reduced; acrostichal hairs in two to four not linear series; prealar seta absent; lower proepimeral seta downward directed; dorsocentral setae 2:3; katepisternals 1: 1: 1, forming an equilateral triangle; lower calypter about twice the upper one, which is truncate; fore tibia with long apical setae; hind tibia with long apical anterior, posteroventral and anteroventral setae; sternite 1 bare; male: hypandrium tubular; female: ovipositor long with microtrichia. References. COURI, 1982a; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. Not yet established. Geographical record. Brazil. Comments. Monobasic genus: Insulamyia inusitata Couri, 1982. Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Neodexiopsis Malloch, 1920 (Figs. 121-134, Plate 2) Type-species. Dexiopsis basalis Stein (orig. des.). Diagnosis. Male dichoptic; 1 pair of reclinate orbital setae; prestomal teeth developed; I abell a reduced; prealar seta absent; lower proepimeral seta downward directed; dorsocentral setae 1:3; katepisternals 1: 1: 1, forming an equilateral triangle; lower calypter about twice the upper one; hind femur with three preapical setae on anterodorsal, dorsal and posterodorsal surfaces (except N. micans and N. crassicrurus which exhibits a considerable modification of the shape and accessory bristling of hind femur); sternite 1 bare; male: hypandrium tubular; female: ovipositor long with microtrichia. References. SNYDER, 1958; COURI & ALBUQUERQUE, 1979; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. COURI & PONT (2000) pointed out one synapomorphy for Cordiluroides + Neodexiopsis + Haroldopsis - hind femur with three preapical dorsal setae. No autapomorphic character was indicated for Neodexiopsis. Geographical record. Mexico, Guatemala, Costa Rica, Panama, Jamaica, Puerto Rico, St. Vincent Is., Colombia, Venezuela, Peru, Bolivia, Ecuador, Galapagos Is., Brazil, Paraguay, Uruguay, Argentina, Chile, Juan Fernandez Is.; Nearctic region. Comments. The genus needs an extensive revision. Bithoracochaeta sociabilis Blanchard, 1937 was transferred to Neodexiopsis by COURI & MOTTA (2000) and not included in the key below. The type-material of Bithoracochaeta sociabilis is lost (MOTTA & COURI, 1999). Recently, PONT (2001) transferred Coenosia diaphana Stein, 1911 and C. incurva Stein, 1911 to this genus. Available keys were transcribed with modifications, and the species that do not appear in any of the key, are listed followed by their known geographical distribution.
Key to species (modified from COURI & ALBUQUERQUE, 1979) 1.
Male with the anal angle of wing forming a prominent lobe (Figs. 126-134); female with the anal angle truncate ..................................................................................... 2 Both sexes with a round anal angle ........................................................................ .4
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2.
Palpus yellow with the basal half darker; antenna brown with apex of pedicel and base of flagellomere yellow; stemite 5 with a row oflong setae, forming a tuft (Fig. 121) (Brazil) ........................................ N. barbiventris Couri & Albuquerque, 1979 Palpus brown; antenna different from above; stemite 5 without any long setae ........... 3
3.
Antenna brown with apex of pedicel shiny in certain lights; mid femur entirely yellow; fore femur on anterior surface without any long seta at base; fore tibia with a long posterior seta (Brazil, Argentina) ................................... .N. neoaustralis Snyder, 1957 Antenna dark brown with apex of pedicel and base of flagellomere yellow; mid femur with the apical third brown, fore femur on anterior surface with a strong seta at base; fore tibia with an submedian anterior seta (Brazil) ...... ..N. parvula Albuquerque, 1958
4.
Apical pair of scutellar setae about half as long as basal pair..................................... 5 Apical pair of scutellar setae as long as basal pair................................................... 9
5.
Fore tibia with short ventral and posteroventral setae apically (Fig. 122); abdomen with a brown median vitta and without lateral clouds (Brazil) .......................... . ............. ....... ....... ...................................... N. pectinata Couri & Albuquerque, 1979
Fore tibia without short ventral and posteroventral setae apically; abdomen with brown lateral round marks and with a brown median vitta................................................. 6 6.
Thorax with a median brown vitta; mid femur on anterior surface with one seta or bare; hind tibia with one anterior to anterodorsal seta and no setae on anteroventral surface .................................................................................................................. 7 Thorax with more than one brown vitta; mid femur on anterior surface with 3-4 setae; hind tibia with 2 anterodorsal setae and one setae on anteroventral surface .............. 8
7.
Proboscis yellow; mid femur totally yellow; fore femur with a row of anterodorsal setae; fore and mid femora and tibiae with short and fine ventral cilia (Peru, Brazil; North America) ............................... ,. ............... .............. .N. calopyga (Loew, 1872)
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Part II - Apical Groups
Proboscis brown; mid and hind femora yellow with brown apices; fore femur with an anteroventral and anterodorsal row of setae; fore and mid femora and tibiae without short and fine ventral cilia (Brazil) ..................N. subtilis Couri & Albuquerque, 1979 8.
General colour yellow; antenna yellow; fore femur totally yellow and with a complete row of anterodorsal setae (Brazil) ................. N. elegans Couri & Albuquerque, 1979 General colour grey; antenna grey with apex of pedicel lighter; fore femur mostly yellow, brown on apical dorsal half; mid femur with no apical anterodorsal seta (Brazil)
.................................................................N. vulgaris Couri & Albuquerque, 1979 9.
Hind tibia without a median or submedian anteroventral seta ................................ .1 0 Hind tibia with a median or submedian anteroventral seta ....................................... 13
10. Mid femur with two anterior setae; hind tibia on anterior to anterodorsal sUlface with long and fine cilia (Fig. 123); mesonotum with 5 brown vittae, the median one reaching apex of scutellum (Brazil) ..........................................N. rustica Albuquerque, 1956 Mid femur with one median or submedian anterior seta; hind tibia with no long anterior setae; mesonotum with vittae not as below............................................................ 11 11. Head and thorax grey pruinose; legs yellow with the dorsal surface of fore femur, apex of mid femur and apical third of hind femur brown; mid femur with a short median anterior seta (Brazil) .................................. N.flavipalpis Albuquerque, 1956 Head and thorax golden pruinose; legs dark brown with trochanters and femora-tibial junctions yellow; mid femur with a long median anterior seta.................................. 12 12. Antenna dark brown, with apex of pedicel and base of flagellomere yellow; flagellomere with round apex; last abdominal segment grey; mesonotum with three brown vittae, the median one reaching scutellum (Brazil) .... .N.fulvifrontis Couri & Albuquerque, 1979 Antenna entirely dark brown (some specimens with apex of pedicel yellow); flagellomere with pointed apex; last abdominal segment yellow; mesonotum with four brown vittae, those coincident with the dorsocentral row of setae reaching scutellum (Brazil)
................................... ........................................ N. paulistensis Albuquerque, 1956 222
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13. General colour dark brown; antenna, palpus and legs dark brown; halter with knob black; calypteres brown in male and yellow with brown margins and fringe in female; abdomen without brown spots (Chile, Argentina) ....... .N. nigerrima (Malloch, 1934) General colour grey, sometimes with yellow pruinescence; halter with knob yellow to light brown; calypteres yellowish-white; abdomen frequently with a pair of brown round lateral spots ............................................................................................... 14 14. Mesonotum without dorsal vittae .......................................................................... 15 Mesonotum with dorsal vittae ............................................................................. 17 15. Integument with golden not shiny pruinescence; legs and halter orange; abdomen without lateral brown spots; costal spine present (Brazil) ................................... .
.......... .............. ..... ........ .............. ...... ........ ..... .....N. itatiaiensis Albuquerque, 1954 Integument covered with grey pruinescence; legs entirely yellow or with the dorsal surface of fore femur brown; halter yellow; abdomen with brown lateral spots; costal spine absent. .......................................................................................................... 16 16. Legs yellowish with the anterior surface offore femur brown; palpus brown; epistoma not prominent; pretarsus witht a basal ventral seta (Brazil) .................................... . ...................................................... .................. .N. albisquama (Albuquerque, 1959)
Legs entirely yellow; pal pus yellow on basal two-thirds and brown on apical third; epistoma prominent; pretarsus with a basal ventral seta (Brazil) ........................... . ....... .............. ..... .... .......... .... '" .................. .... .N. plumbifrons (Albuquerque, 1956)
17. Antenna black with the apex of pedicel lighter...................................................... .18 Antenna entirely dark brown or with another colour.. ............................................. 24 18. Mesonotum with the dorsal vittae coincident with the dorsocentral row of setae reaching the insertion of the basal pair of scutellar setae ....................................................... 19 Mesonotum with the vittae coincident with the dorsocentral row of setae not reaching scutellum .............................................................................................................. 21 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
19. Parafacial, fronto-orbital plate, frons and gena with a grey pruinescence; mid femur without an anterior to anteroventral row of setae on basal half (Brazil) ......................................................................N. alacris Couri & Albuquerque, 1979
Parafacial, fronto-orbital plate, frons and gena with a strong yellow pruinescence; mid femur with an anteriorto anteroventral row of setae on basal halL .......................... 20 20. Antenna entirely dark brown; palpus dark brown with golden pruinescence at apex (Brazil) ............................................................ ..N. tinctifacies (Albuquerque, 1958) Antenna dark brown with apex of pedicel shiny; palpus brown (Brazil) .............................................................. N. novissimo Couri & Albuquerque, 1979
21. Palpus entirely brown, or brown on basal third, legs yellow, except fore femur and apices of mid and hind femora brown .................................................................. 22 Palpus dark brown; legs dark brown, except apices of femora and bases of tibiae yellow.................................................................................................................. 23 22. Antenna with apex of flagellomere round; mesonotum with four large brown vittae; fore tibia with a preapical seta on anteroventral and anterodorsal surfaces (Chile, Juan Fernandez Is.) ..........................................................N. annulipes (Macquart, 1843) Antenna with apex offlagellomere pointed; mesonotum with three fine brown vittae; fore tibia with a preapical anterodorsal seta; no preapical seta on anteroventral surface (Brazil) ..........................................................N. mesofulvata (Albuquerque, 1959) 23. Vertex brown; fronto-orbital plate with grey pruinescence; mesonotum with a median brown vitta reaching half of scutellum; epistoma nor prominent (Brazil, Argentina)
........................................................................................ N. cirratipila Snyder, 1957 Vertex and fronto-orbital plate black; mesonotum without a dorsal median brown vitta; epistoma prominent (Chile, Argentina) ..........................N. dubia (Bigot, 1885) 24. Antenna long and entirely dark brown; flagellomere about 3.6 times as long as pedicel; mesonotum with five dorsal brown vittae (Brazil) .. N latimaculata (Albuquerque, 1956) 224
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Marcia Souto Couri & Claudio Jose Barros de Carvalho
Antenna with pedicel grey and flagellomere entirely yellow; flagellomere about 2.6 times as long as pedicel; thorax with three dorsal brown vittae (Brazil)
.......................................................................N. xanthopoda Albuquerque, 1956
O,4mm
0,4 mm
Figs. 121-123, characters of Neodexiopsis (modified from COURI & ALBUQUERQUE, 1979): 121, N. barbiventris; 122, N. pectinata, fore femur and fore tibia; 123, N. rustica, hind femur and hind tibia. Abbreviation: ar, arista.
Key to species (modified from SNYDER, 1957b) 1.
Apical scutellar setae absent or, if present less than 0.7 as long as the subbasals ...... 2 Apical scutellar setae as long as or longer than the subbasals ................................ 10
2.
Male ................................................................................................................... 3 Female ................................................................................................................ 8
3.
Fore coxa fuscous ...................................................................................................... 4 Fore coxa yellow to fulvous ................................................................................... 5
4.
Hind femur with a distinct anteroventral seta opposite the terminal one in the anterodorsal row; in dorsal view the combined length of abdominal tergites 3 and 4 as long as or longer than either tergite 1+2 or 5; hind tibia with a median seta which is situated more anteroventral than anterior (Argentina) .............. ............ .N. quintivena Snyder, 1957
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Hind femur without an anteroventral seta opposite the terminal one in the anterodorsal row; in dorsal view the combined length of abdominal tergites 3 and 4 distinctly less than the length of either tergites 1+2 or 5; hind tibia with a median seta which is situated more anterior than anteroventral (Argentina) ...... .N. lunatisigna Snyder, 1957 5.
Abdomen entirely darkened; basal terminalia segment with numerous, long, slender hairs which are at least as long as length of tergite 4; palpus and tarsi yellow to fulvous (Argentina) ................................................. ............... .N. intoniclunis Snyder, 1957 Abdomen partly yellow to fulvous at base or beyond; basal terminalia segment with only the usual short setae; palpus or tarsi or both fuscous to brown .......................... 6
6.
Mid femur without an anterior seta on apical 0.25 to 0.33 in addition to the median anterior seta; abdomen with the clothing setulae on the darkened dorsolateral region on tergites 1+2 and 3 minute, upright, very numerous, and closely placed arising from small dark spots (Argentina) .................................... .... .N. setipuncta Snyder, 1957 Mid femur with a distinct anterior seta beyond the median one; abdomen with the dorsolateral clothing setulae on tergites 1+2 and 3 long, somewhat sparse, and either upright or decumbent ............................................................................................... 7
7.
Mid and hind femora without an apical brown spot; thorax not vittate (Argentina)
.................................................................................... .N. uspallata Snyder, 1957 Mid and hind femora with an apical brown spot; thorax with one or three distinct longitudinal vittae (Argentina) ..........................................N. cacumina Snyder, 1957 8.
Hind tibia with a submedian posterodorsal seta (Argentina) ............................ ..
................................................................................. .N. intoniclunis Snyder, 1957 Hind tibia without a posterodorsal seta ................................................................... 9 9.
Fore femur with a dark posterodorsal cloud on apical 0.25 or more (Argentina)
................................................................................... .N. cacumina Snyder, 1957 Fore femur entirely yellow to fulvous (Argentina) .......... .N. setipuncta Snyder, 1957 226
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10. Male: posterior margin of wing with a deep preanal incision which thus produces a thumb-like projection at anal angle; female: anal angle of wing truncate or sub-truncate.... ll Male and female: anal angle of wing evenly rounded and without a prebasal incision .... 12 11. Hind tibia with a median anteroventral seta; palpus largely yellow (Chile, Uruguay, Argentina) ............................................ ....................... .N. geniculata (Bigot, 1885) Hind tibia without a median anteroventral seta; palpus mostly dark (Brazil, Argentina)
,..................................................... ............................N. neoaustralisSnyder, 1957 12. Hind tibia without any anterior or anteroventral setae or long hairs on middle .......... 13 Hind tibia with either a median anterior or anteroventral seta or with a row of long anteroventral hairs ............................................................................................... 16 13. Fore coxa yellow to fulvous ................................................................................ 14 Fore coxa fuscous ................................................................................................... 15 14. Male: fore femur with many closely placed, long, clothing hairs over most of the posterodorsal to posteroventral surfaces (Argentina) .... .N. neomacrocera Snyder, 1957 Male: Fore femur with only the usual short clothing setulae but with a row of distinct posteroventral setae (Mexico) .................................... .N. macrocera (Wulp, 1896) 15. Hind tibia with a median posterodorsal seta; frons with dense golden yellow pruinescence so that the frontal triangle and the fronto-orbital plate are not at all differentiated from the vitta; apex of abdominal tergite 5 fulvous and in sharp contrast to the remaining dark abdominal tergites (Brazil) ....... ......N croceafrons Snyder, 1957 Hind tibia without a median posterodorsal seta; front with sparse yellow to greyish yellow pruinescence, the triangle and fronto-orbital plate clearly differentiated from the vitta; apex of abdominal tergite 5, concolorous with the remainder of abdomen (Argentina) ................................................................ ...N clavacula Snyder, 1957 16. Fore coxa yellow to fulvous (Argentina) .................................N rava Snyder, 1957 Fore coxa fuscous ............................................................................................... 17 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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17. Flagellomere short, with a distinct angular apical protuberance (Argentina)
........................................................................................... ..N. sima Snyder, 1957 Flagellomere long, without a distinct angulate apical process ................................. 18 18. Male: hind tibia with many long apically curled dorsal to posterodorsal hairs which are 0.25 as long as hind tibia; female: basal 0.2 or more of mid femur and hind femur yellow and sharply contrasting with the apically darkened portion; median brown thoracic vitta extending onto middle of scutellum (Brazil, Argentina)
............................................................................... ..... N. cirratipila Snyder, 1957 Male: hind tibia with only the normal decumbent, short, clothing setulae which are less than 0.05 as long as hind tibia; female: mid and hind femora at base concolorous with the remainder of femora; median thoracic vitta not continued onto scutellum (Panama, ?Jamaica, Ecuador, Peru, Brazil, Paraguay, Argentina; North America)
...................................................................................... ..N. rufitibia (Stein, 1911)
Key to species (modified from SNYDER, 1957a, Puerto Rican species) 1.
Apical scutellar setae at least 0.75 as long as subbasals .......................................... 2 Apical scutellar setae usually much less than 0.66 as long as subbasals ................... 7
2.
Postsutural intra-alar seta 1.. ................................................... .N. rex Curran, 1928 Postsutural intra-alar setae 2 .................................................................................. 3
3.
Male ................................................................................................................... 4 Female ................................................................................................................ 5
4.
Femora mostly or entirely fulvous; hind tibia without a submedian anteroventral seta; abdomen with at least the basal portion of abdominal tergite 1+2 yellow to fulvous
........................................................................... ...N. discolorisexus Snyder, 1957 Femora mostly or entirely fuscous; hind tibia with a submedian anteroventral seta or setula; abdomen entirely fuscous in ground colour.. .... ......N. ditiportus Snyder, 1957 228
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5.
Femora mostly dark ............................................................................................... 6 Femora mostly yellow to fulvous ............................................................................. 9
6.
Hind tibia with a median anteroventral seta or setula; anterior ocellar setae subequal to anterior frontals ..................................................... .... N. ditiportus Snyder, 1957 Hind tibia without a median anteroventral seta; anterior ocellar setae less than 0.5 as long as anterior frontals ........................................... N. discolorisexus Snyder, 1957
7.
Postsutural intra-alar setae 2 .................................................................................. 8 Postsutural intra-alar setae 1.................................................................................. 9
8.
Antenna mostly or entirely fuscous .............................. N. maldonadoi Snyder, 1957 Antenna mostly or entirely fulvous .............................. .. N. neojlavipes Snyder, 1957
9.
Male .................................................................................................................. 10 Female .............................................................................................................. 12
10. Mid tibia with only one long anterior seta and one posterior seta slightly beyond middle; most of the apical setae extremely long and hair like; hind tibia without numerous, long, accessory hairs or setae; 2 stigmatal setae ............ .. N. crispiseta Snyder, 1957 Mid tibia with several long anterior and posterior setae or hairs on apical half; hind tibia with numerous long, accessory hairs or setae; single stigmatal seta or the lower one ciliform .......................................................................................................... 11 11. Mid tibia distinctly thickened on apical two-fifths, with numerous, long, curly anterodorsal and posterodorsal setae on this enlarged portion; in frontal view, the frons with dense silvery white pruinescence, the triangle not differentiated; abdomen with the usual dorsal pruinescence and clothing setulae ..................... . ... .... ....................... ................ ......... ....................... ..N. crassicrurus Snyder, 1957
Mid tibia not usually thickened on apical two-fifths, with 1 anterior to anterodorsal seta, several posterodorsal on median half; in frontal view, frons with dense brownish Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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pruinescence, the triangle differentiated; abdomen with the dorsal surface shiny, without or with only sparse clothing setulae .................................... .N. micans Snyder, 1957 12. Palpus, fore coxa and basal four-fifths or more of mid and hind femora fuscous
......................................................................... ...........N. ebenifemur Snyder, 1957 Palpus, fore coxa and basal four-fifths or more of at least mid and hind femora yellow to fulvous ............................................................................................................. 13 13. Stigmatal and propleural setae duplicated; mid femur with the apical posterior to posterodorsal setae ...................................... .................. N. crispiseta Snyder, 1957 Stigmatal and propleural setae not duplicated; mid femur with a single apical posterior to posterodorsal seta ........................................................................................... 14 14. Mid tibia with 2 submedian posterior setae .................N. crassicrurus Snyder, 1957 Mid tibia with 1 submedian posterior setae ........................ .N. micans Snyder, 1957
Key to species (modified from SNYDER, 1958) Male 1.
Fore femur swollen on basal two-thirds; the posteroventral setae on this portion closed placed and with their apices curved towards apex of femur; the basal three-fifths of posterior surface bare, shiny; fore tibia with a basoventral concavity which has short, thorn-like, anteroventral and posteroventral setulae (Figs. 124-125); flagellomere entirely pale, whitish yellow; humeral callus yellow, contrasting with the infuscated thoracic disc (Costa Rica, Panama) ...... ............... N. hydrotaeiformis Snyder, 1958 Fore femur and tibia not unusually swollen or curved; flagellomere entirely infuscated or fulvous only on basal half or less; humeral callus infuscated, concolorous with disc ...................................................................................................................... 2
2.
Hind tibia without an anteroventral seta near middle ................................................ 3 Hind tibia with a weak or strong anteroventral seta near middle .............................. 7
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3.
Apical scutellar setae very short; hind femur with a very slight, basal, ventral swelling bearing numerous short, upright black spinules (Brazil ?) ......N willistoni Snyder, 1958 Apical scutellar setae long; hind femur not swollen near base and with only the usual short clothing setulae ............................................................................................. 4
4.
Thumb-like extension of the anal area of wing relatively short and broad (Fig. 126) (Costa Rica, Guatemala) ........................................... .N. obtusiloba (Malloch, 1934) Thumb-like extension of the anal area of wing relatively long and slender (Fig. 127128) ...................................................................................................................... 5
5.
Longest hairs on arista at least 0.5 as long as the greatest width offlagellomere; mid and hind tarsi mostly yellow to fulvous; glossy spot on sides of abdominal tergite 3 yellow to fulvous (Costa Rica, Panama, Venezuela) ......... .N. emmesa Malloch, 1934 Longest hairs on arista not longer than its greatest diameter; mid and hind tarsi dark; glossy spot on sides of abdominal tergite 3 dark brown to black ............................. 6
6.
Palpus entirely yellow; sides of abdomen yellowish at base; median anterior to anterodorsal seta on mid tibia short or absent; hind femur with clothing setulae between the anteroventral setae very short, not conspicuous (Mexico) .................... .
.................................... ................. ..... .................. ......... .N. punctulata (Wulp, 1896) Palpus brownish to infuscate, occasionally lighter coloured at base; abdominal ground colour uniformly infuscated; median anterior to anterodorsal seta on mid tibia strong; hind femur with clothing setulae between the anteroventral setae long and conspicuous (Brazil, Argentina) ........................................... ........ .N. neoaustralis Snyder, 1957 7.
Fore femur mostly infuscated, only the apices narrowly yellow to fulvous ................ 8 Fore femur mostly fulvous, if dark in parts, the infuscated area confined to basal half or less, or to a linear posterodorsal streak ............................................................... 9
8.
Fore femur with an almost complete row of short but distinct anteroventral setulae; thumb-like extension of wing almost pointed at apex (Fig. 129) (Panama)
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.....................................................................................N. magnicomis Snyder, 1958
Fore femur without distinct anteroventral setulae or with only a few of the clothing setulae at base distinct; thumb-like extension of wing rounded at apex (Fig. 130) (Ecuador) ...................................................... ....................N. equator Snyder, 1958 9.
Thumb-like extension of wing pointed at apex (Fig. 131) (Costa Rica) ...................... . ................................ ..... ............... ............... ....................... .N. preacuta Snyder, 1958
Thumb-like extension of wing rounded at apex ........................................................... 10 10. Fore femur with a row of short, spine-like, anteroventral setulae on apical two- to three-fifths; thumb-like extension of wing as in Fig. 132 (Peru) ........................ .. ..................................................................................... N. peruviana Snyder, 1958
Fore femur without distinct, spine-like, anteroventral setulae on apical two- to threefifths ..................................................................................................................... 11 11. Longest hairs on arista 1.5 to 2.0 times as long as its greatest diameter; thumb-like extension of wing as in Fig. 133 ........................................................................... 12 Longest hairs on arista shorter than its greatest diameter....................................... 13 12. Thoracic vittae absent or much reduced; legs entirely yellow; abdomen with shiny spot on side of tergite 4 fulvous; tergites 3 to 5 without a median vitta (Brazil) ........................................................................................... ...N. cera Snyder, 1958
Thorax with five distinct vittae presuturally; legs with a dark posterodorsal vitta on fore femur and apical spots on hind femur; abdomen with shiny spot on side of tergite 4 black to brownish black; tergites 3 to 5 with a median vitta (Puerto Rico) ...................................................................................N. priscipagus Snyder, 1958
13. Prebasal incision on posterior margin of wing swallow (Fig. 134) (Mexico; Nearctic region) ..................................................................... ........N. arizona Snyder, 1958 Prebasal incision on posterior margin of wing deep (Fig. 127) ................................ 14 14. Abdominal tergite 1+2 with a small, dark area opposite the glossy spot on side of 232
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tergite 3; the spot on tergite 1+2 with a clump of brush-like clothing setulae in line with the apical row of setulae (Bahamas, Caicos Is.; United States of America)
.................................................................................. ... N. peninsula Snyder, 1958 Abdominal tergite 1+2 without a darkened area, the marginal row of setulae not modified in the form of a brush-like tuft opposite the shiny mark on tergite 3 (Chile, Uruguay, Argentina) ...................................................... N. geniculata (Bigot, 1885) Female 1.
Hind tibia without a submedian anteroventral seta ................................................. 2 Hind tibia with a submedian anteroventral seta......................................................... 6
2.
Flagellomere entirely light yellow (Costa Rica, Panama) ...................... .
.......... ....... ......... .......... ..................... ........... ........ N. hydrotaeifonnis Snyder, 1958 Flagellomere mostly or entirely fuscous ................................................................... 3 3.
Apical scutellar setae very short (?Brazil) .. ...................... .N. willistoni Snyder, 1958 Apical scutellar setae long and strong .................................................................... 4
4.
Longest hairs on arista at least 0.75 as long as width of flagellomere (Costa Rica, Panama, Venezuela) ......................................................... .N. emmesa Malloch, 1934 Longest hairs on arista shorter, or scarcely longer, than its greatest diameter.. ......... 5
5.
Palpus yellow (Mexico) ..............................................N. punctulata (Wulp, 1896) Palpus black to fuscous (Brazil, Argentina) ................ ....N. neoaustralis Snyder, 1957
6.
Fore femur with a row of short but spine-like anteroventral setae on apical half ........ 7 Fore femur without spine-like anteroventral setae on apical half, but some of the basal half may be rather spine-like .................................................................................. 8
7.
Femora yellow, at most very faint dark posterodorsal shadow on fore femur (Peru)
............................................................................... .... N. peruviana Snyder, 1958 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Femora mostly fuscous or black; only basal third to half of mid and hind femora and the extreme apices of fore femur yellow to fulvous (Panama) ......................... .
................ .......... ...... ............ ....................... ............. ...N. magnicomis Snyder, 1958 8.
Palpus entirely testaceous yellow........................................................................... 9 Palpus with at least the tip brown to fuscous .......................................................... 10
9.
Flagellomere entirely fuscous (Chile, Uruguay, Argentina) ............................... .
...... ........ .... ...... ........ ...................... ....... ... ............... .... N. geniculata (Bigot, 1885) Flagellomere broadly reddish fulvous at base (Brazil) ............. .N. cera Snyder, 1958 10. Mid and hind coxae mostly dark; femora usually with a variable darkened portion which is most extensive on fore femora in the form of a posterodorsal to posterior cloud and often with a dark anterodorsal to posterodorsal shadow near apex of mid and hind femora or both (Mexico; United States of America) ...................... ..
.................................................................... ...................... N. arizona Snyder, 1958 Mid and hind coxae yellow or with a very limited dark streak near base; femora entirely yellow (Bahamas Is.; United States of America, Caicos Is.) .................. ..
.......................................................................................... N. peninsula Snyder, 1958 Species of Neodexiopsis not included in the key and their geographical distribution [including N. sociabilis (Blanchard, 1937) recently transferred to this genus by COURI & MOTTA (2000) and N. diaphana (Stein, 1911) and N. incurva (Stein, 1911)
transferred to this genus by PONT (2001)]
234
Species
Geographical Distribution
N. antennata Couri, 1987
Brazil
N. brasiliensis (Walker, 1853)
Brazil
N. brevicomis (Malloch, 1934)
Chile
N. cambuquirensis (Albuquerque, 1954)
Brazil
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N declivis (Stein, 1904)
Bolivia
N devia (Curran, 1934)
Galapagos Is.
N diaphana (Stein, 1911)
Peru
Nflavipes (Williston, 1896)
Puerto Rico
N genupuncta (Stein, 1904)
Peru
N incurva (Stein, 1911)
Bolivia
N lanigera (Stein, 1918)
Colombia
N latifrons (Thomson, 1869)
Ecuador
N lineata (Stein, 1904)
Colombia
N microchaeta (Malloch, 1934)
Uruguay
N oculata (Stein, 1911)
Bolivia
N oscillans (Wulp, 1896)
Mexico
N pilosa (Stein, 1904)
Colombia, Peru
N ponti Couri, 1987
Brazil
N recedens (Stein, 1904)
Peru, Bolivia
N rufipes (Mac quart, 1851)
Brazil, Uruguay
N sociabilis (Blanchard, 1937)
Argentina
N sulina Couri, 1986
Brazil
N tenuicornis (Wulp, 1896)
Jamaica
N truncata (Stein, 1911)
Peru
N vitilis (Giglio-Tos, 1893)
Mexico.
N vittiventris (Albuquerque, 1955)
Brazil
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~
124
~d125
128
127
130
E
131
129
~32
134 Figs. 124-134, characters of Neodexiopsis (modified from SNYDER, 1958): 124, N. hydrotaeiformis, male fore femur, anterior view; 125, N. hydrotaeiformis, male fore femur, posterior view; 126, N. obtusiloba, anal margin of wing, male; 127, N. peninsula, anal margin of wing, male; 128, N. neoaustralis, anal margin of wing, male; 129, N. magnicornis, anal margin of wing, male; 130, N. equator, anal margin of wing, male; 131, N. preacuta, anal margin of wing, male; 132, N. peruviana, anal margin of wing, male; 133, N. cera, anal margin of wing, male; 134, N. arizona, anal margin of wing, male.
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Notoschoenomyza Malloch, 1934 Type-species. Notoschoenomyza chrysiceps Malloch (orig. des.). Diagnosis. Male dichoptic; 1 pair of rec1inate orbital setae; prestomal teeth developed; labella reduced; prealar seta absent; lower proepimeral seta downward directed; dorsocentral setae 2: 3; katepisternals 1: 1: 1, forming an equilateral triangle; calypteres subequal in length; hind tibia with at least 2 anterodorsal and 2 posterodorsal setae, one median and the other supramedian; preapical setae on anterodorsal and dorsal surfaces, sternite 1 bare; male: hypandrium tubular; female: ovipositor long with microtrichia., tergites and sternites slender; 3 spermathecae. References. MALLOCH, 1934; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. COURI & PONT (2000) indicated 4 characters for the genus, all considered as homoplasies: enlargement of palpus apex; presence of an anterior to anterodorsal seta on fore tibia; presence of a supramedian posterodorsal seta on hind tibia; posterior concavity on cercal plate absent. Geographical record. Peru, Bolivia, Chile, Juan Fernandez Is., Uruguay, Argentina. Comments. The genus has seven species, all of them restricted to South America. It is likely that the genus also occurs in Central America. Recently, PONT (2001) transferred N. apicalis (Stein, 1911) to Spathipheromyia.
Key to species (modified from MALLOCH, 1934) 1.
Palpus fulvous-yellow; apices of all tibiae reddish-yellow; male: sternite 5 with processes broadly exposed and entirely grey dusted (Chile )........ .N. chrysiceps Malloch, 1934 Palpus black or brownish-black; tibiae not noticeably pale at apices; male: processes of sternite 5 not as broadly exposed nor entirely grey dusted ................................. .2
2.
Pale grey dusted species; abdomen entirely grey dusted and dull, with a dark dorsocentralline on all tergites and two pairs of dark spots on tergites 3-5, only the inner one visible from above, tergite 6 with a large discal black spot (Uruguay)
.................................................................................... .N. suljuriceps Malloch, 1934 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Dark species; abdomen with grey dust broken up on dorsum, surface distinctly shining, the dark central line and lateral spots not sharply defined (Argentina)
.............................................................................. .N. immaculata (Walker, 1836)
Species of Notoschoenomyza not included in the key and their respective geographical distribution Species
Geographical distribution
N. annulata (Stein, 1911)
Peru
N. costata Snyder, 1957
Argentina
N. kuscheliHennig, 1955
Juan Fernandez Is.
N. spinicosta (Stein, 1904)
Peru, Bolivia
Oxytonocera Stein, 1911 (Fig. 135) Type-species. Coenosia nigrohalterata Stein (mon.). Diagnosis. Male dichoptic; 1 pair of reclinate orbital setae; labella not reduced; apex of flagellomere modified (Fig. 135); prealar seta absent; lower proepimeral seta downward directed; dorsocentral presutural setae 2 strong and more or less subequal, postsuturals 3; katepisternals 1: 1: 1, forming an equilateral triangle; lower calypter about twice the upper one; anal vein long; fore tibia with a median seta on posterior surface; hind tibia on median third with a seta on anterodorsal surface and a short one on anteroventral surface; sternite 1 bare. Reference. STEIN, 1911; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. Although belonging to the monophyletic group "Coenosia", COURI & PONT (2000) indicated 6 characters for the genus, all considered as reversals: frontal setae less than 4; labella well developed; dorsocentral pre sutural setae 2; anal vein long; mid tibia on posterior surface with 2 setae on mid third; posteroventral median seta on hind tibia absent. 238
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Geographical record. Peru, Bolivia. Comment. Monobasic genus: Oxytonocera nigrohalterata (Stein, 1904).
Fig. 135, Oxytonocera nigrohalterata, flagellomere, lateral view.
Pentacricia Stein, 1898 (Fig. 136) Type-species. Pentacricia aldrichii Stein (mon.). Diagnosis. Male dichoptic; 2 pairs of reclinate orbital setae; parafacial with setulae on upper part; labella not reduced; prealar seta absent; lower proepimeral seta downward directed; dorsocentral pre sutural setae 2, subequal in length, postsuturals 3; katepisternals 1: 1: 1, forming an equilateral triangle; lower calypter about twice the upper one; scutellum with the basal pair of setae well developed, as long as or longer than apical pair; fore tibia with a median seta on each of the anterior and posterior surfaces; hind tibia with at a median and a supramedian posterodorsal seta, and a strong posteroventral setae; sternite 1 bare; ovipositor short (Fig 136). References. STEIN, 1898; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. A monophyletic genus according to COURI & PONT (2000), who indicated eight characters for the genus, one of them, the shortening of the ovipositor, an autapomorphy. Geographical record. Galapagos Is.; United States of America. Comments. Monobasic genus: Pentacricia aldrichii Stein, 1898.
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Part II - Apical Groups
Fig. 136, Pelltacricia aldrichii, ovipositor, dorsal view (modified from COURI & PONT, 2000).
Pilispina Albuquerque, 1954
Type-species. Pilispina pilitibia Albuquerque (orig. des.). Diagnosis. Male dichoptic; epistoma not produced; arista with short cilia; palpus slender; pro sternum bare; dorsocentrals 1:3, except in P pilitibia (2:3); prealar absent; lateral margin of scutellum bare; sub-basal pair of scutellar setae long; apical pair short and fine; lower proepimeral seta directed downwards; proepisternals 2; katepisternals 1: 1: 1, forming an equilateral triangle; 2 notopleurals; apical scutellar seta reduced; both calypteres developed, the upper one glossiform and the lower one elongated, about 2.3 times longer than the upper calypter, except in Pfumipennis in which both calypteres are very reduced; wing veins bare; anal vein short; sternite 1 bare; male: cercal plate triangular or quadrangular, elongate, hypandrium tubular; female: ovipositor long, with microtrichia, tergites 6 and 7 slender. References. CQURI & PONT, 1999; CQURI & PONT, 2000; COURI, 2000b. Monophyly. CQURI & PONT (2000) indicated three characters for this genus, one a reversal (2 pre sutural dorsocentral setae) and two homoplasies (2 postsutural dorsocentral setae and hind tibia with a supramedian posterodorsal seta). Geographical record. Puerto Rico, Brazil. Comments. Six species are ascribed to the genus (COURI, 2000b), all occurring in South America and only one, P medinai (Snyder, 1957) reaching to Central America (Puerto Rico Is.).
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Key to species (from COURI, 2000b) 1.
Wing long, with anal area very reduced; calypteres also very reduced (Fig. 2 in ALBUQUERQUE 1954b) (Brazil) ................... P.Jumipennis (Albuquerque, 1954) Wing not elongated, anal area of wing and calypteres not reduced ........................... 2
2.
Hind tibia with a suprarnedian posterodorsal seta .................................................. 3 Hind tibia without a suprarnedian posterodorsal seta............................................. 5
3.
Postsutural dorsocentral setae 2 (Brazil) .................... P. pilitibia Albuquerque, 1954 Postsutural dorsocentral setae 3 ............................................................................ .4
4.
Legs brown, greyish polinose; acrostichals with a differentiated prescutelar pair of setae; (Brazil) ............................................... .................. P. paula (Medeiros, 1980) Legs yellow; acrostichals with prescutelar pair of setae not differentiated (Brazil) ...................................................... ................. P. differa (Couri & Carvalho, 1993)
5.
Antenna long, reaching epistorna, and inserted at the upper limit of the eye, in a projected area (Fig. 1 in ALBUQUERQUE, 1957b); hind tibia with more than one anteroventral seta (Brazil) ................................ P. benevenuta (Albuquerque, 1957) Antenna not reaching epistoma, inserted at the mid-level of the eye and not in a projected area; hind tibia without any setae on anteroventral surface (Puerto Rico, Brazil) .............................................. ............................ P. medinai (Snyder, 1957)
Plumispina Albuquerque, 1954 (Figs. 137-139) Type-species. Plumispina longipilis Albuquerque (orig. des.). Diagnosis. Male dichoptic; eyes bare; 1 pair of rec1inate orbital setae; 3 pairs of frontal setae; arista with short plumae on basal third (Fig. 137); dorsocentral setae 1:3; prealar absent; prosternum bare; proepisternals 2; propleurals 2; katepisternals 1: 1: 1, forming an equilateral triangle; lower calypter about 1.8-2.0 times as long as upper calypter; Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
wing veins bare; mid tibia with a median anterodorsal seta; hind femur with one preapical posterodorsal seta; hind tibia with one anterodorsal seta on basal third (Fig. 138), one preapical dorsal and 2 posterodorsal setae; sternite 1 setulose; male: hypandrium tubular (Fig. 139). References. ALBUQUERQUE, 1954b; COURI & PONT, 1999; COURI & PONT,2000. Monophyly. In the cladistic analysis of the tribe, Plumispina was positioned, according to COURI & PONT (2000), in the monophyletic group Coenosia, with two homoplastic characters supporting the genus: hind tibia on posterodorsal surface with a supramedian seta and cereal plate of male without a posterior concavity. Geographical record. Brazil. Comment. Monobasic genus: Plumispina longipilis Albuquerque, 1954.
137 Figs. 137-139, characters of Plumispina longipilis (modified from ALBUQUERQUE, 1954b): 137, head, lateral view; 138, hind leg, lateral view; 139, cereal plate, sursty1i and aedeagus. Abbreviation: ar, arista.
Reynoldsia Malloch, 1934 (Figs. 140-144) Type-species. Reynoldsia pectinata Malloch (orig. des.). Diagnosis. Eye with or without small and sparse cilia; male dichoptic; head subquadrate in profile; epistoma produced (Fig. 140); 1 pair of reclinate orbital setae; antenna with apex of flagellomere acutely pointed; arista with very small cilia; palpus long, slender with a slightly clubbed apex; prealar absent; scutellum bare below and on lateral margins; prosternum bare; proepisternals 2; propleurals 2 directed upwards, posterior 242
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one about 2.5 times as long as the other; katepisternals 1: 1:1, forming an equilateral triangle; lower calypter about 1.5-2.0 times as long as upper calypter; dorsocentrals 2:3; wing veins bare; veins R4+5 and M parallel at apex; legs with claws and pulvilli well developed; hind tibia with at least 2 anterodorsal and 2 posterodorsal setae (Figs. 141 a, 141 b); sternite I bare; male: sternite 5 U-shaped (Fig. 142); cercal plate and surstyli strongly developed (Figs. 143, 144); terminalia produced, phallic complex modified with elongated hypandrium (Fig. 144); female: proboscis with haustellum partly sclerotized; prestomal teeth developed; ovipositor long with microtrichia in all segments; cerci long, surpassing hypoproct; 3 spermathecae; egg: Phaonia type. References. MALLOCH, 1934; COURI, 1995, 1998b; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. The genus is probably monophyletic (CARVALHO & COURI, in press) based on three synapomorphies: presence of2 acrostichals presutural setae; palpus very long and slender; mesonotum with 2 lateral vittae reaching apex of scutellum. Geographical record. Chile, Argentina. Comments: PONT (1972) included Coenosia robusta Stein with reservations among the Reynoldsia and, for this reason the species was not included in the key. It is necessary to exam the type-material of this species which is deposited at the SMT.
Key to species (modified from MALLOCH, 1934) 1.
Mid femur of male with a comb-like series of stout setae on apical fourth of posteroventral surface; mid femur of female with a less pronounced series of more widely separated and weaker setae; both sexes with one or more pairs of moderately strong, irregular presutural acrostichal setae; flagellomere not twice as long as pedicel; halter fuscous (Argentina) .............................................. R. pectinata Malloch, 1934 Mid femur in both sexes without a series of stout setae as above; mesonotum usually without differentiated pre sutural acrostichal setae; flagellomere much more than twice as long as pedicel. ................................................................................................ 2
2.
Male ................................................................................................................... 3 Female ................................................................................................................ 8
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
243
Part II - Apical Groups
3.
Trochanters testaceous-yellow; coxae and femora, except for extreme apices oflatter, black and with grey dust; frons fuscous; scutellum unicolorous brownish-black, slightly shining; mid and hind femora with long, moderately strong, widely spaced anteroventral and posteroventral setae; halter yellow (Argentina) ........R. trochanterata Malloch, 1934 Trochanters dark brown to black; femora either broadly reddish-yellow at apices, or if only narrowly so, then ventral surfaces quite densely furnished with long bristly hair in addition to having the usual two series of long setae or the frons densely white dusted; scutellum partly grey dusted ....................................................................... .4
4.
Mid and hind femora with long fine bristly hairs on ventral surfaces, stronger on anteroventral surface of hind pair; male: sternite 5 with long slightly curled bristly hairs on entire length of the processes; hind tarsus not as long as hind tibia (Argentina)
................................................................................... .R. brevitarsis Malloch, 1934 Mid and hind femora with two series of widely spaced and rather unequal, long, strong setae, one on anteroventral and the other on posteroventral surface and no long bristly ventral hairs; male: sternite 5 with setulose hairs and some quite strong setae, the latter near apex on inner or lower margins; hind tarsus distinctly longer than hind tibia ............................................................................................................... 5 5.
Frontal orbits, parafacial, and gena densely golden-yellow dusted; interfrontalia black, with slight grey dusting; mesonotum black, with slight dark grey dusting and three broad dark brown vittae, central one produced over disc of scutellum; hind tibia with one submedian anteroventral seta (Chile )............................ .R. aurifera (Bigot, 1885) Frontal orbits, parafacial, gena, and interfrontalia white dusted; mesonotum not so distinctly trivittate and scutellum not dark brown on basal half of disc; hind tibia with two or three submedian anteroventral setae ........................................................... 6
6.
Apex of femora very narrowly reddish; halter with orange-yellow knob (Argentina)
........................................................................................... .R. coxata Malloch, 1934 Apex offemora broadly reddish; halter with black knob ......................................... 7 244
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Marcia Souto Couri & Claudio Jose Barros de Carvalho
7.
Anepimeron with a few stiff hairs near center; femora with less than the apical third red (Chile, Argentina) ............................................R. pteropleuralis Malloch, 1934 Anepimeron bare; femora with almost the apical half red (Argentina) ................ .
.............................................................................. ..R. ruJoapicata Malloch, 1934 8.
Halter with a yellow or yellowish-brown knob ........................................................ 9 Halter with a black or fuscous knob ..................................................................... 12
9.
Scutellum entirely fuscous above, very faintly shining; frons fuscous, with very slight brownish dust on interfrontalia (Argentina) ................. R. trochanterata Malloch, 1934 Scutellum largely or entirely grey dusted; frons more distinctly greyish or brownish dusted ................................................................................................................ 10
10. Mesonotum with three quite well-defined and wide dark brown vittae, the central one extending to about middle of scutellum; frontal orbits, parafacial, and gena densely golden-yellow dusted, interfrontalia black (Chile) ................R. aurifera (Bigot, 1885) Mesonotum less conspicuously vittate, central vitta not continued onto scutellum; frontal orbits, parafacial, and gena less distinctly yellow dusted, if rather markedly yellow than the interfrontalia is also quite densely dusted .................................................... 11 11. Scutellum entirely brownish-grey dusted, without dark lateral marks; dust on lateral margin of mesonotum in front of wing base yellowish brown, not markedly paler than the dusted vitta mesad of the sublateral dark vitta; hind tibia with one strong submedian anteroventral seta (Chile )..............................................R. scutellata Malloch, 1934 Scutellum grey dusted, distinctly darkened on sides; dust on lateral margin of mesonotum in front of wing base whitish-grey, much paler than the dusted vitta mesad of the sublateral dark vitta; hind tibia with two strong submedian anteroventral setae (Argentina) ......................................................... .......... R. brevitarsis Malloch, 1934 12. Anepimeron with some short hairs near center (Chile, Argentina) .................... .
............................................................................ .R. pteropleuralis Malloch, 1934 Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Part II - Apical Groups
Anepimeron bare ................................................................................................ 13 13. Mesonotum with at least one pair of well developed pre sutural acrostichals; femora with less than the apical third red (Argentina) ...................R. pectinata Malloch, 1934 Mesonotum with no well-developed pre sutural acrostichals; femora with more than the apical third red (Argentina) .................................... R. ruJoapicata Malloch, 1934
1 em
1 mm
hyp
144 Figs. 140-144, characters of Reynoldsia pectinata (modified from COURI, 1995): 140, head, lateral view; 141a, hind tibia, anterior view; 141b, hind tibia, posterior view; 142, male sternite 5, dorsal view; 143, cereal plate and surstyli, dorsal view; 144, cereal plate, surstyli and hypandrium, lateral view. Abbreviation: hyp, hypandrium.
246
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Schoenomyza Haliday, 1833 (Figs. 145-147) Type-species. SciomyzajasciataMeigen (WESTWOOD, 1840: 143). Diagnosis. Male dichoptic; frons very wide, about one-third of head width (Fig. 145); 1 pair of reclinate orbital setae; antenna short, arista with very small cilia; palpus long, slender with slightly clubbed apex; proboscis with palpus long (Fig. 146); prealar seta absent; scutellum bare below and on lateral margin; prosternum bare; proepisternals 2; propleurals 2, upward directed, posterior one about 2.5 times the length of the anterior setae; katepisternals 1: 1: 1, forming an equilateral triangle; prosternum developed; both calypteres small, upper one roundish, lower one tongue-like and hardly projecting beyond the upper one; dorsocentrals 1:3; wing veins bare; R4+5 and M parallel apically; mid tibia on anteroventral surface with a strong seta on middle; hind tibia (Fig. 147) on anteroventral surface with two setae on middle 1/3, anterodorsal surface with 2 setae on middle third, inserted a little above from the anteroventral ones and longer than those; dorsal surface with a long seta at apical third; anterior surface with two apical setae; sternite 1 bare; male: sternite 5 with few setae on disk; cercal plate large; male: hypandrium tube-like, aedeagus apodeme long, surpassing posterior margin ofhypandrium; female: ovipositor long with rnicrotrichia on all segments. References. MALLOCH 1934; COURI, 1996b; COURI & PONT 1999; COURI & PONT 2000. Monophyly. Probably monophyletic, but according to COURI & PONT (2000), there is no autapomorphy for this genus. By their phylogenetic hypothesis, Schoenomyza is the sister-group of Spathipheromyia. Geographical record. Mexico, Ecuador, Chile, Argentina, Peru, Uruguay; Nearctic, Palaearctic, Afrotropical and Oriental Regions. Comments: Seventeen species occur in the Neotropical region, 16 of them restricted to South America and only one, S. aurifrons Malloch, in Mexico.
Key to species (modified from MALLOCH, 1934) 1.
Postvertical pair of setae very short and fine, shorter longer than the incurved pair external to them and below their level; frons densely greyish-white dusted on orbits,
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
247
Part II - Apical Groups
entire central part chocolate-brown; mesonotum densely whitish-grey dusted on sides and with a broad chocolate-brown central vitta on its entire length that extends over disc of scutellum (Argentina) .......................................... S. univittata Malloch, 1934 Postvertical setae distinctly longer than the incurved pair below their level; frons and mesonotum not coloured as above ........................................................................ 2 2.
Male ................................................................................................................... 3 Female ................................................................................................................ 9
3.
Wings marked with fuscous marks .......................................................................... 4 Wings entirely hyaline or very faintly brownish, without distinct dark marks .............. 5
4.
Wings with a large subquadrate blackish-brown mark from midway on third costal division to apex of vein R 2+3 that extends backward the same width over vein M and partly enclose the crossvein dm-cu (Peru) ............................ S. bella Malloch, 1934 Wing with a fuscous cloud on costa from apex of vein R 2+3 to apex of vein M; frons densely grey dusted, broadly velvety-black on each side (Argentina) ...................... .
........................................................................................... .. S. mallochi Pont, 1972 5.
Frons black; face entirely densely silvery-white dusted; orbits and ocellar triangle densely lavender-grey dusted, the latter acute in front an extending almost to anterior margin of frons; mesonotum with only one pair of minute, widely separated acrostichal hairs that are situated well behind the suture (Argentina) .......... ...... ,S, argyriceps Malloch, 1934 Frons brown on its upper portion and strongly silvery pruinose on lower portion; face and parafacial chocolate brown; gena yellow dusted; mesonotum with acrostichal hairs biseriate over its entire extent (Ecuador) ....................... S. napensis Couri, 1996 Frons yellow or brown dusted; parafacial yellow or yellowish-grey dusted, frontal triangle always shorter, never extended to near to the anterior margin, and coloured differently; mesonotum with the two series of acrostichal hairs restricted to portion behind suture ......................................................................................................... 6
248
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Marcia Souto Couri & Claudio Jose Barros de Carvalho
6.
Frons dark dull brown; parafacial and orbits brownish-grey; ocellar triangle not differentiated; pre sutural acrostichals irregularly biseriate (Argentina)
....................................................................................... .S. biseriata Malloch, 1934 Frons pollen-yellow to golden-yellow dusted; parafacial yellow dusted; ocellar triangle more or less well differentiated ............................................................................... 7 7.
Acrostichal hairs biseriate over their entire extent; legs entirely black; face fuscous, only the edges of the parafacial next to eyes distinctly yellow (Chile) ................ ..
.................... ............ ....... ..................... ....... ..................s. fuscifacies Malloch, 1934 Acrostichal hairs partly uniseriate; legs not entirely black, at least the tarsi partly yellowish; entire face densely yellow dusted ........................................................... 8 8.
Fore tarsus entirely dirty yellow, slightly longer than abdomen and quite slender; hind femur with two strong setae on apical third of anteroventral surface; dark paired spots on abdomen not conspicuous, slightly shining and poorly defined (Argentina)
....................................................................................... .. S. tarsalis Malloch, 1934 Fore tarsus slightly shorter than abdomen, yellowish only apically, and not very slender; hind femur with four or five long strong setae on apical half of anteroventral surface; dark paired spots on abdomen conspicuous, almost glossy (Argentina) .................. .
........................................................................................ .S. armipes Malloch, 1934 9.
Center of frons distinctly white dusted, remainder dark brown; mesonotum with three or five dark brown vittae (Chile ).................................... S. albomedia Malloch, 1934 Center offrons not white dusted; mesonotum not marked as above ....................... 10
10. Face and frons densely golden-yellow dusted, the face slightly paler below; frons hardly darkened on upper third .............................................................................. l1 Face and frons either pale grey or brownish-yellow dusted, if brownish-yellow dusted then upper half of frons dark brown ...................................................................... 12 11. Hind femur with one long and one short seta on apical third of anteroventral surface Muscidae (Diptera) of the Neotropical Region: TAXONOMY
249
Part II - Apical Groups
(Argentina) ..................................................................... S. tarsalis Malloch, 1934 Hind femur with four or five long setae on apical half of anteroventral surface (Argentina)
....................................................................................... S. annipes Malloch, 1934 12. Acrostichal setulae well developed, biseriate on almost their entire extent; frons dark brown or live-brown, whitish or yellow dusted on not more than its anterior half.... 13 Acrostichal setulae lacking or represented by one or two microscopic setulae behind suture; frons not as above ..................................................................................... 14 13. Frons dark brown or olive-brown, whitish or yellowish dusted over less than its upper half (Argentina) ............................................................... S. biseriata Malloch, 1934 Frons grey on its lower portion and brown on upper portion; face and gena yellow dusted (Ecuador) ................................................... ............. S. napensis Couri, 1996 14. Mesonotum with two submedian grey vittae on the anterior margin that do not extend beyond the suture, and only a trace of a broad darker grey vitta on each side between dorsocentrals and intra-alars, part of which is covered with fine brownish dust; frons densely pale grey dusted; legs black, extreme apices of all femora and the entire tibiae brownish yellow (Uruguay) ................................................ S. evittata Malloch, 1934 Mesonotum with a broad chocolate-brown vitta on each side that extends from mesad of the dorsocentrals to or over the bases of the intra-alars, the division in front narrower or obsolete; frons black-brown, orbits greyish dusted; legs black, extreme bases of fore tibiae more or less reddish-yellow (Chile) ................S. lativittata Malloch, 1934
Species of Schoenomyza not included in the key and their respective geographical distribution
250
Species
Geographical distribution
S. aurifrons Malloch, 1918
Mexico
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
S. breviventris Stein, 1911
Peru
S. neobiseriata Snyder, 1957
Argentina
S. neotropica Carvalho & Pont, 1993
Argentina
S. nigrithorax Stein, 1911
Peru
S. willinki Snyder, 1957
Argentina
145 O,5mm
Figs. 145-147, characters of Schoenomyza napensis (modified from COURI, 1996b; MALLOCH, 1934): 145, female head, view frontal; 146, female proboscis, lateral view; 147, tip of hind tibia. Abbreviation: pIp, palpus.
Schoenomyzina Malloch, 1934 (Figs. 148-149) Type-species. Schoenomyzina pallicornis Malloch (orig. des.). Diagnosis. Male dichoptic; frons very wide, about one-half of head width; last frontal pair directed backward; antenna long; flagellomere with a pointed apex; 1 pair of reclinate orbital setae; arista shortly pubescent; palpus long, slender, or slightly enlarged apically; dorsocentral setae 1:2; prealar seta absent; scutellum bare below and on lateral margins; pro sternum bare; proepisternals 2; propleurals 2 directed upward, posterior one about 2.5 times the length of the anterior seta; katepisternals 1: 1: 1, form.ing an equilateral triangle; both calypteres small, the lower one not or very slightly projecting beyond the upper one; wing veins bare; R 4+5 and M parallel apically (Fig. 149); hind tibia with one submedian anteroventral seta, one median anterodorsallonger than the anteroventral one, Muscidae (Diptera) of the Neotropical Region: TAXONOMY
251
Part II - Apical Groups
dorsal surface with a long seta on apical third, anterodorsal surface with a short preapical seta; sternite 1 bare; male: hypandrium tubular (Fig. 148); female: ovipositor long with rnicrotrichia. References. COURI, 1998a; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. COURI & PONT (2000) indicated four characters: three homoplasies (2 dorsocentral setae, sternite 5 triangular and sternite 5 of male with a sclerotized median area) and one reversal (median seta of hind tibia on posterodorsal surface absent). Geographical record. Peru, Argentina, Chile, Juan Fernandez Is. Comments. Six species are ascribed to the genus, all of them restricted to South America.
Key to species (from COURI, 1998a) 1.
Ocellar triangle velvety black, much darker than the rest of frons; mesonotum with a broad central and lateral brown vittae; acrostichals biseriate; wings hyaline with stigmatal region and both crossveins brown (Chile, Argentina) .............. ..
................................................................................... .S. triangularis Malloch, 1934 Ocellar triangle concolorous with rest of frons; mesonotum without a broad central and lateral brown vittae; acrostichals uniseriate or biseriate; wing colour not as above ..................................................................................................................... 2 2.
Palpus yellow; antenna yellow in male, female flagellum with apical two-thirds brown; acrostichals uniseriate over entire extent; hind tibia with the preapical dorsal seta almost as close to apex as the preapical anterodorsal one (Argentina) .............................. .
.................................................................................... S. pallicomis Malloch, 1934 Palpus and antenna dark brown to brown, with or without gray pruinescence; acrostichals biseriate, at least before suture; hind tibia with the preapical dorsal setae not as close to apex as the preapical anterodorsal one ............................................ 3 3.
252
Calypteres similar in length; lunule golden dusted, contrasting with the rest of frons;
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
wings with brownish veins (Juan Fernandez Is.) ..................... S. emdeni Hennig, 1955 Lower ca1ypter very small; colour of lunule not contrasting with the rest of frons; wings hyaline or a little brownish ........................................................................... .4 4.
Wing with no long setae on costal margin (Peru) ..................... s. biseta (Stein, 1911) Wing with long setae on costal margin .................................................................... 5
5.
Wing brownish on costal margin; lower calypter linear, extremely short (Argentina) ................................................................................. .S. Juscicosta Malloch, 1934 Wing hyaline; lower calypter short (Peru) ............................. S. unicolor (Stein, 1911)
.......
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0,5mm Figs. 148-149. characters of Sci1oenomyzina (modified from COURI, 1998a): 148, S. pallicornis, aedeagus, lateral view; 149, S. fuscicosta, wing, dorsal view.
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
253
Part II - Apical Groups
Spathipheromyia Bigot, 1884 Type-species. Spathipheromyia stellata Bigot (mon.). Diagnosis. Male dichoptic; eyes bare; 1 pair of reclinate orbital setae; frons broader than long; dorsocentral setae 1:3; prealar absent; prostemum bare; proepistemals 2; propleurals 2; katepistemals 1: 1: 1, forming an equilateral triangle; dorsocentrals 2:3; wing veins bare; hind tibia with at least two setae on posterodorsal surface, preapical setae on anterodorsal, dorsal and posterodorsal surfaces; stemite 1 bare; male: hypandrium tubular; female: ovipositor long with microtrichia. References. MALLOCH, 1934; COURI & PONT, 1999; COURI & PONT, 2000. Monophyly. The sister-group of Spathipheromyia is Schoenomyza Halliday, 1833 (COURI & PONT 2000), a Neotropical genus with one species occurring in Mexico. These authors indicated four homoplastic characters for the genus: palpus enlarged; fore tibia on anterior to anterodorsal surface with a median seta; hind tibia with a supramedian posterodorsal seta and a preapical posterodorsal seta, at apical fourth. Geographical record. Peru, Bolivia, Chile, Argentina. Comments. The 15 species ascribed to the genus occur only in South America. It was not possible to exam the type-material of allIS species and for this reason, the two available keys in the literature are herein presented. Shoenomyza apicalis Stein, 1911 was recently transferred to this genus by PONT (2001).
Key to species (modified from STEIN, 1911) 1.
Legs black, at most the trochanters yellow (Peru) .............. S.juscipaZpis (Stein, 1911) All tibiae yellow...................................................................................................... 2
2. Mesonotum light grey with a large median vitta velvety black anteriorly and grey posteriorly; scutellum velvety black (Peru) ........ ~ ....................... S. picta (Stein, 1911) Mesonotum not coloured as described above; scutellum never velvety black .............. 3 3.
Mid tibia with one anteroventral seta (Peru) .........................S. minuta (Stein, 1911) Mid tibia without an anteroventral seta ................................................................. ..4
254
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
4. Palpus yellow; body length 4-5 mm (Chile) ........ ;.............. S. auriceps (Stein, 1911) Palpus black; maximum body length 4 mm (Peru) ................... S. nigra (Stein, 1911)
Key to species (modified from MALLOCH, 1934) 1.
Legs including tibiae, largely or entirely black or fuscous, only knees narrowly brownish yellow; antennae black; face, parafacial and anterior part of gena and frons densely golden brown dusted in both sexes (Chile, Argentina) ............... s. atra Malloch, 1934 Legs with at least tip of mid and hind tibiae orange yellow, if only faintly so then the face is greyish white dusted .................................................................................... 2
2.
Male ................................................................................................................... 3 Female ................................................................................................................ 6
3.
Mid femur with a series of strong spines on posteroventral surface that are stouter and closer placed near apex; face, parafacial, gena and frons anteriorly densely golden yellow dusted (Argentina) .............................................. s. magellani Malloch, 1934 Mid femur with no comb-like series of stout setae on apical part of posteroventral surface; face, parafacial, gena and frons different from above ................................ .4
4.
Wing with several yellowish-white patches; most evident when the wing is viewed from the apex against the light, a large one on cell behind vein CuA, bordering cell dm over the greater part of its extent; tarsi largely brownish-yellow, especially their central portion (Peru, Bolivia, Chile, Argentina) ................. S. guttipennis (Thomson, 1869) Wing with or without yellowish white marks, if these are present, then there is at most a small whitish spot behind vein CuA 1 close to base of cell dm; tarsi entirely fuscous above ..................................................................................................................... 5
5.
Face and gena whitish-grey dusted, mush paler than the golden-brown dusted frons, parafacial with a slight golden tinge; pleurae preponderantly bluish-grey dusted, brown
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
255
Part II - Apical Groups
markings much reduced (Chile) ......................................... S. alhiceps Malloch, 1934 Face and gena covered with golden-brown dust and concolorous with anterior half of frons; pleurae almost entirely dull dark brown (Chile )....... S. chilensis Malloch, 1934 Large, robust species, fully 6 mm long; submedian dark vittae on mesonotum broken
6.
by a grey fascia at suture (Argentina) .............................. .S. magellani Malloch, 1934 Smaller species, rarely over 5 mm in length; submedian dark vittae not broken at suture .................................................................................................................... 7 7.
Face, including parafacial and gena densely golden whitish-grey dusted, anterior half offrons less noticeably so; mid and hind tibiae black, grey dusted, with a brownish yellow shade apically, most distinct below; submedian brown vittae on mesonotum incomplete (Chile) .......................................................... S. insularis Malloch, 1934 Face, gena and anterior half or more of frons, densely golden-yellow dusted; all tibia orange-yellow; mesonotal vittae complete (Peru, Bolivia, Chile, Argentina)
................................................................ .............. S. guttipennis (Thomson, 1869)
Species of Spathiphe romyia not included in the key and their respective geographical distribution [including Shoenomyza apicalis Stein, 1911 recently transferred to this genus by PONT, 2001]
256
Species
Geographical distribution
S. apicalis (Stein, 1911)
Peru, Bolivia
S.fenestrata (Bigot, 1888)
Chile
S. stellata Bigot, 1884
Chile
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
-"~, --.~-=
-- -=--
-~-
.-
Marcia Souto Couri & Claudio Jose Barros de Carvalho
Stomopogon Malloch, 1930 Type-species. Coenosia albiseta Stein (orig. des.). Diagnosis. Male dichoptic; epistoma not produced; one pair of backwardly directed orbitals; flagellomere with a pointed apex, not reaching epistoma; arista bare; palpus slender; prostemum bare; dorsocentral setae 1:3; prealar seta absent; lateral margin of scutellum bare; sub-basal and apical pair of scutellar setae long; lower proepimeral seta directed downward; proepisternals 2; intra-alars small, anterior one undeveloped in S.
acuta, or both long (S. albiseta and S. argentina); katepisternals 1: 1: 1 forming an equilateral triangle; both calypteres developed, the upper one glossiform and the lower one elongated, about 2.3 times longer than the upper calypter; wing veins bare; claws and pulvilli large; hind tibia with three anterodorsal and three posterodorsal setae placed opposite one another, 2 of them supramedian and one submedian, anteroventral surface with a submedian seta, anterodorsal and posterodorsal surfaces with preapical setae also placed opposite one an other, dorsal surface with (S. argentina and S. albiseta) or without (S.
acuta) a preapical seta; sternite 1 bare; male: cercal plate quadrangular, elongated, hypandrium tubular (COURI & PONT, 2000: Fig. 55); female: ovipositor long, with microtrichia, tergites 6 and 7 slender. References. COURI & PONT, 1999; COURI & PONT, 2000; PONT, 2001; COURI, in press. Monophyly. According to COURI & PONT (2000), Stomopogon is monophyletic based on the presence oftwo apomorphies: hind tibia with two anterodorsal submedian setae and two posterodorsal supramedian setae. Geographical record. Peru, Bolivia, Chile, Argentina. Comments. Seven Neotropical species are included in the genus. Recently, PONT (200 1) transferred Coenosia capribarba Stein, 1911, C. hirtitibia Stein, 1911, C. garleppi Stein, 1911 and C. inculta Stein, 1911, to this genus.
Key to species (from COURI, in press) 1.
Legs black, sometimes the trochanters and very rarely the median tibia slightly reddish (Bolivia, Chile) ................................................................ .. S. hirtitibia (Stein, 1911) At least hind tibia or part of femur yellow.............................................................. .2
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
257
Part II - Apical Groups
2.
Mid tibia with a long median seta and a shorter submedian anterior to anterodorsal seta ....................................................................................................................... 3 Mid tibia without an anteriorto anterodorsal seta.................................................... .4
3.
Hind tibia with 2 preapical setae, on posterodorsal and anterodorsal surfaces; intraalars small, the anterior one very weak:; parafacial with many short cilia; palpus reddish yellow (Chile) .................................................................... S. acuta (Malloch, 1934) Hind tibia with 3 preapical setae, on anterodorsal, posterodorsal and dorsal surfaces; both intra-alars developed; parafacial with no short cilia; palpus brown with grey pruinescence (Argentina) ............................................... S. argentina (Snyder, 1957)
4.
Mid femur entirely or partly yellow........................................................................ 5 Mid femur entirely black ........................................................................................ 6
5.
Fore tibia black; gena with dense, white setae (beard) (Bolivia, Argentina)
.......................................................................................... s. capribarba (Stein, 1911) Fore tibia yellow; no beard on gena (Peru, Bolivia) ............. S. garleppi (Stein, 1911) 6.
Face, pleurae and legs with many white setae; supra and sub-vibrissal white beardlike setae present (Peru, Bolivia) .................. ...................... s. albiseta (Stein, 1911) Body setae black; no white beard (Chile) .................... ......... S. inculta (Stein, 1911)
258
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
Table 1. Subfamilies, tribes and genera of apical Muscidae, including the number of species in Brazil, South America and the Neotropical Region. Subfamily
Tribe
Cyrtoneurininae
Genus Arthurella Cariocamyia Charadrella Chortinus Cyrtoneurina Cyrtoneuropsis Mulfordia Neomuscina Neomusciniopsis Neurotrixa Pseudoptilolepis Xellothoracochaeta
Mydaeinae
Brontaea Graphomya Hemichlora Mydaea Myospi/a Scenetes stat.rev. Scutellomusca
Coenosiinae
Limnophorini
Coenosiini
Agellamyia
Albe rtine lla Drepanocnemis Limllophora Lispe Lispoides Pachyceramyia Rhabdotoptera Spilogolla Syllimllophora Tetramerillx Thaumasiochaeta Apsil Bithoracochaeta Coenosia Cordiluroides Insulamyia Neodexiopsis Notoschoellomyza Oxytonocera
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
Brazil 0 2
South America 1 2
Neotropical 1 2 3 1
2 0 6 32 0 13 1 1
7 38 3 23 1 1
9 40
6 2 4
6 6 4
3 1 4 4 0 1
11 1 16 6 0 1 1 1
6 6 4 12 1 21 6 1 1 2 1
3 33 9 18 2 1 6
3 37 12 20 4 1 8
33 2 7 10 9 34 4
33
1 1 0 20 5 1 1 0 1 2 0 0 0 7 12 3 1 29 0 1
3 0
1 63 7 1
3 35 1 1
2 7 10 11 35 5 1 89 7 1
259
Part II - Apical Groups
Pentacricia Pilispina Plumispina Reynoldsia Schoenomyza Schoenomyzina Spathipheromyia Stomopogon Total
260
47 genera
0 6 1 0 0 0 0 0 174
I
I
6
6
I
I
9
9 17
16 6 16 7
6 16 7
436
506
Muscidae (Diptera) of the Neotropicai Region: TAXONOMY
Marcia Souto Couri & Claudio Jose Barros de Carvalho
Plate 1
•
•
Cariocamyia maculosa Snyder, 1951
Charadrella malacophaga Lopes, 1938
(6 mm)
(10 mm)
Xenothoracochaeta rufiguttata (Macquart, 1851)
Graphomya mexicana Giglio-Tos, 1893
(8 mm)
(6mm)
•
•
•
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
261
.
.. ",
..
~ ~~""
-~
."
;-1.-
...,
.. .
n iT "
•
Marcia Souto Couri & Claudio Jose Barros de Carvalho
•
.
Plate 2
,
,.
Mydaea plaumanni Snyder, 1941
Limnophora aurifacies Stein, 1911
(6mm)
(6mm)
Cordiluroides listrata Albuquerque, 1954
Neodexiopsis jlavipalpis Albuquerque, 1956
(6mm)
(4mm)
•
•
262'
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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descri~ao
de Dalcyella gen. n. do
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COELHO, S.M.P. 2000. Phaonia Robineau-Desvoidy, 1830 (Diptera, Muscidae, Phaoniinae). II. Revisao das especies neotropicais. Revta bras. ZooI.17: 795-875. COQUILETI, D.W. 1901. Types of AnthomyidGenera. J.N.Y. Entomol. Soc. 9: 134-146. COQUILETI, D.W. 1910. The Type-species of the North American Genera of Diptera. Proc. U.S. Nat. Mus. 37: 499-647. COURT, M.S. 1982a. Insulamyia, gen.n., e notas sobre Spathipheromyia Bigot, 1884 (Diptera, Muscidae, Coenosiinae). Revta bras. Ent. 26: 29-36. COURT, M.S. 1982b. Adenda ao trabalho de Snyder (1954) sobre Cyrtoneurina Giglio-Tos, 1893 (Diptera, Muscidae, Cyrtoneurininae). Revta bras. Ent. 26: 37-53. CO URI, M.S. 1989. Philornis Meinert, 1890 - Key to adults (Diptera, Muscidae, Reinwardtinae). Revta bras. Zool. 6: 569-576. COURI, M.S. 1995. Contribution to the knowledge of Reynoldsia Malloch (Insecta: Diptera: Muscidae). Proc. BioI. Soc. Wash. lOS: 281-291. COURT, M.S. 1996a. Rhabdoptera Stein: redescription of the genus and of R. striatipennis Stein (Diptera: Muscidae). Revta bras. Zool.12 [1995]: 815-818. COURI, M.S. 1996b. Schoenomyza Haliday: redescription of the genus and description of a new species from Ecuador (Diptera: Muscidae). Revta bras. Zool. l2[ 1995]: 819-823. COURI, M.S. 1998a. Revision of Schoenomyzina Malloch (Diptera, Muscidae). Revta bras. Ent. 42: 105-110. COURI, M.S. 1998b. Redescription of two species of Reynoldsia Malloch (Diptera, Muscidae). Revta bras. Ent.42: 101-103. COURT, M.S. 1999a. A revision of Thaumasiochaeta Stein and allied genera (Diptera: Muscidae: Coenosiinae: Limnophorini). Proc. Entomoi. Soc. Wash. 101: 788-803. COURT, M.S. 1999b. Myiasis caused by obligatory parasites Philornis Meinert (Muscidae), pp. 5169. In: Guimaraes, J. H. & N. Papavero, Myiasis in man and animals in the neotropical region. Bibliographical database, Ed. Pleiade, FAPESP, 308 p. COURI, M.S. 2000a. A revision of the genus Apsil Malloch (Diptera, Muscidae, Coenosiinae, Coenosiini). Studia dipterologica 7: 45-57. COURI, M.S. 2000b. A Revision of the genus Pilispina Albuquerque (Diptera, Muscidae, Coenosiinae, Coenosiini). Boi. Mus. naco Rio deJ. (Zoologia) 437: 1-8. COURT, M.S. 2002. A new species of Apsil Malloch from Chile (Diptera, Muscidae). Revta bras. Zool. 19: 105-108.
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COURT, M.S. & D. PAMPLONA. 1992. Cordiluroides Albuquerque, 1954 (Diptera, Muscidae, Coenosiinae): redescric,:5es, notas, sinonfmia e chave para especies. Revta bras. Ent. 36: 255-262. COURT, M.S. & D. PAMPLONA. 1997. Sobre a identidade de quatro especies descritas em Cyrtoneurina Giglio-Tos com diagnoses e redescric,:5es (Diptera, Muscidae). Revta bras. BioI. 57: 411-415. COURT, M.S. & D. PAMPLONA (2000). Sobre as posic;5es sistematicas de quatro especies de Cyrtoneurina Giglio-Tos (Diptera, Muscidae). Revta bras. Zoo!. 17: 1069-1078. COURI, M.S. & A.c. PONT. 1999. A key to the world genera of the Coenosiini (Diptera, Muscidae, Coenosiinae). Studia dipterologica 6: 93-102. COURI, M.S. & A.C. PONT. 2000. A cladistic analysis of the tribe Coenosiini (Diptera, Muscidae, Coenosiinae). Syst. EntomoI. 25: 373-392. CURTIS,1. 1839. British Entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland: containing Coloured Figures from Nature of the most rare and beautiful species, and in many instances of the plants upon which they are found. Volume 16, 12 + 36 + 22 (sem numerac,:ao) pp., London. DODGE, H.R. 1967. Two new metallic species of Helina from Chile (Diptera: Muscidae). Proc. EntomoI. Soc. Wash. 69: 241-243. FIGUEROA, L.A.R. 1999. Sinantropia de dipteros muscoideos caliptrados de Valdivia, Chile. Campinas. 114 p. Dissertac,:ao (Mestrado), Universidade de Campinas (not published). GREENBERG, B. 1971. Flies and Disease. Vol. 1, Ecology. Classification and Biotic associations. Princeton University Press. 856 p. HENNIG, W. 1956. Muscidae [Part, Lieferung 194]. In: Lindner, E. (Ed.). Die F1iegen derpalaearktischen Region, 63b: 97-144, text-figs. 20-42. Stuttgart. HENNIG, W. 1957. Muscidae [Part, Lieferung 197]. In: Lindner, E. (Ed.). Die F1iegen der palaearktischen Region, 63b: 145-192, text-figs. 43-44. Stuttgart. HENNIG, W. 1958. Muscidae [Part, Lieferung 199]. In: Lindner, E. (Ed.). Die F1iegen der palaearktischen Region, 63b: 193-232, text-figs. 45-48. Stuttgart. HENNIG, W. 1959. Muscidae [Part, Lieferung 204]. In: Lindner, E. (Ed.). Die F1iegen der palaearktischen Region, 63b: 233-288, text-figs. 49-58. Stuttgart. HENNIG, W. 1960a. Muscidae [Part, Lieferung 209]. In: Lindner, E. (Ed.). Die Fliegen der palaearktischen Region, 63b: 385-432, text-figs. 71-152. Stuttgart. HENNIG, W. 1960b. Muscidae [Part, Lieferung 213]. In: Lindner, E. (Ed.). Die Fliegen der palaearktischen Region, 63b: 433-480, text-figs. 153-184. Stuttgart. 268
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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Author index ALBUQUERQUE ...................... 49, 57, 58, 64, 68, 80, 126, 128, 135, 172, 175, 187, 188,219,241,242 ALBUQUERQUE & LOPES ................. 50, 51, ISS, 159 AMORIM .................................................................... 7 AMORIM & CARVALHO .......................................... 18 AMORIM & PIRES ................................................... 20 ARAUJO & COURI... ....................................... 19, 136 ARNTFIELD .................................................. 164, 166 AX ....................................................................................7
BERNASCONI .......................................................... 20 BRAUER & BERGENSTAMM .................................. 98 BROOKS & McLENNAN ............................................ 7 CARVALHO ........................... 8, 18-20, 37-40, 58, 62, 63,68,69,74-77,79,80,83,96,99, 100, 104, 119,126, 129, 133, 134, 137, 138, 160, 172, 173 CARVALHO & COELHO .......................................... 19 CARVALHO & COURI..l9, 20, 80, 82, 174, 175,205,206 CARVALHO & PONT.. ..................... 19, 160, 161, 163 CARVALHO et a1... ........................ 17-20, 37, 40, 41, 63, 64, 68, 76, 133, 135, 149, 169, 173, 182, 217 COELHO ............................................ 19, 20, 119, 120 COQUILLET.. .......... 80, lOS, 119, 139, 160, 167, 178 COURI... .......... 19, 88, 94, 95, 189,190,201-206,209, 210,219,240,241,243,246,247,251-253,257 COURI & ALBUQUERQUE ..................... 19, 220, 225 COURI & CARVALHO ................................................ 19, 20, 51, 52, 56, 88, 135, 172, 173, 176, 177 COURI & LAMAS ................................................ 19, 78 COURI & LOPES ........................... 19, 20, 170, 171 COURI & MARQUES ........................................... 211 COURI & MOTTA ................................ 220, 234 COURI & PAMPLONA..19, 52, 140, 142, 150,218,219 COURI & PONT.. ............................................. 19, 20, 134,135,173,174,178,188,205,211,214, 218-220, 237-240, 242, 243, 247, 252, 254, 257 COURI et al .............................................................. 155 CURTIS ....................................................................... 63 DODGE..................................................................................79 FIGUEIROA. ..............................................................................70 GRAZIA et al. ........................................................... 8 GREENBERG .................................................................... 17 HENNIG ................. 8, 18, 19, 37-40, 48, 49, 58-60, 62, 81,98, 119, 135, 164, 173, 178, 183, 184, 191 HERTING ................................................ 39, 160 HUCKETT.. ........................... 63, 64, 74, 75, 187 HUCKETT & VOCKEROTH .................. 18, 80, 119 I.C.Z.N .......................................................... .48, 59, 60 LATREILLE ............................................................... 182 LATREILLE et. al.. ..................................................... 59
LOBANOY. ....................................................................... 164 LOPES ..................................... 19, 59, 61, 135, 156, 167 LOPES & CARVALHO ....................... 78, 156, 157, 158 LOPES & COURI... ............................... 19, 20, 200 LOPES & KHOURI. ............................................. 83 LOPES & MANGABEIRA ........................................ 61 LOPES & MANGABEIRA FILHO ............................ 59 MALLOCH .................................................. 105, 107, 149,164, 177-181, 184, 186, 192, 194,206,210, 214,216,217,237,243,247,251,254,255 McALPINE .......................................... 7, 18, 20, 62 MICHELSEN .................................................. 18,20, 136 MINELLI.. .............................................................. 7, 8 MOTTA & COURI... .......... 19, 211, 212, 213, 220 NELSON .................................................................. 7 NELSON & PLATNICK ............................................. 7 PALKA-ROCHA ......................................................... 64 PALKA-ROCHA & CARVALHO ....... 19, 63, 64, 66, 67 PAMPLONA ........ 19, 40-43, 46, 47, 55, 139-142, 148 PAMPLONA & COURI...19, 46, 47, 64, 70, 73, 83, 139 PONT.. ................ 17, 18, 37, 38, 42, 64, 68, 70, 79, 88, 96, 135, 187, 220, 234, 237, 243, 254, 256, 257 PONT & CARVALHO ................................................. 20 PONT & MAGPAYO......................................... .. .... 38 PONT & PAMPLONA ............................................ 142 PAPAVERO .................................................. 20, 96, 103 ROBACK ............................................................................... 18 RONDANI. ............................................................ 62, 70 SCHUHLI & CARVALHO ................................... 156 SEGUY.................................................................. 74 SIMPSON ............................................................. 7 SKIDMORE ....................................................... 17-19, 37,40,62,64,76,79,81,88,98,105,119,134, 160, 164, 173, 178, 183, 184, 191, 200 SNYDER ..................................................... 62, 74, 75, 81,105, 109, 136, 150, 155, 159, 167, 168, 170, 182, 183, 190, 192, 198, 220, 225, 228, 230, 236 STEIN .............................................................. 105, 178, 179, 184, 185, 191, 192, 214, 238, 239, 254 STEYSKAL. ............................................................... 48 THOMPSON .......................................................... 8, 18 TOWNSEND ........................................................ .41,42 VALERIO & GUIMARA.ES ...................................... 59 VOCKEROTH ...... 19, 20, 63, 70, 74, 75, 99, 136, 164, 167 VOSSBRINK & FRIEDMAN .................................... 20 WESTWOOD ....................................... 59, 163, 214, 247 WILEY. ................................................................ 7 ZINOVIEV......................................................... 119
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
277
Taxonomic index (Synonym are not in italics)
abdita (Giglio-Tos, 1893), p. ................................... 123 abnor11linervis (Stein, 1911), L. ............................... 185 Acanthipterinae ....................................................... 19 Achanthiptera Rondani, 1856 .................................... 136 aerinis Snyder, 1941, H. .................................. lll, 115 acrosticalis Snyder, 1941, H. ................................... IlO acuta (Malloch, 1934), S ................................ 257, 258 aeuta Stein, 1898, H. ............................................ 64, 67 aczeli (Snyder, 1957), L. ........................................... 181 adelpha (Schiner, 1868), H ................................ lll, 115 advena Snyder, 1957, p. .......................................... I23 aenescens (Wiedemann, 1830), 0. ...................... 71-73 aequifrons (Stein, 1898), L. ..................................... 186 affinis Malloch, 1925, M .................................... .45, 47 Agenamyia Albuquerque, 1953 ..... 36, 134, 174, 175,259 aitkeni Dodge, 1963, p. ....................................... 89, 94 alacris Co uri & Albuquerque, 1979, N. ..................... 224 alacris Stein, 1911, L. ............................................. 180 Albertinella Couri & Carvalho, 1997 ......... 29, 176,259 albiceps Malloch, 1934, S ....................................... 256 albiseta (Stein, 1911), S .................................. 257, 258 albisquama (Albuquerque, 1959), N. ........................ 223 albitarsis (Stein, 1898). L.. ...................................... 184 albomedia Malloch, 1934, S ................................... 249 albuquerquei (Lopes & Couri, 1987), L.. .................. 181 albuquerquei Carvalho, 1985, c. ..................... 137, 138 albuquerquei Carvalho, 1989, p. ............................... 84 albuquerquei Couri, 1983, p. ...................................... 93 albuquerquei Lopes, 1985, 0 .............................. 71-73 albuquerquei Pont, 1972, H. .................................. 1 17 aldrichii Stein, 1898, P. ........................................... 239 aliena (Stein, 1911), S............................. 193, 195, 197 al(fusca Couri, 1982, c. .......................................... 140 altaneira (Albuquerque, 1954), L.. ............................. 181 amazonensis Couri, 1983, p. ..................................... 89 alllazonica (Albuquerque & Lopes, 1982), M ....... 68, 69 amazonica Albuquerque, 1951, S....................... 126-128 analis (Macquart, 1851), G.............................. 164, 165 angustiJrons (Loew, 1861), p. .................................... 88 angustlfrons Loew, 1861, p. ....................................... 92 angustifrons Malloch, 1934, S.......................... 195, 196 anguslipennis (Stein, 1911), H. ............................... 106 annulata (Albuquerque, 1957), p. ............................. 121 annulata (Stein, 1911), N ........................................ 238 annulala Stein, 1911, B .......................................... 213 annulipes (Macquart, 1843), N. ................................... 224 anoctiluca (Carvalho, 1983), D ....................... I02, 104 antell/zula Couri, 1987, N. ....................................... 214 antenusi Lopes, 1955, N. ...................................... 60, 61
anthrax (Bigot, 1885), S .......................................... 191 anubes Snyder, 1941, H. .................................. 116, 117 apicalis (Malloch, 1934), p. ................................. 83, 86 apicalis (Stein, 1911), S ........................... 237, 254, 256 apicata (Stein, 1904), N. ......................................... 154 apicala Malloch, 1934, A ................................. 207-210 Apsi/ Malloch, 1929 ........................... 37, 204-210, 259 arcuata (Wiedemann, 1830), N. ............................... 154 argenticeps Malloch, 1934, L. ................................. 186 argentiJrons (Malloch, 1934), C. .............................. 216 argentifrontata Snyder, 1957, S................................ 191 argentina (Snyder, 1957), S .............. ............... 257, 258 argentina Malloch, 1934, L. ...... .............................. 187 argentina Snyder, 1957, H. ...................................... 117 argyriceps Malloch, 1934, S .................................... 248 arizona Snyder, 1958, N. .......................... 232, 234, 236 ar/eriopsis Couri, 1982, C. ....................................... 140 armipes (Stein, 1911), C .................................. 143, 144 armipes Malloch, 1934, S ................................ 249, 250 Artlzurella Albuquerque, 1954 ............. 26, 134, 135, 259 aterrima (Stein, 1904), S......................................... 191 aterrima WUlp, 1896, D ............................................ 63 Atlzerigona Rondani, 1856 ...................... 22, 38, 39, 130 Atherigoninae ....................................... 19, 37, 38, 130 atinNa Snyder, 1949, N. .......................................... 153 atincticosta Snyder, 1949, N. ................................... 154 atra Malloch, 1934, S............................................... 255 atricornis Malloch, 1934, B ............................. 212, 213 atriJrons Stein, 1911, C........................................... 215 atripes Malloch, 1934, A. ................................. 207-210 atrisquama (Stein, 1904), L. .................................... 185 atrovittata (Stein, 1904), S .......................................... 193 auralltiea (Albuquerque & Medeiros, 1980), p. ........... 121 auriceps (Stein, 1911), S ......................................... 255 auriceps Malloch, 1934, G....................................... 165 auricolis Albuquerque, 1980, H. ............................... 117 aurifacies Stein, 1911, L.. ......................................... 179 aurifera (Bigot, 1885), R ................................. 244, 245 auriJera (Malloch, 1934), C. .................................... 216 aurifrons Malloch, 1918, S .............................. 247, 250 australis Carvalho & Pont, 1993, H. ................ I07, 112 Australophyra Malloch, 1923 .............................. 64, 74 Austrocoenosia Malloch, 1934 ................................ 217 autumnalis De Geer, 1776, M .................................... 48 Azelia Robineau-Desvoidy, 1830 .............. 28, 62, 74, 130 Azeliinae .................................... 19, 37, 62, 76, 80, 130 Azeliini ........................................................ 19, 62, 130 bahama Snyder, 1958, L. ......................................... 184 barbitarsis (Stein, 1911), S........................................ 192
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
279
barbiventris Couri & Albuquerque, 1979, N. ....... 221, 225 basalis (Walker, 1853), M ........................................ .43 basicincta (Stein, 1904), p. ........................................ 55 beebei (Curran, 1934), C................................... 142, 145 bella Couri, 1984, P................................................... 92 bella Malloch, 1934, S............................................ 248 benevenuta (Albuquerque, 1957), P. .......................... 241 benjamini (Albuquerque & Medeiros, 1980), P. ........ 121 benoisti Pamp1ona, 1983, D ................................ .41, 42 benoisti Seguy, 1932, C............................................ 217 bequaerti Aldrich, 1932, C...................................... 139 bicolor (Bigot, 1885), C.......................................... 218 bicolor Albuquerque, 1956, p. .......................... 53, 54,76 bicolor Malloch, 1934, C ...........................................79 bigoti (Albuquerque, 1957), P. .................................. 120 bigoti Malloch, 1934, H. ........................... 109, 113, 117 Biopyrellia Townsend, 1932 ......................... 23, 40, 130 bipuncta (Wiedemann, 1830), B ............................... .40 bipunctata (Schiner, 1868), P................................... 123 biseriata Malloch, 1934, S............................... 249, 250 biseta (Stein, 1904), H ............................................ I04 biseta (Stein, 1911), S .............................................. 253 biseta Malloch, 1934, A ................................... 207, 208 biseta Snyder, 1954, C.............................................. 140 Bithoracochaeta Stein, 1911.. .... 34, 134, 211, 213, 259 bivitatta Albuquerque. 1979. 1. ............................. 80. 82 blanchardi Garcia, 1952, P......................................... 90 boliviana Coelho, 1998, P...... ................................ .124 Brachygasterina Macquart. 1851.. ...... 32. 76. 77, 96, 130 brasiliensis (Albuquerque, 1958), D .................. 102, 104 brasiliensis (Walker, 1853), N .................................. 234 brevicornis (Malloch, 1934), N. ...... .......................... 234 brevihirta Malloch, 1934, L. ......... ............................ 180 breviseta (Stein, 1911), S.......................................... 199 brevitarsis Malloch. 1934. R ............................ 244, 245 brevivena Snyder, 1941, H ............. ................... 114-116 breviventris Stein. 1911, S......................................... 251 Brontaea Kowarz. 1873 .................... 33, 160. 163.259 browni Snyder, 1941, H. .......................................... .111 browni Snyder, 1957, S............................................ 199 brunnea (Hough, 1900), C.......................... 27, 141, 145 cacheuta (Snyder, 1957), D .............................. .102, 104 cacumina Snyder, 1957, N ....................................... 226 caerulea (Snyder, 1949), X ...................................... .159 calcitrans (Linnaeus, 1758), S ..................................... 61 callidimera (Bigot, 1887). D ..................................... .42 Calliphoroides Malloch, 1930 ..................................... 98 calopyga (Loew, 1872), N. ...................................... 221 cambuquirensis (Albuquerque, 1954), N. .................. 234 camorinensis Albuquerque, 1956, C.......................... 217 candidifrons (Stein, 1911), S.................................... 193 caneo Snyder, 1941, H. ................ ............................ 114 capalta Snyder. 1949, N. ........ ................................. 152 capensis (Wiedemann, 1818),0 ...........................71-73 capribarba (Stein. 1911), S ............................. 257, 258
280
caras Snyder, 1949. M ............................................. 168 cardini Malloch. 1936, S................ .......................... 172 carinata Dodge, 1968, P............................................. 91 carioca Albuquerque, 1951, S........................... 126, 128 Cariocamyia Snyder. 1951.. ....................... 32. 136, 259 carnifex (Stein, 1911), c. ..................................... 79, 96 carvalhoi (Lopes & Khouri,1996), P. ......................... 83 carvalhoi Coelho, 1998, P. ...................................... 125 castanea Albuquerque, 1955, M .... ............................ 169 catamac!a (Snyder, 1957), D.................................... 102 catharinensis (Carvalho, 1983), D................... 101, 104 centralis Stein, 1911, M ............................................ 169 cera Snyder, 1958, N. ........................................ 232, 236 Chaetagenia Malloch, 1928 ................. 25, 78, 130, 156 chaetosa Malloch, 1934, C....................................... 216 chalcogaster (Wiedemann, 1824), 0 ....................71-73 chalybea (Wiedemann, 1830), P............ .................... 97 Charadrella Wulp, 1896 ...................... 25, 134, 137, 259 chi/ens is (Bigot, 1885), P........................................... 84 chi/ensis Bigot, 1888. G........................................... 165 chi/ensis Malloch, 1934, H. .............................. 108, 112 chi/ensis Malloch. 1934, S........ ................................. 256 chlorogaster (Wiedemann, 1830), P. ......................... 96 Chortinus Aldrich, 1932 .................... 26, 134, 139, 259 chrysiceps Malloch, 1934, N. .................................... 237 Gilitibia (Albuquerque, 1955). M. .... 69 cinnamomina (Stein), P............................................. 88 circulatrix (Walker, 1861), H. .................................. 117 cirratipila Snyder, 1957, N .................... ............ 224, 228 c!avacula Snyder, 1957, N. ..................................... 227 clavitibia (Stein, 1911). S................................. 193, 199 Coenosia Meigen. 1826 .. .36, 200, 214, 217, 238, 242, 259 Coenosiinae ........................ 19, 133-135, 164, 173,259 Coenosiini ... .l9. 134. 135. 173, 174, 178, 200, 205, 206, 259 compressicornis Snyder. 1949. M ............................ .168 compressifrollS (Stein, 1911), S ................................ 194 compressipalpis (Stein, 1911), D ............................. 100 compressitarsis (Stein, 1911), S ................................ 192 compressitarsis (Stein, 1911), T. .......................... 201, 203 concacata Pamplona, 1986, M ............................ .45-47 cOllcilllla (Wulp, 1896), P............................... 53, 55, 56 conjusa Snyder, 1949, P...... .................................... 157 conjusa Snyder. 1954, c. ......................................... 141 COlijus us (Malloch, 1928), P. ..................................... 85 connexa Malloch, 1934, H ....................... 107, 113, 116 consanguinea (Stein, 1911), H ................................ 106 conspersa (Stein, 1911). C............................... 143, 144 convexa (Stein, 1918), P............................................ 88 copiosa (Wulp, 1896), H. ................................ .! 12, 114 Cordiluroides Albuquerque, 1954 .......... .36, 218. 220, 259 cordyluroides (Stein, 1898), P. ........................ 188, 189 cornicina (Fabricius. 1781). N. ................................. .49 Correlltosia Malloch, 1934 ............................ 33, 79, 130 corvina (Giglio-Tos, 1893), L.. ................................. 180 costalis (Walker, 1853), c. ...................................... 140
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
costata Snyder, 1957, N. ......................................... 238 costipennis (Snyder, 1957), P........................... 120, 125 cothumata (Bigot, 1885), C.................................... 217 cotidiana Snyder, 1954, L. ..................................... .184 couriae Pamplona, 1986, M ............................... .45, 47 coxata Malloch, 1934, R ... ...................................... 244 crassicauda Stein, 1911, c. ...................................... 214 crassicrurus Snyder, 1957, N. .................. 220, 229, 230 crepedoseta Snyder, 1940, H. .. ............................... 117 cresa Snyder, 1949, M ............................................. 168 crispaseta Snyder, 1954, 140, 230 crispiseta Snyder, 1957, N. ...................................... 229 crocea Snyder, 1940, H. .......................................... 117 croceafrons Snyder, 1957, N. .................................. 227 cruciata Snyder, 1941, H. ............................... 105, 114 cubana Gregor, 1974, 0 ....................................... 71,72 cubana Johnson, 1919, L. ........................................ 181 currani Snyder, 1949, N. ................................. 152, 154 curvata (Stein, 1904), P........................................... 123 curviventris Albuquerque, 1959, c. ........................... 217 Cyacyrtoneura Townsend, 1931.. ................ 24, 41, 130 cyanea (Macquart, 1843), M .................................... 170 cyanea Townsend, 1931, c. ..................................... .41 cyaneiventris Macquart, 1851, H. .......................... 65-67 cyanicolor (Stein, 1911), H. .................................... 117 cylindrica (Macquart, 1846), C. ............................... .141 Cyrtoneurina Gig1io-Tos, 1893 ..................... 27, 139, 259 Cyrtoneurininae ......................... 37, 133-135, 169, 259 Cyrtoneuropsis Mal1och,1925 .... 27, 134, 141, 142, 148,259 Dalcyella Carvalho, 1989 ............................ 26, 79, 130 dalcyi Pamplona, 1986, M ................................ .44, 47 Dasymorellia Malloch, 1923 ...................... 23,41, 130 dasyophthalma Malloch, 1928, H. .......................... 117 dasyops (Macquart, 1843), H. ................................. 117 debilis (Williston, 1896), B .............................. 161, 163 deceptiva Dodge & Aitken, 1968, P........................... 92 declivis (Stein, 1904), N ...... ..................................... 235 delecta (Wulp, 1896), B ................................... 162, 163 deleta (Wu1p, 1896), L.. .......................................... 179 dendropanacis Pamplona & Couri, 1995, M ....... .46, 47 devia (Curran, 1934), N. .......................................... 235 diaphana (Stein, 1911), N. ....................... 220, 234, 235 diaphanus (Wiedemann, 1817), T.. ...........................75 dichroma (Wiedemann, 1830), H. .......................... .117 differa (Couri & Carvalho, 1993), P......................... 241 dilata Malloch, 1934, A . .................................. 208, 209 diluta (Stein, 1911), L.. .......................................... 185 diminuta Couri, 1984, 1'. ............................................ 93 diminuta Couri, 2002, A .................................. 207, 210 discolor (Stein, 1911), H ................................. 108, 112 discolorisexus Snyder, 1957, N . ....................... 228, 229 discreta (Wulp, 1896), H. ......................................... 117 distincta Couri & Lopes, 1987, P.............................. 54 ditiportus Snyder, 1957, N. ................................ 228, 229 diversipaipis (Rondani, 1863), M .............................. .44
c. ..............................
Dolichophaonia Carvalho, 1993, stat. rev..32, 99, 104, 130 domestica Linnaeus, 1758, M ............................... .48, 49 dorae Stein, 1911, D ... ............................................. 178 dorsilinea (Wulp, 1896), M ........................................ 81 dorsipuncta (Stein, 1918), N. .................................... 151 downsi Dodge & Aitken,1968, p. ........................ 90, 94 Drepanocnemis Stein, 1911.. ....... 35, 134, 177, 178, 259 Drymeia Meigen, 1826 ................................ 29, 63, 130 dubia (Bigot, 1885), N. ............................................ 224 145, 148 dubia (Snyder, 1954), dubitalis Malloch, 1934, S............................... 195, 196 ebenifemur Snyder, 1957, N. .................................... 230 echinogaster (Stein, 1911), H ................................. 106 elegans Couri & Albuquerque, 1979, N. .................... 222 elegans Macquart, 1843, L. .. .................................... 179 elegantula Pont, 1972, L. ........................................ 185 elongata (Albuquerque, 1958), D ................................ 103 emdeni Hennig, 1955, S........................................... 253 emmesa Malloch, 1934, N. .............................. 231, 233 enigmatica Albuquerque, 1955, P............................. 185 enormis Albuquerque, 1956, c. ................................. 217 equator Snyder, 1941, H ................................... 112, 116 equator Snyder, 1958, N .................................. 232, 236 equatoralis Coelho, 1998, 1'. ..................................... 124 equatorialis, Couri & Marques, 2001, B .................... 211 etesia (Giglio-Tos, 1893), H. .................................... 118 eurycephala (Stein, 1911), H. ................................... 106 Euryomma Stein, 1899 .............................................. 20 evecta (Harris, 1780), H . ........................................... 118 evittata Malloch, 1934, S ... ...................................... 250 exotica Carvalho & Couri, 1992, A .......................... 175 exul Williston, 1896, L.. ..... ...................................... 181 fallax (Stein, 1911), H. ............................................ .1 06 falsifica Dodge & Aitken, 1968, P............................. 90 Fannia Robineau-Desvoidy, 1830 .............................. 20 fasciculatus (Malloch, 1934), P................................. 87 fasciventris (Wu1p, 1896), p. ...................................... 89 felsina (Walker, 1849), N. ........................................ 156 femorata (Stein, 1911), D ................................. 100, 104 femorata (Stein, 1911), S......................... 193, 194, 196 fenestrata (Bigot, 1888), S....................................... 256 ferruginea Malloch, 1928, M ................................... 149 flaviantennata (Couri, 1982), c. .............................. 144 flavicomis (Malloch, 1928), N. ............................ 60, 61 flavicoxa Malloch, 1934, B ............................... 211-213 flavidicincta (Stein, 1904),1'. ...................................... 55 flavipalpis Albuquerque, 1956, N. ............................. 222 flavipalpis Malloch, 1934, A .... ............................... 206 flavipalpis Pamplona, 1983, D ................................. .42 flavipalpis Snyder, 1957, S ...................................... 198 flavipes (Williston, 1896), N. .................................. 235 flavithorax (Stein, 1904), P. ................................ 53, 56 fluminensis Albuquerque, 1954, p. .................... 157, 158 jluminensis Couri & Lopes, 1988, M. ........................ 171 frontalis Couri, 1984, P............................................. 93
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
----~.:::--=-----
;--'
----~~-
---:-
c. ...................................
281
julvapoda Snyder, 1940, H. ...................................... 118 julvapoda Snyder, 1949, P.............................. 157, 158 julvifrontis Couri & Albuquerque, 1979, N. .................222 julvipes (Bigot, 1887), D ........................................... .41 julvocalyptrata Malloch, 1934, H ................... .l13, 116 julvohumeralis (Malloch, 1922), 8 ............................77 julvolateralis Stein, 1911, P. ...................................... 96 jumicosta Dodge, 1968, P.....................................90, 95 jumicosta Snyder, 1941, M. ................ .................... .168 jumipennis (Albuquerque, 1954), P................... 240, 241 jumipennis (Wulp, 1896), M. .................................. 169 jumipennis Albuquerque, 1953, A. ............................ 175 juscicosta (Curran, 1934), C............................ .142, 144 juscicosta Malloch, 1934, S..................................... 253 juscifacies Malloch, 1934, S. ................................... 249 juscipalpis (Stein, 1911), S...................................... 254 juscisquama (Snyder, 1954), c. ............................... 147 juscisquama (Wulp, 1868), M., comb.n .................... 69 juscocalyptrata Malloch, 1934, H ............................ 109 juscocalyptrata Macquart, 1855, H. ........................... 64 juscomarginata (Snyder, 1949), x. .......................... 159 gagnei Couri, 1983, P................................................ 94 gallicola (Albuquerque, 1958), D............................. .1 02 gaminarai Carvalho & Pont, 1993, C. .................... 217 garleppi (Stein, 1911), S................................. 257, 258 garrula (Giglio-Tos, 1893), L.. .................................. 181 gemina (Wiedemann, 1830), c. .............. 143, 144, 148 geminata (Stein, 1904), c. ...................................... 140 geniculata (Bigot, 1885), N. .................... 227, 233, 234 geniculata (Macquart, 1851), c. ................... '" ........ 219 genupuncta (Stein, 1904), N. .................................. 235 giacomeli (Carvalho, 1981), D ................................ 103 gigantea Albuquerque, 1956, H. .............................. .l18 glaucinis Dodge & Aitken, 1968, P........................... 92 gluta (GigJio-Tos, 1893), C. ..................................... 142 goianensis Lopes & Khoury, 1995, N. ...................... 152 golbachi Snyder, 1957, S.......................................... 191 golbachi Snyder, 1957, X....................................... .159 gracilis (Stein, 1911), L.. ........................................... 185 gracilitarsis (Stein, 1911), S.................................... 193 grajauensis (Albuquerque, 1957), P.......................... 12l grandis (Couri, 1982), P.......................................... 123 grandis (Schiner, 1868), H ....................................... 118 grandis Couri, 1984, p. ........................................ 91, 95 Graphomya Robineau-Desvoidy, 1830.. .30, 163, 164, 166,259
Graphomyini. ............................................................ 19 griseovirens Malloch, 1928, D................................. 178 guatemala Snyder, 1951, L. ..................................... 187 guttipennis (Thomson, 1869), S...................... 255, 256 Haematobia Le Peletier & Serville in LATREILLE et al., 1828 .................................................. 22, 59, 130
Haroldopsis Albuquerque, 1954 .......................... 218, 220 haustellata (Stein, 1911), T. ............................. 20l, 202 Helina Robineau-Desvoidy, 1830 ............................. 33, 79, 99, 105, 120, 130, 169
282
Hemichlora Wulp, 1893 .............................. 26, 167, 259 hirticeps (Stein, 1911), S.................... ,..................... 191 hirticeps Stein, 1911, D ............................................ 178 hirtitibia (Stein, 1911), S.......................................... 257 hirtitibia Pamplona, 1986, M. .................................... .44 holti Malloch, 1923, x. ........................................ 57, 58 houghi Malloch, 1916, H. .................................... 65-67 hucketti Coelho, 1998, P......................................... 125 hugonis Carvalho, 1989, p. ..................................... 122 humeralis (Stein, 1918), M ................................. .43, 47 Hydrotaea Robineau-Desvoidy, 1830 .............................. .
................................. 24, 63, 64, 66, 67, 70, 74, 130 Hydrotaeini ................................................................ 81 hydrotaeijormis Snyder, 1958, N .............. 230, 233, 236 ignava (Harris, 1780), 0 .......................................70-73 immaculata (Walker, 1836), N. ............................... 238 immaculiventris (Malloch, 1923), S......................... 191 immunda (Stein, 1911), c. ............................... l07, 143 inaequalis Malloch, 1934, c. .................................. 216 inaequifrons Malloch, 1934, L. ................................ 186 incaica (Stein, 1911), T. ........................... 20l, 202, 204 incognita (Snyder, 1954), c. .................................... 147 inconspicua Malloch, 1934, S......................... 196, 198 inculta (Stein, 1911), S.................................... 257, 258 incurva (Stein, 1911), N. .......................... 220, 234, 235 inepta (Stein, 1911), H. .. ................................. 112, 116 inermis (Stein, 1904), M ........................................... 169 inflexa (Stein, 1918), N. ........................................... 152 ingrata (Couri, 1982), P....................................... 52, 55 iniqua (Stein, 1911), L.. .......................................... 181 iniqua Stein, 1911, C .............................................. 215 insons (Giglio-Tos, 1896), M ................................... 169 instabilis Snyder, 1949, N. ......................................... 153 Insulamyia Couri, 1982 ..................... 36, 211, 219, 259 insularis Couri, 1983, P. ............................................. 93 insularis (Williston, 1896), C.................................. 218 insularis Hennig, 1957, L .......................................... 187 insularis Malloch, 1934, S....................................... 256 integra (Stein, 1911), S.......................................... .199 intoniclunis Snyder, 1957, N. ................................... 226 inuber (Giglio-Tos, 1983), c. .................................. .l48 inusitata (Malloch, 1934), c. .................................... 217 inusitata Couri, 1982, I ............................................ 219 irritans (Linnaeus, 1758) H .......................................... 59 itatiaiensis Albuquerque, 1954, N. ............................. 223 Itatingamyia Albuquerque, 1979 ............ 27, 80, 82, 130 jamaicensis (Carvalho, 1983), D .............................. 101 kuscheli Hennig, 1955, N. ....................................... 238 laeta Stein, 1911, L.. ............................................... 180 laevis (Stein, 1911), L.. ............................................ 185 lancifer (Malloch, 1934), P................................... 83, 86 lanigera (Stein, 1918), N. ........................................ 235 lasiosterna Snyder, 1941, H ....................................... ll1 latana Snyder, 1949, L.. ........... ................................ 183 latifrons (Thomson, 1869), N .................................. 235
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
latiJrons Malloch, 1932, N. .................................. 60, 61 latifrons Malloch, 1934, S............................... 195, 197 latiJrons Snyder, 1957, L. ......................................... 187 latiJrontata Albuquerque & Lopes, 1982, N. ............. 155 latimaculata (Albuquerque, 1956), N. ....................... 224 latimana Malloch, 1934, S................................ 196, 199 latimanoides Snyder, 1957, S................................. .199 latinervis (Stein, 1904), p. .......................................... 121 latipalpis (Stein, 1918), P................................ .I56, 157 latipennis (Hough, 1900), M .................................... 169 latipennis (Stein, 1904), H ..................................... .118 latitibia Albuquerque, 1957, 217 lativittata Malloch, 1934, S..................................... 250 latomensis Snyder, 1957, M ......... ........................... 169 lengitinosa Snyder, 1957, P..................................... 119 leucocephala (Wulp, 1896), H. ............................... 118 leucoprocta (Wiedemann, 1830), B ................. 212, 213 leucotelus (Walker, 1853), L.. ................................... .181 leucotrichia Albuquerque, 1956, C........................... 217 levida (Harris, 1780), M ............................................. 81 levis (Stein, 1911), L. ................................................ 184 limbata (Wiedemann, 1830), H . .............................. 118 limbata Bigot, 1885, L.. ........................................... 181 limbinervis (Stein, 1918), D ..................................... 100 Limnophora Robineau-Desvoidy, 1830 ........................... . ..................................... 30, 178, 184, 191, 192,259 Limnophorini.. ........................................................... . ............. 19, 134, 173, 174, 176, 178, 182,200,259 lineata (Stein, 1904), N. ........................................... 235 lisarba Snyder, 1949, L.. .. ....................................... .184 Lispe Latreille, 1797 ..................... '" .26, 182, 184, 259 Lispoides Malloch, 1920 ............. 34, 83, 184, 187, 259 lispomina Hennig, 1955, S...................................... 199 listrata Albuquerque, 1954, C................................... 219 longipalpis (Stein, 1911), T.. ............................ 201-203 longipede Albuquerque, 1956, 217 longipes (Stein, 1911), M .......................................... 149 /ongipila (Stein, 1918), H ................................ 112, 113 longipilis Albuquerque, 1954, P................................ 242 longiseta (Lopes & Khouri, 1989), L. ...................... .181 longiseta (Wulp, 1896), M ...................................... 169 longispina (Malloch, 1923), P................................. 188 lopesae Pamp1ona, 1986, M ............................... ,45, 47 lopesi Albuquerque, 1955, L.. .................................. .187 lopesi Couri, 1983, P................................................... 89 /unatisigna Snyder, 1957, N. .................................... 226 luteola Albuquerque, 1956, H .................................. 118 lyncii (Wulp, 1883), H ............................................. 118 machadoi (Albuquerque, 1958), D ............................ 101 macrocera (Wulp, 1896), N ..................................... 227 macrops Snyder, 1949, N ........................................ 150 macrossoma Wulp, 1896, C ................................ 137, 138 maculata (Scopoli, 1763), G..................................... 165 maculipennis (Macquart, 1843), C ................... 142, 146 maculipennis (Macquart, 1846), p. ....................... 50, 51
c. ..................................
c. ...............................
maculipennis Malloch, 1934, A ....................... 207, 209 maculipes (Stein, 1918), H .... ................................... 118 maculiventris Malloch, 1929, A ................ 207, 209, 210 macu!osa Snyder, 1951, C........................................ 136 macu!osa Stein, 1911, p. ......................................... 122 magellani Malloch, 1934, S.............................. 255, 256 magnicornis Snyder, 1958, N ................... 232, 234, 236 major Albuquerque, 1956, P........ ................................ 54 major Carvalho, 1984, P......................................... 125 major Malloch, 1934, B .............................................77 malacophaga Lopes, 1938, C .................................. 138 maldonadoi Snyder, 1957, N. .................................. 229 mallochi Pont, 1972, S........................................... 248 marginalis (Stein, 1904), M ................ ..................... 169 marginata Stein, 1904, L.. .............. ............................ 181 marginata Stein, 1918, p. ................................. 120, 125 marginipennis (Stein, 1904), H ......................... llO, 114 marginipennis Stein, 1911, L .................................. 179 maricaensis Couri & Motta, 1994, B ........ ............... 213 masoni Couri,1986, P........................................... 91, 95 matogrossensis Couri & Lopes, 1988, M ................... 171 mediana Couri, 1984, P.............................................. 93 mediana Snyder, 1949, N .......................................... 152 medinai (Snyder, 1957), P................................ 240, 241 meditabunda (Fabricius, 1781), M .......................... .170 mega!opyga Albuquerque, 1954, C ............................ 219 Megophyra Emden, 1965 ........................................... 68 mel/ina (Stein, 1904), P............................................. 55 mellina (Stein, 1918), C.......................................... 147 mendozana (Enderlein, 1935), G............................. 165 meraca (Wulp, 1896), H ......................................... 1l8 meridia Snyder, 1941, M ........................................ .I68 meridionalis Townsend, 1892, G............................ .165 mesofulvata (Albuquerque, 1959), N ......................... 224 mexala (Snyder, 1957), P........................................ 124 mexicana (Macquart, 1843), N ................................. 151 mexican a Carvalho, 1984, P........ ............................. 124 mexicana Giglio-Tos, 1893, G................................. 166 micans Snyder, 1957, N. .................................. 220, 230 microchaeta (Malloch, 1934), N .............................. 235 Micropotamia Carvalho, 1993 .......... 29,68,69,74, 130 mima (Townsend, 1927), P. ....................................... 88 mimica (Snyder, 1954), C...................................... .145 mimicola Dodge, 1968, P.............................. 88, 90, 95 minor Albuquerque, 1956, P............................ 52, 54, 56 minuscula (Albuquerque, 1955), M ...................... 68, 69 minuscula (Wulp, 1896), L ...................................... 181 minuscularis Albuquerque, 1956, C.......................... 217 minuta (Stein, 1911), S........................................... 254 minuta Carvalho, 1984, P......................................... 125 mo!esta Meinert, P.................................................... 88 monacha (Schiner, 1868), H ................................... 118 monochaeta Snyder, 1957, p. .................................. 120 monstrata (Wulp, 1896), C ...................................... 148 montanhesa Albuquerque, 1952, X ....................... 57, 58
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
283
Morellia Robineau-Desvoidy, 1830".24, 42, 43, 46, 47, 130 mulcata (Giglio-Tos, 1893), H."""" ........ " .............. I 18 Mulfordia Malloch, 1928 ...... " ................... 25, 149, 259 multomaculata, (Stein, 1904), C. .................... 142, 145 Musca Linnaeus, 1758 ................................. 24, 48, 130 Muscina Robineau-Desvoidy,1830 ........... 32, 80, 98, 130 Muscinae ............................................... 19, 37, 40, 130 Muscini ................................................. 19, 40,58, 130 Mydaea Robineau-Desvoidy, 1830 ............................ . ...................................... 30, 134, 167-170, 172,259 Mydaeinae .................... 19, 80, 130-135, 160, 164, 259 Mydaeini ................................................................... 19 Myospi/a Rondani, 1856 .............. 31, 167, 170, 171,259 napensis Couri, 1996, S............................ 248, 250, 251 nal"Ona (Walker, 1849), L.. ...................................... 180 nasoni (Stein, 1898), L. ............................................. 184 nebulicola (Co uri & Carvalho, 1995), A ........... 176, 177 neglecta (Townsend, 1939), N ................................... .49 Neivamyia Pinto & Fonseca, 1930 ......... 22, 59-61, 130 neoaustralis Snyder, 1957, N ..... 221, 227, 231, 233, 236 neobiseriata Snyder, 1957, S.................................... 251 Neodexiopsis Malloch, 1920 ................................ .. ............................... 36, 134, 218, 220, 225, 236, 259 neoflavipes Snyder, 1957, N. ..................................... 229 neomacrocera Snyder, 1957, N .................................. 227 Neomuscina Townsend, 1919 ..... 27, 134, 149, 155,259 Neomusciniopsis Albuquerque & Lopes, 1982 ............ . .............................................................. 32, 155, 259 Neomyia Walker, 1859 ................................. 23, 49, 130 Neorype/lia Pont, 1972 ............................... 23, 49, 130 neosimilis Snyder, 1942, N ....................................... 154 neosimplex Snyder, 1957, H ..................................... 118 neotrita (Snyder, 1954), c. .............................. 142, 145 neotropica Carvalho & Pont, 1993, C...................... 217 neotropica Carvalho & Pont, 1993, S...................... 251 neotropica Curran, 1934, N. .................................... 154 neotropica Snyder, 1951, H. .................................... 118 neotropica Snyder, 1957, A ....................................... 63 nervicincta Stein, 1918, P................................. 120, 125 Neurotrixa Shannon & Del Ponte, 1926.,,26, 134, 155,259 nicholsoni Curran, 1939, H. .................................. 65-67 nielseni Dodge, 1968, P.............................................. 88 nigerrima (Malloch, 1934), N. ................................. 223 nigerrima Carvalho, 1984, P. ................................... 124 nigra (Lopes & Couri, 1987), S............................... 199 nigra (Stein, 1911), S. .............................................. 255 nigra Couri & Carvalho, 1996, P......................... 53, 54 nigra Dodge & Aitken, 1968, P. ................................ 90 nigrargentata (Albuquerque, 1954), L. ...................... 181 nigribasis (Stein, 1911), L.. ...................................... 186 nigriceps (Stein, 1911), T.. .. ..................... 20 I, 202, 203 nigricornis Malloch, 1934, B ........................... 212, 213 nigricosta Hough, 1900, M. ................................ .45, 47 nigricosta Snyder, 1949, N ..................................... 152 nigrimono (Macquart. 1851), H. ...... 107, 109, 113, 117
284
nigrina (Wiedemann, 1830), H. ................................ 106 nigripes Stein, 1911, T.. ............................................ 200 nigripoda Snyder, 1949, P....................................... 157 nigripuncta Stein, 1911, P....................................... 123 nigrithorax Stein, 1911, S........................................ 251 nigritibia Snyder, 1949, D ........................................ .42 nigriventris (Albuquerque, 1954), P........................... 121 nigriventris (Malloch, 1928), P.................................. 87 nigriventris Malloch, 1934, S.................................. 196 Iligrohalterata (Stein, 1904), 0 ................................ 239 nigromarginata (Stein, 1911), H. ............................ I06 nivaioides Albuquerque, 1956, H. ............................. 118 nobilis Albuquerque, 1956, H. ................................... 118 Iloctiluca (Albuquerque, 1958), D ............................. l02 l10niesmaculata Gaminara, 1930, C......................... 217 normata (Bigot, 1885), B ................................ 161, 163 notha Snyder, 1941, H. ............................................ I 16 Notoschoenomyza Malloch, 1934 ............. 35, 237, 259 l1ovissimo Couri & Albuquerque, 1979, N. .................. 224 l1ubilicosta Malloch, 1923, H. ................................... 64 nubivena Snyder, 1941, M........................................ 168 nudapieura Snyder, 1949, P.................................... 157 nudiseta (Wulp, 1883), S............................................ 98 nudiseta Albuquerque, 1954, A ................................. 135 nudistigma Snyder, 1949, N. ............................. 151, 153 obscllra (Shannon & Del Ponte, 1926), M ............... 170 obscura (Wulp, 1898), P. ........................................... 88 obscurinervis Couri,1984, P....................................... 91 obtusiloba (Malloch, 1934), N ........................ 230, 236 ocasionalis (Couri, 1982), C.................... 141, 142, 146 occidentalis Arntfield, 1975, C ................................. 166 ochricornis (Wiedemann, 1830), M .................... .44, 47 oculata (Stein, 1911), N. ......................................... 235 Ophyra Robineau-Desvoidy, 1830 .... 28,64,70,73,74, 130 oportuna Albuquerque & Lopes, 1979, p. ............. 50, 51 orba Stein, 1911, M ................................................ 169 orbitalis (Stein, 1904), P. .............................. 53, 55, 56 orienta lis Schiner, 1868, A .......................................... 38 oscillans (Wulp, 1896), N. ........................................ 235 ovativentris (Macquart, 1851), L. .................... ......... 181 Oxytonocera Stein, 1911.. .......................... 37,238,259 Pachyceramyia Albuquerque, 1955 ................................ . .............................................. 37, 134, 187-189,259 pacifera Giglio-Tos, 1893, B .................................... 211 pallicornis Malloch, 1934, S............................ 252, 253 pallidicornis (Bigot, 1887), M .................................. 171 pallipes (Stein, 1918), C................................... 142-144 Palpibracus Rondani, 1863 ............. 33, 79, 82, 83, 130 pampiana (Shannon & Del Ponte, 1926), P ................ 97 panamensis Dodge, 1965, C .................................... 166 pansa (Giglio-Tos, 1893), M .................................... 169 pantherina (Wiedemann, 1824), p. .................. 188, 189 Paracyrtoneuropsis Pamplona, 1999 ....................... 142 paraensis Carvalho, 1999, S............................ 127, 129 paralis (Giglio-Tos, 1893), N. .................................. 151
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
paranaensis (Albuquerque, 1954), L ........................ .182 paranaensis Carvalho, 1993, D ...... ......................... I 00 Parapyrellia Townsend, 1915 ................ 24, 50, 51, 130 pararescita (Couri,1995), c. .................................... 146 parsura (GigJio-Tos, 1893), H ................................. 118 parvula (Wulp, 1986), H ......................................... 1l4 parvula Albuquerque, 1958, N. ................................. 221 Passeromyia Rodhain & Villeneuve, 1915 ............. 81, 88 patagoniea Malloch, 1934, L. ...... ............................ 181 paueiseta (Stein, 1904), S ........................................ 199 paueiseta Stein, 1911, M ...... .................................... 169 paula (Medeiros, 1980), P........................................ 241 paulistana (Lopes & Khouri, 1991), L ..................... 182 paulistensis Albuquerque, 1956, N. ........................... 222 paulistensis Pamp10na & Mendes, 1995, M ............... .44 pectinata Couri & Albuquerque, 1979, N ............ 221, 225 peetinata Malloch, 1934, R ............................. 243, 246 pedella (Wiedemann, 1830), H ................................. 106 peninsula Snyder, 1958, N. ....................... 233, 234, 236 pennata Malloch, 1934, A ....................... 206, 209, 210 Pentacrieia Stein, 1898 ..................... 31, 239, 240, 260 peruviana Snyder, 1958, N. ...................... 232, 233, 236 peruvianus (Malloch, 1929), p. .................................. 87 petersoni Couri, 1984, p. .............................................. 91 Phaonia Robineau-Desvoidy, 1830 .................... .
.................................... 31,79,99,105,119,120,130 Phaoniinae ............................................. 19, 37, 98, 130 Philornis Meinert, 1890 ................... 25, 88, 94, 95, 130 pica (Macquart, 1851), L. ........................................ 182 piei (Macquart, 1854), P............................................. 93 pieta (Stein, 1911), S ............................................... 254 pieta Stein, 1911, c. ................................................. 214 pictipennis (Bigot, 1878), N. .... ................................ .153 pietipennis nudinervis (Stein, 1918), N. .................... 153 pictipennis pietipennis (Bigot, 1878), N. .................. .153 pilieeps (Stein, 1911), H. ......................................... 106 piliseta Stein, 1919, L.. .................................... 179, 180 Pilispina Albuquerque, 1954 ....................... 36, 240, 260 pilitarsis Stein, 1911, T. ...................................... 201-204 pilitibia Albuquerque, 1954, p. ........................... 240, 241 pilitibia Stein, 1911, C ................ ............................. 215 pilosa (Stein, 1904), N. ............................................ 235 pilosus (Macquart, 1851), p. ...................................... 84 platystoma (Thomson, 1869), L.. .............. .............. 182 plaumanni (Carvalho, 1983), D ....................... I03, 104 plaumanni Snyder, 1941, M .................................... 168 plumata (Carvalho, 1981), p. ............................... 74, 75 plumata Albuquerque, 1955, B .................................. 213 plumbea Albuquerque, 1954, c. ................................217 plumbijrons (Albuquerque, 1956), N. ........................ 223 plumiseta Stein, 1911, C.................... ..................... 215 Plumispina Albuquerque, 1954 ........... 35,241,242,260 podexaurea (EnderIein, 1935), C ............................ 166 poeeiloptera (Schiner, 1968), H ...................... 1l1, 1 15 Polietina Schnabl & Dziedzicki, 1911.. ........ 25, 51, 130
polystigma (Wulp, 1896), c. ................................... .147 ponti Coelho, 1998, P............................................... 125 ponti Couri, 1987, N. ............................................... 235 ponti Lopes & Khouri, 1995, N. .............................. 150 porteri (Brethes, 1920), L.. ...................................... 182 porteri Dodge, 1955, p. ....................................... 88, 90 Potamia Robineau-Desvoidy, 1830 ....... 29, 74, 75, 130 praeeipua (Walker, 1853), H. ...... ........................... 118 praenubila (Snyder, 1954), c. .................................. 147 praesuturalis (Stein, 1904), p. .................................. 120 preacuta Snyder, 1958, N. ............................... 232, 236 prima (Couri & Machado, 1990), P...................... 52, 53 prima Malloch, 1921, 159 prineipalis (Schiner, 1868), M .................................... 81 priseipagus Snyder, 1958, N. ................................... 232 pro cedens (Walker, 1861), H .................................. 119 proeera Stein, 1911, C. ...........................................215 projeeta (Malloch, 1934), C. ................................... 217 prolatijrons Snyder, 1940, H ................................... 119 propinqua (Stein, 1911), L.. .................................... 185 protosetosa (Snyder, 1954), C................................. .145 pruna (Shannon & Del Ponte, 1926), S ..................... 56 Pseudoptilolepis Snyder, 1949 ....... 25, 78, 156, 158, 259 Psiloehaeta Stein, 1911.. .............................. 32, 96, 130 pteropleuralis Malloch, 1934, R ............................... 245 pteropsila Stein, 1918, P. ................................. 120, 125 pubiceps (Stein, 1911), L. ................................ 185, 187 punetinervis Stein, 1911, p. ...................................... 123 punetulata (Wulp, 1896), N. ............................ 231, 233 punoensis Coelho, 1998, p. ..................................... 123 pura (Stein, 1911), S ............................................... 193 quadristigma (Thomson, 1869), B ..................... .I62, 165 Quadrularia Huckett, 1965 ............................. 105, 120 quercus Coelho, 1998, P........................................ .I22 querula Dodge & Aitken,1968, P............................... 92 quintivena Snyder, 1957, N. ..................................... 225 rava Snyder, 1957, N. .............................................. 227 reeedens (Stein, 1904), N. ........................................ 235 refusa (Giglio-Tos, 1893), H. ..................................... 111 regina Carvalho, 1993, D................................. 100, 104 regobarrosi Albuquerque, 1958, H. ........................... 119 Reinwardtia Brauer & Bergenstamm, 1889 .. 31, 97, 130
x. .........................................
Reinwardtiinae .......................................................... 134 Reinwardtiini... ................................ 19, 62, 76, 80, 130 rescita (Walker, 1861), C ...... ................................... 146 rettenmeyeri Dodge, 1963, P...................................... 89 rex Curran, 1928, N. .. ............................................... 228 reynoldsi (Malloch, 1934), P....................................... 84 Reynoldsia Malloch, 1934 ................. 37, 242, 243, 260 Rhabdotoptera Stein, 1919 ................ 36, 134, 189,259 robusta (Stein, 1911), R ........................................... 243 robusta Carvalho, 1984, P....................................... 124 roppai Pamplona, 1986, M ................................. .4S, 47 rotundiventris Stein, 1911, c. ................................... 215 rubella (Wu1p, 1896), P............................................ 54
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
285
rubripalpis (Wulp, 1896), H.""." ... """" .. """"""" 119 rujiguttata (Macquart, 1851), X .. """""" .. "" .. """" . .159 rujipes (Macquart, 1851), N. .... " .. """"" .. "" .. "" .. ,,,,235 rujitibia (Stein, 1911), N."" .. """ .. " ........ " ...... " .. ",,.228 rujitibia Stein, 1911, T."""" .. """""""" ...... """,,,,,,200 rujitibialis Macquart, 1843, L."""""""" .. """"" .. "" 184 rujoapicata Malloch, 1934, H .. """""""" .. .108-110, 113 rujoapicata Malloch, 1934, R.. """"""""""",,245, 246 rujoscutella Dodge, 1955"""""" .. """"""""""""".154 rujoscutellaris Couri, 1983, P.. "" .. """""""""""""".89 rustica Albuquerque, 1956, N .. """""""""""",,222, 225 sabroskyi Albuquerque, 1957, p."" .. """" ..... "" ........ ,, .. 91 saeva (Wiedemann, 1830), L.. """""""""""""""".179 sanespra Snyder, 1949, N."" .. "" .. """"""""""" .. ".151 sanguinis Dodge & Aitken, 1968, p. ........ " .. " .. "" .. " ... 93 santoamarensis (Albuquerque, 1958), D""" .. ".101, 104 saphirina (Seguy, 1935), D ........ " ............ """ ...... "",,.41 sarcophagina (Wulp, 1896), S.. """"" .. """ .... ",,,, .... 56 Sarcopromusca Townsend, 1927"""""" .. ,,23, 55, 130 scabra (Giglio-Tos, 1893), p. .................... "" ....... 74, 75 Scenetes Malloch, 1936, stat.rev...... 30, 167, 171, 259 schadei Snyder, 1949, N. ...... " .. " ............................ ,,154 schildi Dodge, 1963, p. ............ " .... " .......... "" .......... ".89 Schoenomyza Haliday, 1833 ...... """ ... 35, 247, 254, 260 Schoenomyzina Malloch, 1934""""".34,251,253,260 scordalus (Walker, 1861) H ...... " ...... "" .......... " ...... 167 scutellata (Johnson, 1919), N." .... " ................... 150, 154 scutellaris (Fabricius, 1805) S.. " .............. """ ... I72, 173 scutellata Malloch, 1934, R .... " ........ " .... " ........ " .. ".245 Scutellomusca Townsend, 1931.." .. " .. "" .. ,,25, 172,259 secunda Snyder, 1951, M ...... """"" .... ""."",, .. ,, .... ,,149 seguyi Garcia, 1952, P.. " ...... " .. " .... " .... " .. "" ...... " .. " .. 91 seguyi Pamplona, 1983, D."""" .. "" .. "." .. " .. """" .. ".42 semicinerea (Stein, 1911), S.. """"""" ...... """",,,,.,,191 semimarginata (Stein, 1918), M .. " .. "" ...... " .. "" .. "" . .45 sensitarsis (Carvalho, 1983), D .... " ...... "" ........ "" .... " 101 separatus (Malloch, 1934), P.... " .. """" .... " .. "",, ....... 84 sera (Giglio-Tos, 1893), H .. " .... " ........ " .. " ............ ".119 seriata (Stein, 1911), c." ...... "" .. "" .... "... 141, 143, 144 serotina Wulp, 1896, L. ............ "" .... " .. " ........ " .. "".183 setinervis Dodge, 1963, P...... " .... " .... " .... " .... "" .. " ..... 88 setipuncta Snyder, 1957, N .. ""." .. " .. " ........ """ .. "".226 setitibia (Stein, 1911), S........ " .. " .. """ .... " .. """"",,.193 setiventris Stein, 1911, c. .... ".......... ".. ".. ".... "........ 215 setuligera (Stein, 1911), L. .... """"" .. """"" .. ,, ..... ,,.183 sexpunctata (Wulp, 1883), M .. """ .. " .. "" .. """ ...... ,, 169 shannoni Carvalho & Pont, 1993, P........................ " 121 signatipennis (Wulp, 1896), H .. " .... " .. " .... " ..... III, 115 sima Snyder, 1957, N ............ """" .. " .. "."""" .. ,, .. ,,.228 similata (Albuquerque, 1957), p.""" .. "" .... " ...... "",,.121 simi/ata (Couri, 1982), C........ "." .. " .. "" .... """" .. " .. 146 simi/ata Snyder, 1949, N .. "" .. " .... """"" ..... " .. "",, .. 154 similis (Malloch, 1934), p." .. " .. """""" .... "" .... ,, .... ,,84 simplex (Albuquerque, 1958), D ....... """ ...... "." ..103, 104 simplex Malloch, 1934, H .. """""",,108, 109, 112, 116
286
singularis (Stein, 1919), S........ " .. " .. "" ...... " ... 127, 129 sinopensis Pamplona, 1986, M .. " .. " ... ,,, .... ,, .. ,, ... .45, 47 socia (Wulp, 1896), H" ............ " .... " ........ " .......... ,,115 sociabilis (Blanchard, 1937), N. .. " ................... 220, 235 sociabilis Blanchard, 1937, B. "" .... "" ....................... 234 solitaria Albuquerque, 1958, 0 ...............................71-73 soratiensis Coelho, 1998, P.... """" .................. ,, ...... 123 Souzalopesmyia Albuquerque, 1951.."" ...... 33, 126, 128-130 sparsiplumata (Stein, 1918), N." .... " .. " .. " ........ " ..... 154 Spathipheromyia Bigot, 1884 .. .35, 237, 247, 254, 256, 260 spatulata Malloch, 1934, A. """ ...... " .... "" .. " .. 208, 209 spermophila (Townsend, 1895), P........ " .... " .... " ........ 94 spicatus (Malloch, 1934), p. .. " .. " .... """ .. "" ........ ,, ..... 86 Spilogona Schnabl, 1911..""" ... 34, 178, 190, 191,259 spi/optera (Wiedemann, 1830), C............. 141, 142, 145 spinicosta (Stein, 1904), N .. " .... " .. "" .. " .......... " ....... 238 spinijera (Wulp, 1883), D .. " .. " .... """ ........ "" .. ,, ....... .41 spinuligera (Stein, 1911), S.. " .. " .. " ................ " .. " ... I92 spontanea Carvalho, 1993, D.... """ .... """ .... ,,103, 104 spreta Malloch, 1921, L.. .... "" .. " .. " .. " .. "" ...... " ....... 182 stabilis (Stein, 1911), N. .......................................... " 154 stabulans (Fallen, 1817), M .. " .. " .. """ ...... " .......... ,, ... 81 stein! (Pont, 1972), P........ " .. "" .. " .... " ........ " .......... 88 steini (Enderlein, 1927), P.. " ........ " ...... " ........ 53, 54, 56 stein! (Snyder, 1954), C..... "" ...... " .. " .................. " .. 144 steini Carvalho & Pont, 1993, S...... " ........ " ...... " ..... 199 stellata (Couri, 1982), p. .......... " ........................... 52, 53 stellata Bigot, 1884, S.... " .. "" .. " .. """" .. "" .. " .... ,, ... 256 stigmatica (Lopes & Couri, 1987), S.. " ........ " ......... 199 stigmatica Malloch, 1928, c. .... "" .. "" .............. "........ 78 st!pata (Walker, 1853), G .. "" .. " .. "" ........ "" .. " .... " .. 166 Stomopogon Malloch, 1930 .. " ...... " .. " ....... 35, 257, 260 Stomoxyini." .. " .... " .... """ .. "" .. """",, .. 19, 40, 58, 130 Stomoxys Geoffroy, 1762."""""" .. "" .. """.22, 60, 130 strenua Stein, 1911, C.. "" .. "." .. " .. " .... " .. " ...... " ...... 215 striatipennis (Stein, 1898), R .. "" .... " .. " ........... .189, 190 strigata (Giglio-Tos, 1893), L.. .. """ .. " ........ ,, ...... ,," 182 suavis (Stein, 1911), S....... " .. " .. " .... " .. " ...... " .. " ..... .193 subreptilia Albuquerque & Lopes, 1979, x. ................ 159 subtilis Couri & Albuquerque, 1979, N ....................... 222 suljuriceps Malloch, 1934, N. .. " .. " .. " ...... " .............. 237 sulina Carvalho, 1999, S...... "" ....................... 127, 129 sulina Couri, 1986, N .. " ..... "" ...... "" ....... """,, ........ 235 Syllimnophora Speiser, 1923 .......... " .. " ..... 29, 192,259 Synthesiomyia Brauer & Bergenstamm, 1893 ................ . " ...... """ .... "" ....... """" ... ",, ....... ,, .... 31,81,98, 130 tachinina Brauer & Bergenstamm, 1889, R................. 98 tachnoides (Albuquerque, 1958), D .......................... .102 tarsalis (Stein, 1918), H .. " ... " ........ " .......... "" ...... ",,119 tarsalis Malloch, 1934, S........ " ....................... 249, 250 tarsata (Snyder, 1957), C.................... " ........ "" ........ 217 tauota Lopes, 1984, N .... "." ... " ............ "",, .............. 154 tentaculata (De Geer, 1776), L.""" .......... " ........ ,, ... 183 tenuicornis (Wulp, 1896), N. .. "" ........................ " ... 235 tenuior (Walker, 1853), C... "" ...... """ ......... ,, ......... 217
Muscidae (Diptera) of the Neotropical Region; TAXONOMY
tenuipennis Malloch, 1934, S.......................... 194, 197 tesselata (Stein, 1904), H. ........................................ I06 tetragona Gaminara, 1930, P........................... 120, 125 Tetramerinx Berg, 1898 ............................. 34, 200, 259 texensis (Malloch, 1923), D..................................... lOO Thaumasiochaeta Stein, 1911...24,30,201,203,204,259 Thricops Rondani, 1856 ............................... 28, 74, 130 tinctifacies (Albuquerque, 1958), N. .......................... 224 tinctinervis (Stein, 1918), N. .................................... 154 tinctipenis Albuquerque, 1956, c. ............................. 217 torquans Nielsen, 1913, p. .................................... 92, 95 townsendi Snyder, 1941, H ........ .............................. 113 transporta Snyder, 1949, N ...................................... 153 travassoi Lopes & Mangabeira, 1938, N. .............. 60, 61 triangularis Malloch, 1934, S.................................. 252 trichophthalma Albuquerque, 1959, C...................... 217 trichops (Malloch, 1923), D ..................................... .42 trichops (Stein, 1911), S......................................... .191 trichops (Stein, 1918), H ......................................... 119 trichops Malloch, 1923, D ........................................ .42 trigona (Shannon & Del Ponte, 1926), D ......... 102, 104 trigonata (Wulp, 1896), D ...................................... .101 trinitensis Dodge & Aiken, 1968 .............................. 94 triplex (Stein, 1911), L.. .................. ......................... 186 tripunctata (Wulp, 1896), N. ............................ 150, 155 triseta (Curran, 1931), p. .......................................... 123 triseta Snyder, 1949, N .............................................. 151 trispila (Bigot, 1885), P........................................... 122 trivittatus (Malloch, 1934), p. .................................... 87 trochanterata Malloch, 1934, R ........................ 244, 245 truncata (Stein, 1911), N. .......................................235 tumida Stein, 1911, C................................................ 215 uber Giglio- Tos, 1893, c. ........................................ 141 umanani Garcia, 1952, P. .......................................... 88 umbrosa (Wulp, 1896), H. ...................................... 119 unica (Stein, 1898), T.. ............................................ 200 unica Carvalho, 1993, D .......................................... I02 unicolor (Stein, 1911), S.......................................... 253 uniformis Malloch, 1934, C. .................................... 216 uniseta Malloch, 1934, L. ....................................... 186 univittata Couri & Carvalho, 1996, p. ........ .......... 52, 54 univittata Dodge, 1968, P. ........................................... 90 univittata Malloch, 1934, S..................................... 248 univittatus (Bigot, 1857), P.................................... 83, 86 uspallata Snyder, 1957, N. ....................................... 226 valdiviensis Pamplona & Couri, 2000, P.................... 83
variceps Malloch, 1934, S....................................... 198 varicolor (Hough, 1900), c. ..................................... .147 varicomis (Coquillett, 1900), B ........................ 212, 213 variegata (Stein, 1911), T.. ................................. 201-204 vecta (Giglio-Tos, 1893), N. ..................................... .151 veneris (Bigot, 1888), p. ............................................ 83 venicurva (Stein, 1904), P........................................ 124 veniseta (Dodge, 1967), D .......................................... 80 veniseta (Stein, 1904), c. ................................ 146, 147 vespidicola Dodge, 1968, p. ........................................ 93 vibrissata (Stein, 1918), c. ...................................... 137 vicaria (Walker, 1853), L.. ..................................... .182 vicina Robineau-Desvoidy, 1830, L.. ........................ 182 vierecki Snyder, 1941, H .......... .................................. 110 vilis (Stein, 1911), L.. .............................................. .183 villosa Stein, 1904, H. ........................................... 65-67 viola Malloch, 1934, H ................................... I07, 115 violacea (Robineau-Desvoidy, 1830), M. ................. .44 violaceiventris Macquart, 1851, B........................ ....... 77 violescens (Dodge, 1967), P....................................... 97 virgata (Wiedemann, 1830), L.. ........ ....................... 182 viti/is (Giglio-Tos, 1893), N ..................................... 235 vittata (Macquart, 1851), L. ..................................... 182 vittithorax Stein, 1911, C...................... ................... 215 vittivemris (Albuquerque, 1955), N. ........................... 235 vittiventris Malloch, 1934, S..................................... 197 vittoriae Lopes & Khoury, 1995, N. ......................... 154 vockerothi (Carvalho, 1983), D .......................... 99, 101 vulgaris Couri & Albuquerque, 1979, N. .................... 222 vulgaris Couri, 1984, P.............................................. 93 walkeri (Pont, 1972), c. ................................... 142, 145 walkeri Carvalho & Pont, 1993, H. ........................ .119 willinki Snyder, 1957, S............................................ 251 willistoni Snyder, 1958, N. ................................ 231, 233 wulpi (Snyder, 1954), C........................................... 147 wulpi Couri & Carvalho, 1997, p. ......................... 52, 56 wulpi Pont, 1972, C................................................. 217 xanthopoda Albuquerque, 1956, N. .......................... 225 xanthoptera Pamplona, 1986, M ......................... .45, 47 xena Malloch, 1934, H. .......... ................. 108, 111, 115 Xenomorellia Malloch, 1923 ....................... 23, 57, 130 Xenothoracochaeta Malloch, 1921.. .......... 27, 158,259 zebrina (Bigot, 1885), S.................................. .194, 195 zeketi Dodge, 1963, P................................................ 89 zosteris (Shannon & Del Ponte, 1926), N. ........ .150, 152
Muscidae (Diptera) of the Neotropical Region: TAXONOMY
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• obra • foi • impressa. na • Gnifica • Reproset • Curitiba • PR • Brasil. em • abril • de • 2002 • para. a. Editora • da • Universidade • Federal. do .Parana.
The main scope of this book is to provide means of identifying the Neotropical species of Muscidae. one of the families of Diptera with more than 4000 species in the world. In the Neotropical Region, there are about 840 species distributed in 84 genera. 80 of them in South America. Some species have a great medical and veterinary importance, acting as vectors of many diseases and other species have a considerable ecological importance. Although muscid flies are very abundant in rural or semi-rural environments, most of them are rarely noticed by man. An indirect goal of this book, is also to identify weak points in our knowledge of muscid taxonomic and diversity for the neotropics (e.g taxonomic problems, need of generic revisions, need of more collecting in some groups with few known species. among others).
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