The Hawaiian Honeycreepers: Drepanidinae (Bird Families of the World)

Bird Families of the World A series of authoritative, illustrated handbooks of which this is the thirteenth volume to be published.

Series editors C. M. PERRINS Chief editor W. J. BOCK J. KIKKAWA Freelance ornithologist, artist, and photographer H. Douglas Pratt resides in Baton Rouge, Louisiana where he is a Staff Research Associate of the Louisiana State University Museum of Natural Science. Born in Charlotte NC, he holds a B.S. in biology from Davidson College and an M.S. and Ph.D. in zoology from LSU. A Fellow of the American Ornithologists’ Union, he studies island birds of the tropical Pacific, with special emphasis on Hawaiian honeycreepers.Though without formal art training, Dr Pratt is also renowned as a zoological illustrator. He is author or illustrator of 10 full-length books and a contributor to others ranging from the National Geographic Society’s Field Guide to the Birds of North America to A Field Guide to the Birds of Hawaii and the Tropical Pacific, Mayr and Diamond’s Birds of Northern Melanesia, Phoebe Snetsinger’s memoir Birding on Borrowed Time, and the multivolume Handbook of Birds of the World. He is also widely published in both popular and scientific periodicals. He was recently honored by being asked to complete the final plate for the posthumously published fifth edition of Roger Tory Peterson’s Field Guide to the Birds. Dr Pratt is also a lecturer, ornithological consultant, and leader of ecotours. As a long time resident of the Island of Hawaii, wildlife biologist and photographer Jack Jeffrey is intimately familiar with Hawaii’s remote rainforests and rare birds.As a biologist at Hakalau Forest National Wildlife Refuge, he brings to his images the knowledge from 30 years of observation and study of Hawaii’s endemic species. He combines a naturalist’s curiosity with a photographer’s patience and technical skill to produce beautiful images. He is the recipient of the prestigious National Sierra Club 2002 Ansel Adams’ Award for Conservation Photography and the 1999 Hawaii Audubon Society Conservationist of the Year Award. Jack is co-author, with Doug Pratt, of two books on Hawaiian birds, and has had work featured in magazines including National Wildlife, BBC Wildlife, National Audubon, National Geographic, Smithsonian, Life, Natural History, as well as in numerous text books, calendars and cards. Biologist Sheila Conant grew up in Hawaii, where she became interested in birds at a young age. She earned a B.A. and M.S. in zoology at the University of Hawaii, and an M.S. in botany and Ph.D. in zoology from the University of Oklahoma. Since 1975, she has taught at the University of Hawaii where she is currently Professor and Chair of the Department of Zoology. She teaches ecology and conservation biology. Her research focuses on the life history, ecology, and conservation of Hawaiian birds, especially those of the Northwestern Hawaiian Islands. She has published numerous papers in international scientific journals. Dr Conant is a Fellow of the American Ornithologists’ Union, and a respected and influential authority on Hawaiian birds.

Bird Families of the World 1. The Hornbills Alan Kemp 2. The Penguins Tony D.Williams 3. The Megapodes Darryl N. Jones, René W. R. J. Dekker, and Cees S. Roselaar 4. Fairy-wrens and Grasswrens Ian Rowley and Eleanor Russell 5. The Auks Anthony J. Gaston and Ian L. Jones 6. The Birds of Paradise Clifford B. Frith and Bruce Beehler 7. The Nightjars and their Allies D.T. Holyoak 8. Toucans, Barbets and Honeyguides Lester L. Short and Jennifer F. M. Horne 9. Ratites and Tinamous S. J. J. F. Davies 10. The Bowerbirds Clifford B. Frith and Dawn W. Frith 11. Albatrosses and Petrels across the World Michael Brooke 12. The Grebes Jon Fjeldså 13. The Hawaiian Honeycreepers H. Douglas Pratt 14. Herons James A. Kushlan and James A. Hancock 15. The Cuckoos Robert B. Payne 16. Ducks, Geese and Swans Edited by Janet Kear

Bird Families of the World

The Hawaiian Honeycreepers Drepanidinae H. Douglas Pratt Colour plates, drawings, and photographs by

The author Bird photographs by

Jack Jeffrey With an appendix by

Sheila Conant

1

Great Clarendon Street, Oxford OX2 6DP Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Taipei Toronto Shanghai With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan South Korea Poland Portugal Singapore Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc., New York © H. Douglas Pratt, 2005 The moral rights of the author have been asserted Database right Oxford University Press (maker) First published 2005 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this book in any other binding or cover and you must impose this same condition on any acquirer A catalogue record for this book is available from the British Library Library of Congress Cataloging in Publication Data Data available ISBN 0 19 854653 X 10 9 8 7 6 5 4 3 2 1 Typeset by Macmillan India Ltd Printed on acid-free paper in China

To my mother MAUDE ERWIN PRATT and my aunt EVELYN BROWN CHOATE They planted and nurtured the seeds from which this book grew

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Preface

I first met the Hawaiian honeycreepers on a hot summer day in 1961, when I went with my father to a local discount store in Charlotte, North Carolina. What happened that day would change my life. As I often did when my father was busy looking for things to use in his woodshop, I wandered over to the small book display, not expecting much among the ‘How-to’ books and sports magazines. To my surprise, there was a single copy of the new edition of Roger Tory Peterson’s A field guide to western birds. I had already worn out several copies of A field guide to the birds, the eastern counterpart, and had an earlier edition of the western book.The latter had not been updated for a long time, and seemed rather primitive by comparison with the eastern one. Now, here was the new western guide, with all new artwork, looking every bit the equal of my old favourite. But there was something more: on the last two plates were birds that I had not only never seen but barely knew existed.They were the birds of Hawai‘i. Of course, the book went home with me and I spent the rest of the day immersed in the Hawaiian section. Ever since being given a ukulele for Christmas when I was seven, I had dreamed of these far-away islands. Hawaiian music was all the rage in the early 1950s, thanks to national promotion by The Arthur Godfrey Show, and I learned many songs on my ‘uke.’ During the same period I was becoming an active birder, encouraged by my parents and spurred on by the Bird Study merit badge of the Boy Scouts. But somehow my two passions had never crossed paths. I had previously looked perfunctorily at a book on Hawaiian birds (Munro 1944a), in the Charlotte Public Library but its illus-

trations were not very inspiring. But suddenly, Hawai‘i had real birds! Here they were, painted by my hero Peterson. And what birds they were: scarlet and black birds with long sickle-bills, yellowgreen birds with curved bills varying from short to unbelievably long, yellow-headed grosbeaks, orange birds with short conical bills, and even a yellow one whose bill did not fit together! There were also some conventional-looking warblers, thrushes, and flycatchers, but it was the exotic-looking birds with their odd Hawaiian names that particularly caught my attention. To my surprise, I soon learned that they, in all their varied forms, belonged to a single avian family, the Hawaiian honeycreepers, found only in those islands. I also learned the sad truth that there had once been many other members of the family that were now extinct. But it was a propitious time because several species long thought gone forever had been rediscovered just in time to be included in the new Peterson book. Frank Richardson and John Bowles (1964) had found all of the historically known birds of Kaua‘i in a place called the Alaka‘i Swamp. If those were found, might not others still exist in some remote spot? That night I dreamed of rediscovering extinct birds in idyllic Hawai‘i. The ensuing decade was filled with important distractions: 4 years at Davidson College, 4 years as a high school biology teacher, and a move to Baton Rouge where I began graduate studies at Louisiana State University. Always in the back of my mind was the dream of searching for those wonderful birds in the Alaka‘i.Then, in the fall of 1971, a new student strolled into my office looking for a place to set up his desk. His name was Phillip L. Bruner,

x Preface and he was from Hawai‘i. Hawai‘i! Now all my memories of reading about honeycreepers in the Alaka‘i came flooding back. I made myself a nuisance asking Phil about his birding exploits.Yes, he had been to the Alaka‘i Swamp. Yes, he had seen some great birds, including the fantastically longbilled Kaua‘i ‘Akialoa (it turned out to be the last sighting by anyone). My blood raced as he described this fantastic bird’s movements, how it perched on the upper side of a branch but reached around with its bill to probe the underside! Of course, Phil was more interested in the rich continental avifauna that was all new to him, and I was happy to help him learn my old familiar birds. Phil, his wife Andrea, and I became frequent birding companions and lifelong friends. Andrea was from Tahiti, and Phil had just written a small book on the birds of that part of the Pacific, but it lacked colour illustrations. Eventually, we decided to collaborate on a new Pacific field guide, and when the Bruners returned to Hawai‘i in 1974, they invited me to spend a month with them. Phil and I toured the islands, camping out and staying with his friends and relatives. Everywhere we went, local ornithologists and birders welcomed us with surprising hospitality, a true spirit of aloha. Although we saw some great birds, and gained access to some wonderful native habitats, I still did not get into the heart of the Alaka‘i; this trip would not be my last. As I worked on the first plates for the field guide (eventually published as Pratt et al. 1987), I became dissatisfied with the classification of the honeycreepers, and my major professor, George H. Lowery, Jr., agreed that a review of it would make a good dissertation topic. I began making longer trips to the islands, collecting data on ecology, behaviour, and especially vocalisations of the honeycreepers. The result was my 1979 dissertation ‘A systematic analysis of the endemic avifauna of the Hawaiian Islands.’ I financed my field work in the earlier years primarily by working as an illustrator, and later as a birding tour leader. I have continued both these activities to the present day. Though I have never lived permanently in Hawai‘i, I visit almost annually (I have missed a Hawai‘i visit in only two calendar years since 1974). This book represents the

culmination of my quarter century of study, and is my way of sharing this most intriguing group of birds with anyone who wishes to read on. In 1990, Oxford University Press asked me to contribute a honeycreeper volume to its new series ‘Bird families of the world’. I was delighted with the opportunity to produce a definitive summary of what was known about my favourite birds. The incubation period has been about twice as long as we originally anticipated. The decade of the 1990s proved to be one of great renewal of interest in all aspects of drepanidine biology, with an explosion of important research. Every time I tried to summarise it, another important study would begin that simply could not be ignored. Oxford University Press and I agreed to extend the production time for this book so that this review volume could take these ongoing projects into account and be truly comprehensive. Authors and editors were kind enough to supply me with advance copies of manuscripts or personal communications so that I could include items that would be published nearly simultaneously with this book, the first draft of which was submitted in 2001. Perhaps most important was the American Ornithologists’ Union’s life history series The Birds of North America (BNA), in which the Hawaiian native bird accounts were completed in 2003. The watershed volume Evolution, ecology, conservation and management of Hawaiian birds: a vanishing avifauna (Scott et al. 2001), which grew out of the 1997 meeting of the Cooper Ornithological Society in Hilo, included some of the most recent research in a variety of fields. One new species of honeycreeper was described in late 2003 ( James and Olson 2003) so could not be fully included herein, but mentioned, at least, in appropriate places. Helen James’s (2004) long anticipated study of honeycreeper osteology was published a few months ahead of this volume. It is an ill wind that blows no good. Publication delays after 2001 had the unexpected but beneficial result that no “in press” citations remain.

Taxonomic disagreements Ideally, every scientist looking at the same facts should arrive at the same conclusion. But the ideal

Preface xi assumes that everything is known, and, of course, we will never know absolutely everything. Nowhere is this more obvious than in the area of systematics and taxonomy, which deals with reconstructing events that can never be directly observed. At what exact point a new species exists will always be an arbitrary decision. Systematists must do the best they can with the data at hand to construct a phylogeny and a classification that best fits what is known at the time. Good science is always changing and rethinking hypotheses as new information becomes available. That is not to say, however, that scientists should be mercurial; we should not change our ideas with every whim or every revolutionary new technique. Such things must stand the test of time. Unfortunately, because systematics is inherently less precise and testable than, say, nuclear physics, a certain amount of disagreement among people studying the same organisms is inevitable. In the pages of this book you will see how the ideas of the evolution and relationships of the Hawaiian honeycreepers have evolved over the past century. You will see that taxonomy is strongly influenced by ‘conventional wisdom’ and the customs of the day. The fact that a later observer produces a different interpretation does not necessarily lessen the contribution of previous workers. My goal in these pages is to disagree without being disagreeable.You will see that I differ strongly on many points with Dean Amadon, one of the pioneers of modern research on Hawaiian honeycreepers, but it does not mean that I disrespect his scholarship. It is

clearly unfair to judge his efforts in the light of information he could not possibly have known (i.e. plate tectonics, ages of islands and lineages, recency of the break-up of Maui-nui, and recently discovered palaeontological remains). Likewise, though I disagree on several systematic points with Walter Bock, his contributions to the study of honeycreeper anatomy are undeniable. Currently active systematists, including Robert C. Fleischer and the team of Storrs Olson and Helen James, have proposed phylogenies that are irreconcilable with each other and with mine on several points (see Chapter 4). Such disagreements simply reveal that our knowledge is, as yet, too incomplete to allow resolution of every point. I respect the findings of my colleagues and gladly present them in this book, I hope fairly, alongside my own so that readers can make their own comparisons. Interestingly, not very long ago there were far more differences than will be seen in these pages. Olson and James and I have revised our views over the years as new knowledge has come to light, and a taxonomic consensus, especially at the species level, is clearly developing (Pratt and Pratt 2001). Also, Helen James and Rob Fleischer were members of a team that for the first time presented osteological and genetic data in the same paper (Fleischer et al. 2001). Maybe some day we will know enough that everyone will agree on everything. Until then, our sometimes adversarial methods will take us ever closer to that goal. H. Douglas Pratt

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Acknowledgements

I first want to thank those whose work actually supplements my own in this book. Sheila Conant wrote a very useful appendix on the use of Hawaiian honeycreepers in human artefacts, world-renowned photographer Jack Jeffrey provided the wonderful photographs of birds in various behaviours (as well as numerous ‘culls’ that were invaluable as references for the colour plates), and Jaan Kaimanu Lepson contributed additional photos to illuminate these pages. All of them contributed without compensation, for which I am deeply grateful, and I hope the final product justifies their faith. I also appreciate the co-operation of Marie Morin and Alan Poole of The birds of North America series of bird life history studies, and J. Michael Scott of the monumental Evolution, ecology, conservation, and management of Hawaiian birds: a vanishing avifauna, who gave me advance looks at manuscripts of upcoming papers. Others who gave me ‘sneak previews’ include Thane K. Pratt, Robert C. Fleischer, Jeff Foster, Alan Ziegler, David Burney, and Helen James (who also aided in the choosing of vernacular names for recently described subfossil species). I could not even begin to list all of those who have assisted me in my research over the past quarter century, but some have been so important that I could not fail to thank them. Phillip and Andrea Bruner literally opened the door to Hawai‘i for me by making my first visit possible. Phil and I were in turn ‘shown the way’ on that first expedition by David W. Woodside, then of the Division of Forestry and Wildlife. Others who provided valuable hospitality, advice, information, or opportunities for field work in the right places include (in loosely chronological order) Delwyn

and Francine Berrett, Ron Walker, Rob and Annarie Shallenberger, Jim Jacobi, Sheila Conant, Tonnie L. C. Casey, Charles van Riper, III, C. John and Carol P. Ralph, J. Michael Scott, Cameron Kepler, Storrs Olson, Robert L. Pyle, Andrew Engilis, Reginald and Susan David, Rob Fleischer, Rick Warshauer, David Kuhn, Tom Snetsinger, Michelle Reynolds, Jack Jeffrey,Thane K. Pratt, Cynthia Sallie, Keith Unger, Glenn Klingler, Carol Gentz, Rob Pacheco, Jaan K. Lepson, Mark Collins, Helen James, Carter Atkinson, and Jim Groombridge. For the above as well as access to and assistance with archives and collections I wish to thank Edwin H. Bryan, Alan Ziegler, Frank J. Radovsky, Robert L. Pyle, and Carla Kishinami (Bernice Pauahi Bishop Museum); François Vuilleumier, John Farrand,Walter Bock, Dean Amadon, and Mary LeCroy (American Museum of Natural History); Storrs Olson, Richard C. Banks, and Richard L Zusi (Smithsonian Institution); James L. Gulledge and Greg Budney (Library of Natural Sounds, Cornell Laboratory of Ornithology); Alan Lieberman (Keauhou Bird Conservation Center), Raymond L. Paynter (Museum of Comparative Zoology, Harvard University); George H. Lowery, Jr., and James V. Remsen (Museum of Natural Science, Louisiana State University); Ned K. Johnson (Museum of Vertebrate Zoology, University of California, Berkeley); Frank B. Gill (Academy of Natural Sciences, Philadelphia); Phillip L. Bruner (Brigham Young University Hawai‘i Campus); and James Mejeur (Honolulu Zoo). I could not have completed this project without the assistance of several colleagues at the Louisiana

xiv Acknowledgements State University Museum of Natural Science: Fred Sheldon and Josie Babin taught me the use of PAUP and MacClade programs, Shannon Guidry taught me to scan photographs, and all assisted in countless other ways relating to computer usage; James V. Remsen read parts of the manuscript and made valuable comments about both grammar and content. Prather Warren of LSU University Relations prepared reproduction-quality transparencies of the colour plates.Andrew McClung of the University of

Hawai‘i assisted with scans of the illustrations of feather artefacts supplied by Ron Schaeffer of the Bernice Pauahi Bishop Museum. Judith May got this project off the ground at Oxford University Press, Cathy Kennedy shepherded it in its final stages, and Christopher Perrins oversaw strategic publication decisions throughout. Jiro Kikkawa made valuable suggestions as to content, prompting the addition of some discussions that I hope will broaden the value of this work to biologists generally.

Contents List of colour plates List of abbreviations Plan of the book, names and terms Honeycreeper Topography Frequently mispronounced words of Hawaiian origin

xix xx xxiii xxxi xxxii

PART I General chapters 1 The Hawaiian honeycreepers: evolutionary triumph and ecological tragedy 2 The honeycreepers’ world 3 Discovery and research: historical perspectives 4 Origin and evolution 5 Classification 6 Form and function 7 Behaviour 8 Ecology and breeding biology 9 Diseases and parasites 10 Status, conservation, and the future

3 7 33 44 72 83 110 129 157 172

PART II Species accounts 11

Genus Melamprosops Po‘o-uli Genus Paroreomyza O‘ahu ‘Alauahio Maui ‘Alauahio Kakawahie Genus Telespiza Laysan Finch Nihoa Finch Makawehi Finch Maui-Nui Finch Genus Loxioides Palila Genus Rhodacanthis Greater Koa-Finch Lesser Koa-Finch Genus Chloridops Kona Grosbeak Wahi Grosbeak

Melamprosops phaeosoma Paroreomyza maculata P. montana P. flammea Telespiza cantans T. ultima T. persecutrix T. ypsilon Loxioides bailleui Rhodacanthis palmeri R. flaviceps Chloridops kona C. wahi

187 187 190 191 192 196 197 198 201 202 203 203 204 207 207 209 210 210 212

xvi Contents King Kong Grosbeak Genus Psittirostra ‘O‘u Genus Dysmorodrepanis Lana‘i Hookbill Genus Oreomystis ‘Akikiki Hawai‘i Creeper Genus Vangulifer Kiwi Shovelbill Pololei Shovelbill Genus Aidemedia Straight-billed Gaper Curve-billed Gaper Maui-nui Gaper Genus Loxops ‘Akepa ‘Akeke‘e Genus Magumma ‘Anianiau Genus Hemignathus Subgenus Chlorodrepanis Kaua‘i ‘Amakihi O‘ahu ‘Amakihi Hawai‘i ‘Amakihi Subgenus Viridonia Greater ‘Amakihi Subgenus Akialoa Kaua‘i ‘Akialoa O‘ahu ‘Akialoa Maui-nui ‘Akialoa Lesser ‘Akialoa Hoopoe-billed ‘Akialoa Subgenus Hemignathus Kaua‘i Nukupu‘u O‘ahu Nukupu‘u Maui Nukupu‘u ‘Akiapola‘au Genus Pseudonestor Maui Parrotbill Genus Himatione ‘Apapane Laysan Honeycreeper

C. regiskongi Psittirostra psittacea Dysmorodrepanis munroi Oreomystis bairdi O. mana Vangulifer mirandus V. neophasis Aidemedia chascax A. zanclops A. lutetiae Loxops coccineus L. caeruleirostris Magumma parva

212 213 213 217 217 218 219 221 224 224 224 225 225 225 226 226 226 230 231 232 234 235

Hemignathus (Chlorodrepanis) kauaiensis 235 H. (C.) flavus 238 H. (C.) virens 240 243 H. (Viridonia) sagittirostris 243 244 H. (Akialoa) stejnegeri 245 H. (A.) ellisianus 246 H. (A.) lanaiensis 247 H. (A.) obscurus 248 H. (A.) upupirostris 249 250 H. (Hemignathus) hanapepe 250 H. (H.) lucidus 252 H. (H.) affinis 253 H. (H.) munroi 254 257 Pseudonestor xanthophrys 257 260 Himatione sanguinea 260 H. freethii 263

Contents xvii Genus Palmeria ‘Akohekohe Genus Drepanis Hawai‘i Mamo Black Mamo ‘I‘iwi Genus Ciridops ‘Ula-‘ai-hawane Kaua‘i Palmcreeper GENERA INSERTAE SEDIS Genus Orthiospiza Mauka Grosbeak Genus Xestospiza Cone-billed Finch Ridge-billed Finch

Palmeria dolei Drepanis pacifica D. funerea D. coccinea Ciridops anna C. tenax

Orthiospiza howarthi Xestospiza conica X. fastigialis

264 264 267 267 268 269 273 273 275 275 275 275 276 276 277

Appendix 1: Honeycreepers in Hawaiian material culture by Sheila Conant Appendix 2: Scientific names and families of plants mentioned in the text Appendix 3: Scientific names, families, and subfamilies of non-Hawaiian birds mentioned in the text

278 285 289

Bibliography

291

Index

335

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Colour plates Colour plates fall between pages 184 and 185. Plate 1

Plate 2 Plate 3

Plate 4

Plate 5 Plate 6 Plate 7 Plate 8 Plate 9

Drepanidine finches Greater Koa-Finch, Lesser Koa-Finch, Kona Grosbeak, Palila, Laysan Finch, and Nihoa Finch Heavy-billed honeycreepers ‘Ula-‘ai-hawane, ‘O‘u, Maui Parrotbill, Po‘o-uli, and Lana‘i Hookbill ‘Amakihis and ‘Akialoas: Hemignathus in part Kaua‘i ‘Amakihi, O‘ahu ‘Amakihi, Hawai‘i ‘Amakihi, Kaua‘i ‘Akialoa, O‘ahu Akialoa, Maui-nui ‘Akialoa, and Lesser ‘Akialoa Heterobills, ‘Anianiau, and Greater ‘Amakihi: Hemignathus in part ‘Akiapola‘au, Kaua‘i Nukupu‘u, O‘ahu Nukupu‘u, Maui Nukupu‘u, Greater ‘Amakihi, and ‘Anianiau ‘Alauahios and Creepers: Paroreomyza and Oreomystis Maui ‘Alauahio, O‘ahu ‘Alauahio, Kakawahie, ‘Akikiki, and Hawai‘i Creeper ‘Apapanes and ‘Akepas: Himatione and Loxops ‘Apapane, Laysan Honeycreeper, ‘Akeke‘e, and ‘Akepa Sickle-bills and ‘Akohekohe: Drepanis and Palmeria ‘Akohekohe, ‘I‘iwi, Hawai‘i Mamo, and Black Mamo Adaptive radiation of Hawaiian honeycreepers Honeycreeper feather artefacts

Abbreviations AMNH AOU BNA BPBM BRD BSC CBSC CCH CNPRSU d DLNR DOFAW ESA FR h HAS HFBS IBP ICBP ID in prep. IUCN KBCC LNS LSUMNS Ma mtDNA MVZ NAR(S) NBS NHP NP NPS NWHI NWR pers. comm. pers. obs. PIERC PSC SP sp.

American Museum of Natural History American Ornithologists’ Union The birds of North America (AOU) Bernice Pauahi Bishop Museum Biological Resources Division (USGS) biological species concept comprehensive biological species concept Conservation Council for Hawai‘i Cooperative National Park Resources Studies Unit day(s) Department of Land and Natural Resources Division of Forestry and Wildlife Endangered Species Act of 1970 Forest Reserve hour(s) Hawai‘i Audubon Society Hawaii Forest Bird Surveys International Biological Program International Council for Bird Preservation identification in preparation International Union for the Conservation of Nature Keauhou Bird Conservation Center Library of Natural Sounds (Cornell University) Louisiana State University Museum of Natural Science million years ago mitochondrial DNA Museum of Vertebrate Zoology Natural Area Reserve (System) National Biological Survey National Historical Park National Park National Park Service Northwestern Hawaiian Islands National Wildlife Refuge personal communication personal observations of author Pacific Islands Ecosystems Research Center phylogenetic species concept State Park species (singular)

Abbreviations xxi spp. Str. subsp. subspp. TNCH USFS USFWS USGS

species (plural) Stream subspecies (singular) subspecies (plural) The Nature Conservancy of Hawai‘i United States Forest Service United States Fish and Wildlife Service United States Geological Survey

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Plan of the book, names and terms

Organisation The text is divided into two main sections. Part I is an overview of Hawaiian honeycreeper evolution and natural history in 10 chapters. Part II comprises accounts of each species in systematic order with diagnostic descriptions of the genera. The nine colour plates depict all historically known honeycreeper species and representative artefacts made from honeycreeper feathers.The birds are shown in characteristic poses and in native plants upon which they are known to feed or perch. Almost all were painted directly from specimens, but a few that are not represented in American collections were based on specimen photographs or colour lithographs in a copy of Rothschild (1893–1900) in the John S. McIlhenny Collection of the Louisiana State University Library. At the back are three appendixes: Appendix 1, contributed by Sheila Conant, discusses the importance of honeycreepers in Hawaiian culture; Appendix 2 is a tabular listing of the scientific names and families of plants mentioned in the text, taxonomy following Wagner et al. (1990); and Appendix 3 lists scientific names and families of birds that occur outside Hawai‘i that are mentioned in the text, taxonomy following Sibley and Monroe (1990) or AOU (1998). Native and introduced Hawaiian birds other than honeycreepers are listed in two tables in Chapter 2.And finally, a Bibliography lists all the publications known to the author that deal with Hawaiian honeycreepers in any substantive way, plus literature cited that would not otherwise be included. In Part II, the diagnoses of genera known only prehistorically are based on James and Olson

(1991), while those of historically known genera are based on my earlier work (Pratt 1979a) amplified from more recent sources. Species accounts begin with the full English and scientific names, a selective synonymy of scientific names (major references only), and other vernacular names, both English and Hawaiian. Etymology of the current vernacular and scientific names is given where known. Systematics discusses briefly the species’ evolution, relationships, and taxonomic history cross-referenced to discussions in Part I.Any recognised subspecies are discussed here. The Description includes an overall length measurement (cm) taken mostly from Pratt et al. (1987) and intended only to establish relative sizes among species. Coloration is described in general terms meant to supplement the colour plates. Unlike previous volumes in this series, detailed colour descriptions are not given, nor are the usual linear measurements, both of which are readily available in the recently published BNA accounts. Voice is described, beginning with calls and ending with primary songs. Field identification compares the species with others that might cause confusion for observers. Even extinct species may include this heading to help prevent mistaken ‘rediscoveries’ (see Pratt and Pyle 2000). Distribution, habitat, and status references a range map where feasible, but maps are not given for species known from subfossil remains or from only a few historical specimens. Current ranges are shown in solid tone, historic ranges in cross-hatching. For species that are not mapped, range may be discussed in detail, emphasising the localities where the birds were last found. Diet and foraging, Breeding biology, and Life cycle

xxiv Plan of the book, names and terms and demography complete each species account. For many poorly known species, sections may be deleted or combined to save space and avoid constant repetition of ‘no information’. When using the Bibliography, one should understand several conventions.With a few exceptions, tabular references such as field checklists, museum catalogues, world checklists, and other simple lists are excluded. Only the most recent example is given for some lists, such as the US Federal list of Endangered Species (USFWS 1999a), which are updated periodically. Periodicals are abbreviated following modern conventions but substantive names are not abbreviated, i.e. Journal of the Ni‘ihau Ornithological Society would be abbreviated J. Ni‘ihau Ornithol. Soc. Authors’ given names are listed as initials only. References are cited in the text by use of the ‘name/date’ system. Page numbers within references are given only for direct quotations. Usually only the author’s last name is given, but in cases that would be confusing, initials are given as well. For example, the Bakers, Paul and Helen, are co-authors of two papers published in 2000; on one he is the senior author, while she is the senior on the other.To distinguish them, they had to be cited as ‘P. and H. Baker’ and ‘H. and P. Baker’, respectively. Both my colleague Thane K. Pratt (no relation) and I are senior authors of numerous publications listed. I use his initials with his citations in cases where doing otherwise would cause undue confusion, and use just ‘Pratt’ for my own.

group of birds, so placing them in a family of their own was the only reasonable course. Now that most authors accept that the Hawaiian honeycreepers are a sister group of the cardueline finches (see Chapter 4), maintaining them in a family of their own is less tenable. Raikow (1985), however, presents a cogent analysis of the relationships as currently understood, and recommends maintenance of the family Drepanididae. Koblik (1994), virtually alone among recent authors, agrees with Raikow. Whether the Hawaiian honeycreepers are a separate family or a subfamily or tribe within the Fringillidae, they are widely recognised as a distinctive, monophyletic, geographically isolated group that has undergone a spectacular burst of adaptive radiation. If included in the Fringillidae, they clearly stand apart in their degree of variation from the other members of the family, all of which are clearly ‘finches’ in the general sense (see discussion of names below).Their notoriety is such that packaging them as merely a subgroup of a well-known and widespread family of birds does a disservice to both groups of birds as well as to the naturalists, ornithologists, evolutionists, birders, and conservationists who might benefit from studying them. They clearly are a ‘family’ in a colloquial sense, and their ranking in the taxonomic hierarchy is currently moot. Consequently, this volume in the series ‘Bird families of the world’ is devoted to them unapologetically and without prejudice as to whether they are a ‘real’ family or not.

Honeycreepers or finches? Terminology The family concept Previous volumes in this series have dealt with bird families whose categorical status has not been equivocal. The Hawaiian honeycreepers are an exception: although long regarded as a full-fledged family, Drepanididae, classifications since the 1970s have increasingly moved the group further down the taxonomic hierarchy as a result of our increasing knowledge of origins and relationships. Earlier authors could only speculate about the nearest relatives of this diverse but obviously interrelated

In this volume, ‘Hawaiian honeycreeper’ and ‘honeycreeper’ are used as the general term for members of the Drepanidinae (or Drepanididae or Drepanidini or Drepaninini), and ‘Hawaiian finch’ refers specifically to those honeycreepers that retain finchlike characteristics. Recently, a few authors have rejected the name ‘Hawaiian honeycreeper’ as a group name, preferring to refer to these birds simply as ‘finches’ or ‘Hawaiian finches’, because not all members of the group feed on honey (⫽nectar), many of them do not creep, and they are related to a group of finches. In my opinion, calling these birds finches serves only to obfuscate. Certainly no one

Plan of the book, names and terms xxv using the term ‘honeycreeper’ in this context thinks of it literally, any more than they do for such other bird group names as nuthatch, roller, courser, thrasher, oystercatcher, spiderhunter, nutcracker, warbler, cardinal, bishop, sandpiper, or shearwater. These are simply useful epithets with a long tradition of English usage whose literal meaning is now secondary. Furthermore, few such designations apply to every member of a group. Many warblers do not warble, some flycatchers do not catch flies, relatively few babblers babble, and not all mockingbirds mock. Denying the longstanding name ‘honeycreeper’ on these grounds is pure pedantry. When Hawaiian honeycreepers were first discovered by European scientists, the family designation of many of them was far from obvious, so they were distributed among a variety of avian families. Eventually, the interrelationship of all the thinbilled birds was recognised, but those with finchlike bills continued to be regarded as true finches until the end of the nineteenth century (see Chapter 3). By the time their relationship to the thin-billed (and mostly nectarivorous) birds was established, the name ‘honeycreeper’ was well entrenched. Furthermore, the finchlike honeycreepers were regarded as the result of evolutionary convergence with ‘true’ finches until the late 1970s, when the cardueline finch connection first became widely accepted. Pratt (1979a) used the term ‘Hawaiian finches’ specifically for the finchlike honeycreepers to separate them as a group from the rest of the honeycreepers, and most writers since, e.g. Schluter (1988), have used the term with that restricted meaning. Olson and James (1982b), however, used the term ‘Hawaiian finches’ for the entire subfamily (or tribe in their case) without comment and continue to do so in subsequent writings, as do some others (Kepler 1985). Their usage is not without merit because it calls attention to relationships that differ strongly from the long-held belief that these birds descended from a thin-billed ancestor, and also calls attention to the fact that a disproportionate number of the species known from prehistoric remains are finchlike. However, I agree with Amadon (1986) that tradition and accurate communication far outweigh any perceived instructive value in referring to the

highly varied Hawaiian honeycreepers simply as ‘finches’. A few writers have attempted to compromise by using terms such as ‘honeycreeper-finches’ or ‘finch-honeycreepers’, but such constructs are cumbersome, unnecessary, and, in my opinion, not particularly helpful. The word ‘finch’ has a rather specific general meaning in English that has nothing to do with classification. The new shorter Oxford English dictionary (Brown 1993) defines it as ‘Any of numerous small passerine birds, typically having short, stout beaks adapted for seed eating, belonging to the family Fringillidae and certain related families.’ This definition would certainly be misapplied to an ‘amakihi or ‘I‘iwi. The dictionary’s ‘related families’ include Emberizidae, Cardinalidae, Passeridae, Ploceidae, and Estrildidae, all of which include birds that could be called ‘finches’ (see Zusi 1978 for that specific usage). The term ‘Hawaiian finches’ could even be misinterpreted to refer to members of these families introduced to the islands. A Hawaiian honeycreeper by this other name would sing as sweetly, but its distinctiveness would be lost in nomenclatural ambiguity. Recently, the nickname ‘dreps’ has come into use (Pratt 2002b), and it appears occasionally in these pages. Readers who consult the historical literature will find the basic name of the honeycreeper taxon variously written as Drepaniidae, Drepanidae, and Drepanididae. Mayr (1943) and several major references (Amadon 1950; Baldwin 1953) used Drepaniidae, but Drepanididae is now considered the correct form. In references and quotations from historical texts, I have avoided the use of ‘[sic]’ after earlier name variants because these forms were not incorrect when they were used.

Names of honeycreeper subgroups Virtually every author since the beginning of the twentieth century has found it convenient to subdivide the Hawaiian honeycreepers into smaller species groups for discussion purposes. Some have recognised two groups, some three. Whether these should be recognised taxonomically has been controversial, and is addressed philosophically in Chapter 5. However, I,

xxvi Plan of the book, names and terms

Some genera, e.g. Melamprosops, Paroreomyza, Oreomystis, and Pseudonestor, do not fit comfortably into any of these broad categories, but that does not destroy the usefulness of the groupings for making generalisations.

as English loan-words. As such, they can be (and often are) written without the orthographic conventions (see below) of the Hawaiian language. I have chosen to write bird, plant, and place names of Hawaiian origin with the proper orthography to aid the reader with pronunciation. The ultimate authority on the Hawaiian language, and the source of much of the etymology presented herein, is the Pukui and Elbert (1971) Hawaiian dictionary. We will never know what the people who drove them to extinction called the species of Hawaiian honeycreepers known only from prehistoric remains. Most of their names were probably long forgotten by the time of Cook’s arrival. Nor did James and Olson (1991), the describers, create English counterparts for their Latin names, but Ziegler (2002), a close colleague of Olson and James, proposed several. English names matter in this case, mainly to emphasise the fact that these newly described species are not ancient fossil forms like dinosaurs, but were part of the same modern avifauna as the historically known species. The names used in this book were chosen in collaboration with Helen James (pers. comm.) because I thought it presumptuous to name these birds on my own.

English names of Hawaiian honeycreepers

Peculiarities of the Hawaiian language

Part of the charm and mystique of Hawaiian honeycreepers, for English speakers at least, stems from their exotic-sounding names, nearly all derived from Hawaiian. From the earliest days of European discovery, English speakers adopted the Hawaiian names for native birds in the islands, perhaps because none of the familiar European bird names seemed appropriate. As pointed out by Olson and James (1995), these vernacular names are the only ones that have remained stable through two centuries of taxonomic twists and turns, and one cannot understand the classic honeycreeper literature without knowing them. Most species have no other name in English, so the names can be correctly regarded, as many already have been in a variety of dictionaries, including the Oxford English dictionary,

Hawaiian has a limited alphabet, with only 12 consonants and 5 vowels.Vowels are pronounced as in Romance languages such as Spanish or Italian. Diphthongs are pronounced with more separation of the merged sounds than in English, and are always stressed on the first member of the pair. Most (ei, eu, ou, ai, ae, ao) are pronounced as one would expect, but au always sounds like the ow of how rather than the au of fault. The consonants are pronounced straightforwardly as in English, with the exception of w, which can be pronounced as either w or v. Initially or after u or o, it is w; after e or i, it is a soft v; after a it is correct either way, but in a given word usually is pronounced one way or the other by custom (‘I‘iwi ⫽ eeEE-vee; Kakawahie ⫽ KAH-KAH-wah-HEE-eh;

too, find the need to be able to refer to certain groupings. In my 1979 dissertation, I recognised three subfamilies that I now regard as representing evolutionary grades rather than clades. These correspond more or less to the two subdivisions of Perkins (1903) and Amadon (1950) except that their ‘green birds’ group is split into finchlike and thin-billed divisions. These groupings are still convenient: 1. The ‘red and black’ birds of Perkins and Amadon, which I also refer to as the nectarivores (they could also be called drepanidines, but for obvious reasons to do so would lead to confusion). 2. The finchlike honeycreepers, also called drepanidine finches, Hawaiian finches, and psittirostrines (after the genus Psittirostra). 3. The thin-billed ‘green birds’, which can be called the hemignathine group (after the genus Hemignathus).

Plan of the book, names and terms xxvii Kaho‘olawe ⫽ KAH-ho-oh-LAH-veh; Hawai‘i ⫽ Ha-WYE-ee or Ha-VYE-ee). These examples also include the ‘okina, written as an opening quote mark (‘), which functions as an additional consonant. It represents the stoppage of sound between syllables of the English interjection ‘uh-oh’ or some local British pronunciations of the tt in ‘little’ (‘li‘l’). This ‘glottal stop’ had no equivalent letter in the Roman alphabet, and is often misinterpreted as a diacritical mark or a mark of punctuation.Writing Hawaiian names without the glottal stop makes them difficult to read and pronounce, just as if a letter has been deleted (Pukui and Elbert 1971). An initial ‘okina, as in ‘amakihi, is heard only in a word sequence such as ‘Kaua‘i ‘Amakihi’ (all Hawaiian words end in vowels, so the initial glottal stop breaks the vowel glide between words). The ‘okina is essential to avoid ambiguity and to designate diphthongs. For example, the Hawaiian name for Bulwer’s Petrel is ou, but one of the honeycreepers is called ‘O‘u . Only the ‘okina distinguishes a diphthong from the two-syllable pronunciation. In general, Hawaiian words are accented or stressed on the next-to-last and alternating preceding syllables, with the final stress the strongest (‘amakihi ⫽ AH-mah-KEE-hee). Words of five syllables are stressed on the first and fourth (‘akialoa ⫽ AH-kee-ah-LO-ah;‘alauahio ⫽ AH-lauah-HEE-oh). However, many words have irregular stresses indicated by a horizontal line, called a macron, over the vowel. But this stress is not quite the same as an irregular accent in, say, Spanish, because it is superimposed over the stress that would be present normally. English speakers can approximate the effect by thinking of a syllable with a macron as if it were a separate word (‘akepa ⫽ AH, KEH-pah, not AH-kep-ah). Many words with macrons are actually phrases written as one word. For example Haleakala was originally a phrase (hale a ka la,‘house of the sun’) and is pronounced HAHlay-a-kah-LAH. Several names of Hawaiian honeycreepers have similar etymologies. Stressed syllables (and stressed monosyllabic words such as la, above) originated as repeated vowels with a glottal stop between (Ziegler 2002) that in casual pronunciation became slurred together (as the syllable po in ‘Akiapola‘au, which originated as po‘o (⫽head). Knowing that helps to pronounce words with macrons

correctly. The next section, ‘Frequently mispronounced words of Hawaiian origin’, lists pronunciations of honeycreeper names of Hawaiian origin and a selection of other frequently mispronounced Hawaiian place names and terms.

Language conventions The interplay of English and Hawaiian sometimes produces some linguistic tangles. One involves the fact that Hawaiian has no plural form for nouns, plurality being indicated by context. Depending on personal preference more than anything else, various modern scientific writers may or may not pluralise Hawaiian-derived names. In English, hunters have traditionally dropped plurals, as in ‘We shot three quail, two turkey, and some grouse’, and modern birders, especially in Hawai‘i, have adopted this lingo both in speaking and writing. For this book, I have taken the position that bird names of Hawaiian origin are now English loan-words and can be pluralised. However, I restrict the use of plurals to enumerating kinds rather than individuals. For example, there are several ‘amakihis (Kaua‘i ‘Amakihi, O‘ahu ‘Amakihi, Common ‘Amakihi, and Greater ‘Amakihi), but one might correctly report ‘I saw only three ‘amakihi on the ‘Aiea Trail’. The previous example also demonstrates the widespread practice in ornithological and birding literature of capitalising the specific English names of birds. I follow that practice herein because I believe it reduces confusion. Hawaiian bird names, including those of honeycreepers, are standardised by the American Ornithologists’ Union (AOU 1998), albeit without ‘okina and macrons, so they take on the mantle of proper names that can be capitalised. For those outside the AOU area, I use the names of Sibley and Monroe (1990). A final convention involves tense. Throughout this book I often use the present tense to describe actions of extinct birds, especially when comparing them to living species or when they are part of a list that includes both extinct and living species. It simply reads better and avoids circumlocution. I use past tense when the subject is something that obviously no longer occurs or when present tense would be very misleading. Bear in mind that several

xxviii Plan of the book, names and terms honeycreepers are currently in that limbo of not having been observed for a considerable time but not ‘officially’ declared extinct. Using present tense in those cases at least gives them the benefit of the doubt.

Terminology of moults and plumages I follow the now widely used terminology of Humphrey and Parkes (1959), with the names of moults and plumages capitalised. Nestlings begin with a coat of down, which is replaced before fledging by the Juvenal plumage. Most of the Juvenal plumage is worn for a relatively short time, and is replaced in the First Prebasic moult by the First Basic plumage. The flight feathers from the Juvenal plumage, however, are retained for a full year.These feathers are usually distinguishable from the rectrices and remiges of adults, allowing first-year birds to be distinguished. In birds that have two distinctive plumages each year, Basic plumage is worn during the ‘off-season’, and the Alternate plumage during the breeding season. Hawaiian honeycreepers do not have distinctive breeding plumages, so all plumages after Juvenal are Basic.The moult that precedes this plumage is called Prebasic moult. If the Basic plumage differs progressively as a bird ages, as in several Hawaiian honeycreepers with delayed plumage maturation, it is referred to as First Basic, Second Basic, etc.When a bird reaches a stage in which one plumage looks the same as the next, it is said to be in Definitive plumage. The term immature is used for birds whose exact age is not known but which have not reached the Definitive stage. The term subadult may be used for an immature that is approaching Definitive.

Systematic philosophy and jargon Systematics, the science of evolutionary history, relationships, classification, and naming of organisms has a vocabulary all its own. Some familiar words have specialised meanings in this context, and many terms are used solely by systematists. Laymen often confuse systematics (Chapter 4) with

taxonomy (Chapter 5).The latter is included within systematics and involves only the naming of taxa (defined above) and grouping them with related taxa in a hierarchy.Taxonomy is as old as mankind, but systematics is a post-Darwinian discipline. This book is liberally peppered with systematic terms that may be unfamiliar to the lay reader, so a brief explanation here of evolutionary philosophies and their terminology may be of some help.

Evolutionary systematics When Charles Darwin first proposed the principle of evolution, many of the details, especially those dealing with genetics and the process of speciation, remained to be worked out.The earliest hypothesis of Hawaiian honeycreeper evolution (Perkins 1903) was developed well before these phenomena were thoroughly understood. Eventually, a consensus called the ‘modern synthesis’, as elucidated primarily in the works of Ernst Mayr (1943, 1963, 1969), developed. Mayrian evolutionary systematics dominated studies of honeycreepers from the 1940s to the 1970s.

Phylogenetic systematics The last quarter of the twentieth century witnessed a sea change in systematic philosophy, from evolutionary methods to phylogenetic systematics, which grew mainly from the work of the German evolutionist Willi Hennig (1966). Phylogenetic systematics did not gain wide acceptance among ornithologists until relatively recently, and the different approaches of these philosophies are still being sorted out.The two schools treat some of the same terms differently, and phylogenetic systematics has its own esoteric jargon that may not be familiar to the lay reader. The techniques of phylogenetic systematics are called cladistics. An excellent short summary of cladistic methods in the Hawaiian context is given by Funk (1995), but for a more detailed discussion see Forey et al. (1992) and references cited therein. In cladistics, groupings are defined by one or more shared derived character states or synapomorphies. In

Plan of the book, names and terms xxix other words, all the groupings that share a uniquely derived condition are presumed to have descended from a common ancestor that had that condition. Such a monophyletic group of taxa is called a clade. A clade may or may not correspond to a taxon, but all taxa are clades in phylogenetic classifications. In contrast to evolutionary systematics, clades are defined solely on the basis of synapomorphies, whereas in evolutionary systematics groupings are allowed based on characters that are primitive, i.e. holdovers from an ancestral evolutionary stage. In phylogenetic systematics, such paraphyletic groups may be called evolutionary grades, but they are not recognised taxonomically. Shared primitive characters are called symplesiomorphies. A character that is derived (an apomorphy) at one level becomes primitive (a plesiomorphy) at a higher level. Uniquely derived character states that are found in only one taxon are called autapomorphies. Groups that are each other’s closest relatives are called sister groups, and if they are named, sister taxa.A series of changes in the same character, each derived from the other, is called a transformational series. Such series are the phylogenetic equivalent of Bock’s (1970) ‘microevolutionary sequences’ among Hawaiian honeycreepers. For a transformational series to be used in reconstructing evolution, one must determine the direction or polarity of the changes.The most common method for determining polarity is called outgroup comparison. If a character occurs in two or more forms or states within a group, the state found in the sister group and/or more distantly related taxa is assumed to be the plesiomorphous or primitive condition. An important element of cladistics is the concept of parsimony. The rule of parsimony, or ‘Occam’s razor’ as it is sometimes called (referring to William of Occam, a medieval Englishman who first stated the principle), basically means that one never assumes a complex or convoluted explanation if a simpler one is available. Convergence, now often called homoplasy when it occurs among members of a closely related group, is relatively rare in nature because it usually requires a hypothesis of more complex evolutionary patterns than the alternative that similar things are derived from a common ancestor. Nevertheless, it does happen; Occam’s

razor is a guideline, not a law. Funk (1995: 33) states ‘This principle does not preclude the possibility of convergent or parallel evolution; it simply states that when there is no reason to think otherwise, two characters that appear to be the same are treated as homologous.’ In other words, convergence is never assumed. Nowadays, cladograms are constructed using computer programs, especially one known as PAUP*, an acronym for Phylogenetic Analysis Using Parsimony (Swofford 1999).

Phylogenetic trees A hypothetical reconstruction of evolutionary history is called a phylogeny.Traditionally, a phylogeny is illustrated in the form of a tree or dendrogram, something like a family tree in human genealogy, using speciation events (often hypothetical) as branching points or simply showing a pattern that does not define the branching points at all. In evolutionary dendrograms, taxa are grouped on the basis of overall similarities, and a group is considered monophyletic as long as all of its members share a common ancestor, even if it does not include all the descendants of that ancestor. If a dendrogram is based on numerical data, it is called a phenogram (Sneath and Sokal 1973). A dendrogram that is constructed using cladistics is called a cladogram. Its branching points are based on transformational series of synapomorphies. In phylogenetic systematics, to be monophyletic a group must include all the descendants of a common ancestor. Groups that do not include all descendants are termed paraphyletic. In this book, the term monophyletic is used in the latter sense. In both systems, any taxon that can be shown to be polyphyletic, i.e. comprising descendants of two or more different ancestors, is assumed to be erroneously constituted. Phylogenetic trees are never intended to represent observed fact.They are hypotheses about how evolution may have proceeded. In the earliest systematic studies of Hawaiian honeycreepers, including Perkins (1903), Bryan and Greenway (1944), Amadon (1950), Baldwin (1953), and Richards and Bock (1973), relationships were usually hypothesised based on similarities and differences, with the phylogeny developed later. This methodology represents the evolutionary school of systematics, but

xxx Plan of the book, names and terms in the context of phylogenetic reconstruction is better referred to as the eclectic school (Raikow 1985). In it, a hypothetical phylogeny attempts to express both character transformations and genealogy. In phylogenetic studies, the phylogeny, which is purely genealogical, comes first and is then used to determine relatedness. My own early work (Pratt 1979a) was a product of its times in that I used cladistics to develop a phylogeny, but allowed eclectic thinking to influence my classification. More recently, classifications have been based primarily on phylogenetic principles, but the most widely followed one (AOU 1998) still retains elements of earlier eclectic thinking.The classification proposed in this book (see Chapter 5) is based solely on phylogenetic patterns (Chapter 4). The majority of phylogenetic trees that have been published since 1979 have been based on biochemical or genetic studies.The fact that phylogenies developed by these techniques seem to change in fundamental ways from one study to the next, even in the same laboratory, indicates that results

should still be regarded as tentative. Despite the enthusiasm of the practitioners of these revolutionary techniques, who tend to dismiss contrary evidence from phenotypic characters without serious comment, I believe, like Raikow (1986), that concordance testing with traditional methods is the only reasonable way to evaluate these hypotheses. The earliest phylogenies, such as that of Perkins (1903), were actually drawn to resemble trees, but more recently a horizontal arrangement, with the tree lying on its side, has become the standard. To facilitate comparisons of various trees in this book, I have redrawn those published elsewhere in a common style using the program MacClade (Maddison and Maddison 2000). In no case is the tree topology, i.e. the branching sequence, changed, but other information, such as branch lengths used in some trees, may be lost through this simplification. For the same reasons, I have converted the taxonomy of the originals to modern equivalents and have used vernacular names because they are the only ones that remain comparable across various studies.

Honeycreeper Topography nape

forehead

upper back lower back tertials

malar area wing bars primary coverts secondary coverts scapulars median coverts lesser coverts auricular (cheek)

rump uppertail coverts

eye-ring rectrices (tail feathers)

lore

primaries (flight feathers)

tomium

vent secondaries flank

chin throat breast

thigh

belly hallux

toe claw

postocular stripe crown

side of neck

superciliary (eyebrow)

back

forehead lore maxilla (upper mandible)

secondaries primaries

culmen tail

tomium gonys (lower) mandible

undertail coverts breast

flank belly

tarsus

Frequently mispronounced words of Hawaiian origin English words in phonetic pronunciations (hoe, low, cow, allow, vow, now, and how) pronounced as in English. Strongly stressed syllables in FULL CAPS, secondarily stressed syllables in SMALL CAPS. ‘Akeke‘e ‘Akepa ‘Akialoa ‘Akiapola‘au ‘Akohekohe ‘Alaka‘i ‘Alala ‘alauahio ‘amakihi ‘Anianiau haha‘aiakamanu Haleakala Hanawi haole Hawai‘i Honolulu Hualalai ‘ie‘ie ‘I‘iwi Kamehameha Ka‘u Kaua‘i kanawao kawau Kilauea Koke‘e kokio ke‘oke‘o loulu maile naio naupaka Nene Nukupu‘u ‘ohi‘a ‘ohi‘a-ha

AH-keh-KEH-eh AH, KEP-ah AH-kee-ah-LOE-ah AH-kee-ah-PO-LA OW AH, KO-hay-KO-hay AH-la-KAH-ee rhymes with tra-la-la Allow-ah-HEE-oh ah-ma-KEE-hee AH-nee-ah-NEE-OW HA-HA, EYE-a-ka-MA-noo HA-lay-ah-kah-LAH, not Holly Ockle-a ha-nah-VEE HOW-lay ha-WYE ee or ha-VYE ee HOE-no-LOO-loo, not HA-nah-LOO-lah hoo-ah-LAH-LYE, not walla-lye, not hoo-ah-la-LYE EE-eh EE-eh (4 syllables) ee EE-vee (3 syllables), does not rhyme with kiwi kah-MAY-ha-MAY-ha kah OO, does not rhyme with cow cow-AH ee, (3 syllables), not cow eye kah-nah-WOW kah-VOW KEE-laoo-WAY-ah not killa-WAY-a ko-KEH-eh (3 syllables) ko-KEE-oh KAY oKAY oh LOW-loo MY-lay NYE-o now-PAH-kah nay-nay, not knee-knee NOO-koo-POO-oo OH-HEE ah, not oh-HEEah OH-HEE ah HA

Frequently mispronounced words of Hawaiian origin xxxiii ‘o‘ o ‘O‘u Po‘o-uli pukiawe ‘Ula-‘ai-hawane wiliwili

OH OH OH OO POE-oh OO-lee POO-kee-AH-vay OO-lah EYE-HA-WA-nay willy-willy, not villy-villy

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PART I General chapters

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1 The Hawaiian honeycreepers: evolutionary triumph and ecological tragedy The most important defining fact for Hawaiian honeycreepers is that they are endemic to the Hawaiian Islands in the tropical N. Pacific Ocean. This, the most isolated of the world’s archipelagoes, with relatively large and ecologically diverse islands (Chapter 2), provided a superb natural laboratory. Although Darwin’s famous finches of the Galápagos are better known and arguably better studied, even students of that group agree that the Hawaiian honeycreepers are the most spectacular and most extreme avian example of adaptive radiation on islands (illustrated diagrammatically in Plate 8). No other such closely related group of birds exhibits anything resembling the phenotypic diversity of the honeycreepers, whose variations in bill, plumage, foraging behaviour, and food choice cover the spectrum of passerine (and even a few non-passerine) niches. Within this radiation are quintessential examples of nearly the whole range of microevolutionary phenomena, including allopatric speciation, double invasion, parallelism, convergence, character displacement, co-evolution, and competitive exclusion. Also, the honeycreepers’ interactions with other Hawaiian plants and animals provide a window on distinctive island phenomena. Within the archipelago, the ‘dreps’ provide examples of how birds colonise islands, form new communities, and undergo evolutionary cycles.They are a brilliant and unequalled natural experiment in evolution and biogeography. The honeycreepers’ role as supreme paradigm is not limited to adaptive radiation. Their long isolation from humans predisposed them to become the ‘poster child’ for anthropogenic environmental degradation. Although the history of these birds’

decline and extinction (Chapters 11–12) is not a pretty story, it is an enlightening one.Virtually all of the varied human causes of reduced biodiversity have examples among Hawaiian honeycreepers; virtually all of the techniques used worldwide in species recovery efforts have been tried with them; and nearly all of the problems facing species recovery efforts today, from inbreeding depression to politics, have afflicted recent efforts to salvage what remains. For some subjects, such as the effects of avian disease, the honeycreepers are the world’s best-studied example. The literature of conservation biology is replete with drepanidine models and examples.The following is an overview of the subjects to be discussed in the remainder of Part I. Chapter 2 surveys the Hawaiian environment, including the geological and climatological history of the islands, their habitats and other organisms, and then outlines the changes wrought first by the Polynesians and later by other human cultures. It includes sufficient detail to make the subsequent chapters understandable; more expansive overviews of Hawaiian natural history are given by Carlquist (1970), Culliney (1988), Daws (1988), Beletsky (2000), and Ziegler (2002). Chapter 3 reviews the history of research on Hawaiian honeycreepers, because a knowledge of that subject is essential for understanding the biological concepts in the chapters that follow. Chapter 4, the longest, focuses on the details of evolution of the group, and Chapter 5 presents a taxonomy based on that evolutionary history. Chapters 6–10 review the results of adaptive radiation as they are expressed in functional anatomy, plumage, vocalisations and other behaviour, ecology, and breeding biology.

4 The Hawaiian Honeycreepers The ancestor of the honeycreepers (Chapter 4) was likely a cardueline finch such as a redpoll or siskin. Those finches have bills that are longer and thinner than many others, which perhaps provided a good midpoint for evolution in a variety of directions. Probably the first burst of radiation produced a group of species that would have still been regarded as cardueline finches if no further evolution had occurred. Some, such as those in the genus Telespiza, had thicker bills and more eclectic eating habits than their seed-eating ancestor, but differed relatively little from their ancestors. Others specialised for feeding on particular kinds of seeds of endemic plants: the koa-finches (Rhodacanthis) on the native koa; the Palila (Loxioides) on the dry-forest mamane, another legume; and the native grosbeaks (Chloridops) on the hard seeds of naio, which grows among the mamane (all these plants will be discussed in more detail in Chapter 2). Possibly one of the first deviations from the finch pattern was the development of a thin, warbler-like bill in the alauahios (Paroreomyza). This first experiment away from the finch model apparently did not give rise to any further bill types, but the world was wide open for the others. One branch of the family took a page from the tanagers’ (Thraupidae) playbook, lengthened the finch bill, and gave it a hooked tip for feeding on various soft fruits; the ‘O‘u (Psittirostra) was the result. Related to it is the bizarre hookbill (Dysmorodrepanis), whose feeding habits are unknown and whose bill is so odd that it was long regarded as a freak. The parrotbill (Pseudonestor) exploited wood-boring larvae by using its heavy bill to crush and gouge its way into their hiding places, whereas the Po‘o-uli (Melamprosops) developed a taste (and the anatomical modifications necessary to feed on them) for the then-abundant land snails of the Hawaiian forest.The creepers (Oreomystis) took the niche of continental nuthatches, picking their prey from the bark of trees. But the key to the burst of evolution that produced the varied array we see today was apparently the development of a tongue for feeding on nectar. With that tongue, the true honeycreepers came into being. The first possessor of this unique, tubular, brush-tipped tongue probably had a finchlike bill. The ‘palmcreepers’ (Ciridops), known historically only from one species, probably are the closest

modern equivalent to this ancestor. Aside from the species already discussed, all of the Drepanidinae have this kind of tongue. Because no other passerines in the world have anything exactly like it, we assume that all the birds that possess it share a common ancestor. The nectar-feeding tongue opened up a whole new natural supermarket of food choices. Some descendant species with short slightly curved bills, such as the ‘apapanes (Himatione) and the ‘Akohekohe (Palmeria), specialized on shallow flowers such as those of ‘òhi‘a-lehua.The mamos (Drepanis) evolved along with native lobeliad flowers to such an extent that their sickle-shaped bills exhibit a fingerin-glove fit with the flowers, and the related ‘I‘iwi, probably the quintessential Hawaiian honeycreeper, uses a similar bill in more varied ways. Meanwhile, other birds with the drepanidine tongue were finding novel ways to use it. The ‘Anianiau (Magumma) continued to feed in flowers, but also developed a warbler-like propensity for gleaning insects from leaves. Its bill is short and a bit curved. Before they had lost their finchlike bill, the ‘akepas (Loxops) found that the brush tip worked well for ensnaring tiny insects that lived in the interstices of leaf buds and other crevices. In a move reminiscent of continental crossbills, they developed crossed tips to their bills to facilitate pushing in two directions when the bill was opened. Similar gaping movements of a long nearly straight bill were used by the Greater ‘Amakihi (subgenus Viridonia) to extract prey from leaf bases of the ‘ie‘ie vine. Its close relatives, all of whom look much alike aside from their bills, have curved bills but have modified them in totally unprecedented ways. The ‘akialoas (subgenus Akialoa), with their extremely long sickle-bills and bark-picking habits may be the Hawaiian analogue of Neotropical woodcreepers (Dendrocolaptidae). The familiar ‘amakihis (subgenus Chlorodrepanis) have much shorter sickle-bills and generalist feeding habits that have made them probably the most successful honeycreepers in the face of ecological changes. But the prize for the most unexpected bill morphology must go to the ‘heterobills’, the nukupu‘us and the ‘Akiapola‘au (Hemignathus sensu stricto). Their bills do not even match! The nukupu‘us pick and pry at tree bark with their shorter curved mandible, using the much

Evolutionary triumph and ecological tragedy 5 longer maxilla to winkle out the larvae from the cracks.The ‘ ‘aki’ has taken matters a step further and straightened out the mandible so that it cannot even close with the maxilla without leaving a small gap. As with the aforementioned hookbill, one would have thought it a freak if it were not still performing its distinctive feeding movements in the forests of Hawai‘i. It is, in fact, an ersatz woodpecker (Picidae)! It hammers with its lower bill only, but it gets the job done. Had any small woodpecker managed to reach Hawai‘i (woodpeckers worldwide are poor island colonizers), the ‘aki never could have happened. With the woodpecker niche open, nature filled it with a modification of a honeycreeper that would have made Rube Goldberg proud. Although not the prettiest, the ‘Akiapola‘au is many birders’ favourite Hawaiian honeycreeper precisely because it is so bizarre. Nothing even close to it exists anywhere else (the extinct Huia of New Zealand comes close, but had to involve both sexes to accomplish the same thing). The Drepanidinae have not evolved varieties of coloration nearly to the extent they have bill shapes. None are iridescent, opalescent, or blue.The predominant colours are red and yellow, variously shaded by melanin. With a few exceptions, the members of nine of the aforementioned genera or subgenera could all be called ‘little green birds’ (Pratt et al. 1987). Nevertheless a few species, such as the ‘I‘iwi and ‘Akohekohe, can stand among the world’s most beautiful (Alcorn and Schenone 1991). Chapter 6 discusses all the variations in morphology and coloration in detail. Like their cousin the Island Canary, the Hawaiian honeycreepers are excellent vocalists, but they have not evolved a vocal array nearly as varied as their morphology. Indeed, vocalisations can be considered a fairly ‘conservative’ feature in analysing evolutionary relationships within the group (Chapter 4). The finchlike ones still have canary-like melodic songs, but the ‘little green birds’ sing mostly simple trills or warbles. The nectar specialists have been the most vocally experimental.The song of the ‘I‘iwi, although not melodic, has an eerie, haunting quality reminiscent of songs of the New Zealand Bellbird and Tui or some Neotropical icterids. The ‘Akohekohe sings a unique odd gurgling song, and the ‘Apapane is

among the world’s most versatile vocalists, with hundreds of variations of its lively song. More than any other, it is today the sound of the Hawaiian forest. Chapter 7 discusses honeycreeper songs in detail, along with other aspects of their behaviour. The above review includes only part of the story. Starting in the 1970s, scientists began finding bird bones in all sorts of places in Hawai‘i, from ancient sand dunes to lava tubes. These were not ancient mineralised fossils, but the remains of actual birds that lived only a few hundred years ago. To everyone’s surprise, the bone caches produced remains of many previously unknown native species, including an impressive array of ‘new’ honeycreepers (Olson and James 1982b; James and Olson 1991). In fact, at a minimum these bones double the number of honeycreeper species, and who can say how many additional extinct species remain to be discovered from such deposits? Because these recently named birds were part of the modern avifauna (they disappeared only after Polynesian settlers severely altered habitats in the islands), they are included in the discussion in Part I to the extent possible, and are included in the species accounts of Part II along with historically known species. Of course, we can never know their colours, songs, or tongue structure but a surprising amount of information can be gleaned from the bones and where they were found, including in some cases what the birds ate, what ate them, and what habitats they preferred. The deposits have also produced many bones of historically known species from places where they no longer are found. Most of the new discoveries fall into the category of finches resembling those known historically, including new members of the genera Telespiza, Chloridops, and Ciridops, and a new genus, Orthiospiza. The extinct genus Xestospiza resembles, and may be related to, Melamprosops.Thin-billed birds are represented by an additional ‘akialoa and two genera that further broaden the adaptive array of honeycreeper bills: Aidemedia comprises several species of birds with medium-length, nearly straight bills that superficially look like that of the Greater ‘Amakihi; and Vangulifer with its blunt-tipped bills represents another morphology unique to Hawaiian honeycreepers. These discoveries have been brought to light mostly through the efforts of the husband-and-wife team of

6 The Hawaiian Honeycreepers Helen James and Storrs Olson at the Smithsonian Institution, aided by the late Allan C. Ziegler, formerly of the Bernice Pauahi Bishop Museum. New subfossil sites continue to be discovered, so the story of prehistoric dreps is an ongoing drama. Unfortunately, this introduction as well as this book must end on a sad note.As if the extinction of at least a third of the known species before ornithologists studied them was not bad enough, half of the 38 historically known species are also now extinct and others are likely doomed. One species, the Po‘ouli, is believed to have only three surviving individuals as I write, and may not survive beyond the publication of this book. At least nine more species are threatened or endangered.The surviving species are like the scattered shards of a once great civilization. Consequently, a relatively small portion of this volume is devoted to information on breeding biology and ecology (Chapter 8) because so little of these subjects is knowable. Harvard biologist E. O.Wilson’s (1992) four ‘horsemen of the environmental apocalypse’ (overkill, habitat destruction, introduction of alien organisms, and disease) have produced an avian holocaust in Hawai‘i.The last of these has been especially prominent in the extinction

of Hawaiian birds and is examined in detail in Chapter 9. Overkill probably had less effect on the honeycreepers than on other Hawaiian bird groups. Nothing in honeycreeper history quite equals the slaughter of the Passenger Pigeon or the Great Auk (Fuller 2001), but Hawai‘i’s flightless birds probably were eaten to extinction by the first Hawaiian people (Olson and James 1982a, 1984; James and Olson 1983; Olson 1989a). Nevertheless, honeycreepers were much prized for their feathers, which were a medium of exchange woven into magnificent cloaks and other articles worn by the ali‘i (nobles). Appendix 1 (by Sheila Conant) discusses the importance of honeycreepers in Hawaiian culture and the effect feather-collecting may have had on bird populations. Habitat destruction and introduction of alien organisms are major threats that began with the Hawaiians and continue to the present day. Many and varied conservation efforts (Chapter 10) on behalf of the surviving honeycreepers may provide at least a tiny ray of hope for a better future. Now that you have been properly introduced, explore with me the world of the Hawaiian honeycreepers, their evolution, biology, history, and conservation.

2 The honeycreepers’ world

The physical environment A geological conveyor belt The present day Hawaiian Islands (Fig. 2.1) lie at the northern edge of the tropical Pacific 4000 km from the nearest continent (North America) and 3000 km from the nearest high volcanic islands (the Marquesas in southeastern Polynesia).They are the latest subaerial representation of an ancient long chain of islands and guyots or seamounts (the Emperor Chain) that stretch to the northwest and north almost to the Aleutians. As reviewed most recently by Fleischer et al. (1998), this chain is the result of the movement of the Pacific Plate, which forms the floor of the Pacific Ocean, over a mid-ocean ‘hotspot’.A hotspot is a thin place in the Earth’s outer mantle where a magma plume reaches the surface and produces volcanoes. These begin, like present day Lo‘ihi Seamount, 30 km south of Hawai‘i, as undersea eruptions that may eventually break the surface of the ocean. As eruptions continue, islands increase in size and elevation until plate movement carries them far enough from the hotspot that their connection with it is broken and their volcanoes go extinct (Menard 1986).The process has been likened to a ‘volcanic conveyor belt’ (Carson and Clague 1995; Fleischer et al. 1998). Often, an island includes lavas from several volcanoes that were too close together to remain separate islands as their mass increased. Carson and Clague (1995) reviewed the many geological factors that have operated to produce the landforms we see in the islands today. Island formation and growth is not a regular progression, and eruptive cycles may start and stop several times, with

intervening periods of erosion and subsidence. Young Hawaiian volcanoes tend to take on a classic shield configuration, as seen in today’s Mauna Loa, with a very smooth contour (Fig. 2.2a). In senescence, a shield volcano may become pockmarked on the summit and flanks by cinder cones, as is the case with Mauna Kea (Fig. 2.2b). Even before eruptions cease, the sheer weight of the accumulated lavas produces a sag in the crust under the island causing subsidence that reduces its subaerial elevation. After volcanism ceases, subsidence slows considerably and the effects of erosion become much more important. Erosion eventually may break a large island into several, and these island clusters may go through several cycles of break-up and reunification as a result of changes in sea level. Ultimately, the original volcanic island drops below the surface, but coral growth around the shoreline may form an atoll. Because of cooling currents from the Bering Sea, the waters around the Hawaiian Islands are considerably cooler than would otherwise be the case in the northern edge of the tropics, so coral growth is relatively slow. Thus, in contrast to atolls closer to the equator that may persist indefinitely barring too rapid a rise in sea level, Hawaiian atolls are comparatively short-lived. Even if the island core remains near the surface, plate movement eventually carries it too far north for corals to survive. The present atolls of Kure and Midway at the northwestern end of the archipelago are north of the Tropic of Cancer (Fig. 2.1).Today’s Hawaiian Archipelago comprises islands in nearly every stage of geological evolution, from the still growing Big Island of Hawai‘i to low-lying Kure Atoll. Hawaiian honeycreepers inhabit all island types within this

8 The Hawaiian Honeycreepers 160°

159°

158°

157°

156°

155°W

Hanalei Waimea Canyon

22°N

Waimea Hanapepe

Lihue

Manoa Valley

Pearl Harbor BarberÕ s Point Honolulu

Diamond Head

Kamakou Preserve

Approximate shoreline of Maui-nui

21°

MAUI

Lahaina

Waikamoi Preserve

Lanai City Hosmer Grove Haleakala NP Haleakala Polipoli Springs Ha Hakalau Forest NWR ma ku Kohala 20° a Waimea Co a Plains

st

Midway Kure

Pearl and Hermes Reef

N A K D

NO

Saddle Road

Lisianski RT

Tropic of Cancer

HW

Hanaunau

Laysan ES

TE

Mauna Loa Strip

Gardner Pinnacles RN

HA

Hilo Kaumana Volcano Village

Hawaii Volcanoes NP

19°

French Frigate Shoals WA

IIA

N

Necker ISL

AN

Nihoa

DS

Main Hawaiian Islands

18°

2.1 Map of the Hawaiian Archipelago showing important localities mentioned in the text.

range including still-growing Hawai‘i, the older, highly eroded islands further west, the volcanic fragment of Nihoa, and coralline Laysan. Geographers divide the present archipelago into the main Hawaiian Islands (the high islands from Ni‘ihau eastward), and the Northwestern Hawaiian Islands (the mostly uninhabited small islands northwest of Kaua‘i). For many years, biogeographers debated whether the Hawaiian honeycreepers (and other Hawaiian organisms) might have evolved in the Northwestern Hawaiian Islands (when still high islands), and secondarily colonised the present-day islands as they became habitable. Recent studies (reviewed in more

detail in Chapter 4), however, confirm that the honeycreepers colonised the archipelago and radiated on islands that we know today, and as a group are not older than Kaua‘i. Thus the evolution of the modern high Hawaiian Islands parallels that of the honeycreepers.

Climate and weather Hawaiian weather is dominated by prevailing northeasterly tradewinds that blow almost continuously. Because of the cool surrounding water and nearly continuous breeze, the islands’ climate is regarded as

The honeycreepers’ world 9 one of the most comfortable in the world. During occasional periods of ‘kona weather’, the tradewinds cease and warmer air moves in from the south. During such periods weather may be oppressively hot and humid. Both wind systems may bring rain to the islands. The islands also experience occasional cold fronts from the northwest, usually during the winter months (December–March). Seasonality is not pronounced, but winter is cooler and rainier than the rest of the year.The Hawaiian Islands are not often visited by catastrophic storms, but those that do occur can be devastating. Category 5 Hurricane ‘Iniki in 1992 caused major ecological damage on Kaua‘i, and may have delivered the coup de grâce to several native bird species including some honeycreepers. Depending on configuration and elevation relative to the wind direction, each island has a distinctive pattern of rainfall. In general, areas facing the northeast are wet, and those on the opposite side are dry, but the extent of such ‘rain shadows’ depends on several factors (see island discussions below) and habitats reflecting the full range of edaphic conditions may be found, from the wettest rainforests to the dry desert-like places.Temperatures are, as expected, most strongly influenced by elevation and run the full range from below freezing atop the highest peaks to high daytime temperatures in rain shadow deserts. Although variable from year to year, both Mauna Kea and Mauna Loa may be snow-capped. Snow sometimes also falls on Maui’s Haleakala, but never remains long on the ground.

The islands: a closer look The existing Hawaiian Islands exhibit a wide range of size and elevation (Table 2.1). Each has features that have affected the evolution of its birds, and each must be understood individually as well as in the archipelagic context in order to make sense of the honeycreepers’ history. The following survey takes a detailed look at all of the islands known to have been inhabited by honeycreepers.

Hawai‘i, The Big Island The youngest and by far the largest (more than twice the size of all the others combined) island is Hawai‘i, known locally as The Big Island.The size is close to

Table 2.1 Sizes and elevations of the Hawaiian Islands a. Island Hawai‘i

Maui

Area (km2)

Major peaks

Elevation (m)

10 433.1

Mauna Kea Mauna Loa Hualalai Haleakala Pu‘u Kukui

4205 4169 2521 3055 1764 452 1026 1514 1404 1220 960 1569 275

1883.7

Kaho‘olawe Lana‘i Moloka‘i

115.5 364.0 673.5

O‘ahu

1546.5

Kaua‘i Nihoa Pearl and Hermes Atoll Laysan Midway Atoll

1430.5 0.7

aFrom bTotal

0.3b 4.1 6.4b

Lana‘ihale Kamakou Oloku‘i Ka‘ala Ko‘olau Range Wai‘ale‘ale

3 12 12

Juvik and Juvik 1998. of islets.

that of Jamaica, the State of Delaware, or Wales. It has a number of superlatives: it is the largest purely oceanic island on the planet; Mauna Kea is the world’s tallest mountain (measured from sea floor to summit) and Mauna Loa the most massive; and Kilauea is one of the world’s most active volcanoes. The island was built, beginning about a half-million years ago, by six volcanoes in succession: Mahukona, Kohala, Mauna Kea, Hualalai, Mauna Loa, and Kilauea (Carson and Clague 1995). The first three are extinct, Hualalai was last active in 1808 and is now dormant, and the last two are presently active. The Mahukona Volcano is now subsumed by later lavas and has left little trace in present-day topography. Kohala Volcano has eroded, particularly on its windward side, into deeply incised amphitheatre valleys (Fig. 2.2c) and its volcanic structure has been so obliterated that most residents think of it as a range of mountains rather than a single volcano. Mauna Kea (‘white mountain’) is usually snow-capped from December to March (Fig. 2.2b). Its summit exhibits

10 The Hawaiian Honeycreepers

2.2 Landforms of the Big Island: (a) View from an airliner of the two highest peaks, extinct Mauna Kea (foreground) with typical summit cinder cones of old volcanoes, and active Mauna Loa (behind), a young shield volcano with smooth profile; (b) Mauna Loa looming over Halema‘uma‘u, a deep pit in the summit caldera of Kilauea Volcano; (c) Mauna Kea summit viewed from Pohakuloa, showing erosional ravines formed when late eruptions melted a summit glacier; (d) Pu‘u Wa‘a Wa‘a, a fluted cinder cone on ne. flank of historically active Hualalai, which forms the backdrop.

a series of cinder cones formed by late minor eruptions, and it once had a small glacier, as evidenced by moraines (Stearns 1985). Hualalai (Fig. 2.2c) likewise exhibits the pock-marked irregular summit of an old volcano, but Mauna Loa is a classic smooth shield volcano that may also receive a thin snow cap during the winter (Fig. 2.2a). Kilauea Volcano is unusual in that it has not built a cone of its own. Its caldera sits in a ‘shelf ’ on the south flank of Mauna Loa and its lavas always flow in one direction to the sea (Fig. 2.2a). All of these volcanoes have produced flank eruptions and cinder cones, some of which, like Pu‘u Wa‘a Wa‘a (Fig. 2.2d), are familiar landmarks. Hawai‘i’s youth, massive size, and ongoing volcanism produce features that are unique within the Hawaiian Archipelago. Some of these, such as

recent lava flows and their successional stages, are obvious, others less so. For example, the phenomenon of forest cycling, sometimes referred to as ‘dieback’, is seen primarily on The Big Island. It was long suspected to be another example of environmental degradation (Petteys et al. 1975), but now appears to result from natural phenomena. Lava flows set the ecological clock back to zero. Succession may then produce a uniform-age stand of trees. When all the canopy trees reach senescence nearly simultaneously, the forest ‘dies back’, and the canopy opens for regeneration and the next cycle (Mueller-Dombois 1985, 1987; Gerrish 1989).These diebacks were probably benign during most of the evolution of the honeycreepers, when birds could move from patches of dieback to other

The honeycreepers’ world 11 tracts in different stages of the cycle, but may have played a role in the demise of some species once forests were reduced to remnants. Another forest phenomenon on Hawai‘i is the presence of enclaves of old growth surrounded by younger lava flows. The Hawaiian word kipuka has become the English loan-word for such isolated forests. Some kipukas have acted as de facto refuges for endangered species, including some honeycreepers. Another Big Island phenomenon is the unusual Kona Coast weather system (Carlquist 1970). Here, the expected pattern of wet windward/dry leeward is disrupted by the sheer size of the three largest volcanoes. The Kona Coast is unexpectedly wet, with a major rainforest belt, because the influence of the tradewinds is almost totally blocked. The long slope formed by the leeward (western) flanks of Mauna Loa and Hualalai actually forms its own local weather pattern.At sunrise, the slope is usually clear, but as the sun heats the surface it creates an updraft that draws cool moist air in from the sea.At mid-morning, a cloud bank begins to form in the middle elevations and by afternoon, rain often falls. (The ever-present midday cloud bank facilitated the cultivation of Kona Coffee in this region.) All the while, the beach may remain dry and sunny as would be expected in a rain shadow, so the area is also ideal as a resort. At night, the slope cools more rapidly than the offshore waters, and the draft reverses, carrying the clouds out to sea where they dissipate until the next day. Unless disrupted by kona storms (i.e. those that approach from the southwest) or other irregular events, the Kona weather system is remarkably predictable. Whether it contributed to the fact that the Kona Region was the last refuge of several finchlike honeycreepers, as well as the ‘Alala or Hawaiian Crow, is an intriguing question that probably cannot yet be answered.

The Maui-nui complex ‘Maui-nui’ means ‘big Maui’, and has become the name of choice for a large island that existed just prior to modern times that conjoined the modern islands of Maui, Moloka‘i, Lana‘i, and Kaho‘olawe. Maui-nui was formed by seven volcanoes that correspond to the present islands except for Moloka‘i and Maui, each of

which has an independent eastern and western component, plus the now submerged Penguin Bank that extends westward from Moloka‘i toward O‘ahu.The oldest that has been dated is West Moloka‘i at 1.9 million years, youngest Haleakala at 0.79 million years (Carson and Clague 1995). The channel between Maui and Hawaii, though narrow, is very deep and we can be certain that no Maui/Hawai‘i connection ever existed, although when Hawai‘i first emerged, the channel was narrower than it is today. Until very recently, the same assumption had been made regarding a Maui/O‘ahu connection, but Carson and Clague (1995) found that Penguin Bank probably did provide a dry land connection to O‘ahu when Mauinui was geologically young. The exact date of this connection has not been determined, but is believed to have been before the original Maui-nui first separated into the modern islands. This connection has helped to explain several previously enigmatic distributions of Hawaiian organisms. Maui-nui has gone through several cycles of break-up and consolidation as a result of changes in sea level during glacial periods. It first broke into two islands (Moloka‘i/Lana‘i and Maui/Kaho ‘olawe), then Kaho‘olawe separated, followed by Lana‘i. All have been reunited from time to time, most recently around 12 000 to 15 000 years ago (Menard 1986). During maximal sea levels, East and West Maui were separate islands. Although the historically known avifaunas of Maui, Lana‘i, and Moloka‘i differ, the recency of this connection allows us to reconstruct the avifauna of Maui-nui with some confidence by combining the lists. Present-day Maui is nicknamed The Valley Isle because of the low isthmus connecting its eastern and western parts. East Maui is dominated by massive Haleakala, the islands’ third highest peak. Considered dormant rather than extinct, Haleakala last erupted around 1790 on its flank near the coast. Haleakala has had a complicated geological history, and its current summit crater (Fig. 2.3) is not a true caldera. After a very long period of dormancy, with heavy erosion that carved the original volcano into a series of deep valleys and ridges, eruptions resumed and refilled the central part to produce the floor of the present crater, which is dominated by a line of cinder cones. Lavas eventually flowed to the

12 The Hawaiian Honeycreepers

2.3 Summit crater of Haleakala, formed first by erosion, then refilled by subsequent eruptions along a central rift zone. It is a crater, but not a caldera.

sea through Ko‘olau Gap to the north and Kaupo Gap to the south.The northeast slope of Haleakala strikes the tradewinds head-on, and produces heavy rainfall. The lush rainforests that result have been especially important in the history of Hawaiian honeycreepers, and are today the last stronghold of several critically endangered species. The leeward slope is not as dry as might be expected because it generates a miniature version of the Kona Coast weather system, with some mid-elevation rainfall. The original mid-elevation forests on this slope have largely been replaced by pastures and plantations of alien trees. The central isthmus of Maui is today either devoted to agriculture or supports dry scrub; none of the original lowland vegetation survives. West Maui also receives the full force of the tradewinds, but is not nearly as high or as wet as the northeast slope of Haleakala. West Maui is much older and more deeply eroded, with valley walls so steep that the growth of tall forest is inhibited. Although it harboured several species (none unique), West Maui has played little role in the island’s ornithological history. Despite the fact that the West Maui town of Lahaina was the capital before Honolulu, Maui was visited much later by ornithologists than the other major islands (see Chapter 3), by which time it had lost several species. Thus the fact that its historically recorded passerine avifauna consisted entirely of honeycreepers is an artefact. The uninhabited low island of Kaho‘olawe likewise has essentially no ornithological significance

2.4 Lana‘i/Moloka‘i views: (a) Lana‘i viewed from Kamakou area, Moloka‘i; (b) isolated peak of Oloku‘i, which still harbours a nearly pristine (except for the absence of most native birds) Hawaiian rainforest.

except possibly as a nesting site for seabirds. It lies in Maui’s rain shadow and has no permanent fresh water. For nearly half a century, the US Navy used the island as a bombing target and it is today a devastated wasteland. No honeycreepers are known to have been found there, but searches for prehistoric remains might prove productive. However, such work must await the clearance of unexploded ordnance. Kaho‘olawe has recently been returned to the stewardship of the State of Hawaii, and ideas of how to restore the island have been discussed. It is a potential reintroduction site for Laysan or Nihoa finches (see Chapter 10). Lana‘i (Fig. 2.4a) lies in West Maui’s rain shadow, but the effect is not so pronounced as that of Haleakala on Kaho‘olawe. Consequently, a small area of mesic or near-rainforest is found at the highest elevations. Lana‘i was not permanently

The honeycreepers’ world 13 inhabited until the twentieth century, and its avifauna suffered declines later than those of neighbouring islands. Several elements of the Maui-nui avifauna survived on Lana‘i only and thus appeared historically to be single-island endemics.Today, the forest is heavily invaded by alien plants and the rest of the island is either given over to weeds, now that agriculture has ceased, or nearly barren. Moloka‘i is a long narrow island oriented east to west. East Moloka‘i is much the higher part, and is highly eroded with two deep amphitheatre valleys that isolate the peak known as ‘Oloku‘i (Fig. 2.4b). This peak remained pristine long after much of the island had been ecologically ravaged.The dense rainforests of East Moloka‘i sheltered a particularly rich subset of the Maui-nui avifauna. Today, although forests remain at higher elevations, most of the native birds are gone.West Moloka‘i is low-lying and of little historical interest ornithologically, but is the site of the Mo‘omomi lithified sand dune formation, one of the richest sources of prehistoric bird bones.

O‘ahu The third largest of the main islands, O‘ahu comes closest to giving the appearance of having mountain ranges (rather than eroded single volcanoes). The Wai‘anae Range in the west and the Ko‘olau Range in the east are oriented along a northwest–southeast gradient and slightly offset. Despite their names and superficial appearance, each ‘range’ is the remnant of a single volcano. The western rim of the original Ko‘olau caldera forms the line of cliffs known as The Pali behind Kaneohe Bay, the seaward rim having long ago collapsed into the sea, and the Wai‘anae caldera is similarly obscured by subsequent erosion. The island’s highest peak, Mt. Ka‘ala, lies in the Wai‘anae Range, but it rises almost like a spike above the surrounding peaks, and the Ko‘olau Range actually has a higher average elevation. Although the Wai‘anae Range lies partly in the rain shadow of the Ko‘olaus, the island’s configuration mitigates the effect and both ranges receive abundant rainfall.The broad valley between the two massifs is flooded by the sea at its southern end, forming Pearl Harbor, the only deep water harbour in the archipelago. The southeastern end of the island is

marked by a number of prominent cinder cones including Diamond Head, Punchbowl Crater, Koko Head, and Koko Crater. Because of the presence of Pearl Harbor, O‘ahu has long been the political and commercial centre of Hawai‘i. It was the first island to experience a large number of historical extinctions of birds and remains today the most ecologically altered island. Much of O‘ahu today is urbanised, and the City of Honolulu is home to about 80% of the state’s population.

Kaua‘i and its neighbours Kaua‘i, at 5.1 million years the oldest of the main Hawaiian Islands, is also the most highly eroded. The original configuration of its volcano is difficult to discern.The central peak,Wai‘ale‘ale (Fig. 2.5a), is situated in such a way as to extract maximum rainfall from the tradewinds (over 12 m/year), and was long believed to be the world’s wettest spot. Wai‘ale‘ale is not a well-defined peak, but rather the highest point on the leading edge of the Alaka‘i Plateau, which slopes away to the southwest. Although it reaches roughly the same elevation as much larger O‘ahu, Kaua‘i actually has much more surviving high-elevation forest because of this high plateau. Long known as the Alaka‘i Swamp because of its numerous bogs, this forested wilderness (Fig. 2.5b) provided a refuge for native birds long after they had disappeared from the rest of the island. Drainage to the south from the Alaka‘i has produced spectacular Waimea Canyon (Fig. 2.5c), while the northern edge drops sharply to the sea in a series of knife-like ridges and deep amphitheatre valleys (Fig. 2.5d). Lying to the southwest and in the rain shadow of Kaua‘i are the smaller islands of Ni‘ihau and Ka‘ula. They have all been joined in times past although they represent separate volcanic systems. Neither of these low islands has much significance for Hawaiian honeycreepers (Fisher 1951), although a search for prehistoric remains might be very productive on Ni‘ihau.

The Northwestern Hawaiian Islands The relatively tiny, uninhabited islands that stretch to the northwest from the main islands are entirely different ecologically from the latter. Some are jagged

14 The Hawaiian Honeycreepers

2.5 Kauai landforms: (a) Wai‘ale‘ale, whose summit was once thought to be the world’s wettest spot, from the east on an unusually clear morning; (b) Alaka‘i Plateau viewed from Pihea Lookout, looking eastward toward Wai‘ale‘ale; (c) Waimea Canyon, formed by stream drainage from the Alaka‘i Plateau, with bare earth largely the product of grazing by feral goats; (d) Kalalau Valley, a typical amphitheatre valley of a heavily eroded ancient volcanic island.

volcanic remnants, while others are true atolls. Two of them supported populations of Hawaiian honeycreepers in historic times and others support, or have supported, translocated populations. Nihoa, closest to the main islands, is a tiny volcanic remnant that rises from the sea with nearly vertical high cliffs. It has almost no level land and only shrubby vegetation. The drepanidine Nihoa Finch shares the island with only one other passerine, a sylviine warbler called the Millerbird. Because of its forbiddingly steep terrain and difficulty of landing, Nihoa has seen little disturbance and remains today much as it has for centuries. A population of the Nihoa Finch introduced to French Frigate Shoals in 1967 did not persist. Further west, Laysan is a flat coral island that can be considered a borderline atoll. It lacks a central lagoon, but possesses a central saline lake. Laysan was the site of one of the twentieth century’s great

avian disasters. Commercial attempts to exploit guano produced by millions of seabirds on Laysan ultimately proved the undoing of the island’s ecosystem. In 1902, domestic rabbits were brought to the island by the guano workers.These were eventually liberated, and, in the absence of predators, multiplied geometrically. When the guano works played out, the rabbits were left to themselves. The result was vividly reported by the noted ornithologist Alexander Wetmore (1925) who was a member of the USS Tanager Expedition in 1923: Early accounts of Laysan Island and photographs taken 20 years ago depicted it as a pleasant spot covered with green vegetation. Reports of damage to shrubbery had led us to expect some changes, but had not prepared us

The honeycreepers’ world 15 for the utter desolation that greeted Commander King and me when we landed in the little harbor and walked slowly up the sandy slopes to a point near the tumbledown buildings remaining from the guano workings. On every hand extended a barren waste of sand. Two coconut palms, a stunted hau tree and an ironwood or two, planted by former inhabitants, were the only bits of green that greeted the eye. Other vegetation had vanished. The desolateness of the scene was so depressing that unconsciously we talked in undertones. From all appearances Laysan might have been some desert, with the gleaming lake below merely a mirage . . . In spite of the rabbits, a few dozen Laysan Finches still sang their sprightly songs about the buildings or hopped among the rocks near the lagoon. Three individuals alone of the little Honey Eater [⫽Laysan Honeycreeper] remained on our arrival; these perished during a three-day gale that enveloped everything in a cloud of swirling sand. This may have been the only instance in which humans actually witnessed the death of the last member of a Hawaiian honeycreeper species (although it could happen again very soon). The finch managed to survive the holocaust, apparently by feeding on seabird eggs until vegetation returned.Also surviving was the Laysan Duck, but the Laysan Rail and the island’s only non-drepanidine passerine, the Laysan Millerbird, were lost forever.Wetmore and his associates killed the few hundred rabbits that had not already starved to death, but the ecological damage had been done.Today Laysan probably looks, superficially, much like it did before the rabbit plague, but it is a very different place without three-fifths of its terrestrial avifauna. Laysan Finches have twice been successfully translocated to other islands.They lived successfully for several decades on Midway Atoll, despite extensive human disturbance there, until the Second World War brought rats ashore (Fisher and Baldwin 1946a). Another atoll, Pearl and Hermes Reef, still harbours an apparently established population (AOU 1998). Now that rats have been eliminated

from Midway (R. Shallenberger, pers. comm.), the possibility of reintroducing the Laysan Finch there is being discussed, but will have to await further ecological recovery, including control of mosquitoes (see Chapters 9 and 10).

The living community Because the oceanic islands are younger than the surrounding continents, their flora and fauna had to originate by long-distance transoceanic dispersal. Not all organisms are equally capable of such dispersal, so island communities are characterised by the absence of certain continental groups. Among birds, for example, woodpeckers, trogons, toucans, gallinaceous birds, hummingbirds, bulbuls, babblers, and tanagers are almost never found naturally on oceanic islands, whereas rails, kingfishers, pigeons and doves, swifts, honeyeaters, and sylviine warblers are particularly adept at long-distance colonisation. Among terrestrial animals in general, those that fly or are easily transported on driftwood over long distances are more likely to be found on islands. Thus, mammals other than bats are rare on remote islands, amphibians are almost totally absent, and insects are abundant. This differential vagility (dispersal ability) produces natural communities on islands that are ‘disharmonic’, with a more or less arbitrary and spotty representation of continental organisms. The Hawaiian Islands provide the most extreme example of colonisation across an oceanic filter (Carlquist 1970, 1974), and disharmony is pronounced in both the plant and animal communities.

Hawaiian habitats and plant communities Many authors have attempted to classify Hawaiian plant associations, some in very broad terms (Scott et al. 1986), others in more detail (Wagner et al. 1990).The following classification is a distillation of several others and focuses only on those habitats that have significance to Hawaiian honeycreepers. Other descriptions can be found in Pratt (2002b). For colour photographs of most of the plants mentioned herein, see Pratt (1999a) or Beletsky (2000).

16 The Hawaiian Honeycreepers Scientific names and families of plants are given in Appendix 2.

Rainforests Although they clearly qualify on the basis of rainfall, Hawaiian rainforests are very different from those of the Amazon, the Congo Basin, or tropical Asia, characterised by high tree species diversity, multistoried growth, and numerous lianas (woody vines). Hawaiian rainforests are dominated by a mere handful of canopy tree species, have only three recognisable strata, and support relatively few lianas (Carlquist 1970; Pratt 1999a).They are found on the windward sides of all the main islands. The dominant tree in all Hawaiian rainforests is ‘ohi‘alehua (Fig. 2.6). (The first part of the name refers to the tree, the second to the blossoms, but most naturalists today shorten the name to ‘ohi‘a in all situations.) In many places, this remarkable tree forms the entire canopy. It typically is gnarled and twisted (Fig. 2.6a), but in a few places, e.g. South Kona, grows tall and straight (Fig. 2.6b). It belongs to the Myrtaceae (eucalyptus, etc.) and has the ‘shaving-brush’ flowers typical of that family. The ‘brush’ is formed by stamens and pistils; petals are lacking (Fig. 9.8).The calyx forms a cup at the base of a flower cluster in which nectar collects. Other Metrosideros outside Hawai‘i are primarily wind pollinated, but in the very wet Hawaiian rainforests, birds apparently play the most important role (Carpenter 1976). Most lehua blossoms are scarlet, but may be orange, and a yellow species, lehua mamo, is endemic to O‘ahu (Wagner et al. 1990). Lehua nectar is a major food source for nectarivorous honeycreepers. Interestingly,‘ohi‘a is one of the first pioneers on fresh lava flows, a rather unexpected characteristic for a climax canopy tree. The other major canopy tree of Hawaiian rainforests is koa, a majestic tree (Fig. 2.7a) that only occasionally forms uniform stands. It usually grows as a scattered emergent tree that towers above the main canopy of ‘ohi‘a. Koa tends to grow more abundantly in slightly drier zones such as the upper forest belt on Hawai‘i, above the wettest parts of the forest (Scott et al. 1986), but may originally have had a lowland distribution as well. It is a legume with small yellow pom-pom flower clusters and flattened seed pods. What appear to be sickle-shaped leaves

2.6 ‘Ohi‘a-lehua, Hawai‘i’s dominant canopy tree: (a) usual gnarled growth-form, La‘ie Trail, O‘ahu; (b) straight-trunked variant, Kipuka Puaulu, Hawaii Volcanoes NP; see Figs 6.7, 6.10, 6.13–14, and 8.8 for flowers, Figs 6.5, 8.19–20 for foliage, Fig. 6.6 for buds.

The honeycreepers’ world 17

2.7 Koa, often emergent over the ‘o¯hi‘a canopy in Hawaiian forests: (a) typical forest giant in the edge of a kipuka, Keauhou Ranch, Puna Dist., Hawai‘i; (b) true leaves (compound) and phyllodes (falcate) of koa, Kipuka Puaulu, Hawaii Volcanoes NP.

(Fig. 2.7b) are botanically classified as phyllodes (expanded petioles), with more typically acacia-like double-compound leaves found on saplings and

adventitious branches on old trees. Koa is an important feeding substrate for insectivorous honeycreepers as well as a nectar source, and the seeds were fed upon by the specialised koa-finches. ‘Ohi‘a and koa account for well over 90% of the rainforest canopy on all islands. A few other canopy trees with more restricted distributions are worthy of mention. From Kaua‘i to Maui grows ‘ohi‘a-ha, related to Metrosideros but with very different flowers and growth form, and all islands have one or more species of ‘ohe, characterised by large leathery compound leaves.At least some species of ‘ohe were adapted for bird pollination (Wagner et al. 1990). Native fan palms called loulu were originally much more numerous than today and grew as scattered emergents or in small groves within the broader rainforest. The name of the ‘Ula-‘ai-hawane (q.v.) supposedly alludes to its feeding on native palms (Pratt 2002a). Underneath the canopy grows a subcanopy layer of smaller trees that merges seamlessly into a lower shrub layer. In very wet forests, such as the ‘Ola‘a Tract of Hawai‘i Volcanoes NP, the subcanopy may be a nearly closed layer of tree ferns or hapu‘u. In other places, the subcanopy is fairly open and includes many trees with nectariferous flowers and soft fruits such as ‘olapa (Fig. 8.2b) and lapalapa; kawau, a Hawaiian holly; several species of alani; kolea lau nui (Fig. 8.2d); several species of pilo (Fig. 8.2a), usually with bright orange fruits; and a tree huckleberry, ‘ohelo kau la‘au (Fig. 8.4). These provide food resources for both nectarivores and frugivores (see Chapter 8). Common rainforest understorey shrubs include the native raspberry ‘akala (Fig. 8.3c), whose flowers are popular with nectarivores, especially ‘I‘iwi; pukiawe, which also grows in open scrub habitats; manono, a highly variable plant that can be either a vine, shrub, or small tree; and kanawao (Fig. 8.9d), whose flowers and fruits both attract several species of honeycreeper. One group of shrubs of particular interest, because their evolutionary history is closely intertwined with that of the honeycreepers (see Chapters 4 and 8), are the Hawaiian lobelioids (Campanulaceae) (Fig. 2.8). They represent an example of adaptive radiation comparable to that of the honeycreepers (Givnish et al. 1995; Lammers 1995), and like them have

18 The Hawaiian Honeycreepers

2.8 Growth forms of four genera of Hawaiian lobelioids: (a) rosette form of woody lobelia Lobelia hillebrandii; (b) near rosette of Trematolobelia kauaiensis; (c) multi-branched shrub form in Clermontia fauriei; (d) true rosette of Cyanea leptostegia.

suffered severe reductions and extinctions, perhaps because of the loss of avian pollinators (Cox and Elmqvist 2000). From a herbaceous continental Lobelia evolved woody members of that genus and six endemic Hawaiian genera (Brighamia, Clermontia, Cyanea, Delissea, Rollandia, and Trematolobelia), all of which may be bird pollinated. All except Brighamia are found primarily in rainforests, and their flowers figure prominently in hypotheses of co-evolution with honeycreepers (see Chapters 4 and 8). Most of them are palm-like rosette plants, i.e. a rounded cluster of leaves atop a tall trunk, but Clermontia has a more typical branching shrub growth form (Fig. 2.8c). The close association of birds with these shrubs is even reflected in the native name of Clermontia fauriei, Kaua‘i’s only member of that genus. Members of this genus are called ‘oha or haha, and C. fauriei is called haha‘aiakamanu (Wagner et al. 1990), which roughly translates as ‘bird-food Clermontia’.

Growing through all layers of the rainforest may be found one of Hawai‘i’s few true lianas, the climbing screw-pine or ‘ie‘ie. In some forests, such as those on the Kona slope of Hualalai, it is a prominent feature (Fig. 2.9) but elsewhere it may be quite rare.‘Ie‘ie produces a soft fruit cluster that was a favoured food of the ‘O‘ u, and its sheathed leaf bases harbour insects available to gaping probers such as the Greater ‘Amakihi or long-billed insectivores such as ‘akialoas. Rainforests have other smaller vines including the aromatic maile, and the briarless greenbrier called hoi kuahiwi. Several viny native mints of the genus Stenogyne have flowers adapted for bird pollination. Epiphytes are an important component of Hawaiian wet forests and include many mosses and ferns as well as a few flowering plants such as pa‘iniu, a lily that mimics the growth form of some bromeliads. Bromeliads are poor island colonisers and absent in Hawaii, and there are relatively few native orchids compared to other tropical places.

The honeycreepers’ world 19 Village on Hawai‘i. Elsewhere it has largely been given over to agriculture or urban development. It is a more open forest, with much less epiphytic growth than rainforest. The canopy is again mainly ‘ohi‘a and koa, but on Hawai‘i these are joined prominently by an indigenous soapberry, known locally as either a‘e or manele. It is the understorey that makes this habitat so rich, although many of the plants are rare today. Among the more conspicuous elements are Haswaiian olive or olopua, various species of kopiko, ‘ala‘a, several species of papala, a sandalwood or ‘ili‘ahi, kauila, and several ho‘awa species.Today, the temperate mixed forest is a major conduit for invasive weeds entering the remaining rainforests (Pratt 1999a).

Mamane–naio forest

2.9 ‘Ie‘ie, one of the few true lianas of Hawaiian rainforests.

Temperate mixed forest Despite the historical dominance of rainforest in the lives of Hawaiian honeycreepers, another habitat may have originally been more important. At middle elevations in areas with moderate rainfall, a mixed mesic forest grows that is even richer in plant species than Hawaiian rainforest (Sohmer and Gustafson 1987), and may likewise have originally harboured more bird species. This forest was destroyed, except for a few remnants, fairly early in the islands’ history and we have no way of knowing how important it may have been for the honeycreepers (although prehistoric remains give us some clues; James and Olson 1991).Temperate mixed forest can today be found, in highly altered form, around Koke‘e on Kaua‘i, and around Hawaii Volcanoes NP (especially Kipuka Pua‘ulu) and Volcano

As its name implies, this community is dominated by two shrubby trees, mamane and naio (Fig. 2.10). It is an open, park-like forest with no closed canopy, found primarily on the high leeward slopes of Mauna Kea and nearby areas of the Mauna Kea/ Mauna Loa ‘saddle’. Mountain sandalwood may also have been important at one time, but was largely logged out during the sandalwood trade of the early nineteenth century. Although both of the dominant trees are more widespread, this particular association occurs today only on Hawai‘i.The Palila is restricted to this habitat, and feeds mostly on mamane seed pods. This may also have been the main habitat for the Kona Grosbeak, a naio specialist.When the yellow mamane flowers are in bloom, the forest is visited by the nectarivores, but their use of this habitat is transitory.The Hawai‘i ‘Amakihi, however, reaches its greatest density in mamane–naio forest. Like other Hawaiian communities, this one is now heavily infested with weeds and much damaged by feral ungulates (van Riper 1980b; Pratt 1999a).

Lowland dry forest Probably the most endangered ecosystem in Hawai‘i today is the lowland dry forest, which the prehistoric evidence tells us was of major importance to Hawaiian honeycreepers before the coming of humans to the islands. For reasons that will become apparent later, very few honeycreepers

20 The Hawaiian Honeycreepers tered individuals in disturbed areas. Amazingly, ‘ohi‘a is an important component of this dry habitat, but it is not dominant in most places.That role usually is filled by Hawaiian persimmon or lama. More important for birds is wiliwili, the native coralbean, whose flowers, like those of many other native plants, are adapted for bird pollination. Formerly several species of Hibiscadelphus (Malvaceae) with ornithophilous flowers (Fig. 8.17) grew in this habitat, but all are now mostly maintained in cultivation.

Other habitats

2.10 Montane dry forest: (a) mamane flowers and leaves; (b) naio flowers, leaves, and green fruits; (c) park-like open mamane–naio forest.

live in this habitat today. In overall aspect, the lowland dry forest resembles mamane–naio forest, with no closed canopy.Tattered remnants of it can be seen in the S. Kohala Dist. of Hawai‘i, in Hawaii Volcanoes NP, and in one tiny area on the south slope of Haleakala, Maui. More often, one finds species typical of this habitat growing as scat-

The main Hawaiian Islands have many other habitats in addition to those described above, but none are particularly important to Hawaiian honeycreepers. A fascinating subalpine scrub community, called subalpine dry shrubland by Wagner et al. (1990), can be found on Hawai‘i and Maui, but it does not apparently support a resident honeycreeper community. A few species, such as Hawai‘i ‘Amakihi, ‘Apapane, and Maui ‘Alauahio, often visit this zone, however. Other restricted habitats, such as cliff faces, alpine zones, rain shadow deserts, and shoreline communities are of more interest to botanists than ornithologists (Carlquist 1970; Pratt 1999a). Nihoa’s shrub and grass community (Conant 1985) includes several endemic plants, most notably Pritchardia remota, the only loulu palm that survives in a dry habitat today. Predominant shrubs include ‘ilima and ‘aweoweo, both of which are common coastal plants in the main islands. Following the rabbit plague of the early twentieth century, Laysan’s vegetation recovered to a grass and shrub community that, though it has lost several endemics, resembles the original vegetation (Berger 1981). The dominant shrub is naupaka kahakai or beach naupaka, a common shoreline plant throughout the tropical Pacific. Also common are the bunchgrass kawelu (Fig. 11.3) and two morning-glories, Ipomoea pes-caprae and I. indica.The grass appears to be particularly important for the Laysan Finch.The indigenous puncture vine, nohu, is uncommon on Laysan (Ely and Clapp 1973), but an important component of the vegetation of Pearl and Hermes Reef, where it is fed upon by introduced Laysan Finches (Amerson et al. 1974; Freed et al. 1987a).

The honeycreepers’ world 21

The terrestrial fauna Other birds Other than the honeycreepers, the Hawaiian nonmarine avifauna (Table 2.2) was the result of surprisingly few successful colonisations. Even including birds known only from bones, the avifauna descended from only about 20 lineages (the exact number depends on as-yet undetermined relationships among some of the fossil groups). Non-passerines include the historically known geese, ducks, stilt, moorhen, coot, rails, and buteo (Berger 1981; Pratt et al. 1987).The subfossil record (Olson and James 1991; Ziegler 2002) has so far added additional rails and geese, giant flightless waterfowl called moanalos, flightless ibises, the White-tailed Sea-Eagle (Fleischer et al. 2000), the Wood Harrier that resembled mainland accipiters more than its actual relatives, and several species of long-legged bird-eating stilt-owls that were likewise convergent with accipiters in their morphology.The latter three, plus the Hawaiian Hawk, may have played an important role in the evolution

of cryptic coloration and possibly even chemical defences among Hawaiian honeycreepers (see Chapters 4 and 6). Recently discovered subfossils have not revealed any passerine lineages other than those known historically. None of the other passerine colonisations produced anything like the drepanidine radiation, but they may have closed off a few avenues that would otherwise have been available to honeycreepers (see Chapter 4). They include monarchine flycatchers and honeyeaters from Australasia, solitaires from America, sylviine warblers (restricted to the Northwestern Hawaiian Islands) with South Pacific affinities, and crows of unknown provenance (Fleischer and McIntosh 2001).

Other vertebrates Probably nothing reflects the disharmony of the Hawaiian fauna more than the non-avian animals. The only other vertebrates that occur naturally are bats (the extant hoary bat Lasiurus cinereus semotus and a recently discovered fossil species) and possibly a few lizards. Non-flying mammals were totally absent

Table 2.2 Hawaiian native resident non-marine, non-drepanidine birdsa with their distributions. Lyb Family Plataleidae Moloka‘i Kiwi-Ibis Maui Kiwi-Ibis Family Anatidae Tortoise-faced Moa-nalo O‘ahu Moa-nalo Maui-nui Moa-nalo Pau Moa-nalo Nene (Hawaiian Goose) Kaua‘i Nene-nui Maui Nene-nui Hawai‘i Nene-nui Wetmore’s Flightless Goose Laysan Duck Koloa (Hawaiian Duck) Family Rallidae Laysan Rail Ziegler’s Rail

N

K

O

Apteribis glenos A. brevis Chelychelynechen quassus Thambetochen xanion T. chauliodous Ptaiochen pau Branta sandvicensis B. undescribed B. hylobadistes B. undescribed Geochen rhuax Anas laysanensis A. wyvilliana Porzana palmeri P. ziegleri

Mo

Ln

Ma

H

† † † † † †c †

†

†

† † †c

‡

†

√ ‡

‡

√ †

‡

‡

† † † ‡

22 The Hawaiian Honeycreepers Table 2.2 contd. Lyb Menehune Rail Keplers’ Rail Hawaiian Rail Ralphs’ Rail Severns’s Rail Hawaiian Coot Common Moorhen Family Recurvirostridae Hawaiian Stilte Family Accipitridae White-tailed Eaglef Wood Harrier Hawaiian Hawk (‘Io) Family Strigidae Short-eared Owl Kaua‘i Stilt-Owl Orion Stilt-Owl Moloka‘i Stilt-Owl Erdman’s Stilt-Owl Family Corvidae Alala (Hawaiian Crow) Arch-billed Crow Alala-loa Alala-nui-loa Family Monarchidaee Kaua‘i ‘Elepaio O‘ahu ‘Elepaio Hawai‘i ‘Elepaio Family Sylviidae Millerbird Family Turdidae Kama‘o Oloma‘o ‘Oma‘o Puaiohi Family Meliphagidae Kaua‘i ‘O‘ o (‘O‘ o‘a‘a) O‘ahu ‘O‘ o Bishop’s ‘O‘ o

N

K

O

Mo

Ln

Ma

H

P. menehune P. keplerorum P. sandwichensis P. ralphorum P. severnsi Fulica alai Gallinula chloropus

†

√ √

‡ √

√ √

√

† √ √

√ √

Himantopus knudseni

√

‡

√

√

√

√

† †

† † †

† ‡ †

Haliaeetus albicilla Circus dossenus Buteo solitarius Asio flammeus Grallistrix auceps G. orion G. geleches G. erdmani

†g √

√ †

√

√ √

√

√

† † †

Corvus hawaiiensis C. impluviatus C. viriosus C. undescribed

? † †

√

† †

√

Chasiempis sclateri C. ibidis C. sandwichensis Acrocephalus familiaris

√

†

√ √ √

√

Myadestes myadestinus M. lanaiensis M. obscurus M. palmeri

‡

Moho braccatus M. apicalis M. bishopi

‡

‡

‡

√

† ‡

‡

‡ ‡

‡h

The honeycreepers’ world 23 Table 2.2 contd. Lyb Hawai‘i ‘O‘ o Hawai‘i Kioea Narrow-billed Kioea

N

K

M. nobilis Chaetoptila angustipluma C. undescribed

O

?

Mo

Ln

Ma

? †

H ‡ ‡

√⫽ historical record, † ⫽ subfossil record, ‡ ⫽ historical and subfossil record, ?⫽ occurrence uncertain. aUndescribed subfossil species from Olson and James 1990, James and Olson 1990, as modified by H. James (pers. comm.). bIsland abbreviations: Ly ⫽ Laysan, N ⫽ Nihoa, K ⫽ Kaua‘i, O ⫽ O‘ahu, Mo ⫽ Moloka‘i, Ln ⫽ Lana‘i, Ma ⫽ Maui, H ⫽ Hawai‘i. cReintroduced. dRhymer 2001. eTaxonomy follows Pratt and Pratt 2001. fFleischer et al. 2000. gOlson and James 1997. hHenshaw 1902, Sabo 1982.

before humans. Amphibians are unable to survive long ocean passages.The skinks and geckos that were present at European discovery are most likely to have been hitchhikers in Polynesian canoes, though a few may yet prove to be indigenous (McKeown 1996). Even tortoises, which are surprisingly good island colonisers, failed to reach the remote Hawaiian Islands.These glaring gaps in the fabric of life meant that birds could fill niches normally better suited to grazing mammals, tortoises, or other animals (Olson and James 1991; Ziegler 2002).

Invertebrates Invertebrate groups are well represented in the Hawaiian Islands, but as with the vertebrates, they form a disharmonic fauna. The most important from a honeycreeper’s perspective are the arthropods and land snails, both of which provided major food resources. More significant in the long run, however, was what arthropods were not present, rather than what were. In particular, Hawai‘i had no honeybees (hence the dependence of many plants on bird pollinators), no ants (so that native insects became vulnerable to introduced predatory ants), and especially, no mosquitoes to vector bird diseases carried in by migratory shorebirds and waterfowl. Although Lepidoptera are well represented, including some hawk-moths that

are potential pollinators, only two species of nectar-feeding butterflies made a successful ocean crossing until modern humans introduced the necessary food plants (Howarth and Mull 1992; Beletsky 2000; Pratt 2002b).

Human influences The Polynesians About 1500 years ago (Kirch 1985; Olson 1999), the first human footprints appeared in the Hawaiian Islands. From the first landfall, these Polynesian voyagers began the alteration of the environment. The big, slow moanalos and abundant nesting seabirds undoubtedly provided a ready source of protein (van Riper and Scott 2001) until agriculture could become established. The Hawaiians brought in pigs, dogs, and chickens, which were liberated to forage and build up sustainable populations. Hitchhikers in the canoes such as geckos, skinks, and Polynesian rats Rattus exulans (Burney 1997) likewise found the new land hospitable. As human populations increased, at first slowly and then rapidly (Cuddihy and Stone 1990), lowland forests were cut or burned (Burney et al. 1995) wherever land was level enough to support food crops of taro, yams, sweet potatoes, breadfruit, bananas, and coconut and fibre plants such as

24 The Hawaiian Honeycreepers wauke (paper-mulberry). Diversion of streams produced artificial wetlands for growing taro and fish cultivation, and native riparian vegetation was replaced by aboriginal introductions such as hau and kuku‘i (Newman 1972). Shorelines were dotted with artificial fishponds. Special hunters were employed to raid upland forests for colourful birds (Athens et al. 1991) whose feathers were a muchprized cultural item (see Appendix 1). Eventually a complex society developed whose population may have exceeded that of the present day on several islands (Schmitt 1971; Stannard 1989; but see Juvik and Juvik 1998). Virtually all of the inhabitable lowlands were populated (Tuggle 1979) and their vegetation totally altered (Athens 1997). Cuddihy and Stone (1990) provide a detailed island-byisland account of environmental changes wrought by early Hawaiians. When Europeans first visited the islands in 1778, the Hawaiian civilisation was in a deceptive state of relative equilibrium with its environment, and was viewed as part of the ‘natural’ scene, although early visitors noted that agriculture extended inland for miles to the edge of the forest. The myth of the noble savage was alive and well. Now, even proponents of ethnic Hawaiian values (Burrows 1989) agree that the Polynesian colonisation of Hawai‘i was an ecological disaster (Olson 1989; James 1995; van Riper and Scott 2001). By the time of European contact, the Hawaiians directly or indirectly had driven over half of the islands’ native birds to extinction (Burney 1997; Lövei 2001), including the aforementioned community of flightless grazers and browsers and the predators that depended on them ( James and Olson 1983, 1991; Olson and James 1991; van Riper and Scott 2001). Nesting seabirds were driven to offshore islets. Gone were virtually all of the lowland dry forests and much of the mixed mesic forests with their associated faunas (Athens 1997). Indeed, by 1778 all of Hawai‘i below 800 m had become an entirely artificial ‘ecosystem’, if that term even applies, that supported few native land birds. As van Riper and Scott (2001: 223) point out, ‘what forests and avian habitat that were left at European contact remained only because the limited Polynesian technology prevented them from more fully exploiting their environment’.

The historic period A second (and ongoing) ecological cataclysm began with the voyages of Captains James Cook and George Vancouver. A first and obvious victim was the Hawaiian human population, which plummeted precipitously following outside contact (Stannard 1989; Juvik and Juvik 1999) as a result of diseases such as measles, syphilis, tuberculosis, and influenza, to which the Hawaiians had no immunity.This decline foreshadowed similar catastrophic declines of Hawaiian honeycreepers and other native birds (Chapter 9). So severe was the population decline that, except for the 1815–26 intensive sandalwood trade (Cuddihy and Stone 1990), native Hawaiians played a rather insignificant role in postcontact ecological deterioration, and the need for agricultural labour fuelled huge waves of immigration. These European and Asian peoples were the new agents of change. More often than not, the assault on the environment was indirect through the introduction of alien organisms (Cox 1999) that could do much more pervasive damage than people themselves.

Feral ungulates The pigs Sus scrofa that accompanied the original Hawaiians had very little impact on native ecosystems.They were small, usually kept penned, and did not penetrate far into pristine forests (Stone 1986; Tomich 1986). The huge wild pigs (Fig. 2.11) that today are ravaging the remotest rainforests (Chapter 10) are descendants of European domestic pigs that were continuously released since 1778 (Stone and Loope 1987), perhaps to breed-up the feral stock for better sport. In fact, feral pigs are rarely mentioned in the accounts of nineteenth-century naturalists, and Hawaiian informant David Malo (ca. 1793–1853) made no mention of pig hunting or wild pigs (Malo 1951) in ancient times. The great importance of pig hunting in present-day popular culture in Hawaii (see Chapter 10) appears to be largely a twentieth-century phenomenon. Indeed, Stone and Loope (1987) suggest that pigs have only become an environmental problem in Hawaiian forests in recent decades. Pigs are not only destructive in themselves, but they exacerbate

The honeycreepers’ world 25 other problems: they are the most significant factor in the spread of alien plants in native forests, both by digging up the substrate and by transporting seeds in their hair and droppings (Stone and Loope 1987; Cuddihy and Stone 1990); and their activities produce depressions and cavities that collect water in which mosquitoes breed.The impact of feral pigs on Hawaiian honeycreepers may well have been minimal in the nineteenth and early twentieth centuries, but their destructiveness in the birds’ environment is often catastrophic today (Ralph and Maxwell 1984; Stone and Loope 1987). For example, the remote forests of Kohala Mountain on Hawai‘i have lost their entire understorey to pigs ‘and all that remains are tree-fern skeletons and a quagmire of mud’ (van Riper and Scott 2001: 224). Lobelioids, upon which many honeycreepers fed (see Chapter 8), were once abundant in this forest (Rock 1913), which has also lost most of its native birds (van Riper 1973b). Recently invading feral pigs may be the main reason for the steady decline of the Po‘o-uli since its discovery in 1973 (Mountainspring et al. 1990;T. K. Pratt et al. 1997b). In earlier times, the most damaging of the introduced ungulates arguably were cattle Bos taurus and goats Capra hircus. On voyages in 1793 and 1794, Captain George Vancouver presented Kamehameha I with gifts of both and persuaded the king to place a kapu (taboo) on them for 10 years. The cattle were turned loose in the Waimea Plains of the S. Kohala Dist. (which some writers claim was originally

2.11 Feral pig in native rainforest, showing damaged substrate. Photo © Jack Jeffrey.

wooded), where they thrived, producing ‘immense herds’ within a few decades (Tomich 1986). Carried throughout the islands, cattle formed feral herds everywhere that proceeded to eat their way through the forests and even to become a nuisance for farmers (Brennan 1974).Today’s vast upland pastures were almost all originally forested, but relatively few were purposely logged (logging was a more significant factor in lowland koa forests; Cuddihy and Stone 1990). Many pastures were created by the cattle themselves as they ate every accessible sapling of every tree (Baldwin and Fagerlund 1943). In drier areas or those too rugged for cattle, goats were just as effective in destroying native ecosystems (Yocom 1967). Scott Wilson wrote: The mountains were in olden times densely clothed with tropical vegetation and trees of various kinds, and such is still the case to a considerable extent in most of the islands; though in certain districts, especially of Lana‘i and Oahu, the primeval woods have been completely destroyed by the agency of animals or man. Goats are the chief offenders in Lana‘i, deer . . . in Molokai, and cattle in Hawaii, while in the last-named the ground is being extensively cleared to make room for coffeeplantations. (Wilson and Evans 1890–99: vii–viii.) Hawaiian trees and shrubs had long evolved in an environment relatively free of mammalian browsers (but note that the moanalos probably were browsers; Olson and James 1982b, 1991). For the most part, the plants have lost the alkaloids, thorns, or spines that protect them on continents. In a few generations, the oldest trees died and grasses took over. Even if no one cut down or burned the forest as the Hawaiians had done, it disappeared just as inexorably over vast areas (Fig. 2.12). Mid-nineteenth-century observers noted changes in local weather as a result of destruction of forests. Erosion became widespread and dust storms common. A capsule glimpse of forests in the midst of change as a result of feral cattle is provided by the noted ornithologist R. C. L. Perkins (1893), reporting on his early collecting in Kona:

26 The Hawaiian Honeycreepers From about 1700 to more than 3000 feet the forest has a distinct character. It consists mainly of the lehua-tree, which here is of an enormous size and height, the koa, and several other trees of smaller size. On the rough lava-flows these trees are thickest and most luxuriant; in the spaces between the flows they have largely been destroyed by the cattle, and these spaces, which are up to about 3000 feet waist-deep in the hilo grass, more resemble English parkland than thick woods. Tree-ferns, which were once very numerous, are now much scattered in these spaces, but are more plentiful on the lava . . . Both on the rough flows and in the more open country the Freycinetia (ieie) climbs high up the trees, but except on the roughest lava it shows no leaves or flowers for a considerable height, the cattle eating off the shoots as high as they are able to reach. Even in the now open grass-covered land not many years ago the trees were all united by the ieie vines, while the tree-ferns were in profusion as low as 1400 feet. Perkins’s species list indicates that the most damaged region he describes was a temperate mixed forest, a habitat nearly non-existent today and whose original avifauna can only be surmised. Modern readers may be surprised to learn that some cherished and supposedly ‘natural’ landscapes, such as the exposed red cliffs of Waimea Canyon on Kaua‘i (Fig. 2.5c), are the result of goat-generated erosion (Street 1989), and though smaller than cattle, goats are no less damaging to forests (Spatz and Mueller-Dombois 1973). By the twentieth century, the damage from feral ungulates was so severe that measures had to be taken to save what little forest was left. Judd (1927) referred to the years 1815–1921 as the ‘cattle period’ in Hawaiian 2.12 Progressive forest degradation from grazing: (a) healthy koa forest in a kipuka surrounded by pastures; (b) grazed koa forest with understorey destroyed; (c) open spaces beginning to develop as forest converts to pasture; (d) only ancient forest giants remaining; (e) a lone survivor in a pasture now being invaded by alien weeds such as gorse. Photos (b) and (c) © Jack Jeffrey.

The honeycreepers’ world 27 forests. Feral cattle were eventually eliminated or severely reduced, so that they were no longer a significant factor in the watersheds after the 1930s (Tomich 1986), but the damage had been done. Culliney (1988) gives a more detailed history of this period. Goats are still a problem in places, but they have been eliminated along with feral pigs in most parts of Hawai‘i’s two national parks with the aid of massive fencing projects (Stone and Loope 1987; Katahira et al. 1993) and vegetation is beginning to recover (Scowcroft and Hobdy 1986; see also Chapter 10). The ravages of other ungulates were more localised. On Moloka‘i, axis deer Axis axis, descended from a herd given to Kamehameha V in 1868, contributed greatly to forest degradation and by the turn of the twentieth century had to be controlled by organised hunts (Tomich 1986). Perkins (1903) reported the loss of two-thirds of the trees in a single Moloka‘i tract between 1896 and 1902 as a result of grazing and browsing by feral cattle and axis deer. Axis deer were transferred at various times to O‘ahu, Lana‘i, and Maui, but have been much more closely managed than the initial herd on Moloka‘i. Nevertheless, they are now becoming a threat to upland rainforests in such places as Waikamoi Preserve (TNCH) adjacent to Haleakala NP (Tummons 1999c; C. Gentz, pers. comm.). Sheep, both feral Ovis aries and mouflon O. musimon introduced for hunting purposes, had their greatest impact in the mamane–naio forests of Mauna Kea (Warner 1960; Scowcroft 1983; Scowcroft and Giffin 1983; Scowcroft and Sakai 1983). They were first brought to Hawai‘i by Captain James Colnett in 1791 (Wyllie 1850), and Vancouver brought more on his voyages. They existed for many years as managed herds, but the sheep industry essentially ended in 1963 (Brennan 1974). Unchecked, their impact would have doomed many endemic plants along with the Palila ( Juvik and Juvik 1984; Scott et al. 1984), which has the distinction of being the only Hawaiian honeycreeper to have been the successful plaintiff in a precedent-setting lawsuit (Palila vs. Hawaii Department of Natural Resources, Ninth Circuit Court of Appeals; Bean and Rowland 1997) upheld on appeal in 1985 (Turner 1988; Scott and Conant

2001). Maintenance of feral sheep, both domestic and mouflon, for hunting on Mauna Kea was found to violate Section 9 of the Endangered Species Act. Despite the efforts of a powerful and wellorganised, but numerically small, group of hunting advocates and their allies in the state government, sheep of both species are now being removed from the wild, albeit grudgingly (Tummons 1999b, d) and two decades later than they should have been, under the Federal court order. At least some progress is being made (Stone 1985a; Scott and Conant 2001).

Rats and other mammals According to Atkinson (1977), large rats, in particular the black or roof rat Rattus rattus, may have played a major role in the declines of many species of Hawaiian honeycreeper.This hypothesis will be discussed further in Chapter 9. Other rodents that reached the islands as stowaways include Norway rats R. norvegicus and house mice Mus musculus. Norman (1975) believed the role of rodents as predators was exaggerated, but their effect on birds at Midway Atoll was dramatic (Fisher and Baldwin 1946a). Small Indian mongooses Herpestes auropunctatus, first introduced by sugar planters in the 1880s and now established on all main islands except Kaua‘i and Lana‘i, apparently have not been implicated in any direct negative impact on Hawaiian honeycreepers (Pratt 1994a). They are certainly detrimental to Nene and other groundnesting birds (Tomich 1986), and have recently been discovered to be a significant factor in survival of fledgling Hawaiian Crows ( Johnston and Banko 1992). Mongooses do not readily climb trees or swim, but are certainly capable of both (Zimmer 1980; pers. obs.). Feral cats have been in Hawai‘i since 1778, and are known to feed on Hawaiian honeycreepers (Perkins 1903: 394; Snetsinger et al. 1994), but few data have been gathered on their full impact. Notably, cats were abundant and attacking native birds on Lana‘i in the 1890s (Rothschild 1893–1900; Perkins 1903) two decades before the major declines of honeycreepers there (Munro 1960). Like mongooses, cats are much more dangerous to ground-dwelling and ground-nesting birds than to most Hawaiian

28 The Hawaiian Honeycreepers honeycreepers, but may be a serious predator on Palila (T. K. Pratt et al. 1997a).

Alien birds As a result of attempted introductions of hundreds of species of alien birds to the Hawaiian Islands, today’s avifauna has more non-native than native species (Pratt 1994a), the largest proportion of introduced species in any modern avifauna. When we consider that the vicinity of Honolulu had become an almost birdless zone, we can better understand the sentiments expressed by an anonymous writer in The Polynesian for 22 October 1853 (quoted in Caum 1933): The introduction of new animals, plants &c., by residents returning from their visits to other countries, is highly praise-worthy and commendable; and we hope persons visiting foreign countries and returning, will bear in mind that they may thus contribute to the general good of the islands, by the introduction of many things that will contribute to the pleasure or profit of themselves or the residents generally. And this from the Commercial Advertiser for 16 August 1860 (same source): ‘Owners of vessels leaving foreign ports for Honolulu, will confer a great favor by sending out birds, when it can be done without great expense. We need more songsters here.’ Indeed, from 1850 to 1867 the Royal Hawaiian Agricultural Society made great efforts to import foreign birds, but detailed records were either lost or never kept (Caum 1933).As the native birds continued their retreat, renewed interest in importing alien species led ultimately to the formation in 1930 of the Hui Manu (⫽ Bird Club, not to be confused with modern clubs of birders), a society dedicated to the introduction of foreign birds to Hawai‘i. By that time, importation of birds was no longer quite so ‘commendable’, but even those with contrary inclinations were given to rationalisation. After condemning the general practice of introducing alien birds, Caum (1933) nevertheless thought that some carefully selected species could be introduced without harm to natives, which by his time were virtually entirely absent from the lowlands.

The Hui Manu persisted in its activities, with the full co-operation and participation of territory and state authorities, until 1962, when their last introduction (Mariana Swiftlet) occurred. During the same period, the Hawaii government pursued a program of introduction of game birds for hunting purposes.As reviewed by Berger (1981), enthusiasm for introductions continued into the 1970s, and resulted in the total transformation of Hawai‘i’s avifauna (Pratt 1994a).A continuing legacy of the misguided government and private policies is that Hawai‘i, unique among the 50 United States, provides the same legal protection for introduced birds as endemic ones, greatly complicating any effort to control the aliens. Admittedly, most introduced species have filled niches and life zones, e.g. lowlands and coastal areas, that had been vacated by native birds (Mountainspring and Scott 1985; Ralph 1990b; Shehata et al. 2001). Because the current ranges of introduced birds and those of surviving Hawaiian honeycreepers are often mutually exclusive, the impact of competition with alien birds is thought to be relatively minor (Moulton and Pimm 1983; van Riper and Scott 2001). Nevertheless, a few species, e.g. Japanese White-eye, Red-billed Leiothrix, Japanese Bush-Warbler, Northern Cardinal, Kalij Pheasant, Red-vented Bulbul, and Hwamei (Melodious Laughing-thrush), have penetrated into the deepest rainforests (Scott et al. 1986; pers. obs.).The Kalij Pheasant probably plays a negative role in carrying seeds of invasive plants into native habitats. Recent studies (Cann and Douglas 1999; Shehata et al. 2001) have revealed that the introduced birds vary widely in their ability to serve as disease reservoirs, with some surprising culprits. On O‘ahu, the species with the highest prevalence of avian malaria (see Chapter 9) are the bulbuls, shama, and, surprisingly, Nutmeg Mannikin and Spotted Dove. The muchmaligned and ubiquitous white-eye may not be as important a disease reservoir as many have believed. One major honeycreeper habitat, mamane–naio forest, has been invaded by more alien birds, including both passerines and game birds, than any other. Table 2.3 (amplified from Pratt 1994a) lists the alien species that have been added to Hawai‘i’s avifauna.

The honeycreepers’ world 29 Table 2.3 Alien birds naturalised in the Hawaiian islands with their distributionsa. Year e Family Ardeidae Cattle Egret Family Anatidae Mallard Family Phasianidae Chukar Grey Francolin Black Francolin Erckel’s Francolin Japanese Quail Red Junglefowl Kalij Pheasant Common Pheasant Green Pheasant Common Peafowl Wild Turkey Family Odontophoridae California Quail Gambel’s Quail Family Columbidae Rock Pigeon Spotted Dove Zebra Dove Mourning Dove Family Psittacidae Rose-ringed Parakeet Mitred Parakeet Red-crowned Parrot Family Tytonidae Barn Owl Family Apodidae Mariana Swiftlet Family Alaudidae Sky Lark Family Paridae Varied Tit Family Pycnonotidae Red-vented Bulbul Red-whiskered Bulbul Family Sylviidae Japanese Bush-Warbler Family Turdidae White-rumped Shama

Mi b

K

O

Mo

Ln

Ma

H

Bubulcus ibis

1959

√

√

√

√

√

Anas platyrhynchos

?

√c

√

?

√

√

√ √ √ ? ? ⫻

√

1923 1958 1959 1957 1921 ⬎1788 1962 1865 ⬎1900 1860 1788

√ √ √ √ √ √

Callipepla californica C. gambelii

1865 1928

?

Columba livia Streptopelia chinensis Geopelia striata Zenaida macroura

⬎1893 1865 1922 1964

Alectoris chukar Francolinus pondicerianus F. francolinus F. erckelii Coturnix japonicus Gallus gallus Lophura leucomelanos Phasianus colchicus P. versicolor Pavo cristatus Meleagris gallopavo

√

√ √ ⫻ √ ? √

√ √ √ √ √ ⫻

√ √

√

√ ?

√ √

√

?

√ √ √ √

√ √ √

√

Psittacula krameri Aratinga mitrata Amazona viridigenalis

? ? ?

√

Tyto alba

1958

√

Aerodramus bartschi

1962

Alauda arvensis

1865

?

√

Parus varius

1890

⫻

⫻

Pycnonotus cafer P. jocosus

1965 1966

Cettia diphone

1929

√

Copsychus malabaricus

1931

√

√ ? √ √

√ √ √ ⫻ √ √ √ √ √

?

√ √

?

√ √

√ √ √

√ √ √

√ √ √ √

√ √ √ √

√ √

√ √d

? √

√

√

√

√

√

√

√

√

√

√

√

√

√

√

√d

√

√ √

30 The Hawaiian Honeycreepers Table 2.3 contd. Year e Family Timaliidae Greater Necklaced Laughing-thrush Grey-sided Laughing-thrush Hwamei Red-billed Leiothrix Family Zosteropidae Japanese White-eye Family Mimidae Northern Mockingbird Family Sturnidae Common Myna Family Emberizidae Yellow-faced Grassquit Saffron Finch Red-crested Cardinal Yellow-billed Cardinal Family Cardinalidae Northern Cardinal Family Icteridae Western Meadowlark Family Fringillidae House Finch Yellow-fronted Canary Island Canary Family Passeridae House Sparrow Family Estrildidae Red-cheeked Cordonbleu Lavender Waxbill Orange-cheeked Waxbill Black-rumped Waxbill Common Waxbill Red Avadavat African Silverbill Nutmeg Mannikin Chestnut Munia Java Sparrow aAmplified

Mi b

K

O

Mo

Ln

Ma

H

√ √

√ √

Garrulax pectoralis

1919

√

G. caerulatus G. canorus Leiothrix lutea

1947 1900 1918

√ ⫻

? √ √

√ √

Zosterops japonicus

1929

√

√

√

√

√

√

Mimus polyglottos

1928

√

√

√

√

√

√

Acridotheres tristis

1865

√

√

√

√

√

√

√

√ √ √

√

√

√

Tiaris olivaceus Sicalis flaveola Paroaria coronata P. capitata

√

1970 ⬎1960 1928 1930

√

Richmondena cardinalis

1929

√

Sturnella neglecta

1931

√

1859 ⬍1965 1910

Carpodacus mexicanus Serinus mozambicus S. canaria Passer domesticus Uraeginthus bengalus Estrilda caerulescens E. melpoda E. troglodytes E. astrild Amandava amandava Lonchura cantans L. punctulata L. atricapilla Padda oryzivora

1871 ⬍1965 ⬍1965 ⬍1965 ⬍1965 ⬍1965 1900 ⬎1960 1865 1936 ⬍1965

√

√

√

√

√

√

√

√ √

√

√

√

√ √

√

√

√

√

√

√

√

⫻ ⫻ √

√ √ √ √ √

√ √ √ √ √ √

√

√ √ ? ?

√ √

√ √ √ √

from Pratt 1994a. abbreviations as in Table 2.2, except Mi ⫽ Midway Atoll. c√ ⫽ naturalised (AOU 1998), ⫻ ⫽ established for a time, now extirpated, ? ⫽ status unknown or questionable. dPratt 2002b. eYear first introduced; italics indicate approximations. bIsland

√ √ ⫻ √ √ √ √ √

The honeycreepers’ world 31 Alien invertebrates As might be expected, insects and other invertebrates form a large percentage of post-contact introduced organisms in Hawai‘i. Some, such as honeybees, were introduced purposely but most were inadvertent hitchhikers.Whether competition with bees for nectar has affected populations of drepanidine nectarivores has not been studied. Howarth (1985) reviewed the effects of these foreigners on native ecosystems. In most cases, direct impacts on honeycreepers have been more hypothesised than demonstrated, but at least some researchers (Banko and Banko 1976; Banko et al. 2001) believe depletion of native invertebrate food resources by alien predators and parasites has had a negative effect. Likewise, whether predatory alien snails have impacted food resources of the severely endangered Po‘o-uli, a snail-eater, has not been determined. But the overarching impact of the night-biting mosquito Culex quinquefasciatus is now well documented (Warner 1968; van Riper et al. 1986; Cann and Douglas 1999; van Riper and Scott 2001).This vector of avian malaria and avian pox is probably the single most devastating alien organism for native birds, especially the drepanidines, and will be discussed in detail in Chapter 9.

Agriculture and forestry At the time of European contact, the Hawaiian lowlands were given over to small farms growing subsistence food and fibre crops. That pattern persisted until The Great Mahele of 1848, when private ownership of land was widely established (Tabrah 1980). Prior to that time, the concept of real estate was unknown in Hawai‘i, all land being held in trust by the ali‘i. Commoners had always been tenants, and did not benefit from the mahele, but now the nobles could sell their small holdings to larger conglomerates, facilitating the formation of huge ranches and commercial plantations. By the end of the century, sugar had become king in Hawaii, and the fields extended right up to the lower edge of the remaining native forests (Cuddihy and Stone 1990). Those few lowland forests that survived the Hawaiians and feral cattle were cut down to make way for cane or to fire the mill boilers ( Judd 1927). In an odd twist, this conversion

to sugar monoculture actually may have been a good thing for some Hawaiian honeycreepers. Sugar production requires huge quantities of water, and the planters realised early on that their watersheds had to be protected.At the same time, the loss of forests above Honolulu was seen as causing major erosion, siltation, and failure of wells (Cuddihy and Stone 1990). The result of this concern was the establishment of a territorial (now state) Forest Reserve System. Major programs were instituted to remove feral animals, fence the reserves, and plant trees in deforested areas. All of these meant that at least a few threats to remaining honeycreeper habitat were reduced. Unfortunately, the tree-planting programs rarely included native species. Huge tracts were planted in eucalypts such as blue gum, swamp mahogany, and paperbark. Early reforestation efforts favoured fast-growing tropical species that had no potential timber value (to discourage future logging of the watershed), many of which have now become invasive weeds in remaining native forests (Cuddihy and Stone 1990; Pratt 1999a). Prominent among these invasives are firetree, Brazilian pepper (known locally as Christmasberry), black acacia, and strawberry guava (Vitousek 1992). The latter two have been particularly invasive following 1992 Hurricane Iniki on Kaua‘i, and have prevented regeneration of extensive tracts of former ‘ohi‘a/koa forest in the Koke‘e area (pers. obs.). Although most honeycreepers eschew these foreign trees, the nectarivores readily feed on their flowers, and a few alien plantations, e.g. Polipoli Springs and Hosmer Grove on Maui (Pratt 1995, 2002b), support populations of up to four drepanidines including some insectivores. For better or worse, alien trees are now a permanent part of the honeycreepers’ world.

Weeds In addition to the invasive species planted for forestry, many plants have become naturalised in the islands inadvertently, often as escapes from ornamental plantings (Cox 1999). Many of these, though attractive, are now threatening even remote montane rainforests. The major problem species currently include kahili ginger, blackberry, Koster’s curse, and banana poka, with others, such as the

32 The Hawaiian Honeycreepers ‘green cancer’ Miconia calvescens just gaining a foothold (see Chapter 10). Banana poka (Fig. 8.12), in addition to its obvious effect in smothering native trees, has been implicated in a particularly insidious perturbation of natural processes. Its flowers attract large numbers of ‘I‘iwi,‘Apapane, and ‘amakihis and, because its corollas are not the ‘right’ shape, may ‘teach’ them a technique of nectar-stealing that they then use on native flowers (Conant et al. 1998; see Chapter 8 for details), thereby avoiding their role as pollinators.

Clearly, the Hawaiian honeycreepers that survive today inhabit a world very different from the one in which they evolved. The many species now extinct were not able to cope with the changes, but a few have been able to adapt to new plants (Chapter 8), new prey items, and even new diseases (Chapter 11). Unfortunately, today’s honeycreepers live in a dynamic world that will continue to change rapidly and inexorably. Undoubtedly, some of these future changes will overwhelm the adaptive ability of some species.

3 Discovery and research: historical perspectives

The century of discovery Cook and other British explorers When naturalists with Captain James Cook’s third voyage came ashore at Waimea, Kaua‘i in January 1778, they bartered with local villagers for a bundle of bright red birds. We now know that they were ‘I‘iwi, the first Hawaiian honeycreeper to be seen by scientists or non-Hawaiians. Later, Cook spent more time at Kealakekua Bay on the Kona Coast of Hawai‘i, where he collected more ‘I‘iwi along with specimens of Lesser ‘Akialoa, Hawai‘i Mamo, ‘Apapane, Hawai‘i ‘Amakihi, ‘Akepa, and ‘O‘u (Stresemann 1950; Medway 1981; Olson 1989b). These birds were taken back to Europe and described and named, but little was known about them. Over the ensuing century, other species were described by various authors based on specimens similarly collected during brief visits or by island residents who gathered little information about the birds. Relatively few large expeditions visited the islands, and new species came to light willy-nilly.The following is a brief overview of discovery and description of honeycreeper species. For more detailed accounts, see Newton (in Wilson and Evans 1890–99: xi– xix), Kay (1972a), and Olson and James (1994a). After Cook, the next important expedition to Hawai‘i was the voyage of HMS Blonde in 1825 under the command of George Anson seventh Lord Byron, successor to the famous poet. The main purpose of the voyage was to return the bodies of King Kamehameha III and his queen to their native land after both had died of measles shortly after arriving in England on a mission to learn about European legal systems.The ship’s naturalist,

Andrew Bloxam, made a number of important discoveries and recorded valuable natural history information, but the specimens were dispersed and much of his work remained unpublished or unappreciated until Olson (1996b) brought much of it together for the first time. One of the more interesting observations of Bloxam was that the Hawaiian nectarivores were not, as had been previously supposed, related to the creepers of the genus Certhia, and were instead rather finchlike. Thus he may have been the first to hint at the existence of a unique taxon of Hawaiian honeycreepers.Although he also briefly visited Hawai‘i, Bloxam’s specimens and notes on drepanidines all came from O‘ahu, and were the first from that island. In addition to several widespread species already known, he found the first O‘ahu ‘Amakihi and O‘ahu ‘Akepa.

American visitors Americans John Kirk Townsend and Thomas Nuttall visited O‘ahu in January 1835, and Townsend and Ferdinand Deppe visited O‘ahu, Kaua‘i, and Hawai‘i in January 1837 (Townsend 1839). It is solely from their specimens that we know of the historical existence of the O‘ahu ‘Akialoa and O‘ahu Nukupu‘u, and Deppe collected the first O‘ahu ‘Alauahio (Wilson and Evans 1890–99). From September 1840 to April 1841, ships of the United States Exploring Expedition (Stanton 1975) were in the Hawaiian Islands, although the expedition’s naturalist,Titian Peale, was present for only a short time. Few bird specimens from that expedition survive, and no new honeycreepers were discovered (Olson and James 1994).

34 The Hawaiian Honeycreepers

3.1 Cotype of Ciridops anna, from the Mills Collection, now at BPBM.

Local naturalists For several decades, Hawaiian ornithology languished save for the efforts of a few resident amateur collectors. In 1851, J. D. Mills emigrated from England and settled in Hilo, where he became a merchant (Manning 1979; Olson 1994). Over the next 36 years he collected birds on the island, including specimens of ‘Ula-‘ai-hawane (Fig. 3.1) from which Sanford B. Dole (see species account for ‘Akohekohe for biographical notes), also an amateur ornithologist, later named the species (Dole 1878b). Théodore Ballieu, the French Consul at Honolulu from 1869 to 1878, collected specimens on Hawai‘i that included the first of the Palila. Norwegian immigrant Valdemar Knudsen settled on Kaua‘i in 1857 (Knudsen and Noble 1999) and began sending bird specimens to the Smithsonian Institution as early as 1866 (Olson and James 1994a). Based on his collections, Stejneger (1887a) described two new endemic Kaua‘i honeycreepers, the ‘Anianiau and the ‘Akikiki.

The classic research period, 1887–1919 The Newton and Rothschild expeditions The pace of ornithological research in Hawai‘i changed completely with the arrival of Scott B. Wilson from Great Britain in April 1887. Under

the sponsorship of Alfred Newton, he remained in the islands until the end of 1888. Wilson was the first ornithologist to visit Hawai‘i for the sole purpose of making a thorough collection of native birds. He added many new species and gathered some of the first natural history information ever published on Hawaiian birds, presented along with colour engravings by F. W. Frohawk in the classic Aves Hawaiiensis: the birds of the Sandwich Islands (Wilson and Evans 1890–99), the colour plates from which were recently reprinted (Wilson and Buff 1989). Wilson’s work came at a time of intensive, at times bitter and acrimonious, competition among British ornithologists, and Hawaiian ornithology was one beneficiary. Having heard of Wilson’s discoveries, Newton’s rival Walter Rothschild sent Henry Palmer to Hawai‘i to collect birds from December 1890 to August 1893. Palmer was assisted by George C. Munro, a young New Zealander who later settled in Hawai‘i and eventually became the patriarch of Hawaiian natural history (Amadon 1964; Mearns and Mearns 1992), and by E. Wolstenholme, whose history is somewhat obscure (Olson and James 1994a). Palmer was an extremely thorough and energetic collector, who was accused by Newton of ‘pillaging the avifauna’ (Olson and James 1994a). His work was summarised in a massive tome oddly titled The avifauna of Laysan and the neighboring islands with a complete history to date of the birds of the Hawaiian possessions (Rothschild 1893–1900), illustrated with beautiful colour lithographs by Frohawk and J. G. Keulemans that are today valuable collector’s items, the Hawaiian equivalent of Audubon prints. These and the previously mentioned Frohawk plates are the only contemporary colour illustrations of many now extinct Hawaiian honeycreepers.

R. C. L. Perkins Although Wilson visited the islands from time to time after 1888, he made no new discoveries and collected few specimens. To respond to Rothschild’s challenge, Newton shepherded the formation of a Joint Committee for the Zoology of the Sandwich Islands, involving the Royal Society and

Discovery and research: historical perspectives 35 the British Association for the Advancement of Science. Because of delays over how the work was to be funded, it was not until 1892 that they sent R. C. L. Perkins to Hawai‘i. As a result, Perkins was in several instances ‘scooped’ by Palmer, leading Wilson to comment:‘The loss of the season of 1891 was unfortunate for the credit of the Joint Committee; for many discoveries which its collector, had one been sent out in that year, could not have failed making fell to the lot of the persons employed by Mr. Rothschild’ (Wilson and Evans 1890–99: xx). Perkins made two long trips to Hawaii, the latter financed in part by the Trustees of the Bernice Pauahi Bishop Museum in Honolulu. Perkins was both an ornithologist and an entomologist, and although he discovered only one new species (Black Mamo), his contributions to our knowledge of Hawaiian birds (Perkins 1893, 1895, 1901, 1903) are monumental. He was a keen and detailed observer whose writings are still useful today. Even a century later, anyone who wishes to study Hawaiian natural history effectively must read Perkins. Manning (1986) gives a detailed account of Perkins’s Hawaiian research. In their competitive rush,Wilson, Palmer, and Perkins named 21 species (as herein delineated) of Hawaiian honeycreepers, more than doubling the known total in less than a decade.At the close of the nineteenth century, only three living species still awaited discovery.

Henry W. Henshaw In 1894, the American naturalist Henry W. Henshaw moved to Hilo for health reasons (Nelson 1932). By 1900, he had regained enough strength to return to field work and began amassing a major collection of Hawaiian birds. Although he found no new species, his collection of skilfully prepared skins, mainly from windward Hawaii, proved extremely valuable. His specimens were eventually dispersed among several institutions, but approximately half of them went to BPBM. He was one of the last to observe several species of honeycreeper that have since gone extinct, and was clearly alarmed and puzzled by the ongoing population crashes of native birds (see Chapter 9). Henshaw had an interest in avian biology as well as simple collecting, and he urged others

to gather such information while such efforts were still possible. He published a summary of his work (Henshaw 1902) that is exceeded only by the work of Perkins (1903) in the depth of its natural history observations.

The Bernice Pauahi Bishop Museum The staff of the BPBM, founded in 1889 by Charles Read Bishop in memory of his late wife, a Hawaiian princess (Fig. 3.5), was very active in ornithology around the turn of the twentieth century. The first director, William T. Brigham, had been an amateur bird collector himself (Olson 1992). William Alanson Bryan’s (1901a) Key to the birds of the Hawaiian Group was a major contribution that brought together much previously scattered information. Bryan and Alvin Seale followed Henshaw’s admonition and were active in finding and describing nests and eggs of native birds (Bryan and Seale 1900; Bryan 1905a,b,c,d), and in gathering other natural history information, particularly on O‘ahu (Seale 1900; Bryan 1901b), which had been neglected by the British collectors. Bryan (1908) also made the last observations on Moloka‘i when it still had populations of most of its historically known birds.

Exploration of the Northwestern Hawaiian Islands After Palmer first reported the distinctive avifauna of Laysan, a number of ornithologists became interested. Schauinsland (1899) spent 3 months on the island in 1896.The steamer Albatross of the US Fish Commission called there 16–23 May 1902, leading to two reports by Walter K. Fisher (1903, 1906). W. A. Bryan also visited during that year, and returned 9 years later (Bryan 1912a). Based on the deterioration of the habitat by 1911, he made dire predictions about the fate of the island unless certain measures were taken. None were, and his predictions largely came true (see Chapters 2 and 10). Because of the difficulty of landing there, Nihoa had been bypassed by the expeditions that visited Laysan. Bryan, however, suspected that a land bird might be present and persuaded the captain of the

36 The Hawaiian Honeycreepers US revenue cutter Thetis to look for one on a 1915 inspection tour. The captain did find a bird, and even secured specimens but they were lost in a storm (Bryan 1916). The following year the ship had better luck, and brought back the type specimens of the Nihoa Finch (Bryan 1917), which Bryan named Telespiza ultima because he thought (wrongly as it turned out) it would be the last Hawaiian native bird discovered. Not only was another bird, the Nihoa Millerbird, subsequently discovered on that island by the 1923 Tanager Expedition (Wetmore 1925; Olson 1996a), but two more historical Hawaiian honeycreepers were eventually to come to light, not to mention those identified from prehistoric bones.

George C. Munro on Lana‘i One last footnote to this period in the history of Hawaiian honeycreepers occurred when Munro collected an unusual bird on Lana‘i in 1913. After the Rothschild expeditions were completed, Munro had taken a succession of jobs as a cattle ranch foreman, including a 6-year sojourn on Lana‘i before it became a pineapple plantation (Mearns and Mearns 1992). Munro was convinced, because he saw others in subsequent years, that his odd bird was a new species and sent it to Perkins (1919), who named it a new genus and species Dysmorodrepanis munroi. But hardly anyone noticed! By that time, ornithological research in Hawai‘i had virtually ceased and was to remain dormant for a quarter century. To add insult to injury, Greenway (1939) dismissed Munro’s bird as a freak, and Amadon (1950) concurred. Finally, after a thorough examination of the single specimen (Fig. 3.2), James et al. (1989) concluded that the Lana‘i Hookbill was, after all, a valid species of drepanidine finch endemic to Lana‘i (and Maui-nui).

The quiescent years The ‘quiet period’ of drepanidine research was indeed unfortunate because we will never know the exact pattern, timing, or nature of the population declines that must have occurred over those years. Birds that were still common when reported

3.2 Unique type of Dysmorodrepanis munroi compared with a female specimen of Psittirostra psittacea, BPBM.

in the early years of the century were often not heard from again for decades, by which time many had become extremely rare (Richards and Baldwin 1953). Others simply disappeared without a trace. It is now obvious that they were not reported because no one was looking for them.

Reawakening and rediscovery Munro and the renaissance Things began to change when Munro, assisted by Walter Donaghho and others, conducted the Hawaiian Bird Survey from 1935 to 1937. The survey covered all of the forested islands, and confirmed that at least two of the long-lost honeycreepers, the Palila and the ‘O’u, still survived (Munro 1944). In retrospect, Munro’s surveys must be seen as rather superficial, because he missed many species that still survive today, but at very least they started the ornithological ball rolling again in Hawai‘i. In 1939, the Hawaii Audubon Society was founded. Its journal, ‘Elepaio, though always rather unassuming in appearance, was to become a major vehicle for the dissemination of knowledge about Hawaiian birds. In 1940, an expedition from the Philadelphia Academy of Sciences visited Nihoa (Vanderbilt and Meyer de Schauensee 1941), and found that its birds had fared much better than those of Laysan in the years since the 1923 Tanager expedition. In 1944, Munro published the first

Discovery and research: historical perspectives 37 edition of Birds of Hawaii, which included the results of the bird surveys and much previously unpublished information about the Rothschild expeditions of the 1890s. It remains a classic today and is still in print in a second edition (Munro 1960) published shortly before Munro’s death at the age of 97 in 1963 (Amadon 1964). Also in 1944, Bryan and Greenway published the first taxonomic revision of Hawaiian birds since the development of the ‘modern synthesis’ of evolutionary theory (Mayr 1942). Their list set new generic and species limits for the Hawaiian honeycreepers, but could not be regarded as a thorough revision.

Dean Amadon The 1941 attack on Pearl Harbor focused national attention on Hawai‘i, and the Second World War brought many mainlanders to the islands who might never otherwise have been there. Among them was Dean Amadon, who had already developed an interest in Hawaiian honeycreepers while curating the Rothschild collection, recently acquired by the American Museum of Natural History (AMNH) in New York. From April 1944 to August 1945, Amadon was stationed with the US Army in Hawai‘i, mostly on O‘ahu but including 2 months on Hawai‘i. When not involved in military duties, he was able to study the collections of the BPBM (Amadon 1944b) and do some field work (Amadon 1945). After the war, Amadon returned to Cornell University, where he produced a doctoral dissertation (Amadon 1950) that, when published, became one of the landmarks of drepanidine research. The Hawaiian honeycreepers (Aves, Drepaniidae) (Amadon 1950) was the first complete revision of the group since Perkins (1903), and brought their taxonomy into line with contemporary thinking that favoured large polytypic species and broadly defined genera; it was the basis of honeycreeper classification for the next three decades.

Rediscovery Amadon had been assisted on The Big Island by Paul H. Baldwin, a naturalist with the National Park Service at Hawaii (now Hawaii Volcanoes) NP

(Baldwin 1944). Baldwin likewise was conducting research on Hawaiian honeycreepers, but his focus was on their life history and ecology rather than taxonomy. His pioneering dissertation (Baldwin 1953), focusing on three of the more common species (‘Apapane, ‘I‘iwi, and Hawai‘i ‘Amakihi), was the first modern biological study of any Hawaiian honeycreeper. Baldwin, along with Lawrence P. Richards, was also active in looking for the ‘lost’ honeycreepers of Maui and Hawai‘i that had so long been unreported.Their field work confirmed the existence of the ‘O’u and the Palila on Hawai‘i, as previously reported by Munro (1944a). On Maui they ‘rediscovered’ two supposedly extinct species, the ‘Akohekohe, last reported on Moloka‘i in 1907, and the Maui Parrotbill, which had not been seen since the 1890s (Richards and Baldwin 1953).They also published a second-hand report of the Maui ‘Akepa, not seen since 1894. However, their searches of the type localities of Greater ‘Amakihi, Kona Grosbeak, and the koa-finches were unsuccessful. Moloka‘i had not been visited by ornithologists since 1907 (Bryan 1908), when most of its birds (including all of its known honeycreepers) were still present. Munro’s surveys found only the ‘Apapane (Munro 1944a), and Manning Richards (1946) may have seen Hawai‘i ‘Amakihi there. Frank Richardson (1949), assisted by David W. Woodside, made a more thorough search in 1948, visiting parts of the island missed by Munro, but was unable to find any but these two previously reported honeycreepers. Thus it was exciting when Noah Pekelo, the District Fish and Game Warden, reported 1961 to 1963 sightings of the Kakawahie, then considered extinct (Peterson 1961). But the most spectacular rediscoveries were to occur on Kaua‘i. In 1960, Richardson and John Bowles, with assistance again from the intrepid Woodside, made an island-wide survey of birds on Kaua‘i.They penetrated deep into the swampy and forbidding Alaka‘i Plateau, and made three separate week-long hiking trips there. As a result, they made the astounding discovery that all of Kaua‘i’s historically known birds still survived, including the drepanidine Kaua‘i ‘Akialoa and Kaua‘i Nukupu‘u, as well as the ‘O‘ o‘a‘a and Kaua‘i’s two

38 The Hawaiian Honeycreepers solitaires, the Kama‘o and Puaiohi, all of which had been thought possibly extinct (Richardson and Bowles 1964).A highly fortuitous coincidence was that these discoveries came just in time to be included in Roger Tory Peterson’s (1961) new edition of A field guide to western birds. His colour plate and the news of the fantastic discoveries of ‘extinct’ birds probably did more than anything else to bring Hawaiian birds to the attention of mainland birders and ornithologists. In 1964, Huber (1966) saw an ‘akialoa and an ‘O’u in the same ‘ohi‘a tree, close to where Richardson and Bowles had seen them. In the same area in 1968, Richard Gauthey, Ian Atkinson, and Colin Huddleston found all of the rarities except the ‘akialoa (Gauthey et al. 1968). Fortunately, the party carried an early cassette tape recorder with which they obtained what turned out to be the only recordings ever made of the song of the ‘O’u as well as other valuable recordings (Pratt 1996b). In 1969, Phillip L. Bruner (pers. comm.) visited a part of the Alaka‘i not visited by Richardson and Bowles, where he saw a Kaua‘i ‘Akialoa, the last ever reported. During the late 1960s and early 1970s, the Alaka‘i birds were monitored by John L. Sincock of the US Fish and Wildlife Service. In 1975, Sheila Conant, Robert Shallenberger, and I spent a week in one of Sincock’s study areas and made some of the last observations of several species, but, like Sincock, failed to find the ‘akialoa (Conant et al. 1998). Back on Maui, the rediscovery period had a few more surprises, including a really big one. In August 1967, The Nature Conservancy sponsored an expedition to remote Kipahulu Valley (Fig. 3.3), which the organisation was interested in adding to Haleakala NP (Matthiessen 1970). Among the party was Winston Banko, an employee of what was then called the US Bureau of Sport Fisheries and Wildlife.While separated from other members of the party, Banko (1968) reported seeing three Maui Nukupu‘u, a bird widely regarded as extinct (Greenway 1968) that had not been seen since 1896. Subsequent to the expedition, National Park Ranger George Morrison reported another nukupu‘u in Kipahulu Valley (Banko 1968).

3.3 Remote Kipahulu Valley, where the Maui Nukupu‘u was reported in 1967, for the first time in the twentieth century.

The Hana Rainforest Project No doubt encouraged by these and other recent rediscoveries, a group of undergraduate students at the University of Hawai‘i organised the 1973 Hana Rainforest Project with a grant from the National Science Foundation under the Student Originated Studies Program. The team, representing a wide variety of disciplines, set out to study the ecology of the rainforest on the north-eastern slope of Haleakala (essentially the opposite slope from Kipahulu Valley). These inexperienced but dedicated observers surprised everyone when they reported sightings of a previously unknown bird. David Woodside, then with the Hawai‘i State Division of Forestry and Wildlife (DOFAW), collected two specimens, and Dean Amadon and Walter J. Bock assisted the team of Tonnie L. C. Casey and James D. Jacobi in writing the description of the Po‘o-uli, a previously unknown species of Hawaiian honeycreeper (Casey and Jacobi 1974).This discovery really caught the world’s attention. If entirely new species remained to be discovered, then maybe all those wonderful ‘extinct’ honeycreepers and ‘o‘os were still out there in some remote forest just waiting for an enterprising ornithologist or lucky birder to find them. It was the impetus for a flood of new research on Hawaiian birds in general, and especially the Hawaiian honeycreepers.

Discovery and research: historical perspectives 39

The era of modern research The age of modern scientific research overlapped somewhat with the period of rediscovery. Some researchers, like Baldwin, Richards, and the Hana Rainforest Project team, looked for possibly extinct birds while conducting scientific studies, but more often scientists were understandably more focused on common species.Why take on a project involving a bird one can never find when we know so little about ones we can? So the two communities did not interact very much until the mid-1970s, when large government-sponsored programs would bring them all together (see below). Research sponsored by universities or museums was more likely to focus on biology than on exploration, and introduced a wide variety of new avenues of investigation to honeycreeper research.

Universities and museums Richards was a student in the Department of Zoology, University of Illinois when he conducted the aforementioned field work (Richards and Baldwin 1953) as part of a study of functional anatomy of Hawaiian honeycreepers (Richards 1957). The results of that study were not immediately published but eventually were combined with additional research by Walter J. Bock (Richards and Bock 1973), of Columbia University and AMNH, who published several other important drepanidine anatomical studies (Bock 1970, 1972, 1978). Another avian anatomist, Robert J. Raikow of the Carnegie Institute, carried out research on Hawaiian honeycreeper limb morphology (Raikow 1976, 1977a) that led eventually to new theories of the group’s origins and relationships (Raikow 1977b, 1978). Richard L. Zusi of the Smithsonian Institution made further contributions based on cranial anatomy and jaw musculature (Zusi 1978, 1989; James et al. 1989). In 1964, Andrew J. Berger, already a noted ornithologist as co-author of a major textbook (Van Tyne and Berger 1959), became professor of zoology at the University of Hawaii. Although he participated in the Kipahulu Valley Expedition (Banko 1968), Berger was less interested in searching for

rarities than in learning more about the native birds that still had viable populations. He also had an interest in the historical record, and included lengthy quotations from the writings of Perkins, Henshaw, Bryan, and other early naturalists, whose works had become difficult to find, in Hawaiian Birdlife (Berger 1972a), a valuable update of Munro’s (1944) book. Berger (1969, 1970b) and his students (Berger et al. 1969) pioneered life history and breeding biology studies of Hawaiian honeycreepers. Robert Eddinger’s (1970; quoted extensively in Berger 1972a, 1981) work on the ‘Apapane, ‘I‘iwi, Kaua‘i ‘Amakihi, and ‘Anianiau in the Koke‘e area were the first such thorough life history studies of any drepanidine, and he also discovered the first nests of the ‘Akikiki (Eddinger 1972a) and ‘Akeke‘e (Eddinger 1972b). Charles van Riper, III studied life history and ecology of the two most common honeycreepers in mamane–naio forest, the Palila (van Riper et al. 1978; van Riper 1980a; Scott et al. 1984) and Hawai‘i ‘Amakihi (Kamil and van Riper 1982; Kern and van Riper 1984; van Riper 1984, 1987), and contributed another ‘first nest’, that of the Maui ‘Alauahio (van Riper 1972). Sheila Conant wrote her dissertation on the O‘ahu ‘Elepaio, but studied Hawaiian honeycreepers in the course of distributional and ecological studies (Conant 1980) in Hawaii Volcanoes NP under the International Biological Program (IBP). Another Berger student, Lawrence Hirai (1978), surveyed the birds of Lana‘i in 1975–76 as part of a US Department of Agriculture environmental study, but found little good news for the island’s honeycreepers. Berger (1980) also pioneered in the maintenance of Hawaiian honeycreepers in captivity. Shortly before he retired, he published a completely revised edition of Hawaiian birdlife (Berger 1981), in which a new classification of Hawaiian honeycreepers, based on my own work (Pratt 1979a), was first published. Upon graduation, Sheila Conant joined the university faculty and has encouraged her own students, notably Maile Stemmermann Kjargaard and Marie Morin, to continue life history studies as well as research dealing with pressing conservation matters. When Berger retired from the Department of Zoology at the University of Hawaii in the early 1980s, he was replaced by

40 The Hawaiian Honeycreepers Leonard A. Freed, who has continued a graduate program of life history studies among other subjects. He and his student Jaan K. Lepson conducted important studies of the ‘Akepa and the ‘Akeke‘e (Freed et al. 1987b; Freed 1988; Lepson and Freed 1995, 1997; Lepson and Pratt 1997). In 1968, Richard E. Warner of the Museum of Vertebrate Zoology at the University of California, Berkeley published his landmark study of the role of disease in the extinction of Hawaiian honeycreepers and other native birds. His work fostered follow-up research on avian diseases and parasites by former Berger student van Riper (1975, 1991a,b; van Riper and van Riper 1985; van Riper et al. 1986, 1987, 1994) that eventually confirmed most of Warner’s (1968) startling findings (see Chapter 9) and set the stage for much ongoing research. Rebecca L. Cann of the John A. Burns School of Medicine, University of Hawaii at Manoa, is currently applying new molecular techniques to the study of disease in Hawaiian honeycreepers (Feldman et al. 1995; Cann et al. 1996; Cann and Douglas 1999). From 1973 to 1975, F. Lynn Carpenter and Richard E. MacMillen, then at the Department of Ecology and Evolutionary Biology at the University of California, Irvine, conducted field work on the slopes of Mauna Kea and Mauna Loa that resulted in the first publications to deal with ecology and community structure of Hawaiian honeycreepers (Carpenter 1976, 1978, 1987; Carpenter and MacMillen 1976a,b, 1980; MacMillen and Carpenter 1980). MacMillen (1974, 1981) also conducted the first drepanidine physiological studies. Their work complemented the previously cited ecological studies of van Riper. John H. Carothers of the MVZ later conducted somewhat similar studies on Maui (Carothers 1982, 1986a,b).

Collection and archival of honeycreeper sounds Beginning in 1950, William V. Ward, a Honolulu banker and a field associate of the Cornell Laboratory of Ornithology, made the first tape recordings of Hawaiian honeycreeper vocalisations. Using rather bulky and primitive equipment, he succeeded

in collecting tapes of the more common species, some of which were included in A field guide to western bird songs (Kellogg et al. undated). Ward’s (1964) paper on the songs of the ‘Apapane was the first on the vocalisations of any Hawaiian honeycreeper. Despite these early efforts, many honeycreeper songs had never even been described adequately, much less recorded prior to the 1970s (Pratt 1996b). By then, technology had made bird song recording much more feasible in remote areas, and Robert J. Shallenberger, then with the DOFAW, began collecting Hawaiian bird recordings. In 1975, working with Shallenberger and James L. Gulledge of the Library of Natural Sounds, Cornell Laboratory of Ornithology, I began an attempt to record all living species of honeycreeper as part of my graduate research at the Louisiana State University Museum of Natural Science. Vocalisations proved to be a highly informative new avenue of research into the relationships and evolution of the honeycreepers, and played a major part in my (Pratt 1979a) dissertation (see also Chapters 4 and 7). Shallenberger and I encouraged other researchers to collect and archive Hawaiian sound specimens. Partly as a result, J. Michael Scott, C. Fred Zeillemaker, and Thane K. Pratt of the USFWS,Timothy A. Burr of DOFAW, and Sheila Conant and Marie Morin of the University of Hawaii made valuable recordings in the late 1970s and early 1980s. Conant and Morin’s tapes from the Northwestern Hawaiian Islands were particularly significant. Eventually, all but three honeycreeper species (O‘ahu ‘Alauahio, Maui Nukupu‘u, and Kaua‘i Nukupu‘u) believed to be extant into the 1970s were added to the LNS, and selections were published in cassette form (Pratt 1996a).

Government surveys and studies One result of the Shallenberger/Pratt collaboration was a series of training tapes for the landmark USFWS Hawaiian forest bird surveys.This massive project (Scott et al. 1986), begun in 1975 under the direction of J. Michael Scott, reflected the newfound interest of the US government in conservation of Hawaiian birds, and was one of the most ambitious surveys of its kind ever attempted anywhere. Preliminary work involved developing

Discovery and research: historical perspectives 41 field identification techniques for difficult species (Scott et al. 1979), differentiating and learning previously unappreciated species-specific call notes (in which I assisted), searching the literature to determine the extent of current knowledge (Scott et al. 1980), preliminary field studies on Moloka‘i (Scott et al. 1977) and Maui (Scott and Sincock 1977), and fine-tuning of field survey techniques (Ralph 1981a,b; Ramsey and Scott 1981; Scott and Ramsey 1981; Scott et al. 1981a,b). Next came the actual surveys in which intrepid teams of ornithologists, botanists, ecologists, and trail-cutters backpacked down regular transects through previously untrodden wilderness of rainforest, deep ravines, lava tubes, and thickets of ‘ie‘ie and ‘uluhe fern. So athletic was the endeavour that it was even featured in Sports Illustrated (Moore 1978)! The list of participants in the field work (Scott et al. 1986: 385) reads like a Who’s Who of previous and subsequent workers in Hawai‘i, and includes an amazing total of 23 authors cited in the Bibliography.The participants in this ‘training academy for Hawaiian ornithology’ gathered data on bird distributions and densities at regularly set sampling stations using a variable circular-plot method (Reynolds et al. 1980), a technique that involves counting all birds heard or seen during a set count period, estimating their distances from the observer, and developing population estimates using sophisticated statistical methods. (For discussions of problems and techniques associated with this method, see Ralph and Scott 1981.) The teams also gathered botanical data used to map vegetation types in the study areas, which included all native forests above 1000 m on Hawai‘i, Maui, Moloka‘i, Lana‘i, and the ‘known distributional area for endangered forest birds’ on Kaua‘i. Kaua‘i was treated differently because of the previous work there of John L. Sincock, who had spent years studying the birds of the Alaka‘i Plateau (Sincock et al. 1984). The surveys detected all but two of the honeycreepers thought to have survived into the 1960s on the islands studied. Neither the Kakawahie nor the Kaua‘i ‘Akialoa were found, and must now be presumed extinct. The high hopes for further rediscoveries were not fulfilled, but the surveys produced an invaluable body of baseline data for future management. Many

species were shown to exist at vanishingly low levels, and though these results were discouraging, at least we finally knew for certain where the hands were pointing on the extinction clock. While the Scott surveys were ongoing, other government agencies were also active in drepanidine research. C. John and Carol Pearson Ralph with the Institute of Pacific Islands Forestry of the US Forest Service conducted an intensive investigation of the ecology of forest birds, mostly honeycreepers, on the island of Hawai‘i (Ralph 1977). The National Park Service funded distributional studies of birds on park lands conducted by Conant and her students at the University of Hawaii (Conant and Stemmermann 1979; Conant 1980, 1981b), as well as the aforementioned avian disease research by van Riper. The US Army and the Hawai‘i Department of Transportation funded environmental impact studies in the Ko‘olau Range on O‘ahu (Shallenberger 1976a,b; Shallenberger and Vaughan 1978) that helped fill in the gap for that island in the Scott surveys, and yielded some of the last observations of the O‘ahu ‘Alauahio (Shallenberger and Pratt 1978). The various state and federal agencies co-operated to a remarkable and uncharacteristic degree in the decade 1975–85, and conducted a highly efficient and well coordinated research program that produced numerous reports authored by multi-agency teams, e.g. Ralph and van Riper (1985), Scott et al. (1984, 1987a, 1988a), and especially the Hawai‘i forest bird recovery plan (Scott et al. 1983). One result of these efforts was the establishment of Hakalau Forest National Wildlife Refuge on the windward slope of Mauna Kea, Hawai‘i, the first refuge created solely for the conservation of forest birds.

Palaeontological discoveries Richards and Bock (1973: 1) wrote that ‘fossils of these birds have not yet been discovered, and are not likely ever to be found’, a statement they probably came to regret, but which reflected the conventional wisdom of the day that volcanic oceanic islands were unlikely sources for fossil remains. Almost contemporaneously with the above prediction, the ornithological world was

42 The Hawaiian Honeycreepers astounded by the discovery of the skeleton of a previously unknown flightless goose in lithified sand dunes at Mo‘omomi on the island of Moloka‘i (Stearns 1973). Joan Aidem, an observant local amateur naturalist, found the bones and sent them to Alan Ziegler at BPBM, who in turn sent them to the Smithsonian Institution.Then in 1974 two entomologists, Frank Howarth and Wayne C. Gagne, and botanist Betsy Harrison Gagne, all of the BPBM, found unusual bird bones in a lava cave near Hana, Maui. These turned out to represent two flightless rails and a flightless ibis (Olson and Wetmore 1976) whose existence could never have even been suspected. Soon, other ‘fossil’ sites were found on O‘ahu, Kaua‘i, and Moloka‘i, that produced thousands of bones of passerines, many of which turned out to be previously unknown Hawaiian honeycreepers. The Smithsonian husband-and-wife team of Storrs

160°

159°

158°

Olson and Helen James, aided by Ziegler, began the tedious work of sorting out the bewildering array of new Hawaiian birds (Olson and James 1982b). Subsequently, additional deposits of bird remains were found on Maui (Madeiros et al. 1989; Olson and James 1991). Prehistoric but recent species described from these deposits ( James and Olson 1991) included four ‘new’ genera and 14 additional species of drepanidines. An important point about these ‘fossils’ is that they do not, for the most part, represent the ancient mineralised bones usually meant by the term, but rather are actual bones of birds that died very recently in geological terms and that lived alongside the historically known species until the arrival of humans in the islands.They are, in other words, extinct members of the modern avifauna, not ancestral forms, and they have revolutionised thinking about virtually every aspect of Hawaiian

157°

156°

155°W

22°N

Makawehi Dunes ÔUlupau Head

BarberÕ s Point

ÔIlio Point

Approximate shoreline of Maui-nui

MAUI

21°

Crystal Cave

20°

19°

3.4 Map showing localities that have yielded subfossil remains of Hawaiian honeycreepers.

Discovery and research: historical perspectives 43 biogeography, evolution, and ecology. James (1998) has recently done an osteologically based phylogenetic analysis of the entire honeycreeper radiation, including the subfossil species, that will likely be formally published about the same time as this book (H. James, pers. comm.). The prehistoric record of Hawaiian honeycreepers continues to grow, with many species as yet undescribed ( James and Olson 1991; H. F. James, pers. comm.) and new deposits turning up (Fig. 3.4), including rich new ones on Hawai‘i (Giffin 1993) and Kaua‘i (Burney et al. 2001).

Molecular and biochemical studies The first biochemical method to be applied to honeycreepers was Sibley’s (1970) study of egg-white proteins, soon superseded by investigations using DNA–DNA hybridisation (Sibley and Ahlquist 1982; Bledsoe 1988). These studies were important in elucidating the relationships of the Hawaiian honeycreepers, but were not refined enough to reveal much about the pattern of relationships within the group. Johnson et al. (1989) used starchgel electrophoresis of proteins to investigate both

extra-group and intra-group relationships of the honeycreepers. Since then, more and more precise techniques have been employed. Restrictionsite variation in mitochondrial DNA (Fleischer et al. 1991, 1994;Tarr and Fleischer 1993, 1995) and gene sequencing studies (Fleischer et al. 1998, 2001) revealed evolutionary patterns in considerable detail, but produced phylogenetic trees with large gaps and unanswered questions because of the lack of genetic material from extinct historical taxa and fossils. Some gaps can now being filled thanks to the discovery (Saiki et al. 1988) of the polymerase chain reaction (PCR), which permits data extraction from very small fragments of DNA such as can be obtained from study skins and subfossil bones (Pääbo 1989; Cooper 1993, 1997). Results of a few such studies of other taxa have been published (Fleischer and McIntosh 2001), but results of ongoing studies involving drepanidine ancient DNA (R. C. Fleischer pers. comm.) have not yet been published. Interestingly, molecular techniques are proving to have important applications far beyond the arcane world of systematics (e.g. Fleischer et al. 1994; Cann and Douglas 1999; Jarvi et al. 2001; Shehata et al. 2001).

4 Origin and evolution

Are Hawaiian honeycreepers a natural group? When the various Hawaiian honeycreepers were first discovered, the idea that they could all belong to a single interrelated group must have seemed preposterous.They were birds that ran the gamut of passerine variation, and most of them did not have any obvious close mainland relatives. By the end of the nineteenth century, a consensus had formed that the thin-billed forms were an endemic Hawaiian family, Drepanididae, but most authorities still considered the finchlike species to be just island representatives of the Fringillidae, which at the time included virtually all the world’s ‘finches’ that today are allocated among several families. Noted anatomist Hans Gadow (1899) at first concluded that the Hawaiian finches were not related to the honeycreepers. He soon modified his opinion at the urging of Perkins, who had become convinced from his ‘study of the habits, the voice, and the peculiarly strong and disagreeable scent of the birds’ (Gadow 1899: 243) that both the finchlike and the thin-billed birds belonged to the Drepanididae. Perkins’s ideas were actually an early example of the concept of synapomorphy. If the finchlike and thinbilled honeycreepers both had some characteristic that was unique among passerines, i.e. a shared derived character state, then they had to be related.

Drepanidine odour To Perkins, the single most important character that tied the finches to the honeycreepers was an unusual odour found in both groups.

Perkins (1893: 108–9) wrote: I cannot liken this scent to any other that I know; but I should certainly call it disagreeable. In [the Hawai‘i ‘Amakihi] it is strongest of all, so much so that when a small company of these birds was overhead in the trees the whole air was full of it; both my native assistant and myself noticed it again and again. Certain nests I could readily recognise as belonging to [the ‘amakihi] by the overpowering scent that still clung to them after the young had flown. Henshaw (1901: 156) agreed, but attributed the strongest odour to a different species when he referred to: the powerful musk-like, but not unpleasant, odour which attaches to the feathers of most of them. Perhaps this odour is more marked in Ou than in any other species. It is so strong in this species that I am sure I have detected it from living birds when near by on low trees, although my sense of smell is anything but acute. In a freshly killed specimen this odour is simply overpowering, and is much stronger in the early morning than later in the day. Almost everyone who has handled Hawaiian birds has noticed ‘drepanidine odour’, which to me resembles the smell of earwax or old army canvas, and which Berlin and VanGelder (1999) called ‘musty sweet’. I do not find it as unpleasant as

Origin and evolution 45 Perkins did, but that it exists is undeniable. Colleagues at LSUMNS have remarked that they can smell it throughout the rather large bird range whenever the case housing our relatively small collection of Hawaiian honeycreepers is opened.At the BPBM, Andrew Engilis and I tested each other’s ability to detect the odour by sniffing randomly arranged specimens of various passerine birds placed in opaque cloth bags. We both consistently picked out the drepanidine specimens on smell alone (Pratt 1992a). As with the nests mentioned by Perkins, drepanidine odour is easily transferred to the fingers and thence to other specimens, so investigations of it have to be done carefully to avoid cross-contamination. This problem surfaced early, and allowed Perkins (1901: 571–2) to get in a ‘dig’ at Rothschild: So far as Hawaiian birds are concerned, this odour is absolutely restricted to the Drepanines. Mr. Rothschild in his work on Laysan makes the astonishing statement that the Meliphagine [‘o‘ o] has a similar and even more powerful odour; but this is only one of those errors which, for want of due care, the museum naturalist is liable to make in opposing facts ascertained and proven in the field. The explanation is very simple: the [‘o‘o] . . . freshly killed or alive has no such odour. The specimens supposed to possess it had no doubt been enclosed in boxes with Drepanines, or when collected in the field had been placed in a bag with them, and had thus become impregnated with their odour. Drepanidine odour is very long-lasting, and, as stated above by Perkins, can be detected in old nests apparently indefinitely. Intensity varies among individuals of a species and, according to Henshaw (1902), possibly with the seasons. It may be totally lacking in some individuals, especially in those species, such as the Laysan Finch, that inhabit dry habitats (Fisher 1906). The odour is apparently contained in the oils that birds use to dress their plumage (Pratt 1992a), because I found only a faint scent in Hawai‘i ‘Amakihi from the mamane–naio forest on leeward Mauna Kea, but rainforest ‘amakihi from only a few miles away had

a very pronounced odour. Rainy weather requires more frequent application of water-repellent oils. Henshaw (1901: 156) had the same idea but was ‘able to detect only a slight odour from this oil when freshly squeezed from the oil-gland.’ Lepson and Woodworth (2002) found the odor stronger around the uropygial gland than elsewhere on individual Hawai‘i Creepers. My experience has been that ornithologists tend to denigrate odour as a useful taxonomic character. Amadon (1950), for example, failed to mention it even though he based his hypothesis of monophyly on Perkins’s ‘field work’. Perkins was not alone in attributing phylogenetic significance to drepanidine odour. Henshaw (1901: 156) wrote: If this characteristic odour originated after the ancestors of the present species reached the islands, and if it is in any way beneficial to its possessors, it seems singular that it should not be shared by all the woodland species whose habits are analogous. Several species are, however, wholly without it. It is possible, as I believe Mr. Perkins has suggested, that what at first seems to be of trivial significance may be found to have a deeper meaning, and that this odour may point to the ancestry and to the ancestral home of some of the island birds. As the American Cœrebidæ, according to Dr. Gadow, are the most likely group from which the Island Drepanididæ are derived, it would be most interesting to discover if the plumage of any of the former have the same characteristic scent. In this connection it is interesting to note that the Oo, Omao and Elepaio are believed by Dr. Gadow to have a nonAmerican origin and not to be Drepanine. It is significant that the feathers of these species, together with Io, do not possess the peculiar odour which is shared, I believe, by all the Island Drepanine forms, certainly by all of them resident upon the Island of Hawaii. Some scientists have suggested to me that odour is useless without a chemical analysis. But I contend (Pratt 1992a) that:

46 The Hawaiian Honeycreepers if we can say that a bird is ‘red’ without a spectrographic analysis, then we can say that it smells like an old tent without subjecting it to gas chromatography (the challenge of doing so in this case has so far proven insurmountable). Non-quantitative characters are not necessarily invalid, and this one is particularly important because it is the historic foundation of the hypothesis of drepanidine monophyly. Perhaps someone reading this book will take on the challenge of performing the chemical analysis of drepanidine odour, a task that is outside my expertise. Does drepanidine odour define the honeycreeper taxon? Perkins (1903) reported it in Hawai‘i ‘Amakihi, Kona Grosbeak, Greater Koafinch, Palila, ‘O‘u, Maui Parrotbill, ‘Akepa, ‘Anianiau, Greater ‘Amakihi, Lesser ‘Akialoa,‘Akiapola‘au, ‘Akohekohe, and Black Mamo. From my own field work I can add ‘I‘iwi, ‘Apapane, and ‘Akikiki to the list, and Lepson and Woodworth (2002) have reported it in the Hawai‘i Creeper. Most specimens of Laysan and Nihoa Finch lack the odour, but I can detect it faintly in some.This list includes at least one member of every drepanidine genus but two: Paroreomyza (‘alauahios and Kakawahie) and Melamprosops (Po‘o-uli). In the case of the ‘alauahios, Perkins suggested that their lack of the disagreeable odour made them more susceptible to predation by the Short-eared Owl (i.e. the odour was an adaptation that made its possessors unpalatable to predators). He had no observations of ‘alauahio being taken by owls, but based his hypothesis on the fact that these birds, alone among the honeycreepers he knew, react strongly to the presence of this potential predator (see Predator response, Chapter 7). My own experience with live Maui ‘Alauahio in the hand supports Perkins’s contention that it lacks drepanidine odour, and I have certainly witnessed several examples of mobbing behaviour by this species, directed at feral cats as well as owls. Perkins may really have been onto something with regard to the function of drepanidine odour. However, neither he nor any other scientist could have put the whole picture together without the recent discoveries of prehistoric bird remains in

Hawai‘i ( James and Olson 1991; Olson and James 1991). Henshaw (1901) considered the Short-eared Owl a recent coloniser in Hawai‘i, and Olson and James (1991) believe it arrived after the Polynesians. It probably never was a major predator on Hawaiian forest birds. A recent survey of odours in birds (Weldon and Rappole 1997) suggests that such odours may evolve as predator defences. We now know that pristine Hawai‘i was far from ‘a land without predators’ as Willis (1972), like most of his contemporaries, believed.The endemic Wood Harrier, described from bones by Olson and James (1991), was so different from typical harriers that it was at first thought to be an accipiter (Olson and James 1982b). Its morphological modifications are clearly adaptations for taking small birds. Likewise, the prehistoric endemic stilt-owls were long-legged diurnal bird predators (many of the passerine bird bones in prehistoric deposits are from owl pellets) with morphologies convergent with that of the Wood Harrier (Ziegler 2002). Clearly selection pressure for predator defences existed, and it may have been severe. It is perhaps noteworthy that finchlike honeycreepers, especially Telespiza finches which have drepanidine odour only faintly if at all, are disproportionately represented in the subfossil samples, while the nectar-feeders, many of which have very strong drep odour, are inexplicably nearly absent from these deposits ( James and Olson 1991). Drepanidine odour may have been an early adaptive response to strong selection pressure from a community of bird predators, and loss of predator mobbing behaviour may characterise those species that possess the odour, because they no longer need to mob. Exactly how it functions in warding off predators could be fertile ground for further research.

Evidence from tongue morphology Nearly all passerine tongues have a pair of rearward projections at the posterior end called lingual wings (Gardner 1925; Pratt 1992a). The tongues of most Hawaiian honeycreepers (see also Chapter 6) are peculiar in that they are squared off or truncate at the base, with no such rearward projections. In fact, the tongues of finchlike honeycreepers differ from those of cardueline finches solely in this

Origin and evolution 47

Carpodacus

Psittirostra psittacea

Leucosticte

Telespiza cantans

Pipilo

4.1 Comparison of tongues of drepanidine (right) and other finches (left). From Pratt (1992b), after Raikow (1977b).

feature (Fig. 4.1). But members of the genera Paroreomyza and Melamprosops have deeply notched tongue bases exactly like those of most passerines (Fig. 6.15). (For a discussion of the evolution of this character, see the section on adaptive radiation at the end of this chapter.)

Anomalies Is it merely coincidence that the same two genera that stand apart from most honeycreepers in lacking the odour also stand apart in tongue structure and the lack of mobbing behaviour (as well as in primary songs, cranial morphology, and breeding biology, to be discussed later)? Or does this mean that these two genera are not really Hawaiian

honeycreepers, and that the group is, after all, polyphyletic? I suggested as much in two papers on the subject (Pratt 1992a,b), and Tarr and Fleischer (1995) supported my suggestion that Paroreomyza, at least, might not be a drepanidine genus (they had not yet studied Melamprosops). If these two genera are excluded, then the Hawaiian honeycreepers form a neat, discrete taxon definable on the basis of three synapomorphies: drepanidine odour, loss of predator responses, and loss of lingual wings. But nature is not always neat, and if these aberrant species are not dreps, then what are they? The ‘alauahios and Kakawahie are particularly enigmatic because they also lack the cranial synapomorphies (see Skeleton below) that unite cardueline finches and Hawaiian honeycreepers, they are the only

48 The Hawaiian Honeycreepers supposed dreps to bind their nests with spider silk, and they are the only ones that routinely have nest helpers (see Chapter 8). Nevertheless, they have the ‘jizz’, as birders would say, of the honeycreepers. They clearly are an aberrant part of the adaptive radiation of the group, but superficially they do not seem out of place in it, so much so that they were once considered conspecific with birds that do possess the ‘defining characters’.Their distinctive songs could be seen as derived from a finchlike one, but their other aberrant features could be the clue that their resemblances to honeycreepers are the result of convergence. An early genetic study ( Johnson et al. 1989) indicated that Paroreomyza was part of the earliest line to diverge from the ‘main line’ of honeycreeper evolution after the colonisation of Hawai‘i (for their phylogeny and others based on molecular data, see Fig. 4.6). These results are problematical, however, because the Johnson team grouped Paroreomyza with Oreomystis as the sister group to all the other Hawaiian honeycreepers. On phenotypic grounds, such a finding is cladistically untenable because Oreomystis has all of the defining drepanidine synapomorphies that Paroreomyza lacks. Tarr and Fleischer (1995) presented molecular evidence that Paroreomyza was indeed outside the main radiation of the honeycreepers (but that Oreomystis belonged among the main line genera and was not the closest relative of Paroreomyza), and suggested that Paroreomyza might be an independent derivative of the Carduelinae (but that is contradicted by the fact that ‘alauahios lack many of the synapomorphies that unite the honeycreepers with this finch group). Shortly afterwards, however, the same laboratory’s analysis of allozymes (Fleischer et al. 1998) produced results close to those of Johnson et al. (1989), with Paroreomyza and Oreomystis forming a sister clade to the rest of the honeycreepers. James’s (1998) study of cranial osteology also recognised the Paroreomyza/Oreomystis clade and placed it in an unresolved trichotomy with the Po‘o-uli (see below) and a thin-billed honeycreeper clade, these three in turn the sister group of the drepanidine finches. Clearly, excluding Paroreomyza from the Hawaiian honeycreepers based on Pratt (1992b) and Tarr and Fleischer (1995) would have been

premature, but the full story remains to be told. Therefore, I retain Paroreomyza as a drepanidine genus herein, but I consider it a basal offshoot from the main line of evolution, before the defining synapomorphies appeared. When the Po‘o-uli was discovered, it was classified as a Hawaiian honeycreeper essentially by default (Pratt 1992a). Aided by Dean Amadon and Walter J. Bock, the describers (Casey and Jacobi 1974) cited nothing specific that proved the bird drepanidine. Although unstated, geographic probabilities likely played a role in this classification, because Amadon (1986) saw ‘no good reason to hesitate to place Melamprosops in the Drepanidinae’, as if to say that it had to be a honeycreeper unless proven otherwise. Amadon still did not supply any reason not to hesitate in calling the Po‘o-uli drepanidine. Bock (1978) stated that features of the tongue skeleton and musculature of Melamprosops not only support its placement among the drepanidines, but ‘provide further support for the monophyly of the family’. However, most of the ‘drepanidine features’ he cites are ‘shared with cardueline finches and some with other nineprimaried oscines’ and thus do not unequivocally show the Po‘o-uli to be a honeycreeper. Furthermore, his drawings reveal that the tongue has prominent and well developed lingual wings. Both Andrew Engilis and I have examined the two existing specimens and agree that they lack drepanidine odour (Pratt 1992a). Of course, if they had the odour it would not be very meaningful after years of handling by ornithologists who did not appreciate the cross-contamination problem cited above, but absence of odour is important. Engilis and I also examined the two Po‘o-uli nests in the BPBM and found that they, too, lack the scent, whereas some nearly century-old drepanidine nests clearly retain it. Unlike the three Paroreomyza, the Po‘o-uli, with its highly distinctive plumage and a song totally unlike that of any of the other living drepanidines, does not by any means immediately suggest a honeycreeper to most observers. Nevertheless, it, too, seems more at home among the dreps than with any other possible family, and geographic probabilities cannot be ignored. Perhaps when we know

Origin and evolution 49 more about recently described prehistoric species ( James and Olson 1991), the Po‘o-uli will seem less aberrant. Its bill shape is superficially, at least, similar to those of the newly described genus Xestospiza, but the resemblance could easily represent a shared primitive condition or convergence. From James and Olson’s comparisons, the interorbital septum (about which more later) of Melamprosops appears to be derived from the distinctive cardueline/drepanidine type (Zusi 1978), and some of Bock’s (1978) tongue features might turn out to be synapomorphies if analysed from a cladistic perspective. Those features would at least put it in a clade with the honeycreepers and their nearest relatives (see below), but like Paroreomyza it would have to have diverged very early from the ‘main line’. That is exactly what Fleischer et al. (2001) found in a study that used both molecular and osteological characters.They placed it among the honeycreepers, but in a basal position with a nearly 4-million-year history of independent evolution. One of the recently described prehistoric species, Orthiospiza howarthi, is also enigmatic. Although it was originally described as drepanidine (Olson and James 1991), it fell outside the honeycreeper clade in James’s (2004) more recent study, and appeared allied instead with the cardueline bullfinches of the genus Pyrrhula. It was excluded from her detailed osteological analyses. On the other hand, James (2004) found “no statistical support” for excluding Orthiospiza from the Hawaiian honeycreepers, and it is regarded as drepanidine herein. Despite one recent study that considered the group polyphyletic (Chu 2002), a possible result of limited sampling (James 2004), little doubt now remains that the Hawaiian honeycreepers, including Paroreomyza and Melamprosops, are a natural group even though no single phenotypic character can be cited to unify them. However, the colonisation event itself can be viewed as a defining moment in the group’s evolution. The original coloniser apparently gave rise to at least two, and more likely three, clades: one comprising the majority of species defined by drep odour and loss of lingual wings; and one or two others that gave rise to the ‘alauhios and the Po‘o-uli before those defining features appeared. The next important

evolutionary question is: Where do we put the Hawaiian honeycreeper branch on the avian tree?

Whence came the honeycreepers? Gadow (1899) first proposed that Hawaiian honeycreepers were derived from Neotropical honeycreepers (including such birds as the Bananaquit, Dacnis spp., flowerpiercers, and Chlorophanes and Cyanerpes honeycreepers), a group later shown to be polyphyletic (Beecher 1953) but at the time classified as the family Coerebidae. Gadow was proceeding from his belief at the time that the Hawaiian finches were not related to the thin-billed honeycreepers. In fact, he made a strong case for classifying the Hawaiian finches in the Fringillidae (note that in Gadow’s day, this family was a catchall for birds with seed-eating bills, and included both the emberizine finches and the cardueline finches, later shown to be independent of each other). In discussions about the relationships of the Hawaiian honeycreepers, he even stated that the drepanidine tubular tongue could be derived from a fringilline one, a statement that would prove prophetic. But when he later conceded that the Hawaiian finches and honeycreepers were related, he failed to make the logical connection that if the drepanidine finches were fringilline, then the thin-billed birds might be also. Instead, he thought the Hawaiian finches evolved from a thin-billed ancestor and came to resemble cardueline finches by convergence. In 1929, the Russian anatomist P. P. Sushkin published a short but important paper in which he compared skulls of a representative sample of Hawaiian honeycreepers in the genera Himatione, Vestiaria, Hemignathus (subgenus Akialoa), Oreomystis, Loxioides, and Telespiza, with those of cardueline finches, ‘coerebids’, and meliphagids (honeyeaters such as the Hawaiian ‘o‘os). He found several cranial features common to cardueline finches and both thick-billed and thin-billed drepanidines, in which both groups differ from other passerines, and especially from tanagers. In cladistic terminology, he discovered a suite of synapomorphies that showed the carduelines and drepanidines to be sister

50 The Hawaiian Honeycreepers groups. In so doing, Sushkin established a second hypothesis about honeycreeper relationships. Amadon (1950: 231) dismissed Sushkin’s findings almost out of hand, although he admitted that skulls of Hawaiian finches ‘are like those of cardueline . . . finches, not those of tanagers’. He supported Gadow’s coerebid hypothesis (and specifically excluded tanagers as well as cardueline finches as potential ancestors), although he was aware of Beecher’s soon-to-be-published findings that there was really no such thing as a coerebid (some are specialised tanagers,Thraupidae; some are specialised wood warblers, Parulidae). He presented no positive evidence of a coerebid affinity other than vague similarities, but stated that derivation of Hawaiian honeycreepers ‘from a nectar-feeding coerebid-like ancestor offers no particular difficulties’ (Amadon 1950: 231). He firmly held to the belief that thin bills were generalised and thick ones specialised. After describing a hypothetical evolutionary sequence in which drepanidine finches are secondarily derived from thin-billed ancestors, he wrote (Amadon 1950: 232): If, on the other hand, the Drepaniidae evolved from tanagers or finches, we must suppose the sequence was just the reverse: modification of a heavy-billed type like Psittirostra or Ciridops that terminated eventually in more generalised forms like Loxops or Himatione. This process is rather difficult to envisage in terms of functional anatomy. Dedifferentiation and ‘reversal of evolution’ do occur, but as a rule evolution is from less to more specialised forms. In particular, a heavy, seed-crushing bill would seem to be something of an evolutionary dead end. The Galapagos finches might be considered an exception, but it is by no means proved that the thinner-billed geospizids . . . are not nearest the ancestral type. Moreover bill variation in the Geospizinae is much less radical than in the Drepaniidae. In his study of the Fringillidae, Tordoff (1954a: 31) took strong exception to Amadon’s belief and stated:

If heavy bills are indeed evolutionary dead ends, then for a hypothetical ancestor of the fringillids, one must visualize some kind of bird with a bill at least as thin as the most needle-beaked living fringilline . . . If one grants that bills of seed-eating birds can become a little thinner, then it is fruitless to argue that they cannot become much thinner. I see no justification for considering a heavy, seed-crushing bill an evolutionary dead end. Gadow’s second choice as an ancestral group had been the tanagers (Thraupidae), which now have subsumed most of the coerebids.This is an important point, because Amadon (1950) considered coerebids to be generalised nectar-feeders and finches of all types to be specialised derivatives. Placing the coerebids among the tanagers makes them specialised, not generalised, members of their group and reverses Amadon’s unsupported hypothesis that thin bills could not be derived from thick ancestors. Amadon also ignored (or was unaware of ) the fact that the most ecologically generalised Hawaiian honeycreeper is the Laysan Finch, an eclectic feeder on everything from seeds and fruit to insects to seabird eggs (see Chapter 8), and that many cardueline finches are ecological generalists (Newton 1973). Although Amadon (1950) acknowledged that the nectar-feeding tongue of many Hawaiian honeycreepers (Chapter 6) is unlike that of any Neotropical honeycreeper, he considered it another variation on the theme within the group. Beecher (1951) thought the Hawaiian honeycreepers were ‘almost certainly of tanager origin’ but cited evidence that was apparently never published. In this he essentially agreed with Amadon, because he considered the Neotropical honeycreepers to be tanagers specialised for nectarivory. He was apparently somewhat troubled by the appearance of adaptations for gaping in the ‘I‘iwi, and failed to note that many cardueline finches, such as goldfinches, siskins, and redpolls are also gapers (Newton 1973). He also made the remarkable statement (Beecher 1951: 285): ‘That the drepanidids have a tanager origin is supported also

Origin and evolution 51 by the identity of muscle-patterns in the Hawaiian finch, Psittirostra cantans [‘O‘u], and the cardueline finch, Carpodacus mexicanus [House Finch]. This may be interpreted as two independent origins of nearly identical finches from a common tanager ancestor.’ Although his myological findings seemingly corroborated Sushkin’s osteological ones, he dismissed probable synapomorphies as parallelism. The next author to weigh in on the controversy was Bock (1960), who at that time simply called attention to Sushkin’s work, apparently to prevent the unquestioned general acceptance of the tanager ancestry hypothesis. Later, Richards and Bock (1973) presented a rather enigmatic analysis that included elements from both sides of the debate. Although they accepted a cardueline ancestry for the group, they placed Ciridops, a finchlike nectarfeeder with a drepanidine tubular tongue (Bock 1972) at the base of the radiation. (Note that James and Olson (1991) found Ciridops to be only superficially finchlike; it is apparently a highly derived member of the nectarivorous honeycreeper clade.) Enigmatically, Richards and Bock (1973) accepted Amadon’s assertion that drepanidine finches are secondarily derived rather than primitive. This fact seems inconsistent with acceptance of a cardueline ancestor, but their support for the cardueline hypothesis was rather lukewarm. They stated (Richards and Bock 1973: 123–4) that even though their conclusion was: that the Hawaiian honeycreepers evolved from cardueline finches or from their immediate ancestors, we wish to emphasise the dangers of its wholehearted acceptance. Much of the cited supporting evidence for this idea is really no better than the evidence support-ing the Coerebidae-Thraupidae origins of the Drepanididae. Uncritical acceptance of the cardueline origin of the drepanidids has the same serious disadvantages as had the general acceptance of coerebid-thraupid origins of the Drepanididae. So, as late as the 1970s, the jury was still out with regard to the closest relatives of the Hawaiian honeycreepers.

It was in this equivocal atmosphere that I began my own critical evaluation of the various lines of evidence for drepanidine ancestry. I broadened the investigation to include various kinds of information that were unavailable to the avian anatomists or to Amadon. Unlike Richards and Bock (1973), I did not take Amadon’s classification as a default position but rather started from the beginning with no prejudice either way. I reviewed the historic evidence and gathered new information that might shed light on the controversy. The following is a summary of my review of traditional characters along with new information I was able to turn up from both before and after my graduate work (Pratt 1979a).

Skeleton Most investigators who have looked at the osteology of Hawaiian honeycreepers have focused on the skull (cranium). Sushkin’s observations (confirmed by Amadon) revealed that a solid bony palate, rare in passerines generally, is found in both cardueline finches and Hawaiian honeycreepers. Beecher (1951) thought the skull showed tanager affinities despite some obvious cardueline/drepanidine similarities. Amadon reviewed the literature available at the time and presented a tabular summary. It showed many points common to carduelines and drepanidines, and many contrasts between drepanidines and tanagers, but he concluded that cranial osteology held little phylogenetic information. I agree that the features in his table are difficult to interpret, and probably cannot tell us much without a more detailed analysis. Bock (1960) surveyed a skull feature called the palatine process of the premaxilla in passerines. He found that cardueline and drepanidine finches shared a derived state (called the lateral flange condition). Richards and Bock (1973) reported several other features common to carduelines and thinbilled honeycreepers, but whether these characters represent derived conditions is difficult to determine from the authors’ comments. One telling feature is the presence of ‘a slight lateral flaring of the palatine process of the premaxilla’ in the ‘Akepa, indicating a possible evolutionary trend of reduction of the lateral flange in thin-billed honeycreepers.

52 The Hawaiian Honeycreepers Zusi (1978) discovered a diagnostic characteristic of the interorbital septum of cardueline finches.The interorbital septum in most passerines is a thin plate of bone that separates the eye sockets in the middle of the skull. It is usually a single layer of bone and often has openings or ‘windows’ called fenestrae. In carduelines and birds related to them, the interorbital septum is unusually thick, is double-walled with supporting struts called trabeculae between, and usually unfenestrated (⫽lacking openings). Other birds with ‘finch’ bills resemble most passerines in having thin septa with large openings.The only other ‘finches’ to have the thick unfenestrated interorbital septum are Hawaiian honeycreepers. Zusi (1989) later reported that the thin-billed honeycreepers have thick interorbital septa but have a thin single-walled portion in the centre, sometimes with a small fenestra. The thinbilled drepanidines are the only non-finch-billed passerines with thick septa (Zusi 1978, 1989). Richards and Bock (1973) illustrated the thinner but still cardueline septa of ‘Akepa, Hawai‘i Creeper, and Hawai‘i ‘Amakihi, but their figure of Maui ‘Alauahio is quite distinctive and appears at first to be a noncardueline type. Richard L. Zusi and I together examined skulls of Maui ‘Alauahio and Kakawahie (both Paroreomyza) and found that neither has the solid bony palate typical of Hawaiian honeycreepers, and that both have a very thin septum with a very large foramen and with the cranial fenestra (an opening that lies above and usually slightly behind the interorbital septum) extended forward so that all that remains of the septum is a thin band of bone. This condition is similar to that of emberizine finches as illustrated by Zusi (1978). This difference could be seen as another indicator that Paroreomyza may not really be drepanidine (Pratt 1992b). One character, however, shows that the aberrant septum of Paroreomyza could have evolved from the cardueline type. One of the less obvious features of the cardueline septum is that the floor of the cranial fenestra is broad and flat rather than narrow (Zusi 1978, 1989), and appears straight in profile, without a hump or upward protrusion. The lower contour is straight in Paroreomyza as well as Melamprosops. In the latter, the septum is also only a single layer of bone, and is fenestrated, so it might otherwise be considered nondrepanidine in form except for the flat lower margin

of the cranial foramen ( James and Olson 1991).This one seemingly minor character may be the only certain synapomorphy that can place Paroreomyza, Melamprosops, and many of the recently described fossil genera ( James and Olson 1991) in the cardueline/ drepanidine clade. The most detailed study of drepanidine skeletons is that of James (1998, 2004), which became available to the author too late for a thorough review here. Suffice it to say that her results show overwhelming evidence of a close relationship between carduelines and Hawaiian honeycreepers. Indeed, her evolutionary topology suggests that the dreps are embedded within the cardueline radiation, rather than being a sister group to it of equivalent taxonomic rank. However, the sister-taxa hypothesis could not be rejected on the basis of her data.

Myological evidence Myological studies of Hawaiian honeycreepers have involved the muscles of the jaw and tongue (Beecher 1951, 1953; Bock 1972, 1978; Richards and Bock 1973; James et al. 1989; Zusi 1989) and the limbs (Raikow 1976, 1977a,b). Beecher’s (1953) findings on jaw musculature parallel those already discussed for the bones of the skull. Bock stated unequivocally that Hawaiian honeycreeper jaw muscles were like those of cardueline finches and unlike those of thraupine honeycreepers. Raikow discovered two derived conditions that indicate a cardueline/ drepanidine connection. The first is a tibial head of the shank muscle M. peroneus brevis, the second a coracoid head of the upper forelimb muscle M. deltoideus minor. Both of these conditions are present in all of the honeycreepers he studied and in some carduelines, but are rare among passerines generally and absent in thraupine honeycreepers (Raikow 1977b). As with the skeleton, the muscles certainly point to a cardueline ancestry for the drepanidines.

Nasal operculum As discussed in more detail in Chapter 6, fruit- and nectar-feeding honeycreepers have a protective operculum over their nostrils.Amadon (1950) considered this to be evidence that the drepanidine

Origin and evolution 53 finches evolved from nectar-feeding ancestors, but Raikow (1977b) disagreed, showing that the nostrils of most drepanidines did not differ significantly from those of cardueline finches.

Plumage similarities Some aspects of Hawaiian honeycreeper plumage (Chapter 7) may shed light on drepanidine ancestry. Recent publications that illustrate all species of cardueline finches (Clement et al. 1993) and tanagers (Isler and Isler 1987) in colour enable easy comparisons. Among Hawaiian honeycreepers, the predominant colours are melanin (black or brown), red, yellow, and a dull green produced by a combination of yellow and melanin. Structural colours such as bright green, blue, or violet are lacking, as are opalescent and metallic or iridescent effects. Exactly the same pigmentation is present in cardueline finches, although a very few show a bit of iridescence. Although this fact may not constitute a synapomorphy, it shows that both groups have the same genetic raw materials for coloration. On the other hand, tanagers are noted for their brilliant plumages that often include structural colours, iridescence, and opalescence. Reds and yellows, the predominant colours of Hawaiian honeycreepers, although common among typical tanagers, are almost non-existent among the Neotropical honeycreepers, the tanager subgroup to which some have considered the Hawaiian honeycreepers related. Cardueline plumages often exhibit sparrow-like streaks, wing-bars (especially in females and juveniles), and black face masks. Sparrow-like streaks are found in Laysan and Nihoa finches, and many Hawaiian honeycreepers have black faces that are reminiscent of carduelines such as citrils and some canaries (Serinus), as well as goldfinches, siskins, and redpolls (Carduelis). The Akeke‘e in particular has a very siskin-like or citril-like appearance that extends even to the profile of the bill and the deeply notched tail. Amadon (1950: 231) dismissed this resemblance as superficial, with the circular reasoning that ‘akepas (including ‘Akeke‘e) are congeneric with birds that are not cardueline-like, a classification that was original with his revision. Richards

and Bock (1973) were impressed by a supposed close resemblance of the ‘Ula-‘ai-hawane to the cardueline rosy-finches, but I find it less compelling than the aforementioned similarities to Serinus. Indeed, the two historically known Telespiza finches might well have been classified in the genus Serinus if their drepanidine affinities had not been known. Amadon attributed their streaked patterns to convergence that resulted from adaptation to the low grassy habitats on Laysan and Nihoa, but recent discoveries by James and Olson (1991) show that their restriction to those islands is relictual and that they once were found throughout the archipelago. They could just as easily have inherited their plumage pattern from a cardueline ancestor and represent the primitive condition among the Hawaiian honeycreepers. A few tanagers have wing-bars or black masks, but streaks are rare, confined mainly to the genus Tangara and to female Cyanerpes honeycreepers, but in no case do these features closely resemble their expression in Hawaiian honeycreepers. Birders often comment on the superficial similarity of the colour pattern of the ‘I‘iwi to that of the Scarlet Tanager, but because the Hawaiian bird is a highly specialised nectarivore by virtually any reckoning, I regard the resemblance as at best convergence and more likely just coincidence. Evolutionary shifts from red to yellow among sister species or congeners has occurred several times among Hawaiian honeycreepers (‘Akepa and ‘Akeke‘e, Greater and Lesser Koa-Finch, ‘alauahios and Kakawahie) and the Maui ‘Akepa is dimorphic for red and yellow. Such shifts are also frequent among cardueline finches.The development of yellow and orange variants in the introduced Hawaiian population of House Finches is well known (Grinnell 1911; Pratt et al. 1987).The Red Siskin of Venezuela and the Saffron Siskin of Ecuador differ solely in that the former is red where the latter is yellow. Somewhat similar red/yellow shifts have occurred in some typical tanagers, but none among the thraupine coerebids, among whom red and yellow feathers are unusual. One interesting feature that may be just coincidence, but worth mentioning nevertheless, is the odd truncation of the primaries of the ‘I‘iwi and ‘Apapane. This feature apparently evolved to

54 The Hawaiian Honeycreepers enhance an audible ‘wing note’ produced by these birds and their close relatives. No such feather modification is found among the tanagers, but the cardueline Hawfinch has truncate and curled tips of the inner primaries, apparently used in courtship (Newton 1973).

Evidence from vocalisations As detailed in Chapter 7, most Hawaiian honeycreeper songs fall into three main categories (which do not necessarily correspond to taxa), with Paroreomyza and Melamprosops each in a separate category of its own. The mostly red and black nectarivores have very distinctive songs that distinguish them as a group from the rest of the honeycreepers.They are highly derived birds, whether one considers the ancestor to be a finch or a coerebid.Therefore, their songs, which Amadon aptly (1950) compared to those of American blackbirds (Icteridae) and which also closely resemble those of honeyeaters (Meliphagidae), probably cannot tell us much about honeycreeper ancestry. The other two groups have songs that have been compared by virtually every author and many modern observers to those of cardueline finches. Songs of the ‘O‘u, Palila, Laysan Finch, and Nihoa Finch have all been described as canary-like or linnet-like, both allusions to cardueline finches.The thinner-billed ‘green’ honeycreepers have canary-like whisper songs, but their primary songs are short trills or warbles that sound like short pieces of a complex canary-like song.The Maui ‘Alauahio has a very distinctive song that, although hardly canary-like, sometimes resembles that of the cardueline House Finch.All of this makes evolutionary sense only if the complex or canarylike songs were inherited from a cardueline ancestor, reduced to whisper songs on the one hand, and modified to sound like honeyeaters on the other.To reverse the sequence, with complex, strikingly cardueline-like songs (coincident with other finchlike characteristics) evolving by convergence in the isolation of the Hawaiian Islands, is much less plausible. Call notes tell much the same story but unite all of the song groupings. In particular, a loud upslurred whistle, closely resembling whistles of several species of Carduelis, is characteristic of a broad array of Hawaiian honeycreepers from the ‘O’u and

Laysan Finch to the ‘amakihis, ‘akepas, Maui Parrotbill, and ‘Akiapola‘au, to the ‘I‘iwi and ‘Akohekohe. Among cardueline finches, it is a social/alarm call, which Mundinger (1979) regards as a synapomorphy uniting the Carduelinae and Fringillinae as a clade. Could this call be the evolutionary thread that ties the whole radiation, from fringillines to honeycreepers, together? We may never be able to answer that question for certain, but we can certainly say unequivocally that Hawaiian honeycreepers often sound a lot like cardueline finches and never sound anything like tanagers.Tanagers as a group, and the coerebids in particular, have comparatively simple songs (Isler and Isler 1987) with nothing approaching the complexity heard among carduelines or Hawaiian honeycreepers. Some behaviours associated with vocalisations also reflect the cardueline/drepanidine relationship. Many species of both groups exhibit flight songs as part of their courtship displays. Newton (1973: 164) describes ‘butterfly flights’ of carduelines in the open, ‘accompanied by continual calling and singing’, and with ‘slow deliberate wingstrokes . . . interspersed with long glides.’ High song flights above the forest, with periods of hovering and gliding, are conspicuous in the ‘Apapane, ‘Akohekohe, and ‘I‘iwi, and less showy flights are performed by Hawai‘i ‘Amakihi, Palila, ‘Akiapola‘au, and Maui ‘Alauahio among others (Chapter 7). Song flights of any kind are unknown in tanagers and Neotropical honeycreepers (Isler and Isler 1987). Another oddity shared by the two groups is the tendency to sing in other than territorial or reproductive situations. Newton (1973: 163), contrasting the carduelines with the fringilline Chaffinch and Brambling, writes: In the carduelines, although the song is most frequent and vigorous during breeding, it is not restricted to this time, nor to the territory, but occurs at all seasons except during the autumn moult . . . Its function, moreover, is less obvious, as it is uttered in social, as well as in aggressive and sexual contexts. He could just as well have been describing Hawaiian honeycreepers. Some species in each ‘vocal grouping’ mentioned above seem to sing all

Origin and evolution 55 year, whether breeding or not. Interestingly, the species for which Newton’s description would be most appropriate are the ‘Apapane and ‘I‘iwi, members of the group that, in the quality of their songs, are less like carduelines than most honeycreepers. The fact that they resemble honeyeaters in tone quality but carduelines in pattern and seasonality supports the idea that the meliphagid-like songs of the nectarivorous dreps are derived from those of a cardueline ancestor.

(Philippines), and Hispaniola. Among the tanagers, only a few species, none of them coerebids, migrate long distances (Isler and Isler 1987), and they do not do so in flocks. Furthermore, they are poor island colonisers. Although aberrant tanagers are found in the West Indies, they have not colonised Cocos Island or the Galápagos, Pacific islands much closer to possible source areas than Hawai‘i.

Evidence from breeding biology

The first biochemical study to look at Hawaiian honeycreepers was Sibley’s (1970) electrophoresis of egg-white proteins. It indicated a close relationship between cardueline finches and drepanidines. The more sophisticated DNA–DNA hybridisation technique (Sibley and Ahlquist 1982) reinforced that finding and produced a phylogenetic diagram that made the honeycreepers and carduelines sister groups, and they together the sister group of fringilline finches. Similarly, Bledsoe (1988) diagrammed the cardueline and fringilline finches and the Hawaiian honeycreepers as the branches of an unresolved three-way split (trichotomy). These results confirmed the conclusions from traditional methods reached earlier by Raikow (1977b), Pratt (1979a), and Olson and James (1982b). A contrary opinion was published by Johnson et al. (1989) as the result of a biochemical study of allozymes, that concluded, among other things, that the Hawaiian honeycreepers are genetically closer to Emberizidae (including tanagers) than to the Carduelinae; and that the ancestor of the group was neither a nectar-feeding coerebid nor a seed-eating cardueline, but rather a ‘generalised’ bird with intermediate feeding habits and structures. The Johnson team understood the anomalous nature of their own findings and included some possible explanations in their discussions. A. H. Bledsoe (whose 1988 paper had supported the cardueline hypothesis), provided at least two hypothetical explanations for the anomalous results ( Johnson et al. 1989). All subsequent studies have assumed the Carduelinae and Drepanidinae to be each other’s closest relatives, Johnson et al. (1989) notwithstanding. In a phylogenetic analysis of the mitochondrial cytochrome b gene, with the cardueline House Finch and a Neotropical honeycreeper as outgroups, Fleischer et al. (1998)

Territoriality (Chapter 8) in cardueline finches and Hawaiian honeycreepers is very similar, and quite different from that of most songbirds. Males initially defend a moving territory centred on the female. Once the nest site is chosen, the territory remains quite small around the nest and the owners will tolerate unusually close approach by conspecifics.Territories also may be clustered into small ‘colonies’. A particularly noteworthy feature is that, among passerines, only some cardueline finches and some Hawaiian honeycreepers allow faecal sacs from their young to accumulate on the outside of the nest near the end of the nestling period. This rare trait must have been inherited in both groups from a common ancestor, and later altered secondarily in some species of each.

Geographic likelihood Bock (1960) was the first to point out that the ancestor of the Hawaiian honeycreepers would likely be some kind of bird that (a) wanders or migrates long distances; (b) travels in flocks; and (c) settles down to breed in places to which it has wandered. Among the suggested ancestors of Hawaiian honeycreepers, only the cardueline finches have all these qualifications. Many carduelines undergo periodic irruptions (Newton 1973), when they may wander great distances and appear in totally unexpected places. Also, the wanderers may remain in the new locality to breed, often producing permanent colonies. The crossbills are the best known exemplars of this behaviour (Nethersole-Thompson 1975).These characteristically boreal birds have produced tropical offshoots in Indochina, Luzon

Biochemical and molecular evidence

56 The Hawaiian Honeycreepers upheld the cardueline/drepanidine relationship, as did Fleischer and McIntosh (2001) with a larger sample of outgroups including tanagers, both New World and Old World warblers, and a New World blackbird. Groth’s (1994, 1998) cytochrome b sequence analyses of all finchlike passerines indicated that Hawaiian honeycreepers are not just close to cardueline finches, they are cardueline finches because they are closer to the genus Carduelis than to other cardueline genera in the study (e.g. Carpodacus). The close relationship of Hawaiian honeycreepers and cardueline finches can no longer be reasonably questioned, and the issue now is whether they can even qualify as a named subgroup.

Evolutionary history Historical development Except for the fact that he lumped together many of Perkins’s (1903) genera, Amadon’s (1950) dendrogram is very similar to the earlier one. It envisions two main branches from an unspecified common ancestor (Fig. 4.2), which in his text is stated to be a thin-billed member of the Neotropical tanager/honeycreeper group. One branch groups the thin-billed ‘green’ birds with the finchlike honeycreepers, with the latter as the most highly evolved members, while the other branch groups the ‘red and black’ birds with Ciridops as the

most divergent (from the ancestor) genus. In the latter respect, Amadon (1950) differs slightly from Perkins (1903), who regarded Drepanis as the most derived genus. Richards and Bock (1973) presented an abbreviated dendrogram that placed Ciridops at the base of the honeycreeper radiation but still regarded the drepanidine finches as specialised derivatives.

Phenotypic studies Raikow (1977b) was the first to conduct a cladistic analysis of the Hawaiian honeycreepers.At the time, such analyses were done ‘by hand’ but by the 1980s computers had entered the scene, and Raikow (1986) reprised his earlier work with a simple program that did not alter his earlier topology. Raikow’s cladogram (Fig. 4.3) suffers, in my opinion, from his uncritical acceptance of Amadon’s taxonomy. Because Amadon’s genera were so broad, Raikow failed to examine ‘akepas, nukupu‘us, ‘alauahios, and all of the drepanidine finches except Laysan Finch and ‘O‘u. Ciridops was unavailable for dissection and also was not included, so his phylogeny is far from comprehensive. But it was revolutionary in being the first to accept unequivocally a cardueline ancestry for the honeycreepers and the first to recognise the importance of the drepanidine tubular tongue in defining a major clade.

Laysan Finch drep finches heterobills/'akialoas

Maui Parrotbill −− 'O'u

parrotbill

'Akikiki

'Greater Loxops'

Kaua'i 'Akialoa

'l'iwi mamos 'Apapane 'Akohekohe − 'Ula-'ai-hawane

Hawai'i 'Amakihi − 'Akohekohe 'Apapane 'l'iwi Mamo (sp.)

4.2 Phylogeny developed by Amadon (1950), long regarded as the ‘standard’; redrawn with modern nomenclature substituted.

4.3 Cladistic phylogeny of Raikow (1986) redrawn in modern style with nomenclature standardised.

Origin and evolution 57 Laysan/Nihoa Finch koa-finches Palila Kona Grosbeak −− 'O'u Maui Parrotbill 'Akikiki/creeper 'akepas heterobills 'akialoas 'amakihis − 'Ula-'ai-hawane mamos + 'l'iwi − 'Akohekohe 'Apapane

4.4 Phylogeny based on a broad range of phenotypic characters from Pratt (1979a).

The phylogeny in my 1979 dissertation (Fig. 4.4) was broadly similar to that of Raikow despite its rearrangement of species and genera. Again, the honeycreepers fell into two main groups, with the finches in one and the thin-billed taxa in the other. However, I grouped the parrotbill with the thinbilled birds (see Chapter 5).The genera Paroreomyza and Melamprosops were not included because of my belief at the time that they were non-drepanidine, and I was still uncritically following Greenway (1939) and Amadon (1950) in regarding Dysmorodrepanis as a freak. Unfortunately, the classification based on this phylogeny was, like that of Raikow (1977b), eclectic rather than cladistic and produced some paraphyletic taxa.

Biochemical and molecular studies Until the recent development of PCR technique, which has allowed extraction of DNA from old study skins and even prehistoric remains (Saiki et al. 1988), phylogenies of Hawaiian honeycreepers based on biochemical or molecular data suffered

from a lack of comparable research material and often differed in the taxa studied. Nevertheless, we can make some comparisons.The study by Johnson et al. (1989), discussed previously in the context of the origins of the honeycreepers, is equally enigmatic with regard to relationships within the group. They presented several different trees, the first of which (Fig. 4.5a) was produced by a numerical program called UPGMA (Sneath and Sokal 1973), which produces a phenogram based on a measure of genetic distance.The most surprising hypotheses of that tree include the following: (1) as in previous studies, the honeycreepers divide into two clades, but in this case the two ‘creeper’ genera Paroreomyza and Oreomystis form one, and the rest of the honeycreepers the other; (2) the Kaua‘i ‘Amakihi is more closely related to ‘akepas (at least Loxops caeruleirostris) than to the other ‘amakihis; and (3) the ‘Anianiau is in a clade with the drepanidine finches, whose finchlike characters are derived secondarily from a thin-billed, tubular-tongued ancestor. With the same data, Johnson et al. (1989) conducted a cladistic analysis using PAUP (Fig. 4.5b) that produced different, but in my opinion equally counterintuitive, results. In the latter, Paroreomyza and Oreomystis are sister taxa but fall in the middle of the drepanidine radiation, the two drepanidine finch genera are not sister taxa (meaning that drepanidine/cardueline similarities evolved by convergence not once but twice), the ‘I‘iwi is sister group to the Laysan Finch and not to the ‘Apapane (even though no other ‘red and black’ honeycreepers were included), and the ‘Anianiau is not in that cluster. A different study of allozymes (Fleischer et al. 1998) produced a fundamentally different tree (Fig. 4.5c), but one that had at least a few similarities to those of Johnson et al. (1989). Again, the closest outgroup to the honeycreepers is not the expected cardueline finches, but a clade that includes the carduelines and American blackbirds (not quite the same as Johnson et al., but close). Again, the Oreomystis creepers and the ‘alauahios form a clade that is sister to all the other honeycreepers. Although Fleischer et al. (1998) made much of that point of agreement, their other hypothesised relationships, though not well resolved, differed strikingly from

58 The Hawaiian Honeycreepers Carduelini Laysan Finch 'l'iwi Palila 'Anianiau 'Apapane 'Akeke'e 'O'ahu 'Amakihi Hawai'i 'Amakihi Maui 'Amakihi Kaua'i 'Amakihi 'Akikiki (a)

(b)

Maui 'Alauahio Purple Finch Red Crossbill Saffron Finch Scarlet Tanager 'Apapane 'Anianiau 'Akeke'e Kaua'i 'Amakihi O'ahu 'Amakihi Maui 'Amakihi Hawai'i 'Amakihi Palila Laysan Finch 'l'iwi 'Akikiki Maui 'Alauahio 'Akikiki Maui 'Alauahio _ 'Akepa 'Akeke'e _ 'Akohekohe 'Anianiau 'Apapane Kauai' 'Amakihi Maui 'Amakihi Hawai'i 'Amakihi 'l'iwi Laysan Finch Palila Purple Finch Pine Siskin

(c)

House Finch Red-wing, Blackbird

those suggested by Johnson et al. (1989). For example, the Kaua‘i ‘Amakihi clusters with the other ‘amakihis; the ‘akepas form an unresolved clade with ‘Anianiau, ‘Apapane, and ‘Akohekohe that does not include ‘I‘iwi; and Laysan Finch and Palila form an unresolved trichotomy with the clade that includes ‘I‘iwi, ‘amakihis, and the ‘akepa/‘apapane clade. Importantly, this topology keeps all the tubular-tongued taxa in a single clade that requires no reversals of this important character. The earliest use of DNA in honeycreeper systematics was that of Sibley and Ahlquist (1982), but DNA–DNA hybridisation does not provide sufficient resolution to be useful below the family or subfamily level. In the past decade, the most influential centre for DNA research on Hawaiian honeycreepers has been the laboratory of R. C. Fleischer and his colleagues. Their earlier research used restriction-fragment analyses to develop ‘mitochondrial gene trees’ (Tarr and Fleischer 1995) which ‘may differ from the actual phylogeny among closely related organisms.’Tarr and Fleischer (1993) used mitochondrial DNA to investigate the ‘amakihis and ‘Anianiau at the species level and later (1995) expanded their analysis to include representatives of most living genera (as recognised at the time).The species-level conclusions of the earlier study will be discussed in Chapter 5, but noteworthy here is that the ‘Anianiau in that study consistently formed a clade with the Kaua‘i and O‘ahu ‘amakihis that is sister group to the Hawai‘i ‘Amakihi. The 1995 study presented two different trees (Fig. 4.6a, b), one a cladogram produced by PAUP, the other a genetic distance tree. In the distance tree (a), the ‘Anianiau is sister group to other ‘amakihis as before, but in the parsimony tree (b) it is far removed from ‘amakihis as the sister group to all thin-billed honeycreepers except ‘Akikiki and Maui ‘Alauahio. Unlike the two allozyme studies discussed above, Tarr and Fleischer (1995) did not cluster ‘Akikiki and ‘alauahio together in either tree; in both, the former is sister group to Telespiza 4.5 Phylogenies based on allozymes; redrawn in common style with nomenclature standardised: (a) Johnson et al. (1989, fig. 1, UPGMA method); (b) Johnson et al. (1989, fig. 4, PAUP method); (c) Fleischer et al. (1998, fig. 2b, allozymes).

'Akeke'e _ 'Akepa

Hawai'i 'Amakihi Maui 'Amakihi Kaua'i 'Amakihi 'Anianiau _ 'Akepa

Hawai'i 'Amakihi

'l'iwi _ 'Akohekohe

Kaua'i 'Amakihi

Hawai'i Creeper

'Akeke'e

'Apapane _ 'Akohekohe

Hawai'i 'Amakihi _ 'Akepa

'l'iwi

(a)

'Apapane

Maui 'Amakihi

'Apapane _ 'Akohekohe

'l'iwi

Maui Parrotbill

'Anianiau

Maui 'Alauahio

'Akikiki

Laysan Finch

'Akikiki

Laysan Finch

'Akikiki

Crossbill

Maui 'Alauahio

Maui 'Alauahio

Rosefinch

House Finch

(b)

House Finch

(c)

Titmouse Carduelini Nihoa Finch Laysan Finch

'Apapane

Palila

'l'iwi _ 'Akohekohe

Maui 'Amakihi Hawai'i 'Amakihi

Kaua'i 'Amakihi O'ahu 'Amakihi

Kaua'i 'Amakihi

Hawai'i 'Amakihi

O'ahu 'Amakihi

Maui 'Amakihi _ 'Akepa

'Apapane _ 'Akohekohe

'Akeke'e

'l'iwi

Hawai'i Creeper

Hawai'i Creeper

Palila Laysan Finch

'Akeke'e _ 'Akepa

'Anianiau

'Anianiau

Maui Parrotbill __ 'Akiapola'au

'Maui Parrotbill __ 'Akiapola'au

Nihoa Finch

'Akikiki

Po'o-uli

Maui 'Alauahio

Maui 'Alauahio

House Finch (d)

Green Honeycreeper

(e)

'Akikiki

4.6 Phylogenies based on molecular data; redrawn in common style with nomenclature standardised: (a) Tarr and Fleischer (1995, fig. 9.2, genetic distance); (b) Tarr and Fleischer (1995, fig. 9.3, PAUP); (c) Feldman (1997, as shown in Freed 1999, fig. 7.3); (d) Fleischer et al. (1998, fig. 2a, mtDNA); (e) Fleischer et al. (2001, fig. 1).

60 The Hawaiian Honeycreepers (the representative finch-billed genus), whereas the latter is either sister to all other honeycreepers (parsimony) or to the cardueline House Finch (distance) which, if true, would mean Paroreomyza is not a drepanidine genus. Using DNA sequence data and a different subset (n ⫽ 9) of taxa, Feldman (1994; phylogeny reproduced in Freed 1999) developed a topology (Fig. 4.6c) that presents somewhat different groupings. In this phylogeny, the ‘red and black’ group is sister to the Hawai‘i Creeper rather than the ‘amakihis, and that group forms an unresolved trichotomy with the ‘amakihis and ‘akepas.That cluster is then sister to the parrotbill, whose position is difficult to interpret because no drepanidine finches were included in the study.The ‘Akikiki and Maui Creeper cluster as a clade that is the sister group to all other honeycreepers, as in the allozyme studies of Johnson et al. (1989) and Fleischer et al. (1998). The first DNA sequence results from Fleischer’s laboratory (Fleischer et al. 1998) resulted in a phylogeny (Fig. 4.6d) that was roughly similar to Feldman’s but which included 18 taxa. Important characteristics of this phylogeny are: (1) the ‘Anianiau is nowhere near the ‘amakihis, but instead forms a clade with Maui Parrotbill and ‘Akiapola‘au, two species not included in previous studies; (2) that clade is sister group to the finchlike genera; (3) the O‘ahu ‘Amakihi forms a clade with Hawai‘i ‘Amakihi rather than with the Kaua‘i ‘Amakihi, contra the earlier restriction-fragment study (Tarr and Fleischer 1993); (4) the ‘red and black’ honeycreepers form a natural group that is sister to the ‘amakihis; and (5) the Hawai‘i Creeper, herein considered congeneric with the ‘Akikiki and not included in previous DNA studies, clusters instead with ‘akepas as sister group to the ‘amakihi/ red bird clade. The latest results from Fleischer’s team (Fleischer et al. 2001) adds Po‘o-uli to the species studied but rather than, as one would expect, that species simply appearing at the proper place in the previous topology, several of the fundamental hypothesised relationships are inexplicably different (Fig. 4.5e). This time, the drepanidine finches are sister to the rest of the honeycreepers (including Po‘o-uli), but do not form a single clade. The ‘Anianiau/parrotbill/ Akiapola‘au clade remains, but is sister to the

‘akepa/Hawai‘i Creeper clade, which had previously been sister to an ‘amakihi/red bird cluster. Also, the pattern of relationships among the ‘amakihis, which had been fundamental to the ‘conveyor belt’ premise of their 1998 paper, reverts to the topology of the earlier study (Tarr and Fleischer 1993) in which Kaua‘i and O‘ahu ‘amakihis are clustered as a sister group to the Maui/Hawai‘i species. Such flip-flops give pause to non-molecular systematists. To be fair, the differences between studies may not be as great as they at first appear. Several terminal clades (i.e. ‘Akikiki ⫹ ‘alauahios, ‘akepas ⫹ creeper, the ‘red’ birds, and, surprisingly, parrotbill ⫹ ‘Anianiau ⫹ ‘Akiapola‘au) are consistent in these studies, although how these clusters are interrelated seems as yet unresolved. The last clade is particularly interesting because its members exhibit very disparate phenotypes, but the ‘Akiapola‘au/parrotbill relationship, at least, has been upheld in new analyses of traditional data (see below). Robert Fleischer (pers. comm.) himself cautions that the honeycreepers have radiated so quickly that the details of their interrelationships may be more difficult to discern that those of other groups of birds, and the current level of resolution in DNA studies may not be up to the task.

New phenotypic studies Using a PAUP analysis of osteological characters, James (1998) developed a phylogeny (Fig. 4.7) that includes many of the prehistoric genera described by James and Olson (1991), making it the most comprehensive study to date in terms of taxa covered. However, as might be expected from fragmentary remains, many data points are missing from the matrix. Her topology (published in Fleischer et al. 2001 without subfossil taxa) does not differ strikingly from earlier ones based on phenotypic characters (Raikow 1977b, 1985; Pratt 1979a), but a basal split between the ‘green’ and the ‘red and black’ honeycreepers is not evident. As in the earlier studies, the drepanidine finches are considered basal to the radiation, but the Maui Parrotbill is grouped with them. Uniquely in this study, the Hawai‘i Creeper is basal to a large clade that includes the genus Hemignathus as delineated by the AOU (1998) plus the ‘red’ group. Within that clade, the Kaua‘i

Origin and evolution 61 ‘Amakihi remarkably does not cluster with the other ‘amakihis, but is the sister group to the rest of the clade. The ‘alauahios are considered a sister group to the ‘Akikiki (which is not shown as closely related to the Hawai‘i Creeper). I believe many of these results may be the result of misinterpretation of convergence as synapomorphy (Pratt 2001). For example, the ‘red’ birds have bills superficially shaped like some of those in Hemignathus, but the defining synapomorphies of the group are distinctive features of plumage and vocalisations. Likewise, the parrotbill may cluster with the finches simply as a byproduct of secondary thickening of its bill, whereas behavioural, ecological, genetic, and plumage data all indicate that its relationships lie elsewhere (see below and Chapter 5). Finally, the topology clearly shows that the enlarged genus Loxops of James and

House Finch Pine Grosbeak Nihoa Finch Laysan Finch Makawehi Finch Palila Maui-nui Finch Koa-Finch sp.? King Kong Grosbeak Kona Grosbeak Wahi Grosbeak Lesser Koa-Finch Greater Koa-Finch Ridge-billed Finch Po'o-uli _ _ O'u _ Maui _ Parrotbill Lana'i Hookbill Kiwi Shovelbill Pololei Shovebill 'Akikiki _ _ Kakawahie Maui 'Alauahio Greater 'Amakihi Maui-nui Gaper Straight-billed Gaper _ 'Akepa 'Akeke'e Hawai'i Creeper Akialoa _ _sp? 'Akiapola'au Nukupu'u 'Greater 'Akialoa' Giant Akialoa sp. Lesser 'Akialoa Kaua'i 'Amakihi 'Anianiau Hawai'i 'Amakihi 'l'iwi Hawai'i Mamo Black Mamo 'Apapane Laysan Honeycreeper _ 'Akohekohe Kaua'i Palmcreeper O'ahu Palmcreeper sp. _ 'Ula-'ai-hawane

4.7 James’s (1998) osteological phylogeny including subfossil taxa.

Olson (1991) and Olson and James (1995), as well as the subfossil genus Vangulifer, are polyphyletic. Because so much new information had accumulated in the ensuing two decades and methods of phylogenetic analysis (such as the computer program PAUP) had improved considerably, I decided to reprise my earlier work (Pratt 1979a).The result (Pratt 2001b; Fig. 4.8) is a much more robust phylogeny based on 39 highly varied phenotypic characters that are known for most species. Its hypotheses can be summarised as follows (Pratt 2001b): (1) Hawaiian honeycreepers, including Po‘o-uli and the ‘alauahios, are monophyletic and a subgroup of the Fringillidae; (2) the parrotbill is related to the heterobills rather than to drepanidine finches; (3) the large genus Hemignathus used herein is strongly supported by phenotypic characters; (4) Chaffinches Cardueline finches Telespiza finches Palila Kona Grosbeak Koa-finches _ _ 'O'u _ Lana'i Hookbill __ 'Akiapola'au Nukupu'us Maui Parrotbill 'Akialoas 'Amakihis Greater 'Amakihi _ 'Akepa/'Akeke'e 'Akikiki Hawai'i Creeper 'Anianiau _ 'Ula-'ai-hawane _ 'Akohekohe 'Apapane 'l'iwi Hawai'i Mamo Black Mamo Po'o-uli _ _ Alauahios/Kakawahie

4.8 New phylogeny based on a broad spectrum of phenotypic characters (Pratt 2001, fig. 1c).

62 The Hawaiian Honeycreepers although the parrotbill may belong in Hemignathus, the ‘Anianiau does not; and (5) the Hawai‘i Creeper and ‘Akikiki are closely related, based on a large suite of phenotypic synapomorphies, but together they are not closely related to the ‘alauahios (Paroreomyza). Results from DNA studies that show a relationship between the Hawai‘i Creeper and ‘akepas, or from osteological studies that suggest a relationship to ‘amakihis (Olson and James 1995), could result from a past hybridisation event (Pratt 2001b), although none has been demonstrated. The taxonomic implications of these findings are discussed in the next chapter. Since 1998, much new subfossil material has come to light (H. James, pers. comm.) that has helped to fill in much of the ‘missing data’ from James’s (1998) matrix. James’s (2004) new topology, which became available too late to be shown here, differs from her 1998 one in several important respects. In the update, the problematical Hemignathus/red bird clade remains, but the Kaua‘i ‘Amakihi clusters with the others as sister group to the ‘red’ species, and this pairing is sister to an Akialoa/heterobill clade; the Hawai‘i Creeper is now in a clade with the ‘akepas (which brings the osteological topology into agreement with the Fleischer team’s DNA one, at least on this point) that is sister to one that clusters the Greater ‘Amakihi with the subfossil genus Aidemedia; and the fossil genus Xestospiza is shown to be polyphyletic, with X. conica related to the Hawaiian grosbeaks and koafinches. Clearly, we are far from consensus on the phylogeny of Hawaiian honeycreepers. For nonmolecular systematists, a phylogeny generated from a combined data matrix of osteological and other phenotypic characters would be a logical next step, one likely to be highly instructive. Ultimately, I believe DNA studies may provide definitive answers to most, if not all, phylogenetic questions; but until the results from the labs become more consistent (to the layman, they seem to change in fundamental ways with each new study even from the same lab), we cannot base a classification solely on them. I also believe DNA researchers should pay more attention to phenotypic data as an indicator of where laboratory results may be in error or where past hybridisation may have produced misleading results. On the

other hand, those of us who practice systematics based on more traditional data ignore the molecular results at our peril. If ornithologists have any ‘article of faith’, it is that eventually all avenues of investigation will reach the same conclusions.

Timing of honeycreeper evolution When the colonisation of Hawai‘i and the subsequent radiation of honeycreepers actually occurred has been the subject of considerable debate. Most early naturalists, in light of the great morphological diversity among drepanidines, assumed the group’s history to be ancient, perhaps beginning in the Eocene (Perkins 1913). Zimmerman (1948), however, appreciated how rapidly evolution could proceed in an isolated archipelago. With little geological information at his disposal, Amadon (1950: 237) surmised that life in the Hawaiian Islands could be as ancient as others had suggested, but presciently stated that ‘some of the data indicate that adaptive radiation in this family . . . has also taken place more rapidly than might be expected.’ Following the development and general acceptance of plate tectonics theory in the 1960s, new hypotheses began to emerge. The discovery of the Emperor Chain of seamounts stretching northward almost to the Aleutians from Kure Atoll led many to surmise that the honeycreepers might have evolved well before the present main islands existed, and island-hopped to their present locations. In any case, the initial colonisation has to have been over distances comparable to those we see today because when the Emperor Seamounts were high islands, they were in the position of the present Hawaiian Islands (Carson and Clague 1995). Schlanger and Gillett (1976) suggested that the Laysan Finch and Laysan Honeycreeper, as well as the only other endemic passerine, the Millerbird, were relicts from a time when Laysan was a high, forested island. To me (Pratt 1979a), morphological evidence suggested otherwise, and Olson and Ziegler (1995) argued persuasively that the honeycreepers known from Laysan and Nihoa colonised them secondarily from the main islands.

Origin and evolution 63 Based on biochemical studies, Sibley and Ahlquist (1982) estimated that the honeycreeper lineage originated 15–20 Ma, and Johnson et al. set the beginning at 7–8 Ma, both dates well before any of the current high islands emerged from the sea, and Givnish et al. (1995) believed the earlier date coincided with the estimated age of Hawaiian lobelioids with which the honeycreepers apparently co-evolved (but see below). However, recent geological data (Carson and Clague 1995) reveal some major problems for the hypothesis that the Drepanidinae evolved before the emergence of Kaua‘i 5.1 Ma. Nihoa and Necker, the next islands up the chain, technically would still have been ‘high’ islands when Kaua‘i emerged, but even at their maximum elevations of 1300 m and 1100 m, respectively, would never have been as high as Kaua‘i at its maximum (2600 m). By the time Kaua‘i was capable of supporting a diverse flora and fauna, they would have been much lower and smaller. Furthermore, the distance between Nihoa and Kaua‘i is roughly twice that existing today between Kaua‘i and O‘ahu. All of these conditions make the older islands much less likely as source areas for organisms colonising the main Hawaiian Islands. As Carson and Clague (1995: 18) state: ‘It seems hardly possible that propagules arising from these low islands could be a source of a significant number of suitable colonists for the higher-altitude areas of the newer islands.’Thus a geological bottleneck apparently occurred that broke the chain of island hopping between Nihoa and Kaua‘i. Recently, more sophisticated methods of lineage dating demonstrate a much younger age for the honeycreeper radiation than some had suggested, but one correlated with the geological history of the current high islands (Tarr and Fleischer 1993, 1995; Fleischer et al. 1998; Fleischer and McIntosh 2001). These studies indicate that the drepanidine radiation did not occur outside the extant high Hawaiian Islands, and that it cannot be older than about 5 million years. Interestingly, the other avian lineages known from the main Hawaiian Islands also have ages compatible with a pre-Kaua‘i bottleneck (Fleischer and McIntosh 2001).These findings are an indication of how severe the constriction of inter-island colonisation was at that point and show

that the islands from Kaua‘i eastward were nearly a blank slate with regard to birds. The most recent estimates are that the drepanidines split from a cardueline ancestor 5–6 Ma, and experienced their first ‘basal split’ about 4 Ma (Fleischer and McIntosh 2001). They colonised the islands as each in turn emerged from the sea, as if on a ‘geological conveyor belt’ (Fleischer et al. 1998). Groth (1994) and Fleischer and McIntosh (2001) offered several possibilities, including the use of ‘continental biogeographic points in their calibration’ and the use of outgroups too distantly related, to explain the apparently inflated age estimate of Sibley and Ahlquist (1982).

Evolution demonstrated by Hawaiian honeycreepers Hawaiian honeycreepers are the supreme avian example of adaptive radiation, far exceeding the degree of variation seen in any other bird group of equivalent taxonomic rank. Except for the Cocos Finch and the Warbler Finch, all of the famous Galapagos finches look like finches (Bowman 1961; Castro and Phillips 1996; Lack 1947). The New Zealand wattlebirds (Callaeidae) exhibit a wide variational range (Amadon 1950), but have few known species.The only real challenges to the honeycreepers’ pre-eminent position come from the vangas of Madagascar (Langrand 1990) or the birdsof-paradise of New Guinea (Frith and Beehler 1998), both of which include sickle-billed species among short-billed ones. But from a zoogeographic perspective, these taxa represent a continental rather than an archipelagic situation. The honeycreeper radiation has produced variations that include many examples of parallel and convergent evolution. Sometimes the attempt to fill a particular ecological niche resulted in unique and even bizarre drepanidine adaptations that would have amazed Rube Goldberg. Other honeycreeper adaptations simply have no close mainland analogues. This section begins with a survey of microevolutionary processes among Hawaiian honeycreepers, then moves on to broader questions about adaptive radiation.

64 The Hawaiian Honeycreepers

Speciation, double invasion, and character displacement The fundamental driving force of all evolution, as Bock (1970) pointed out using honeycreepers as his model, is repeated cycles of (1) allopatric speciation followed by (2) secondary sympatry, which produces (3) further divergence, called character displacement. Bock (1970) believed that this fundamental model, as promulgated by Mayr (1963) and others, was sufficient to account for large (macroevolutionary) changes and well as those at the species (microevolutionary) level. Schluter (1988), using drepanidine finches as one test case, not only refined the concept of ecological character displacement and discounted other hypotheses for adaptive divergence, but showed that the lack of competition from other taxa on remote archipelagos could greatly enhance its effects to produce the spectacular radiations observed in Hawai‘i and the Galápagos. For many years debate (reviewed by Gosline 1968 and Grant 1998b) raged as to whether speciation events are driven by random events such as the ‘founder effect’, in which the small genetic subset represented in a founding population leads to a ‘genetic revolution’ and immediate differences not driven by selection (Mayr 1942; Carson and Templeton 1984), or whether the changes following colonisation are adaptations to the new environment (Barton and Charlesworth 1984).The 1967 introduction of a small (59 males, 51 females) population of Laysan Finch to Pearl and Hermes Reef (Conant 1988a,b) inadvertently provided an experiment to test both hypotheses.Within the short span of two decades, the finches at Pearl and Hermes developed significant differences in bill shape that corresponded to size differences in the fruits of the puncture vine nohu among the various islands and islets the birds inhabited. Nohu is a minor component of the flora (and finch diet) on Laysan, but a major one at Pearl and Hermes. The rapidly evolved bill differences seem clearly the result of adaptation to available foods rather than to genetic drift or founder effects (Freed et al. 1987a; Conant 1988a,b). In a follow-up study, Fleischer et al. (1991) found that although the parent population on Laysan, which experienced a crash earlier

in the century, did have reduced allozyme variation as compared to other birds, the translocated populations surprisingly had greater variability than the source population. Founder effects were apparently negated by selection favouring heterozygosity. While these studies did not eliminate founder effects as a force in allopatric speciation, they are consistent with the finding of Grant and Grant (1998) that selection is far more important, at least in birds (Grant 1998b). Speciation patterns in Hawai‘i are distinctive because of the linear formational history of the islands themselves, the ‘geological conveyor belt’ of Fleischer et al. (1998). Species arise by successive colonisations of new islands as they become habitable, followed by adaptation that produces genetic isolation. Once speciation is achieved, reverse colonisations from younger to older islands are possible. Such back-colonisations are often called double invasions, but in the Hawaiian context true double invasions are not as easy to discern as in the case of a nearby island being repeatedly colonised from a continental source (Grant and Grant 1998). Such patterns of linear speciation with subsequent reverse colonisations have been shown for a wide variety of Hawaiian organisms (Wagner and Funk 1995). These Hawaiian patterns may represent what has been called the ‘taxon cycle’ (Greenslade 1968; Ricklefs and Cox 1972; Pregill and Olson 1981), the insular version of Brown’s (1957) ‘centrifugal speciation’.The taxon cycle, as summarised by Grant and Grant (1998), is ‘the evolutionary trend from a single, broadly adapted generalist species that colonises an archipelago to several, locally adapted, specialist, and restricted species.’ ‘Recycling’ can occur if a specialist species becomes a generalist and expands secondarily throughout the archipelago. Pratt (1979a) made the first attempt to apply the concept to Hawaiian honeycreepers, and recent molecular studies (Tarr and Fleischer 1993, 1995; Fleischer et al. 1998) have shown that at least some speciation patterns are consistent with the taxon cycle. Ricklefs and Cox (1972) divided the cycle into four stages clearly recognisable among Hawaiian honeycreepers: (1) an undifferentiated species is found throughout the archipelago (‘Apapane, ‘I‘iwi, and

Origin and evolution 65 ‘O‘u); (2) each island has its own subspecies (or allopatric species) and some island populations may be lost (‘amakihis, ‘akialoas, heterobills, ‘akepas, Paroreomyza); (3) well-differentiated species have fragmented or disjunct distributions (Oreomystis, Ciridops, Chloridops, koa-finches, mamos, Telespiza); and (4) only single-island endemics remain (Po‘o-uli, Maui Parrotbill, ‘Anianiau, ‘Akohekohe). The following honeycreeper complexes were selected to emphasise double invasions and/ or character displacement at various stages of the taxon cycle. The aberrant genus Paroreomyza provides perhaps the most exemplary demonstration of how speciation, double invasion, and character displacement work together to produce diversity. It is a simpler case than others because it involves only O‘ahu and Maui-nui. The ‘alauahios are known both historically and as subfossils from O‘ahu (P. maculata) and Maui-nui (P. montana), where the historical distribution was only on Maui and Lana‘i, but with similar-sized prehistoric remains from Moloka‘i ( James and Olson 1991). The red, large-billed Kakawahie P. flammea is not represented in the palaeontological record and was historically found only on Moloka‘i, so must therefore have been sympatric with the yellow, small-billed P. montana on Maui-nui (Pratt 1992b). The O‘ahu ‘Alauahio is also yellow (males only) and its bill is intermediate in size. The obvious scenario is that the original Paroreomyza underwent allopatric speciation on O‘ahu and Maui-nui, and then the O‘ahu species reinvaded Maui-nui, where sexual selection led to differences in plumage colour and other selective forces resulted in character displacement in bill size (Pratt 1979a, 1992b; Pratt and Pratt 2001). (Note: H. F. James (in H. and P. Baker 2000) stated that the report ( James and Olson 1991) of fossils of P. montana from Moloka‘i was overstated and that, though they are smaller than P. flammea, the remains are too fragmentary for positive identification. But unless a third species inhabited Mauinui, the above scenario remains reasonable.) The nectarivorous sickle-bills Drepanis spp. show how the taxon cycle might begin anew. The two mamos are specialised single-island endemics that are in Stage 3 approaching Stage 4.The ‘I‘iwi,

however, is a Stage 1 generalist that occurs undifferentiated throughout the main islands. It may be the most recently evolved species in the complex or it may be an old species that recently made an adaptive ecological shift, perhaps to feeding in ‘ohia (Pratt 1979a), which enabled it to restart the taxon cycle. However, an equally plausible hypothesis for the lack of inter-island differentiation in the ‘I‘iwi (as well as the other two undifferentiated species) is that these birds’ habits of making long, high flights, possibly associated with searching for their nectar (‘I‘iwi, ‘Apapane) or fruit (‘O‘u) resources, predisposes them to frequent inter-island dispersal (Freed et al. 1987a) that prevents their proceeding further than Stage 1 of the taxon cycle. The ‘amakihis are apparently in Stage 2 of their taxon cycle, with each island (the Maui-nui group considered as one) having a distinctive form. They are all now considered separate species except that the Maui-nui and Hawai‘i forms have diverged less and are ranked as subspecies (Pratt and Pratt 2001; also see Chapter 5). Fleischer et al. (1998) used molecular data to demonstrate that the ages of the island forms corresponded with the ages of the islands they inhabit. The degree of phenotypic differentiation follows an identical pattern. Two supposed double invasions by ‘amakihi, one on Kaua‘i and one on The Big Island, have been frequently mentioned in the context of character displacement (Amadon 1947; Bock 1970), but neither of the ‘doubles’ may actually belong to this complex (Pratt 2001b). On Kaua‘i, the representative ‘amakihi has a much larger bill than the others along with numerous other differences that show it to have reached the species level (Pratt et al. 1987;Tarr and Fleischer 1993; Conant et al. 1998; Pratt and Pratt 2001). Bock (1970) and Pratt (1979a) attributed its distinctive features to character displacement induced by the presence of the ‘Anianiau or ‘Lesser ‘Amakihi’. Between them, the Kaua‘i ‘Amakihi and the ‘Anianiau subdivide and expand the ‘amakihi niche as expressed in bill size and shape. Recent studies (Tarr and Fleischer 1995; Conant et al. 1998; see also Chapter 5) have shown the ‘Anianiau to be a singleisland endemic in Stage 4, rather than a doubleinvading ‘amakihi, but character displacement may nevertheless be the cause of the observed divergence.

66 The Hawaiian Honeycreepers Schluter (1988) showed that character displacement can occur even when the competing species are not closely related.A better candidate for a double invasion event in this complex is at the opposite end of the archipelago on Hawai‘i, where the ‘typical’ Hawai‘i ‘Amakihi is joined by the Greater ‘Amakihi. Here, instead of subdividing the niche, the larger species has developed a distinctive bill morphology, but at least it is similar enough to be retained tentatively in the genus Hemignathus with the other ‘amakihis (but see Chapter 5; James 1998; Pratt 2001b). Whether it actually belongs to that clade, and therefore represents a true double invasion, is still problematical. The subgenus Akialoa is another Stage 2 complex with several double invasions. Each major island had its own endemic historically known species, but these were not strongly differentiated and were often considered conspecific (Bryan and Greenway 1944; Olson and James 1982b; Pratt et al. 1987) or divided into two species (Amadon 1950) until the recent analysis of prehistoric remains.The discovery of remains of sympatric second species of Akialoa on Kaua‘i, O‘ahu, Maui-nui, and Hawai‘i (the latter two undescribed) forces the recognition of all forms as full species (Olson and James 1995). The Hoopoe-billed Akialoa, the second species on Kaua‘i and O‘ahu ( James and Olson 1991), was only slightly larger than sympatric counterparts but had qualitative differences that indicate character displacement of a different sort. The sympatric ‘akialoas may have had different diets or feeding methods even though their bills were of similar length. Interestingly, the undescribed form from Hawai‘i was a larger one sympatric with the smallest member of the complex, almost surely an example of character displacement. Until recently the heterobills (nukupu‘us and ‘Akiapola‘au) were assumed to be a simple example of Stage 2, with four allopatric species, one of which has a distinctive bill morphology.The latter now can be seen as a drastic example of qualitative character displacement, because a subfossil giant nukupu‘u, the largest drepanidine yet discovered, has recently been unearthed on Hawai‘i ( James and Olson 2003.), too late to be fully included in this book.

The finches of the genus Telespiza present an intriguing case of Stage 3 of the taxon cycle, marked by multiple invasions and fragmentary distributions. However, because the main-island distributions are based on prehistoric remains, the record is likely to be incomplete.Thus the following generalisations are tentative.All species are qualitatively similar, and species limits are based solely on measurements ( James and Olson 1991). Subfossils that have been identified as Laysan Finch and the Nihoa Finch have been found on O‘ahu (T. cantans), and Moloka‘i (both species), the latter being the only island where these two are known to have been sympatric. (The specific identity of these remains should, in my opinion, be regarded as tentative; plumage and vocal differences could easily have served as reproductive isolating mechanisms between species that might be osteologically indistinguishable. Perhaps future studies of ancient DNA (Pääbo 1989; Cooper et al. 1996; Cooper 1997) will confirm the identifications.) A smaller species, T. ypsilon, is also known from Moloka‘i and Maui, and Maui had an even smaller one, as yet undescribed because remains are too fragmentary, so Maui-nui may have had four sympatric Telespiza finches with a stepwise size hierarchy ( James and Olson 1991; see also Chapter 6). Another small species, T. persecutrix, was present on Kaua‘i and O‘ahu, and sympatric with T. cantans on the latter. The measurements on which these five species are based approach each other but do not overlap.The circumstantial case here for character displacement following multiple invasions is compelling. In their close measurements and extensive sympatry, these finches resemble Darwin’s finches more than any other Hawaiian honeycreepers. The koa-finches Rhodacanthis of The Big Island are unquestionably an example of double invasion and character displacement.They were closely sympatric, even syntopic (Munro 1960), and differed only in bill size and male coloration (Olson 1999). The bill differences are undoubtedly the result of natural selection for differential feeding, but the colour differences are more likely attributable to sexual selection to reinforce genetic isolation (Ziegler 2002). Subfossils of Rhodacanthis have been found on O‘ahu and Maui, but their specific identity cannot

Origin and evolution 67 be determined on the basis of the limited material available for comparison. Specimens resembling both historically known species are known from Maui, but the O‘ahu population has ambiguous characters and could be a third species not yet described ( James and Olson 1991). Consequently, we cannot say at present which of the two Hawai‘i species was the second invader. The Hawaiian grosbeaks Chloridops spp. appear to be a Stage 3 complex with one or two possible examples of double invasion and character displacement, as described by James and Olson (1991).Two allopatric species, the Kona Grosbeak on Hawai‘i and Wahi Grosbeak from Maui and O‘ahu, have been described and a third form is known from Kaua‘i but not yet named. A smaller unnamed species is also known from Maui. The King Kong Grosbeak, named for its massive bill which is the largest of any drepanidine finch, was sympatric with the Wahi Grosbeak on O‘ahu and may represent a double invasion, but James and Olson (1991) suggest that it might eventually prove to belong to a different lineage.

Convergence, homoplasy, and parallelism These three processes are actually nuances of a single phenomenon by which one organism comes to resemble another to which it is unrelated or only distantly related genetically. Parallelism is a special case in which similar (i.e. parallel) adaptations appear in different, but closely related lineages. For example, the development of sickle-shaped bills in both the ‘red and black’ lineage and the hemignathine complex of Hawaiian honeycreepers would be parallelism. Because of differences in structural details, the falcate bill shapes probably were independently derived even though these two lineages may be sister groups. The selective force (co-evolution with plants; see Chapter 9) was the same for both.At a higher level, the development of a similar array of bill morphologies in cardueline and drepanidine finches (Fig. 6.1) can be viewed as parallelism. At more distant levels of relationship, the distinction between parallelism and convergence is mostly semantic, but a fairly compelling

example involves the development of olive green birds with long sickle bills used for both insectivory and nectarivory in both the Drepanidinae (‘akialoas) and the Meliphagidae (New Guinea longbills; Beehler et al. 1986), both probably derived from nectarivorous ancestors. Other insectivorous passerines with bills that resemble those of ‘akialoas, including Afro-palaearctic hoopoes (Upupidae), African woodhoopoes (Phoeniculidae), Neotropical scythebills, Asian spiderhunters, and the Sicklebilled Vanga of Madagascar, have similarly shaped bills but somewhat different, though overlapping, feeding habits. For example, spiderhunters take insects and nectar but are not bark-pickers ( Jeyarajasingam and Pearson 1999), while scythebills (Hilty and Brown 1986) and Sickle-billed Vanga (Morris and Hawkins 1998) pick bark but do not take nectar. These are quintessential examples of convergence as discussed by numerous authors (e.g. Amadon 1950; Olson and James 1982b; James and Olson 1991; Ziegler 2002). Homoplasy is essentially a synonym for convergence that has somehow gained favour in the jargon of laboratory and computer age systematists, although it is most often used when comparing fairly closely related taxa. Most convergence/ homoplasy (including parallelism) is the result of similar selective forces acting in similar habitats. Nearly all Hawaiian honeycreepers have some counterpart elsewhere in the world with which they could be considered convergent to a greater or lesser extent. Some similarities, like that of Hawaiian grosbeaks to other large-billed finches in several families, or the convergence of crossbills and ‘akepas (see Chapter 6), are so obvious as to be unremarkable, but others are more subtle. Often convergence involves only one feature, as in the development of a nasal operculum (see Chapter 6) in many nectarivorous but otherwise dissimilar birds, but it can involve everything from plumage colour to bill morphology to vocalisations, or any combination thereof. Birders familiar with Hawaiian honeycreepers are often struck by the superficial resemblance between the ‘Apapane and Myzomela honeyeaters of Micronesia and Australasia. Even within the archipelago, some honeycreepers seem to be convergent with

68 The Hawaiian Honeycreepers honeyeaters.The mostly black plumages, not seen in any other fringillids (Clement et al. 1993), of the mamos and ‘Akohekohe are clearly convergent with coloration of the ‘o‘os, and the ‘I‘iwi and ‘Akohekohe are even convergent with meliphagids in vocalisations (see Chapter 7 and species accounts). Fairly close resemblances to ‘amakihi bill shape can be found among honeyeaters of the genera Oedistoma and Lichmera, the Neotropical Bananaquit, and, interestingly, the geospizine Cocos Finch. The latter is isolated from the other Darwin’s finches on tiny Cocos Island, where it uses its short, sharp, decurved bill in an amazing variety of ways; it is a quintessential generalist (Werner and Sherry 1987; Weiner 1994), as are the ‘amakihis (see Chapter 9). The ‘Ula-‘ai-hawane seems to have morphological counterparts (Chapter 6) among both the tanagers (Thraupidae) and flowerpeckers (Dicaeidae), but whether the resemblance was also ecological (Chapter 8) cannot be determined. The three members of Paroreomyza remind most observers of American wood-warblers (Parulidae) in colour, bill shape, and behaviour, while the two Oreomystis are often compared to nuthatches (Sittidae) in behaviour (but the match is not very precise; see Locomotion in Chapter 8).The prehistoric drepanidine gapers Aidemedia spp. have been compared with both meadowlarks Sturnella spp. and starlings Sturnus spp., and the Xestospiza finches resemble American cowbirds. James and Olson (1991) demonstrate that this resemblance is an example of convergence because other features of the bill and skull (discussed above) show Xestospiza to be drepanidine or cardueline rather than icterine. One prehistoric ‘akialoa bill resembles that of a Hoopoe so closely that it was named upupirostris (‘hoopoe-billed’), although whether it also had the hoopoe’s terrestrial feeding habits may never be known (remains do not include identifiable leg bones). Virtually all of the previous comparisons have been made by many observers, but some rather striking drepanidine convergences have not been previously noticed. Alone among major island groups of the tropical Pacific, Hawai‘i lacks fruit-doves (Pratt et al. 1987), the small frugivore niche being filled by Hawaiian solitaires and the drepanidine ‘O‘u. The latter may, in fact, be the honeycreeper equivalent of

a fruit-dove, although any apparent convergence does not involve bill morphology. A surprisingly close parallel is found in the Whistling Dove of Fiji, a similar-sized, sexually dimorphic green bird with a yellow head in males. Even its vocalisations, which are decidedly not dove-like (pers. obs.) resemble the whistled calls of the ‘O‘u! Whether these vocal and colour similarities are really the result of convergence or are merely coincidence no one can say. One supposed example of convergence often cited previously (small parrots and the Maui Parrotbill) is probably not very apt. As discussed in more detail in Chapter 6, the bill of Pseudonestor is parrot-like only in profile, and its manner of use in crushing twigs and mangling bark is quite unlike anything seen in small parrots (although both use the bill in locomotion). More likely convergences with the Maui Parrotbill are found in the vanga genus Xenopirostris, in which the bill is thickened vertically, laterally compressed, and used to strip bark in search of insects (Langrand 1990; Morris and Hawkins 1998), and the Wattled Ploughbill of New Guinea, an insectivore that acts like a ‘chubby parrot-billed creeper’ that ‘digs and hammers on bark’ and ‘hops about, occasionally hanging upside down to check undersides of limbs’ (Beehler et al. 1986: 188), behaviour that closely resembles that of Pseudonestor (Simon et al. 1997). Another supposed example of convergence that, in my opinion, cannot be so characterised is the suite of striking phenotypic resemblances of the ‘Akikiki and the Hawai‘i Creeper. Those who believe these two species are not congeneric (see Chapter 5) must attribute their similarities to homoplasy, but, as I have pointed out elsewhere (Pratt 2001b: 96), if the Hawai‘i Creeper and Oreomystis bairdi prove not to be closely related, ‘the large number and varied character of apparent synapomorphies of these two species will represent one of the most remarkable and noteworthy examples of convergence ever demonstrated.’

Competitive exclusion When the ancestor of the monarchine flycatcher that became what we know as the ‘elepaios colonised Hawaii and spread through the archipelago in the

Origin and evolution 69 first stage of its taxon cycle, it inexplicably skipped Maui-nui.‘Elepaios are known in the fossil record for all three islands they still inhabit, but not from Mauinui ( James and Olson 1991).The explanation may be that the flycatcher niche on Maui-nui was already filled to capacity by Hawaiian honeycreepers that effectively prevented colonisation by Chasiempis.The only historically known honeycreeper that regularly captures insects on the wing is the Maui ‘Alauahio (H. and P. Baker 2000). But the presence of one ‘alauahio, which mainly feeds in other ways, would probably not have been enough (and was not on O‘ahu) to exclude a true flycatcher. James and Olson (1991) have recently described two species of Vangulifer from Maui, the Pololei Shovelbill and the Kiwi Shovelbill. Although they have no close analogy in any other flycatching birds, their unique blunt-tipped bills combine features of several, including todies (Todidae), tyrant flycatchers (Tyrannidae), and monarchs (Monarchidae), and both may have been aerialsally feeders on insects (see Chapter 6). Another possibility is that the exclusion has nothing to do with feeding. ‘Elepaios characteristically use spider silk in construction of their nests (Berger 1981), and the only honeycreepers to do so all belong to Paroreomyza (H. and P. Baker 2000; P. and H. Baker 2000), yet another way in which this enigmatic genus differs from other drepanidine genera. Could spider silk for nest construction be a limiting factor that resulted in competitive exclusion? These interactions may also explain why neither Vangulifer nor Paroreomyza are known from the smaller island of Kaua‘i. O‘ahu is the only island where an ‘elepaio and a Paroreomyza are found together.

Adaptive radiation Although virtually every author on the subject of adaptive radiation has at least mentioned the Hawaiian honeycreepers, which represent by far the most diverse group of birds to have evolved from a common ancestor in a relatively short time span, few have addressed the means by which such variation comes about. Most attention has focused on the result rather than the process. In contrast, Darwin’s finches (Geospizinae) of the Galapagos Islands have been studied in considerable detail

(Lack 1947; Grant 1986; Weiner 1994), in part because the Galápagos offer a much less disturbed environment and a smaller array of species that have not been so decimated by extinctions. Darwin’s finches represent a much earlier stage of the process, when most species still look more or less like their common ancestor (Baptista and Trail 1988) and intermediate steps between major adaptational shifts are still evident (Grant 1986). It has long been assumed that the finch radiation in the Galápagos is younger than that of the honeycreepers, but more recent data show that they may be of comparable ages. The differences probably instead result from the very different climates of the two archipelagoes, the Galápagos being much more arid.The Hawaiian honeycreepers have diverged so much that we now see them as an array of disparate morphologies, with many ‘missing links’. But if Darwin’s finches are ‘Act I’ of the adaptive radiation scenario, then the honeycreepers are clearly ‘Act II’, and enough information has now accumulated, with recent palaeontological discoveries filling the gaps, that we can begin to analyse them in comparable ways. All of the evolutionary phenomena discussed so far have combined to produce the varied forms we see today.We understand the microevolutionary processes fairly well, but two questions remain: (1) why did the cardueline finches produce the only Hawaiian lineage to radiate adaptively, and (2) what were the keys that opened the adaptive doors along the way? On the first question, conventional wisdom once held that the drepanidines’ cardueline ancestor must have arrived earlier and had a head start on the other colonisers. But we now know that was not the case (Fleischer and McIntosh 2001). Honeyeaters were already represented in Hawai‘i when the founder of the honeycreepers arrived, yet they could not even close off the adaptive avenue to nectarivory.We do not yet know the arrival times for the other passerine groups. The solitaires at least seem to have prevented the development of a huge array of frugivorous honeycreepers, and the ‘elepaios kept the flycatcher niche mostly to themselves, but neither gave rise to any species adaptively different from its ancestor.

70 The Hawaiian Honeycreepers If the honeycreepers are not the oldest Hawaiian passerine lineage, then perhaps there is something about finches that predisposes them to wide divergence. In the Galápagos Islands, Darwin’s finches are joined by the widespread Vermilion Flycatcher, endemic subspecies of Yellow Warbler and Southern Martin, the endemic Galapagos Flycatcher, and four endemic allopatric mockingbirds (Castro and Phillips 1996), a species complement comparable in numbers to Hawai‘i’s but with much less endemism. As in Hawai‘i, only the Galápagos finches have undergone adaptive radiation. Among the other groups, only the mockingbirds have advanced beyond Stage 1 of their taxon cycle, and they only slightly. The remote Tristan da Cunha group in the South Atlantic, which Lack (1947) referred to as a ‘mini-Galápagos’, has been colonised by a thrush and an emberizine finch. The Tristan Thrush has formed two subspecies, but the finch produced two species, Tristan Bunting and Wilkins’s Bunting, which are sympatric on one island (Watson 1975). And there end the examples of avian adaptive radiation on islands! Worldwide, only these three exist (Lack 1947), and all started with a thick-billed, seedeating ancestor. With only three examples, generalisations are a bit strained, but with all examples finch-derived, the idea that finches have a predisposition to adaptive radiation gains credibility. Perhaps the reason no adaptive radiation occurred in Micronesia or central and eastern Polynesia (Olson 1990) is the fact that no carduelines or emberizines managed to reach those remote islands. The only finches in either region are estrildid parrot-finches (Pratt et al. 1987). (Finches have reached the North Atlantic island groups, but because of migratory patterns and prevailing winds, those islands are not as isolated as they would appear on the basis of distance alone, and their avifaunas are not strikingly distinct from that of Europe (Lack 1947).) Amadon’s (1950) supposition, quoted above, that heavy seed-crushing bills are an ‘evolutionary dead end’, obviously could not have been more wrong. But the fact that finches from two different lineages produced adaptive radiations argues against any hypothesis that involves mysterious and vague terms like ‘genetic capability’ or ‘greater plasticity in the genome’ to explain the

finches’ advantage. A much simpler explanation is that in the context of all passerine variations, a generalised finch bill, far from being a specialisation, is quintessentially intermediate; it is of average length, width, and depth. It is easy to see how various other bill shapes can be derived from it by variations in one or more of these parameters (Tordoff 1954; see above quotation). The typical long, slightly curved bills of meliphagids, for example, are probably more difficult to modify into anything different and immediately useful. It is the thin-billed passerine taxa that may be the evolutionary dead-ends, not the heavy-billed ones we call finches. Moving on to the second question, the drepanidine radiation seems to have been produced by a surprisingly small number of key adaptations, as suggested by Simpson (1953). According to the phylogeny presented above (Pratt 2001b), the initial drepanidine adaptive array may have been a group of finches plus a warbler-like derivative (Paroreomyza), a situation that parallels what we see in Darwin’s finches today. The finches at this stage apparently included the ancestors of Xestospiza and Melamprosops, both of which appear to be relicts in Stage 4. Among them also was one that lost its lingual wings and developed drepanidine odour. Which of these was the ‘key’ is difficult to say.The adaptive basis for the loss of lingual wings (see Chapter 6 for details) is not known and though it appears to predispose its possessor for nectarivory, exactly how is by no means obvious. The only other passerines with truncate tongue bases are the sunbirds (Nectariniidae; Gardner 1925; Scharnke 1932), but the V-shaped notch is very shallow in some honeyeaters, including the Hawaiian ‘o‘os (Scharnke 1931; Dorst 1952). Perhaps the loss of lingual wings was selectively neutral and just a marker for the odoriferous lineage. The odour at least has hypothetical selective advantages.Whatever their functional basis, these two key adaptations made the finch that possessed them the ancestor of all the other modern drepanidines. Most of that species’ progeny remained essentially finchlike (Telespiza, Rhodacanthis, Chloridops, etc.), but, as in the first burst of differentiation, one became a thin-billed derivative, possibly the ancestor of Oreomystis. One of the finchlike species,

Origin and evolution 71 perhaps resembling the partly nectarivorous (but not tubular-tongued) ‘O‘u, apparently took a liking to nectar.This bird founded a lineage that perfected the nectar-feeding apparatus, eventually producing another key adaptation, the drepanidine tubular tongue (described in detail in Chapter 6). The intermediate stages between a finch tongue and the tubular tongue can only be surmised, but the tongue of the Maui Parrotbill may resemble one of the steps (Chapter 6). Because the tubular tongue is so widespread among a varied array of honeycreepers, earlier researchers wrongly assumed that it was the ancestral type for the family. Raikow (1977b) rightfully pointed out that this highly derived structure has no counterpart outside the Drepanidinae and does not resemble tongues of other passerine nectarivores, each of which is independently derived in its own way (Gardner 1925; Parker 1977). Initially, the possessors of the drepanidine tongue elaborated into an array of nectar-feeders, represented today by the relict ‘red and black’ group of honeycreepers, with disparate morphologies and large adaptational gaps. One line’s bills began co-evolving with curved flowers (see Chapter 9). Ehrlich and Raven (1964) considered such co-evolution a major driving force in adaptive radiation in butterflies. Eventually one species, probably resembling a modern ‘amakihi, became a more generalised feeder with a diet that included many insects.And it did so without having to modify its tongue in any way! The seemingly highly specialised, nectar-adapted tongue proved to be equally adept at snaring insects. From that ‘amakihi-like ancestor arose ‘akepas with crossed bills for prying apart leaf buds, the generalist warblerlike ‘Anianiau, and one lineage that became increasingly insectivorous. Among those, one again became caught up in plant/pollinator co-evolution. In the

‘akialoas, the bill became longer and longer until it reached the practical limits of bill length. One of those long-billed birds may have found that its bill was more useful in probing for insects others could not reach than in feeding on long-tubed flowers. It could then have developed the habit of tapping the bark to disturb its prey, and eventually one population, perhaps by a fortuitous deformity, found that such action worked better if the lower mandible was shorter; thus appeared the heterobills. One of those, the ‘Akiapola‘au, is now poised for a new adaptive burst (if it can survive the ravages of mankind) by strengthening and straightening the shorter lower mandible to become the drepanidine analogue of sapsuckers.The immediate ancestor of that bird may also be the ancestor of Pseudonestor, which modified the original heterobill in a different way. An equally plausible scenario (Pratt 1979a) would have the heterobills and parrotbill in one lineage and the ‘amakihis and ‘akialoas in another. A third idea was suggested by Bock (1970), who presented what amounts to a transformational series beginning with the generalised bill of the O‘ahu and Hawai‘i ‘amakihis and proceeding through the larger bill of the Kaua‘i ‘Amakihi with increased bark-picking, to the heterobills, with their shortened lower mandible.The heterobills then give rise on the one hand to ‘akialoas, in which the lower mandible is lengthened and the entire bill becomes nearly as long as the rest of the bird, and on the other hand to the parrotbill, in which the lower mandible becomes secondarily thickened. Bock (1970) did not believe this to be an actual transformational series, but it may not be far from it. The burst of adaptation in the hemignathine group probably happened relatively recently, because the forms involved have very few ‘missing links’.This is, of course, only a scenario, but what a grand one it is!

5 Classification

The most significant overall taxonomic revisions of Hawaiian honeycreepers have been those of Perkins (1903), Amadon (1950), Pratt (1979a), Olson and James (1982b), and James and Olson (1991). Innovations made by Greenway (1944, 1968), Raikow (1977b), Berger (1981), AOU (1983, 1998), Pratt et al. (1987), and Sibley and Monroe (1990) are all refinements of previous revisions. The most widely used taxonomy today is that of the AOU (1998), which has its roots in Pratt (1979a) via Berger (1981), with refinements based on numerous papers published subsequently on various subgroups. The classification of Hawaiian honeycreepers used in this book does not coincide exactly with any previous classification. Although relatively few of its distinctive features are novel, it can be regarded as a new overall revision based on a large body of recent research (see Chapter 4). I do not, however, regard it as the final word, but as the best classification for summarising current knowledge. Taxonomies that have been, and continue to be, controversial are discussed below.

Genera Generic limits within the Drepanidinae have long been the most controversial level of the classification.Although species limits are approaching a consensus, as discussed below, the genera remain the source of considerable disagreement. Nor is the disagreement simply a philosophical matter of lumping or splitting, as in the case of the differences between the generic limits of Amadon (1950) and Greenway (1968). The genera of James and Olson (1991) and the AOU (1998) are essentially different

in their statements of relationships, but they also differ in some cases because no biological basis exists for determining where in a phylogeny generic lines should be drawn. Even with rigorous phylogenetic analysis, the decision as to where to draw lines between named taxa is largely still a matter of taste, as long as the taxa are monophyletic. However, hypotheses as to which clade a given species belongs can be tested by both observation and experiment. Some molecular systematists have suggested that generic limits might be based on lineage age, but no consensus exists as to what that age should be. In fact, the guidelines proposed by Mayr (1969) are still as good as any. Initially, nearly all drepanidine species were placed in monotypic genera. Amadon (1950) consolidated many of these but, in my opinion (Pratt 1979a) as well as that of Greenway (1968) and Banks and Laybourne (1977), he went too far. My earlier classification essentially followed Mayr’s (1969) guidelines by dividing genera at obvious morphological gaps, a totally different approach from that of Amadon (1950), who believed that in a group undergoing rapid adaptive radiation, genera should be defined more broadly than in more ‘conservative’ groups, but provided little justification for that opinion.Amadon (1986) complained that ‘the genera of the Hawaiian honeycreepers have been bandied about in rather cavalier fashion’, and Olson and James (1995) decried the wide acceptance of my classification ‘among non-taxonomists without any consideration having been given to its merits’, but subsequent analyses (see previous chapter for a discussion) have shown that the classification has considerable merit. I still subscribe to the philosophy used in my earlier

Classification 73 classification (Pratt 1979a) that genera should be clearly diagnosable, with the added proviso that they should form a monophyletic group. The genera recognised herein are diagnosed in Part II of this book, but several particularly controversial genera require a closer look here.

includes new specimen data from Kaua‘i ( James, 2004) suggests that Xestospiza conica might also belong in an enlarged Chloridops. New fossil discoveries and ongoing studies based on ancient DNA extracted and amplified from prehistoric remains (R. L. Fleischer, pers. comm.) may clarify the interrelationships of the drepanidine finches.

Drepanidine finches Most of the following discussion is paraphrased from Pratt (2001b). Amadon (1950) placed all the drepanidine finches (except the Lana‘i Hookbill, which he regarded as an aberrant specimen) in the single genus Psittirostra, reflecting his hypothesis that the birds’ finchlike characters were secondarily derived from a thin-billed ancestor and therefore likely to be monophyletic. Greenway (1968) used Psittirostra for the ‘O‘u alone, with the others placed in Loxioides. Banks and Laybourne (1977) thought Amadon’s Psittirostra was morphologically too broad, and advocated a return to the original five genera. Once a cardueline ancestry for the honeycreepers became established, Amadon’s large Psittirostra could well have represented a paraphyletic assemblage based on shared primitive characters (Pratt 1979a). At first, Olson and James (1982b) maintained Amadon’s Psittirostra but recognized five subgenera. Later ( James and Olson 1991) they recognized all five as genera plus Dysmorodrepanis and several new genera of drepanidine finches. James and Olson’s (1991) discovery of new species of Chloridops and Telespiza effectively removes Amadon’s (1950) discomfort with too many monotypic finch genera. My phylogeny (Pratt 2001b; Chapter 4 herein), based on phenotype and including relatively few characters to differentiate the finch genera, could be interpreted as supporting Greenway’s (1968) classification (depending on where one draws the lines), but recognizes the same genera as James and Olson (1991). Bill similarities suggested possible mergers of Chloridops and Loxioides (Pratt 2001b), but James and Olson (1991) thought Telespiza closer to Loxioides. James’s (1998) topology, based solely on osteological characters, supports lumping Telespiza with Loxioides as well as the merger of Rhodacanthis and Chloridops (which has priority), and a more recent analysis that

Loxops Amadon’s most controversial innovation was the lumping of several genera into a broadly defined Loxops, which Conant et al. (1998) considered ‘the most extreme example of alpha-level taxonomic over-lumping in the annals of avian systematics.’ ‘Greater Loxops’ included the ‘akepas (Loxops sensu stricto), the ‘amakihis (Hemignathus spp.), the ‘Anianiau (Magumma), the creepers (Oreomystis), and the ‘alauahios (Paroreomyza). According to Conant et al. (1998) the enlarged Loxops ‘has no defining synapomorphies or even any general similarities to unite its disparate members’ other than the fact that all have relatively short bills (Pratt 1979a). The dismemberment of Amadon’s Loxops began with Raikow’s (1977b) separation of Paroreomyza (including Oreomystis). However, Raikow did not combine Greater Loxops with the ‘red and black’ birds in a single clade on the basis of the lost plantaris, as would have been reasonable at that point. Johnson et al. (1989), Pratt (2001b), and James (2004) reviewed this issue and all concluded that the presence or absence of the plantaris has little or no phylogenetic significance. Not only is its presence variable among passerines generally, but it varies among cardueline finches, and among carduelines it even varies within the genus Serinus (canaries, serins, etc.), which is otherwise rather uniform. Furthermore, it is a condition that appears to have reversed itself independently so many times in the course of evolution that its absence can hardly be regarded as a uniquely derived condition even within a group like the Hawaiian honeycreepers. Raikow (1975) and Raikow et al. (1979) presented examples of ‘lost’ muscles that reappear in evolutionary lines, and Raikow was quoted by Johnson et al. (1989) as considering the reappearance of the plantaris quite possible among drepanidines.

74 The Hawaiian Honeycreepers Presumably on osteological grounds, Olson and James (1982b, 1995) agreed with Raikow that Paroreomyza and Oreomystis were separate, but maintained a ‘Greater Loxops’ that included akepas, ‘amakihis, ‘Anianiau, and the Hawai‘i Creeper. However, if James’s (1998) topology (Fig. 4.7) is correct, their Greater Loxops would be polyphyletic. Pratt (1979a) suggested that the cross-billed ‘akepas and the curve-billed ‘amakihis should not be considered congeners, and Berger (1981), and the AOU (1983) restricted Loxops to the akepas, placing the ‘amakihis in Hemignathus (Pratt 1979a; see below). James’s (1998), phylogeny is compatible with restriction of Loxops to akepas, but recent DNA studies have suggested that the Hawai‘i Creeper (see next paragraph), which lacks the drepanidine tubular tongue found in both ‘akepas and ‘amakihis, might belong there as well (Fleischer et al. 1998, 2001).

Oreomystis and Paroreomyza Originally, Paroreomyza was a subgenus of Oreomystis (Perkins 1903), but Amadon (1950) lumped all into Greater Loxops. As noted above, Raikow (1977b) separated Paroreomyza (then including Oreomystis) from Loxops and Olson and James (1982b) did also but divided it three ways, leaving the Hawai‘i Creeper in Greater Loxops, restricting Oreomystis to the ‘Akikiki, and reserving Paroreomyza for the members of Perkins’s original subgenus. Pratt (1979a) retained the Hawai‘i Creeper in Oreomystis. Recent molecular studies have mostly supported the distinctiveness of Oreomystis and Paroreomyza, although the degree to which they are related is still being debated. Johnson et al. (1989) and Fleischer et al. (1998) consider them sister groups outside the main line of drepanidine evolution; James (1998) and Fleischer et al. (2001) consider them sister groups within the ‘core clade’; but Tarr and Fleischer (1995) and Pratt (2001b) considered them not closely related, with Paroreomyza outside the main line and Oreomystis embedded within it. Noteworthy is that Oreomystis has both drepanidine odour and truncate lingual wings that define the core honeycreeper clade, whereas Paroreomyza lacks both. Did it lose them secondarily?

Whether the Hawai‘i Creeper is an Oreomystis also remains controversial. I recently reviewed the rationale for keeping the Hawai‘i Creeper in Oreomystis (Pratt 2001b, from which the following summary is taken). The plumage of the Hawai‘i Creeper, although superficially similar to some plumages of ‘akepas and ‘amakihis, actually more closely resembles that of O. bairdi. Both lack sexual dichromatism as adults, never have wing-bars, and possess a distinctive Juvenal plumage involving pale facial coloration. The bill shape of the Hawai‘i Creeper differs from those of both ‘akepas and ‘amakihis but is virtually identical to that of O. bairdi except for being a little thinner, which may reflect the fact that the creeper chooses slightly smaller branches for foraging. The Hawai‘i Creeper’s foraging movements are virtually identical to those of the ‘Akikiki, with both being the only honeycreepers that could be described as ‘nuthatch-like’. Both ‘Akikiki and Hawai‘i Creepers form close-knit post-fledging foraging parties, with the juveniles uttering distinctive, syncopated begging calls. These calls are not only nearly identical in the two species (for sonogram comparison see Pratt 2001b), but are unlike any other calls described for Hawaiian honeycreepers. The ‘Akikiki and Hawai‘i Creeper have nearly identical tongues that are narrow, non-tubular, and notched at the slightly frayed tip (Richards and Bock 1973), and that differ strikingly from the drepanidine tubular tongue (see Chapter 6) found in all other birds proposed as relatives of O. mana. The tubular tongue is a defining element of a major drepanidine clade (Raikow 1977b), and the Hawai‘i Creeper cannot belong to it unless its tongue is secondarily derived from a tubular one and resembles that of O. bairdi through convergence. I believe attributing so many resemblances to convergence strains credibility and that the Hawai‘i Creeper is, indeed, a second species of Oreomystis.

Hemignathus and its subgenera Perkins (1903) used Hemignathus for the ‘akialoas only, placing the heterobills in Heterorhynchus, but both genera were later shown to have been based on the same type specimen, so Hemignathus

Classification 75 was applied to the heterobills and the new genus Akialoa was named (Olson and James 1995). Amadon (1950) united the heterobills and ‘akialoas under Hemignathus. Perkins (1903) considered ‘amakihis more closely related to these groups than to ‘akepas, with which Amadon (1950) grouped them, but Pratt (1979a) was the first to place ‘amakihis in Hemignathus (cf. Olson and James 1988, who retained them in Greater Loxops). ‘Greater Hemignathus’ is easily diagnosed with a variety of plumage and morphological characters (see Part II). With the exception of the ‘Anianiau, which I no longer place in this genus (see below), all of its members are so similar in coloration and habits that field identifications must include observation of the bill to be credible (Pratt et al. 1987; Conant et al. 1998; Pratt and Pyle 2000). Amadon (1986) objected to the inclusion of short-billed and long-billed birds in the same genus, but many other avian genera such as Calidris, Phaethornis, Nectarinia, Toxostoma, and Geospiza exhibit variations in length and shape of bill of similar magnitude. Long ago, Bock (1960) pointed out that the evolutionary plasticity of bill shape made it a poor genus-level character. In James’s (1998) osteological phylogeny (Fig. 4.7), the ‘amakihis, ‘Anianiau, ‘akialoas, and heterobills (the ‘green’ birds) belong to a single clade which, surprisingly, also includes the ‘red and black’ honeycreepers. Pratt (2001b) tentatively attributed this anomaly to convergence in that both the ‘red’ group and the ‘green’ group include birds with short, down-curved bills (and note that James also groups Pseudonestor with Psittirostra probably for similar reasons). James (2001) distorted my meaning with the statement that her ‘red’ clade included no ‘green’ birds, which is true as far as it goes but misses the point. Her ‘red’ clade (Drepanis/‘Apapane/‘Akohekohe) is but one branch imbedded within the larger clade to which I referred. That major group is an unresolved trichotomy, the branches of which are: (1) Kaua‘i ‘Amakihi; (2) an ‘akialoa/heterobill clade; and (3) a two-pronged clade of the red birds on the one hand and the remaining ‘amakihis and ‘Anianiau on the other. With the red birds left in place, a new genus would have to be erected for the

Kaua‘i ‘Amakihi or the entire clade (red birds ⫹ heterobills ⫹ ‘amakihis ⫹ ‘Anianiau) would have to be combined in a huge genus (which would be Drepanis by the Law of Priority)! With the red bird branch pruned, that larger clade would precisely delimit Hemignathus as recognised by the AOU (1998). Alternatively, if the trichotomy could be resolved so that the first two branches above became sister groups, then the placement of at least one ‘amakihi in Hemignathus (whether ‘greater’ or lesser) would be unavoidable. After so many years of decrying my placement of ‘amakihis and ‘akialoas in the same genus, James and Olson can hardly be expected to admit that their own data now support such an arrangement. I suspect that a combined data matrix of osteological and other phenotypic characters would resolve the anomalous placement of the ‘red’ group in the middle of Greater Hemignathus, and indeed James (2004) could not statistically reject my alternative hypothesis. Greater Hemignathus is not, however, supported by Fleischer et al.’s (1998) mtDNA study (nor, for that matter, does that study support Greater Loxops). But R. C. Fleischer (pers. comm.) considers the branching sequence revealed in their analyses (which did not include ‘akialoas or the Greater ‘Amakihi) to be poorly defined as yet. Until better resolution and broader coverage is achieved with molecular techniques, generic limits based on phenotypic characters (osteology included) are probably the most reliable. Four subgenera of Hemignathus can be delineated: Hemignathus, Akialoa, Chlorodrepanis, and Viridonia. The latter two were combined by Greenway (1968), but my recent phylogenetic analysis (Fig. 4.8) and James’s (1998, 2004) topology both suggest not only that Viridonia ⫹ Chlorodrepanis would be paraphyletic, but that Viridonia might well deserve full generic rank. I consider ‘akialoas and ‘amakihis to be sister groups (Pratt 1979a) that could well be combined, but pending further input from molecular biologists, I keep them separate. The membership, or even existence, of Greater Hemignathus will be challenged if recent molecular (Fleischer et al. 1998) and phenotypic (Pratt 2001b) findings on relationships of Hemignathus sensu stricto

76 The Hawaiian Honeycreepers and Pseudonestor (see below) pan out. R. C. Fleischer (in T. K. Pratt et al. 2001) suggests that the heterobill/parrotbill lineage may have arisen separately from the ‘amakihi/‘akialoa one early in honeycreeper evolution. Simply adding a fifth subgenus to Hemignathus would thus obscure relationships within the group, and at that point Greater Hemignathus would probably best be broken up. If that were done, then new generic limits would restrict Hemignathus to the heterobills and parrotbill, leaving ‘amakihis and ‘akialoas (and Viridonia if it is really related to ‘amakihis) in Chlorodrepanis.

Pseudonestor The Maui Parrotbill’s namesake feature has obscured its relationships for many years. Latham, first describer of the ‘O’u, called it the ‘Parrot-billed Grosbeak’, and Wilson (Wilson and Evans 1890–99) referred to Pseudonestor as the ‘small ‘O’u’. Raikow (1977b) considered Pseudonestor a sister group to Psittirostra on the basis of the strongly hooked or overhanging maxilla in both. However, he did not include in this supposed synapomorphic grouping other honeycreeper taxa (e.g. nukupu‘us) that have overhanging upper mandibles. In the context of the honeycreeper radiation, a down-curved maxilla hardly constitutes a synapomorphy, and virtually everything else about the bird indicates a relationship to the thin-billed, mostly insectivorous honeycreepers, in particular the heterobills. Perkins (1903) was probably the first to recognise this relationship, and Bock (1970) analysed it in some detail. However, although Pratt (1979a) placed Pseudonestor among the hemignathine group, Berger (1981) and the AOU (1983) continued to associate it with the drepanidine finches. My more recent analysis (Fig. 5.1; Pratt 2001b) shows even more clearly that the parrotbill is not a drepanidine finch, and that it is allied with one subclade of Greater Hemignathus. James (1998, 2004) still regards the parrotbill as allied with drep finches, but an experimental manipulation of the data allied it with Hemignathus wilsoni. Once the conflation of parrot-like and finchlike bills, which James (1998) continues to do, is overcome, the similarities between the heterobills and the parrotbill are striking, from a modified jaw

muscle they share (Zusi 1989), to the way the bill is used in feeding (Chapter 6), to the songs and calls of both juveniles and adults (Chapter 7). Bock (1970) considered the derivation of the parrot-like bill from the thinner heterobill quite reasonable. Furthermore, DNA studies also indicate a close relationship between parrotbill and ‘Akiapola‘au (Fleischer et al. 1998, 2001). The parrotbill’s unique tongue could have easily been secondarily derived by neoteny; it looks like a drepanidine tubular tongue that has been unrolled (or never rolled), much like the tongues of young ‘Anianiau (Bryan and Seale 1900) _ _ 'O'u Maui 'Alauahio O'ahu 'Alauahio _ _ Kakawahie Maui Parrotbill O'ahu Nukupu'u Maui Nukupu'u __ 'Akiapola'au Kaua'i Nukupu'u O'ahu 'Amakihi Hawai'i 'Amakihi Kaua'i 'Amakihi Lesser 'Akialoa 'Greater 'Akialoa' Greater 'Amakihi _ 'Akepa 'Akeke'e 'Anianiau Hawai'i Creeper 'Akikiki

5.1 Species-level phylogeny (50% majority rule consensus tree) of hemignathine ‘little green birds’ plus Psittirostra, from Pratt (2001b).

Classification 77 that have tubular tongues as adults but lack them as nestlings (the tongue curls into a tube with age; see Chapter 6). Despite the fact that without the parrotbill included, Hemignathus would be paraphyletic in my phylogeny (Pratt 2001b), I have kept them separate pending further confirmation of the relationship from molecular and other data.

Magumma Perkins (1903) assigned the ‘Anianiau, sometimes referred to as the ‘Lesser ‘Amakihi’, and the ‘amakihis to Chlorodrepanis. Mathews’s (1925) Magumma was rarely used because virtually all authors, including myself (Pratt 1979a), uncritically considered the ‘Anianiau congeneric with ‘amakihis, wherever placed. A re-examination of characters (Conant et al. 1998; Pratt 2001b) showed that this ‘default’ position was probably incorrect. Although as a ‘little green bird’ (Pratt et al. 1987), the ‘Anianiau resembles ‘amakihis in overall plumage colour and general behaviour, the similarities prove superficial when examined in detail (Fig. 5.1). Magumma lacks essentially all of the defining synapomorphies of Greater Hemignathus. Sexual dimorphism and dorsoventral contrast are greatly reduced, dark lores are completely lacking, immatures lack wing-bars, the bill is only slightly decurved and flesh-coloured with a dusky culmen, and feet are pale rather than dark. Consequently, Conant et al. (1998) and I (Pratt 2001b) recommended placing the ‘Anianiau in its own monotypic genus. Some DNA studies also support that treatment. Tarr and Fleischer’s (1995) phylogeny showed the ‘Anianiau widely separated from the ‘amakihis in a clade of its own, but Fleischer et al. (1998, 2001) enigmatically grouped it with the ‘Akiapola‘au and the Maui Parrotbill. Either way, it is not a close relative of the ‘amakihis (but in the latter case could be retained in Hemignathus if Pseudonestor were lumped with it). Note, however, that James (2004) maintains the alliance of the 'Anianiau with the ‘amakihis.

Drepanis and Vestiaria The only other point at the generic level on which Berger (1981) and the AOU (1983) disagreed with

Pratt (1979a) was in not merging these two nominal genera. I still maintain, as I did in 1979, that the monotypic genus Vestiaria for the ‘I‘iwi cannot be distinguished from Drepanis except on the basis of relatively slight species-level colour differences. A red-to-yellow shift is hardly the basis for a separate genus when it was long considered insufficient reason for splitting species of several honeycreepers (e.g. ‘Akepa and ‘Akeke‘e, Kakawahie and ‘alauahios). Amadon (1950: 166) stated that ‘variation from yellow to red is obviously accomplished readily and need not be considered as necessarily indicating specific difference.’ Amadon (1986) felt that if Drepanis and Vestiaria were merged, Himatione and Palmeria would have to be also. He may be right (although he did not support either merger), but Palmeria’s distinctive plumage features make it look, superficially at least, rather different from Himatione. In every variation of my phylogenetic analysis (Pratt 2001b), Drepanis and Vestiaria were very close sister groups, but that was not the case with Palmeria and Himatione. James’s (1998) osteological phylogeny also supported merger of Drepanis and Vestiaria, but not Palmeria and Himatione.These points may all be moot if R. Fleischer (pers. comm.) is correct that all of the ‘red and black’ genera are so close genetically that they might well all be merged into Drepanis.

Species Limits The long-standing biological species concept (BSC) popularised by Mayr (1942), the definition of a species that every biology student has memorised for decades, has recently been challenged by several new ones (Hollocher 2001). The proponents of phylogenetic systematics have offered the phylogenetic species concept (PSC) in which a species is a population or cluster of individuals ‘diagnosably different from other such clusters, and within which there is a parental pattern of ancestry and descent’ (Cracraft 1983). Diagnosability can be based on ‘any feature or set of features, ranging from single fixed nucleotide substitutions to major phenotypic (but genetically based) features’ (Zink and McKitrick 1995).The fundamental difference is that the BSC is based on ability to interbreed whereas genetic isolation is irrelevant

78 The Hawaiian Honeycreepers in the PSC.The BSC seeks ‘isolating mechanisms’ that function in preventing interbreeding between species, whereas the PSC seeks detectable differences to delimit species. For support of the PSC, see Hazevoet (1996) and Zink and McKitrick (1995), and for the BSC see Mayr (1992), Collar (1996, 1997), Snow (1997), and Pratt and Pratt (2001). The PSC has not so far achieved wide acceptance, even among phylogenetic systematists. I subscribe to most of the tenets of that school, but continue to believe that the BSC is best for most scientific and environmental purposes. The PSC has the potential to elevate all island subspecies to species, so it is worth considering in the drepanidine context. In my opinion, the biggest problem with the BSC has been poor application of the concept in insular situations in the past. Many very distinctive allopatric island populations were ranked as subspecies when they should have been considered species. Amadon and Short (1976) tried to introduce the concept of the ‘megasubspecies’ for such forms, but the idea did not gain wide acceptance. Pratt and Pratt (2001) found that in the case of Hawaiian honeycreepers, proper application of the BSC, with attention to potential isolating mechanisms such as vocalisations and especially plumage colour, resulted in a list of biological species that differed only slightly from one formulated under the PSC. We found that plumage colour was a far better indicator of species limits than had been appreciated in the past. Amadon (1950) particularly denigrated coloration as a criterion, but recent molecular studies (see discussion of ‘amakihis below) have revealed that even slight (but consistent) colour differences between island populations can indicate genetic differences of the same order of magnitude as seen in continental species with more striking phenotypic differences. Furthermore, fossil discoveries have shown that some species formerly lumped by Amadon (1950) were actually sympatric until extinctions changed the picture (see ‘alauahios below).We now know that on islands, at least, birds that look different usually are different. Recently, Johnson et al. (1999) have offered a ‘a new comprehensive biologic species concept (CBSC)’ that will, in my opinion, alleviate many of the past

problems in application of the BSC in island situations. It is stated as follows: An avian species is a system of populations representing an essentially monophyletic, genetically cohesive, and genealogically concordant lineage of individuals that share a common fertilisation system through time and space, represent an independent evolutionary trajectory, and demonstrate essential but not necessarily complete reproductive isolation from other such systems. The species limits for Hawaiian honeycreepers proposed by Pratt and Pratt (2001) could well have been drawn under the new CBSC, although they were established before publication of that concept (publication dates notwithstanding; Pratt and Pratt 2001 had an unusually lengthy ‘2004’ period).The following review of honeycreeper groups whose species limits have been controversial is mostly paraphrased from Pratt and Pratt (2001).

Telespiza finches Laysan and Nihoa finches provide a good example of how coloration often flags other isolating mechanisms. They have obvious differences in overall size as well as bill size and shape. Although similar, their plumages are diagnostically different with females differing more than males. Amadon (1950) considered them conspecific, but Banks and Laybourne (1977) split them based on differences in moults and number of plumages required to reach Definitive. Ely and Clapp (1973) and Clapp et al. (1977) reported differences in breeding biology, and Conant (in Pratt 1996) documented vocal differences. With so many potential isolating mechanisms, all recent authors have recognised the Laysan Finch and Nihoa Finch as biological species. They are also obviously phylogenetic species. Biological species status was proven by James and Olson’s (1991) discovery of fossil remains of both species in the same sites on Moloka‘i. Fleischer et al. (1998) found that the two differ genetically to the same degree as other biological species pairs.

Classification 79

‘Amakihis Amadon (1950) and others of the period considered all of the ‘typical’‘amakihis to belong to a single polytypic species, but all realised that the Kaua‘i representative of the complex had a much larger and heavier bill than the others. Conant et al. (1998) showed that bill measurements of the Kaua‘i ‘Amakihi do not overlap those of other forms.The larger bill is used differently as well, with a much greater proportion of bark-picking in the bird’s feeding repertoire. Songs and calls also differ significantly (Pratt et al. 1987; Pratt 1996a). Despite the many potential isolating mechanisms, summarised by Conant et al. (1998), Pratt’s (1979a) split was not accepted until biochemical data ( Johnson et al. 1989; Tarr and Fleischer 1993) indicated the same thing (AOU 1995). Olson and James (1988) found that placement of the ‘amakihis in Hemignathus caused the epithet stejnegeri, long used for the Kaua‘i ‘Amakihi, to be preoccupied, so Pratt (1989a) offered H. kauaiensis as an alternative. Interestingly, Tarr and Fleischer’s (1993) study revealed that the O‘ahu ‘Amakihi not only differed genetically within the complex to the same degree as the Kaua‘i bird, but was its sister group, and therefore not conspecific with the morphologically more similar ‘amakihis of the Maui-nui group and Hawai‘i.Thus the AOU (1995) recognised both the Kaua‘i and O‘ahu birds as full species. However, Fleischer et al. (1998) later produced a different branching sequence in which the O‘ahu ‘Amakihi was, after all, sister to the Hawai‘i ‘Amakihi. Hemignathus flavus could still be accepted, however, on the basis of its genetic distinctiveness. The only evidence that the O‘ahu ‘Amakihi might be a separate species before the DNA studies was its distinctive plumage. On plumage alone, O‘ahu birds can be distinguished with near 100% accuracy from those of other islands, and the females are especially unusual in having wing-bars as adults.Virtually all publications in the twentieth century had used the epithet chloris for the O‘ahu ‘Amakihi, but Olson (1996b) showed that the name H. flavus Bloxam, 1827 has priority. ‘Amakihi populations on Moloka‘i, Lana‘i, and Maui were originally given separate scientific names,

but neither Amadon (1950) nor I (Pratt 1979a) could find sufficient differences to warrant even subspecies status. However, collectively the Maui-nui birds differ from those on Hawai‘i sufficiently to be recognised as Hemignathus virens wilsoni. Consequently, practitioners of the PSC would recognise four, rather than three, species of ‘amakihi.

‘Akialoas All ‘akialoas are extinct, so biological species limits must be determined by inference from other data. Each of the main islands or complexes had at least one ‘akialoa in historic times, but those of O‘ahu and Maui-nui (Lana‘i only except as fossils) are known from only a few specimens. A recently extinct second species Hemignathus upupirostris was present on Kaua‘i and O‘ahu, and as yet undescribed species were sympatric with known ones on both Maui-nui and Hawai‘i (Olson and James 1995). Bryan and Greenway (1944) lumped all of the historically known ‘akialoas, but Amadon (1950) recognised two species, Kaua‘i Akialoa (H. procerus emended to H. stejnegeri by Olson and James 1995) and Hawai‘i ‘Akialoa H. obscurus, because of the striking bill length and size differences between them. However, probably because he had examined no O‘ahu specimens and only two from Lana‘i, Amadon mistakenly concluded that those forms belonged to the smaller species. Pratt (1979a) and Olson and James (1982b) found that they actually were much closer to the larger form, and lumped all ‘akialoas as a single species again, but the AOU (1983) maintained Amadon’s (1950) taxonomy. Pratt et al. (1987) pointed out that the main problem with Amadon’s classification was that the line between the two species had been drawn in the wrong place. They recommended, and the AOU (1997) eventually adopted, a two-species classification of Lesser ‘Akialoa H. obscurus for the Hawai‘i bird and Greater ‘Akialoa H. ellisianus for the other three. The discovery of the prehistoric species complicates the situation because their relationships are as yet unresolved. For example, the as-yet-undescribed larger Hawai‘i species might be H. ellisianus, H. upupirostris, or a new species entirely. Until things

80 The Hawaiian Honeycreepers are better sorted out, I endorse the recommendation of Olson and James (1995) that all described forms be considered full species. Plumage variation among the historically known forms is comparable to that seen among ‘amakihis and is non-clinal, suggesting that these forms really are separate biological species. They certainly qualify as phylogenetic species.

Heterobills Each of the four largest islands had one historically known heterobill. Bryan and Greenway (1944) combined all as a single species, but even Amadon (1950) recognised the distinctiveness of the ‘Akiapola‘au Hemignathus munroi1 and separated it from the nukupu‘us. Although described as separate species, lucidus (O‘ahu), affinis (Maui), and hanapepe (Kaua‘i) have long been considered subspecies of H. lucidus. However, this taxonomy overlooks striking differences in size and coloration among the three forms (Pratt and Pratt 2001), which indicate that more than one species is involved. They certainly represent three phylogenetic species, but considering the predictive reliability of plumage colour in other cases of inter-island variation among closely related drepanidines, they are probably good biological species as well. DNA researchers have very recently discovered rather large genetic distances between the forms that enhance this probability (R. Fleischer, pers. comm.). T. K. Pratt et al. (2001) tentatively kept the three as a single species following current practice, but recognize the strong case for splitting them (T. K. Pratt, pers. comm.). I therefore recognise three species of nukupu‘u herein.

‘Alauahios The genus Paroreomyza has an anomalous distribution that includes only the central islands of O‘ahu and the Maui-nui complex. Dismissing the striking colour variation among them, Amadon (1950) lumped the four Paroreomyza with the two species of Oreomystis (see above) ‘in what may be the best example of how far astray taxonomists who ignored colour differences could go’ (Pratt and Pratt 2001). However, Amadon (1950) could not have known

the geological history of the Maui-nui group and his taxonomy was not as unreasonable when proposed as it appears today. If the Kakawahie and the Maui ‘Alauahio are conspecific, then their striking differences had to have evolved since the break-up of Maui-nui, a period we now know to have been as little as 10 000 years ago. Because the Kakawahie is the largest and heaviest-billed Paroreomyza, and the Maui/Lana‘i form the smallest and smallestbilled, they are a good example of character displacement (see previous chapter), which must have occurred during a period of sympatry on Maui-nui (Pratt 1979a). When Olson and James (1982b) found remains of a smaller form on Moloka‘i, where only the larger one was previously known, sympatry was essentially confirmed and they agreed with Pratt (1979a) that at least two species of Paroreomyza existed. They at first lumped the O‘ahu and Maui ‘alauahios, but later split them ( James and Olson 1991). The O‘ahu ‘Alauahio’s bill is intermediate between those of the two Maui-nui species, but the coloration of both males and females is distinctive and diagnostic (Pratt et al. 1987). The two named forms of the Maui ‘Alauahio, montana from Lana‘i and newtoni from Maui, are barely distinguishable in coloration and are at most biological subspecies. Pratt and Pratt (2001) suggested that they ‘may represent fragments of a former inter-island cline, such as that shown by ‘elepaios on Hawai‘i (Pratt 1980), in which paler birds inhabited the lower and drier parts of Mauinui and darker ones the rainforests of Haleakala, and found that the few specimens from West Maui did show some intermediacy. Such problems as where to draw species lines among the fragments of a former cline demonstrate that the PSC is not as free of subjective judgements as its proponents claim (Collar 1997; Snow 1997; Pratt and Pratt 2001).Thus the question of whether the Maui and Lana‘i ‘alauahios are phylogenetic species is moot.

‘Akepas Bryan and Greenway (1944) were the first to delineate the two species of Loxops recognised today, but Amadon (1950) disagreed and combined them as the ‘Akepa. My suggestion (Pratt 1979a) of

Classification 81 a return to Bryan and Greenway’s classification was ignored by Berger (1981) and the AOU (1983), but following publication of additional research (Pratt 1989b), the AOU (1991) recognised the ‘Akeke‘e as a separate species. This case is another example wherein striking plumage differences predicted potential isolating mechanisms in other aspects of the birds’ biology and, ultimately, species limits (Pratt and Pratt 2001). The males of the three island representatives of the ‘Akepa are easily distinguished, but the females less so, and whether the colour differences are sufficient to warrant recognition of three biological species of ‘Akepa (i.e. O‘ahu ‘Akepa Loxops wolstenholmei,2 Maui ‘Akepa L. ochracea, and Hawai‘i ‘Akepa L. coccineus) is as unclear today as it was a decade ago (Pratt 1989b; Pratt and Pratt 2001). One possible isolating mechanism between the Maui and Hawai‘i forms is that the latter is an obligate cavity nester (Lepson and Freed 1997), but the only nest ever observed of the Maui bird was apparently being built in the open (Perkins 1895). Because the O‘ahu bird is extinct and the Maui one nearly so, additional relevant biological information is unlikely to become available. Preliminary results of ongoing DNA studies (R. Fleischer, pers. comm.) are revealing fairly large genetic distances between these forms, however, and I suspect that ultimately three species will be recognised.The case for three phylogenetic species is a bit stronger, but still unclear because the plumage differences between the Maui and Hawai‘i birds are not completely discrete (although male bill colour seems to be).

‘Apapanes The ‘Apapane exhibits no geographic variation among the main Hawaiian islands. A close relative, the Laysan Honeycreeper, was long regarded as a subspecies, but Olson and James (1982b, 1991) listed it as a full species without comment. Schlanger and Gillett (1976) considered it an ancient relict of the days when Laysan was a high, forested island, but recent molecular studies (Fleischer et al. 1998) suggest that the entire lineage of Hawaiian honeycreepers is too young to have inhabited Laysan at the time. Olson and Ziegler (1995) believed it colonised

Laysan from the main islands long afterward, and gave it species status because of unspecified distinctive cranial features. Pratt and Pratt (2001) discussed a variety of potential isolating mechanisms including ‘distinctive song and song phenology (Rothschild 1893–1900); different feeding behaviour (including often walking on the ground to forage among flowers; Fisher 1903); different nest placement and structure (Schauinsland 1899; Bailey 1956); and, most obviously, totally different habitat.’Also, in the course of preparing the illustrations for this book, I noticed that the Laysan Honeycreeper lacks the truncation of the primaries that produce the ‘Apapane’s ‘wing note’ (Chapter 7). Pratt and Pratt (2001) considered it ‘inconceivable that these birds, adapted to two different worlds, could successfully interbreed, much less do so freely’, and chided systematists for not paying more attention to the obvious differences and ‘automatically lumping on the basis of vague similarity’. The Laysan Honeycreeper is unquestionably a valid phylogenetic species, and this case demonstrates that better application of the BSC often achieves the same results.

Higher Categories Although long regarded as a family Drepanididae, we now know that the Hawaiian honeycreepers, despite their unique history and wide divergence, are but a subgroup in evolutionary terms (although Koblik [1994] continues to argue for family status). When they were a family, subdivisions seemed natural. Perkins (1903) recognised two ‘Divisions’ that were the basis for Amadon’s (1950) subfamilies Drepanidinae and Psittirostrinae. Raikow (1977b) was more impressed with the underlying uniformity of the group and downplayed the subdivisions. Pratt (1979a) ranked the honeycreepers as a subfamily of Fringillidae, but Berger (1981) did not accept this aspect of my work and continued to refer to them as a family. Amadon’s subfamilies were now tribes, and I divided the Psittirostrinae into Psittirostrini and Hemignathini so that three subdivisions could be recognised (AOU 1983; but note that the parrotbill, as discussed above, was misplaced in the Psittirostrini, contra Pratt 1979a).

82 The Hawaiian Honeycreepers Olson and James (1982b) considered the whole taxon a tribe Drepanidini within the Carduelinae and did not further subdivide (i.e. no subtribes). The latest AOU (1998) checklist dropped the tribes, but retained the honeycreepers’ status as one of the three subfamilies of Fringillidae (with Carduelinae and Fringillinae). I now agree with Raikow (1977b) that subdivisions of the group are not very meaningful and in any case would not follow traditional lines. My own phylogenetic analysis supports continued recognition of the subfamily Drepanidinae, a position not philosophically different from that of the Olson/ James team, who simply draw the lines at a different place in the hierarchy. However, if Groth’s findings (1994, 1998) and James’s (2004) most recent topology are

correct, the Hawaiian honeycreepers may be subsumed within the cardueline finches at whatever rank that group holds, and will no longer have a group-name of their own. We can be grateful that such a conclusion was not reached before the decision to include this volume among a series titled ‘Bird Families of the World’! 1. The longstanding name Hemignathus wilsoni Rothschild became preoccupied when the ‘amakihis were placed in the same genus and Pratt (1979a,b) offered this replacement. 2. The name Loxops rufa (Bloxam 1827), which had ‘been in all but universal use for the Oahu bird’ since 1895 was shown by Olson (1986), following Stejneger (1900), to be unavailable. The correct epithet is wolstenholmei Rothschild, 1893.

6 Form and function

Postcranial anatomy The postcranial (body) anatomy of Hawaiian honeycreepers is surprisingly uniform for such a diverse group. Early workers found little in the body skeleton of Hawaiian honeycreepers that differed in any noticeable way from that of most perching birds. James and Olson (1991), in fact, report that most of the postcranial bones found in prehistoric deposits cannot be identified to species unless part of an associated skeleton.

Skeleton The only variations of note among the honeycreepers involve the relative length and thickness of the femur and tarsometatarsus of the hindlimb ( James and Olson 1991). Drepanidine finches tend to have a relatively short, thick tarsometatarsus, but the thin-billed species have longer and thinner ones. These differences may be related to how active the birds are. My own observations and those of early observers indicate that the finches tend to be more sedentary or inactive and to have a rather dumpy or heavy-bodied appearance as compared to the active, lighter-bodied, thin-billed species.

Musculature Raikow (1976, 1977a) made a detailed study of the limb musculature of Hawaiian honeycreepers and those of other passerines (Raikow 1978). As a result, the drepanidines are among the best known passerines in this respect. Probably Raikow’s most important finding was that these muscles are, like the bones they move, remarkably uniform within

the Drepanidinae. Those myological features that indicate a relationship between the cardueline finches and Hawaiian honeycreepers were discussed in the Chapter 4.Among the honeycreepers, only a few muscles exhibit any significant variation, and the wing muscles vary less than those of the leg (Raikow 1977a). In his discussion of honeycreeper evolution, Raikow (1977b) found one muscle variation to be of enough significance to prompt the only change he made in Amadon’s (1950) classification. A tiny muscle of the shank called M. plantaris (or colloquially ‘the plantaris’) is present in some passerines, absent in others. From Raikow’s (1976) study, we know the plantaris is present in Laysan Finch, Nihoa Finch, ‘O‘u, ‘Akiapola‘au, Kaua‘i ‘Akialoa, and ‘Akikiki and absent in Hawai‘i ‘Amakihi,‘Apapane, ‘I‘iwi, and ‘Akohekohe. Subsequent dissections (S. Olson, pers. comm.) revealed that the Hawai‘i Creeper lacks the plantaris. Unfortunately, Raikow (1976) included neither an ‘akepa nor any of the ‘creepers’ (which were all then considered conspecific) other than the ‘Akikiki.We also still do not know the status of this feature in several other systematically strategic groups including ‘alauahios, Po‘o-uli, Maui Parrotbill, and Greater ‘Amakihi. The significance (or lack thereof ) of the plantaris in reconstructing the evolution of honeycreepers is discussed in Chapter 4.

Internal organs Internal organs of Hawaiian honeycreepers have been little studied, primarily because earlier researchers failed to find any promising avenues of

84 The Hawaiian Honeycreepers investigation. Most internal organs are very similar to those of other passerines and even non-passerines (Gadow 1899). A crop is present in some drepanidine finches and some of the nectarivores, but absent in others. This feature varies almost randomly among passerines generally, and Amadon (1950) attributed no phylogenetic significance to it. Gadow also examined intestinal convolutions of Hawaiian honeycreepers, but found that they did not differ from those of other passerines.

Cranial anatomy As discussed in Chapter 4, Hawaiian honeycreeper skulls have many features, most of them synapomorphies, in common with cardueline finches. Amadon (1950) concluded that cranial osteology held little phylogenetic information, but his own observations can easily be interpreted as supporting a cardueline ancestry. In a table that compared five skull modifications in possibly related passerines, Amadon listed two possible concordances of Hawaiian honeycreepers with coerebids and only one with tanagers, but resemblance in all characters between drepanidines and carduelines. Whether such resemblances are phylogenetically significant is the issue, of course, but we must remember that the whole concept of synapomorphy (see Chapter 4) was not widely appreciated at the time. Possible cardueline/drepanidine synapomorphies that have been noted by several authors include a solid bony palate, lateral flange condition of the palatine process of the premaxilla (Bock 1960), and a thick, double-walled interorbital septum (Zusi 1978). All of these conditions show evolutionary modification within the Hawaiian honeycreepers, with the thinner-billed species showing less resemblance to cardueline finches. The consensus today is that these lesser expressions are derived from the cardueline condition rather than that the cardueline resemblances evolved by convergence. For example, the ‘lateral flange’ condition is fully developed in Hawaiian finches and identical to that seen in carduelines (Bock 1960), but is less developed in the ‘Akepa and reduced to a tiny remnant in the Common ‘Amakihi (Richards and Bock 1973). This transformational series could be read in either

direction, but progressive loss of this feature seems more believable and parsimonious. The interorbital septum has been studied in most species of Hawaiian honeycreeper, allowing some definite patterns and trends to be discerned. As reviewed in the previous chapter, the interorbital septa of cardueline and Hawaiian finches are identical. More importantly, they of the same type in the ‘O’u and the Maui Parrotbill, two heavy-billed but not finchlike honeycreepers (Zusi 1989), as well as in the ‘akepas (Richards and Bock 1973). Some of the drepanidine finches may have small fenestrae in the septum, a feature that can vary individually as it does in the Palila (two of Zusi’s six specimens had small fenestrae). In the other species of honeycreepers the septum is thinner and has ‘a single-walled, flat, translucent, central portion of the septum. Many of these species have a fenestra in the single-walled portion.The presence and size of both of these features . . . are variable within and between species’ (Zusi 1989: 717–18). Another previously discussed feature of the cardueline/drepanidine interorbital septum is that it forms a broad flat floor of the cranial fenestra, which lies anterior to the braincase and above the septum. In other passerines, the floor of the cranial fenestra tends to be narrow or knife-like, and to have a hump or upward-pointing profile.This may be the most consistent cranial feature within the Hawaiian honeycreepers, including even the aberrant genera Paroreomyza and Melamprosops. James and Olson (1991) found it to be the one feature that showed some of their fragmentary bones to be drepanidine. It also provides support for the idea that the thinner septa are derived from the thicker type rather than the reverse. A feature of avian skulls in general that is particularly relevant to any discussion of the feeding apparatus of Hawaiian honeycreepers is known as cranial kinesis. Unlike mammals, birds have upper jaws (maxillae) that are movable against the main part of the skull. Dorsally, the maxilla attaches to the front of the braincase by a flexible bony attachment called the nasofrontal hinge. At either side, a long narrow bone called the jugal bar extends rearward from the proximal end of the maxilla to an articulation with the quadrate bone. The quadrate

Form and function 85 has been fused with others in mammals, so that the lower jaw or mandible articulates directly with the skull. In birds, the mandible articulates with the quadrate, which then articulates with the skull. Thus a forward movement of the quadrate moves both the upper and lower jaw.The presence of the quadrate also means that the bills of birds open much wider than would be the case if the jaw articulated directly with the skull as in mammals (note that cartoonists, and unfortunately many bird artists, have often lost this point to anthropomorphism). This high degree of movement, or kinesis, enables birds, and especially Hawaiian honeycreepers, to do some remarkable things with their bills, as the following review of the feeding apparatus will show. Kinesis is particularly important for the bill movement known as gaping. In the course of feeding, many honeycreepers gape, or forcefully open the bill, to spread apart leaf buds, flower corollas, leaves cemented together by insects, leaf axils, cracks in bark, or other such things. Gapers not only have considerable kinesis of the maxilla, but a characteristic feature of the mandible called the retroarticular process. This is a rearward projection of the mandibular ramus past the point of articulation with the quadrate. It serves as the attachment for muscles that, using the quadrate as a fulcrum, pull the lower jaw downward. Many honeycreepers, with a wide array of feeding adaptations, have prominent retroarticular processes.

Feeding apparatus: the bill Finchlike bills The Hawaiian finches have a range of variation on the theme that parallels that of the cardueline finches (Fig. 6.1).Variations involve the length and thickness of the bill, the degree of overhang of the maxilla, the shape of the tomia (cutting edges), the curvature of the culmen (upper contour) and gonys (lower contour), and the ridges and grooves on the underside of the maxilla ( James and Olson 1991). The latter feature received little attention until recently, but has proven important in working out relationships of recently discovered prehistoric Hawaiian finches ( James 1998). These ridges and

grooves are present in both the rhamphotheca and the underlying bone, and apparently function in holding and processing various food items. The members of the genus Telespiza have the least specialised bills and the most generalised feeding habits among drepanidine finches.The bills are similar to those of carduelines in such genera as Serinus and Carpodacus, which also have relatively unspecialized feeding habits. In overall appearance, members of this genus probably are the closest surviving species to the ancestor of all Hawaiian honeycreepers (Pratt 1979a).The two surviving species are known to feed on everything from grass seeds to hard pericarps of nohu (puncture vine) to eggs of seabirds (Morin and Conant 2002; see also Chapter 9). Because as many as four species of Telespiza may have been sympatric on a single island ( James and Olson 1991), the question arises of how they subdivided such a broad feeding niche. The four described species exhibit a symmetrical stepwise cline in bill size, reminiscent of size variation within the genera Geospiza and Camarhynchus of the Galápagos finches (Bowman 1961). Historically known species differ not only in size, but also in shape of the bill.The Laysan Finch has a longer bill with a prominently overhanging maxilla, the Nihoa Finch a shorter, more compact bill with little maxillary overhang (Fig. 6.1).These features are difficult to assess in the subfossil species because the horny rhamphotheca is absent. The bill of the Palila is similar overall to those seen in Telespiza but is considerably foreshortened, resembling the bill of the cardueline Bullfinch (Fig. 6.1). The resemblance is, however, superficial because the Bullfinch feeds primarily on flowers and flower buds and does not use its feet to hold food (Newton 1973), whereas the Palila, like many honeycreepers, uses its feet to hold food items, in this case mamane seed pods that it rips open by pulling out a seam zipper-like (Fig. 6.2; Plate 1) to extract the seeds (Berger 1981, pers. obs.), against a substrate. Interestingly, the Nihoa Finch extracts seeds from pods of ohai in a virtually identical manner (Morin and Conant 2002). The term ‘grosbeak’ has been used for heavybilled finches in several avian families, including the Fringillidae.The Kona Grosbeak had the largest bill

86 The Hawaiian Honeycreepers

6.1 Cardueline and drepanidine finch profiles showing varieties of bill shape, from Pratt (1979): (a) Telespiza ultima; (b) T. cantans; (c) Rhodacanthis palmeri; (d) Chloridops kona; (e) Loxioides bailleui; (f ) Psittirostra psittacea; (g) Pyrrhula pyrrhula; (h) Coccothraustes coccothraustes; (i) Pinicola enucleator; ( j) Serinus sulphuratus; (k) Carpodacus mexicanus; (l) Carduelis sinica; (m) Acanthis flammea.

Form and function 87 placement in a monotypic genus ( James and Olson 1991), and its exact position among the Hawaiian finches is uncertain ( James 1998). Nothing is known of how it used its heavy bill.The koa finches Rhodacanthis palmeri and R. flaviceps had large, comparatively elongated bills with evenly curved maxillary tomia (Fig. 6.1). Both species, which differed primarily in size, were specialised feeders on the bean-like seed pods of the koa tree. They used the cutting edges of the bill to chop up young koa pods into smaller pieces or to tear into more mature pods to extract the green seeds (Olson 1999).

Elongated or hooked thick bills

6.2 Feeding by Palila: (a) holding mamane pod in feet to rip out side seam and extract seeds; (b) discarded pods showing characteristic openings along one side. Photos © Jack Jeffrey.

of the historically known Hawaiian finches, and the prehistoric King Kong Grosbeak had the heaviest of all ( James and Olson 1991).The upper bill in Chloridops is distinctive in having a curving (sinuated) cutting edge or tomium (Fig. 6.1) and large blunt ventral ridges ( James and Olson 1991). The Kona Grosbeak fed, as far as known, on the seeds of naio, which are encased in a hard endocarp. Cracking these fruits produced a very loud noise, which lured early collectors to the feeding birds (Munro, in Olson 1999). As Munro described, the fruits were cut into several pieces that were further processed by holding them to one side of the bill above a projecting ‘tooth’ on the lower mandible. Whether the other species of Chloridops were similarly specialised is not known. The prehistoric Mauka Grosbeak had a number of unique features, including unusually large nasal openings, that justify its

Several Hawaiian honeycreepers have bills that are somewhat finchlike, but elongated or otherwise modified so that the birds can no longer be called ‘finches’ in the colloquial sense. Clustered here are several superficially similar bills that are not phylogenetically related. Each is unique in its structural details and manner of use. James and Olson (1991) consider the birds grouped here to represent an evolutionary grade among the Hawaiian honeycreepers in which a basically finchlike bill is modified in certain ways that make its possessor no longer a true finch. The ‘O’u has an elongated, vaguely finchlike bill with a hooked and strongly overhanging maxilla (Fig. 6.1). It has no analogue in other passerine families, although the bills of mountain-tanagers, also mainly frugivores, show some resemblance (Isler and Isler 1987). The ‘O’u uses the bill to pluck small fruits (Fig. 8.2) whole, to take individual fruits from composites such as ‘ie‘ie (Fig. 8.1) or to bite into larger fleshy ones. One of the most bizarre bills among Hawaiian honeycreepers is that of the Lana‘i Hookbill Dysmorodrepanis munroi, known from a single specimen and long thought to be a deformed ‘O’u (Greenway 1939, but see James et al. 1989).The upper bill resembles a slightly more decurved ‘O’u maxilla, but the lower one is sharply curved upward so that the tip contacts the midpoint of the upper in resting position.Thus the closed bill exhibits a permanent gap or diastema between the mandibles. The tomia of both mandibles are turned inward so that

88 The Hawaiian Honeycreepers the bill lacks sharp cutting edges. As James et al. (1989) show, kinesis allows the tips of the mandibles to ‘oppose each other like ice tongs’ when the bill is fully open, and the tips to be brought together when both mandibles are depressed.The only specimen collected had berries in its stomach, but James et al. (1989) thought that such a bizarre bill adaptation could not have evolved simply for berrypicking because so many birds take such foods without unusual bill morphology. They speculate that the bird may have specialised on native land snails by carrying them in the diastema and using the hook to extract them from their shells in the manner of Snail Kites.Although this scenario could be dismissed as fantasy, it is an intriguing possibility that has to be considered until real information comes to light (but note the hazard of such speculation in the absence of field observations as discussed below under ‘crossbills’). The exact purpose of the hooked bill will likely never be known. Another elongated but thick bill is that of the Po‘o-uli Melamprosops phaeosoma.Again, certain tanagers have vaguely similar bills in profile, but there the resemblance ends.The closest analogue known to me is the bill of the Pink-billed Parrotfinch endemic to Viti Levu, Fiji, which also forages over the bark of trees but in a rather different manner (Pratt et al. 1987). As stated above, the Po‘o-uli is known to feed on snails, among other animal foods (Baldwin and Casey 1983) gleaned from tree limbs. A somewhat similar bill shape occurs in the prehistoric genus Xestospiza, but in that genus the bill is more conical and has a rather striking, but superficial, resemblance to bills of American cowbirds ( James and Olson 1991; Ziegler 2002). Another bill that falls into this grouping, but whose resemblance to the others is unquestionably superficial, is that of the genus Ciridops. It is a thick bill, longer than high, that resembles somewhat the bills of small tanagers (Thraupidae) in such genera as Tangara and Chlorophonia in the Americas (Isler and Isler 1987), and bills of dicaeid flowerpeckers and berrypeckers of the Indo-Pacific (Beehler et al. 1986). Richards and Bock (1973) considered Ciridops the most primitive drepanidine genus because of its rather generalised bill shape and what James and Olson (1991) properly regard as ‘a fancied

resemblance in plumage to the cardueline genus Leucosticte’. However, both upper and lower bills of Ciridops are unlike the bills of cardueline and drepanidine finches and are instead shortened versions of the comparatively weak bills found in other members of the ‘red and black’ clade ( James and Olson 1991). The feeding habits of the two described species can never be known for certain. The ‘Ula-‘ai-hawane is known from only five historical specimens, all of which were taken by Hawaiian collectors in the employ of Europeans who unfortunately denigrated the information that could have been gained from native informants.We can gain some inference from the Hawaiian name, which literally translates ‘red, eats palm fruit’ (it probably was preceded by the word manu, ‘bird’), but perhaps better translates as ‘red palmcreeper’. The palms referred to are the native loulu, the soft fruits of which are called hawane (Fig. 11.14, Part II).

Parrot-like bill The aptly named Maui Parrotbill has a unique feeding apparatus that, although superficially convergent with those of parrots, differs from them in important respects. Whereas most psittacine bills are bulbous and broad as viewed from the front, that of the parrotbill is laterally compressed and is thus more parrot-like in profile than in a head-on view (Fig. 6.3). The tips of both upper and lower mandibles are sharp, and the cutting edge in the middle of the lower mandible is flared downward,

6.3 Female Maui Parrotbill showing lateral compression of bill. Photo © Jack Jeffrey.

Form and function 89 apparently for a better grip during feeding manoeuvres (Mountainspring 1987). Parrotbills are the most strikingly sexually dimorphic honeycreepers in bill size, with sex easily determined in the field by bill size alone (Mountainspring 1987; Simon et al. 1997). Because the bill is thick in lateral aspect, it has often been considered to be directly derived from a finchlike bill and has particularly been compared to the bill of the ‘O’u (Raikow 1977b), although recent evidence indicates it is secondarily thickened from possibly a heterobill ancestor (Pratt 2001b). Bill function of the Maui Parrotbill is totally different from that of finches, undoubtedly because the bird is almost totally insectivorous (Simon et al. 1997).The thick bill notwithstanding, the parrotbill forages in ways strongly reminiscent of the feeding

of the thin-billed ‘Akiapola’au (see below) with upper and lower bills often used independently (Fig. 6.4). Like the ‘Akiapola’au, the Maui Parrotbill has no exact mainland equivalent in use of the bill. The closest parallels are the Wattled Ploughbill of New Guinea (Beehler et al. 1986) and some of Madagascar’s vangas (see also Chapter 4). Mountainspring (1987) classified bill manoeuvres of Pseudonestor in five categories in descending order of importance: excavation (peeling away bark and digging into wood); twig-splitting (crushing a twig between the mandibles); gleaning from surfaces; probing into flowers or leaf clusters; and plucking fruit. To this I would add a sixth, slicing open or crushing fruit, which Mountainspring included in excavation. As described by Mountainspring (1987), an excavating bird digs its upper mandible

6.4 Foraging methods of Maui Parrotbill: (a) crushing dead twigs to extract larvae; (b) pulling away pieces of bark; (c) wrenching large chunk of dead wood; (d) gouging into substrate with lower mandible. Photos © Jack Jeffrey.

90 The Hawaiian Honeycreepers into a crack or crevice, then grips and peels away pieces of bark to expose insect tunnels. These are then further exposed until the prey is discovered. The upper mandible may be used as a plough, scraper, or hammer. Mountainspring (1987) reported a single instance in which a bird opened its bill fully and hammered with the lower mandible in a manner identical to that of the ‘Akiapola‘au. I have observed this method of excavation at least twice. I have also observed a variation on this manoeuvre not reported by Mountainspring, but recently described by Baker and Baker (1997a), in which the tip of the maxilla grasps a small branch on the opposite side from the bird’s body and the mandible then chisels at the wood. The incredible strength of the bill and jaw muscles of Pseudonestor are most evident in twig-splitting (Fig. 6.4a), wherein the bird simply bites down on a brittle twig at right angles to break it and expose burrowing larvae. Particularly tough twigs require strong wrenching movements of the head to complete the process. Mountainspring (1987) regarded gleaning as an opportunistic use of the bill, particularly when young were being fed on caterpillars. Probing manoeuvres are used to capture insects in unaccustomed sites, and are also associated with occasional nectar feeding (Simon et al. 1997).Whether the Maui Parrotbill actually eats fruit is subject to some question. Numerous observers have seen them pluck fruits or slice into them attached to the plant. Plucked fruits are held against the upper bill and sliced by movement of the lower mandible. Simon et al. (1997) consider these activities to be searches for insect larvae. However, Hopper et al. (1993) and I have observed feeding on fruit that may not have involved insects (see Chapter 8 and species account).

reminiscent of spruce cones (Fig. 6.6), which may explain the convergence. Loxops bills are only slightly crossed and the feature is not readily discernible in the field. Bill tips may cross to either left or right in about equal proportions (Hatch 1985).

Crossed or twisted bills Only two examples of crossed bill tips are known in birds: the true crossbills and the Hawaiian ‘Akepa and ‘Akeke‘e. Crossbills pry open cones of various kinds of conifers, and thus have very stout rhamphothecae (Newton 1973; Nethersole-Thompson 1975). The ‘akepas feed among leaf buds and leaf axils (Fig. 6.5) and have much weaker bills. The imbricated buds of ‘ohi‘a are structurally

6.5 Foraging in terminal leaf clusters of ‘ohi‘a by members of genus Loxops: (a) ‘Akeke‘e; (b), (c) ‘Akepa. Photos © Jack Jeffrey.

Form and function 91 1997; Conant et al. 1998). A feeding Loxops inserts its closed bill between surfaces that are to be spread apart, such as scales of ohi‘a buds, and then gapes. The opposite-pointing bill tips spread the surfaces apart, and the tongue is extended and retracted between to snare small prey items.

Warbler-like bills

6.6 Imbricated scales on leaf buds of ‘ohi‘a mimic morphology of spruce cones as shown in the unusually large buds of endemic Kaua‘i variety dieteri, which perhaps influenced selection for larger bill size in the ‘Akeke‘e.

In overall aspect, Loxops bills are rather finchlike, especially as compared to the sharp conical bills of carduelines such as siskins and redpolls. The asymmetry of the mandibles is reflected in the underlying bone and muscle structure (Richards and Bock 1973).Those authors used these anatomical features to develop a complex scenario of how the bill might be used in foraging. Readers of that lengthy discussion (Richards and Bock 1973: 81–94), embellished with four fanciful figures, should be cautioned that it is entirely speculative and that virtually none of it has been confirmed by subsequent field observations (Conant et al. 1998). Particularly prominent in the Richards/Bock scenario are twisting movements of the head, which have never been observed despite lengthy observations from Perkins (1903) to the present (Lepson and Freed 1997; Lepson and Pratt 1997).The lesson here is that imaginative speculations, based solely on interpretations of anatomical features, even by reputable scientists, should not be given much credence (is anyone in palaeontology listening?).The temptation to speculate is great, especially when the only information we may ever have is anatomical, as in the case of subfossil Hawaiian honeycreepers. At least Richards and Bock (1973) had the courage to speculate about birds that could still be observed. The actual manner of bill use by ‘Akepa and ‘Akeke‘e is now fairly well documented (Benkman 1989; Lepson and Freed 1997; Lepson and Pratt

Like the term ‘finch’, the name ‘warbler’ has been bestowed on a variety of unrelated birds around the world.All of them have bills that are long, fairly thin, and straight or with slight curvature. They tend to exhibit a straight or convex (as opposed to concave as seen in ‘amakihis) gonys.Typical warbler-like bills are found in the Old World Sylviinae and the American wood-warblers (Parulidae). Birds with such bills most often feed by gleaning prey from leaf surfaces, but most also have other feeding methods, including some examples of frugivory and nectarivory, and relatively few could be considered specialised. Among the Hawaiian honeycreepers there are two examples of warbler-like adaptations. The bill of the ‘Anianiau is an all-purpose tool that allows the bird to feed in a variety of ways, although leaf-gleaning (Fig. 6.7b) is the most important (Lepson 1997). Secondarily it feeds on nectar (Fig. 6.7a), but because its bill is only slightly decurved, it chooses mostly open flowers such as those of ‘ohi‘a and various shrubs (see also Chapter 8). The nasal operculum is well developed as would be expected in a frequent nectar-feeder. The other warbler-like honeycreepers belong to the genus Paroreomyza, but their bills are distinctive (Fig. 6.7c, 6.9) in several structural details and are not homologous to the superficially similar bill of the ‘Anianiau. Behaviourally, they are rather similar to that species, and the fact that they do not occur on Kaua‘i could be an example of competitive exclusion (Chapter 4). The O‘ahu ‘Alauahio, Maui ‘Alauahio, and the Kakawahie are primarily gleaners of leaves and branches. The Maui ‘Alauahio, by far the best known of the three, also catches insects on the wing in aerial sallies and feeds occasionally on nectar (Carothers 1982; H. and P. Baker 2000).These feeding behaviours were never reported in the other species, but observations are too few to exclude the

92 The Hawaiian Honeycreepers ‘medial slot with parallel lateral ridges’ which, with the simple tongue as the lower part, forms a nectar tube (Richards and Bock 1973). Despite these apparently nectar-specific adaptations, Maui ‘Alauahio are less efficient in foraging even on flowers with short corollas when compared with ‘I‘iwi and ‘Apapane (Carothers 1982).

Short bark-picking bills Two Hawaiian honeycreepers that forage in the manner of nuthatches comprise the genus Oreomystis and are commonly referred to as ‘creepers’. The Hawai‘i Creeper and the ‘Akikiki forage on trunks and branches of trees (Fig. 6.8) somewhat in the manner of nuthatches, Holarctic creepers and Australian treecreepers. However, they are not an exact parallel to any of these groups. Unlike nuthatches and holarctic creepers, they do not brace their tails

6.7 Hawaiian honeycreepers with warbler-like bills: (a) ‘Anianiau taking nectar from lehua blossom; (b) ‘Anianiau foraging in leaves; (c) Maui ‘Alauahio. Photos © Jack Jeffrey.

possibility that they may have also fed in these ways. Importantly, the adaptations for nectarivory in the Maui ‘Alauahio are entirely different from those of other nectar-adapted honeycreepers, including the ‘Anianiau. The horny palate has a longitudinal

6.8 Bark-picking by ‘Akikiki vertically (a), and head down (b). Photos © Jack Jeffrey.

Form and function 93

6.9 Profiles of Hawaiian ‘creepers’, Paroreomyza at left, Oreomystis at right. From Pratt (1979, 1992).

against tree trunks, and they differ in bill shape from both of them. They are closer to Climacteridae in these respects, but have heavier, more finchlike bills. Bills of Hawaiian creepers are slightly downcurved, with a concave gonys, but are fairly heavy at the base (Fig. 6.9).The fundamental difference in contour of the gonys as compared to that of ‘alauahios was one of the first clues that Amadon’s (1950) “Creeper” was an amalgam (Pratt 1979a). The bill is used to pick and pry into cracks and crevices of tree bark, to probe under lichens and mosses, and to peck, sometimes strongly, at the bark surface in search of arthropod prey. Fairly prominent retroarticular processes of the mandible show that the creepers also use gaping movements to expose prey (Richards and Bock 1973).

Probing and gaping bills A few honeycreepers appear to use probing and gaping motions as their primary feeding manoeuvre.

The Greater ‘Amakihi was the only historically known species in this category. According to Perkins (1903) it fed primarily by probing into the wrap-around leaf bases of ‘ie‘ie vines (Fig. 2.9). Secondarily it sought its large beetle prey under loose bark of ‘ohi‘a and among ferns. The bill was relatively long and nearly straight, but with a very slight downward curve. Gaping was an important manoeuvre, as shown by the very long retroarticular process (Richards and Bock 1973), relatively the longest in any honeycreeper except the following. Among recently described prehistoric species are three members of the new genus Aidemedia that appear to be very similar in general morphology to the Greater ‘Amakihi ( James and Olson 1991). Their bills are longer and more decurved, but have the same general shape and long retroarticular processes seen in the Greater ‘Amakihi. Indeed, that species, Maui-nui Gaper, Straight-billed Gaper, and Curve-billed Gaper form a smooth morphocline in degree of curvature that is reminiscent of Bock’s (1970) ‘microevolutionary sequences’, and might actually form a transformational series. James and Olson (1991) describe several structural details that led them to separate the Greater ‘Amakihi from Aidemedia generically, but James (1998) put them in the same clade.The bills of Hawaiian gapers superficially resemble those of meadowlarks, or the European Starling (Ziegler 2002), but details ( James and Olson 1992) show Aidemedia to be drepanidine.

Shovel-tipped bill One Hawaiian honeycreeper bill type is known only from remains of prehistoric species. It is long and delicate, deep and broad, with a very blunt, rounded tip.The somewhat shovel-like tip inspired the name Vangulifer (‘little shovel-bearer’) for the genus ( James and Olson 1991). The Pololei Shovelbill and Kiwi Shovelbill differ in degree of curvature of the bill and in a few other features. James and Olson (1991) consider them congeneric because they share the blunt tip and short retroarticular processes associated with a long delicate bill, but James (1998) considered the genus paraphyletic.This peculiar bill has no apparent equivalent outside Hawai‘i. It is ‘too long and weak to be designed for seed cracking; too deep and broad to be suited for probing in bark; too blunt

94 The Hawaiian Honeycreepers for a nectarivore; and . . . unlikely to have been used for forceful gaping’ ( James and Olson 1991: 63).The describers suggest that Vangulifer might have been a fly-catching honeycreeper, comparing it to todies and tyrant flycatchers, other birds that capture prey in aerial sallies.Those groups have much flatter bills than Vangulifer, but when monarchine genera such as Monarcha and Myiagra, or for that matter Chasiempis, which have deeper and blunter bills than most tyrannids, are included in the mix, the case for shovelbills as generalised flycatchers is strengthened.

Insectivorous and multipurpose sickle-bills So many Hawaiian honeycreepers have sickleshaped, down-curved, or falcate bills that some writers (e.g. Lack 1947) refer to the whole group as ‘Hawaiian sicklebills’. Several variations on this theme occur, and curved bills have apparently evolved independently within the honeycreepers at least twice, with both apomorphies giving rise to short and long versions. Most of the falcate bills are highly specialised, but that of the ‘amakihis has enabled its possessors to exploit a variety of foods and habitats (Fig. 6.10), and to become arguably the most successful, adaptable, and resilient of Hawaiian honeycreepers. Their bills are fairly heavy at the base, short, down-curved and tapering evenly to an acute tip, with a very slightly overhanging maxilla. The three species differ among themselves in the percentages of various feeding activities, but all are considered ecological generalists (Lindsey et al. 1998; Pratt 1999b). All ‘amakihis use the bill to extract nectar from both open flowers and those with short curved tubular corollas. All species are also insectivores, but the Kaua‘i ‘Amakihi has a much larger bill than the others and spends a greater proportion of its time prying and probing bark (pers. obs.; Lindsey et al. 1998) for arthropods. The bill is also suitable for leaf and twig gleaning as well as probing in clumps of dead leaves (Lindsey et al. 1998). The ‘akialoas are generally considered to be primarily insectivores, but accounts by Perkins (1903) emphasise the importance of nectar in their diet (see Chapter 8). At least three major variations of

6.10 Foraging by ‘amakihis: (a) Hawai‘i ‘Amakihi taking nectar from lehua blossom; (b) Hawai‘i ‘Amakihi feeding on ornithophilous mamane flowers, whose corollas curve to match the bird’s bill; (c) Kaua‘i ‘Amakihi picking at bark for insects. Photos © Jack Jeffrey.

the ‘akialoa bill can be discerned. Most species exhibit a very long, shallowly curved bill, but the Lesser ‘Akialoa had a shorter and much more

Form and function 95 sharply decurved bill.The Hoopoe-billed ‘Akialoa, sympatric with other large species on Kaua‘i and O‘ahu in pre-human times, differs from the others in lacking the lingual trough, a groove in the mandible into which the long tongue rests in the other ‘akialoas.This qualitative difference may indicate that this bird had a shorter or differently shaped tongue ( James and Olson 1991), and may have fed less or not at all on nectar. If that were the case, it would help to explain the broad sympatry with a partly nectarivorous relative. Perkins (1903) considered the feeding habits of ‘akialoas intermediate between those of ‘amakihis and nukupu‘us ‘since they are greater nectar-eaters than [nukupu‘us] and hunt more persistently, creeper-like, on the limbs of forest trees for wood- and bark-eating insects than does the [Hawai‘i ‘Amakihi.]’ (see also Chapter 9). In contrast, Henshaw (1902), who made his observations shortly after Perkins, when ‘akialoas and many other Hawaiian forest birds were becoming rare, never saw them feed on nectar. His observations of insectivorous feeding habits of H. obscurus bear repeating here (Henshaw 1902: 38): The stout legs and sharp claws of this bird enable it to pass rapidly over the large limbs of koa and ohia, and to cling in any [position] and to seize from their hiding places in the tangles of ferns and mosses various sorts of grubs, beetles and their larvae which are its chosen food . . . The akialoa is especially fond of probing into the deep, cup-shaped leaf cluster of the ieie vine. These are usually full of dead leaves and fallen trash, in the recesses of which insects are perfectly safe from all birds except the akialoa and its relative, the akiapolaau, the latter being furnished with a similar hooked probe. I have no doubt that the peculiar leaf clusters of the ieie have had more to do with the development of the extraordinary bills of these two birds than anything else. Perkins (1903: 423–4) also compared the insectivory of the ‘akialoas with that of the nukupu‘us: The insects on which [the nukupu‘us and ‘Akiapola‘au] feeds are sought for mainly

in similar situations to those affected by [‘akialoas], namely in or beneath the bark of trees and in decaying wood; and in fact the very same species are frequently found in the stomach of each of these birds, but those of the first named . . . have a more varied diet. In the large Koa trees of open woods in Kona from 2000–3000 ft. above the sea the Akialoa could be traced by its audible tapping on the bark, the sound resembling that produced by the strokes of the beak of the Nukupuu, except that it was less loud. In the excessively wet forests of windward Hawaii it was sometimes seen on the stems of the treeferns, or amongst the masses of climbing Ieie (Freycinetia arborea), exploring with its long beak in the first case the cavities of the stems of old fronds, broken off close to the caudex, and in the second the bases of the stiff clasping leaves and the debris found there, in both of which situations insects habitually harbour. In the subgenus Hemignathus, the heterobills, the lower mandible is about half the length of the upper, and the two parts are often used independently. In the three species of nukupu‘u the lower mandible is curved to fit into the upper. The Akiapola‘au differs from them in having the lower mandible straight. It does not follow the contour of the maxilla, and a slight gap or diastema may be present in the closed bill. The independent use of the mandibles reaches an extreme in the ‘Akiapola‘au, although cranial kinesis allows the tips of the mandibles to be brought together for picking up and manipulating prey. Early observers report that the feeding movements of the nukupu‘us and the ‘Akiapola‘au were essentially the same, except that those of nukupu‘us were weaker and less vigorous. The ‘Akiapola‘au is perhaps the most remarkable of all Hawaiian honeycreepers in its feeding apparatus. Even early naturalists realised that it is Hawai‘i’s ecological equivalent of a small woodpecker, a niche unfilled in the islands because woodpeckers are such poor transoceanic colonisers. Had any small woodpecker been present, the makeshift adaptations of the ‘Akiapola‘au undoubtedly

6.11 Bizarre foraging movements of ‘Akiapola‘au: (a) raring back to strike a blow with lower mandible; (b) position for oblique hammering at large branch; (c), (d), (e) bracing with lower mandible while inserting upper to probe for larvae; (f ) pulling away a piece of bark (compare to Maui Parrotbill, Fig. 6.4c); (g) using mandible tips together to pick something from a twig; (h) manipulating prey with opposed bill tips; note in latter two that lower mandible can swing out quite far. Photos © Jack Jeffrey.

Form and function 97 could never have evolved. Evolution is only as good as it has to be. The woodpecker analogy is most obvious in the bird’s pecking movements, but it even extends to the formation of rows of sap wells (see also Chapter 8) on tree trunks (Fig. 8.6), reminiscent of those made by North American sapsuckers.The ‘Akiapola‘au uses its combination-tool bill in a variety of ways (Fig. 6.11), and because of fairly large bill size differences between the sexes, males and females differ somewhat in feeding behaviour. Probably the most improbable feeding mode involves opening the bill to its widest gape, and pecking vigorously at the bark of a tree, whether obliquely on small branches with the upper mandible held out of the way, or directly on large trunks. Perkins ( 1903) believed that both mandibles struck the substrate in the latter case, but Munro (1960) disagreed and recent film footage (T. K. Pratt, pers. comm.) shows only the mandible making contact. Even if the maxilla did strike the surface, its strength and flexibility (Perkins 1903) would preclude damage. Females use similar movements to hack into clusters of dead koa leaves. Perkins (1903: 428) describes the amazing vigour of this activity: Into these blows it throws its whole weight, swinging backwards from the thighs to renew its stroke. In some cases these blows, if not for the purpose of driving out hidden insects, at least have that result, for several times I saw the bird after a stroke make a sudden dart, sometimes even taking an insect on the wing . . . Not infrequently it lays hold of a projecting piece of bark or the stump of some small broken branch, and shaking its head from side to side and pulling in all directions, endeavours to tear it out. If unsuccessful in the attempt, it will alternate this treatment with a shower of blows from the points of its gaping bill, till either the stump gives way or the labourer is convinced of the futility of further efforts. In this passage, he mentions a wrenching motion reminiscent of that described earlier for the Maui Parrotbill. Henshaw (1902) also refers to

such movements and remarks that ‘the skull is unusually thick, and the muscles of the neck and the maxilla are remarkably developed so as to permit this double function of the bill as a hammer and as a wrench.’ Perkins’s (1903) description of the movements of the bird and the independent use of the upper bill remain as apt as when first written: Their climbing movements are more perfect than those of other birds, and they creep over the tree-trunks either upwards or downwards with equal ease, and hunt both along the upper and under surface of the branches . . . in the high forest of Kona I first had the opportunity of watching one of these birds in pursuit of food . . . It was visiting one after another a number of fallen tree-trunks, large but smooth-barked examples of Acacia koa.Along each of these it proceeded from one end to the other, peering now over the right side of the trunk and now over the left, so that in a single journey it searched both sides of the tree without retracing its steps. The upper mandible is thrust into small holes or cracks in the wood, while the point of the lower presses on the surface of the bark, and in this manner the burrows of wood-boring insects are opened out. So too it thrusts its upper beak under loose pieces of bark, resting the lower one on the surface, and breaks off fragments of bark, under which its food is concealed.The upper mandible, though so slender as to be slightly flexible, is very strong, and this flexibility of the head aided by the extreme flexibility and strength of the neck no doubt greatly assists the bird in exploring and opening out the burrows. Fortunately, although endangered, the ‘Akiapola‘au is still common enough to be observed with some frequency. I can confirm all of the early naturalists’ descriptions from my own work and Jack Jeffrey has captured many of them on film (Fig. 6.11). Additional details are given in the species account.

98 The Hawaiian Honeycreepers

Flower-adapted sickle-bills The three primarily nectarivorous sickle-billed honeycreepers are often regarded as the quintessential representatives of the group. They include not only the first species described (Hawai‘i Mamo) but the species most familiar in modern iconography (‘I‘iwi). Their bills differ from those of the thinner-billed sicklebills in that they are supported by bone throughout their length (Baldwin 1953), and thus probably represent an independent adaptation. All of them have fully developed nasal opercula characteristic of exclusive nectarivores (Raikow 1977b). The Hawai‘i Mamo and the Black Mamo were highly specialised to feed on the curved flowers of arborescent lobelioids. Their bills fit the corollas of certain species quite closely, and are likely coevolved with them (see Chapter 8). Bryan (1908) watched a Black Mamo inserting its bill ‘up to the nostrils, in the flower, while the bird balanced itself on the branches, assuming almost every imaginable attitude in its operations.’ Both mamos also fed occasionally on ‘ohi‘a nectar, but a far more common visitor to those flowers is the ‘I‘iwi. Despite the seeming clumsiness of the larger sickle-bills, these species are surprisingly more efficient foragers at short flowers than nectarivores with shorter bills that ‘fit’ better (Carothers 1982). The ‘I‘iwi has far more generalised feeding habits (Fig. 6.12) and appears far more adaptable than the mamos, although its bill apparently evolved in the same way to take advantage of curved lobelioid flowers (Perkins 1903). It feeds mostly in ‘ohi‘a flowers (Fig. 6.13), but includes a wide variety of native and introduced plants in its diet (see Chapter 8) and also takes some insects (Fancy and Ralph 1998). When feeding in ‘ohi‘a, ‘I‘iwi employ a very different approach and posture as compared to the short-billed nectarivores (below). They tend to approach the flower clusters from below and reach upward, placing the tip of the curved bill into the calyx cups almost like a siphon (pers. obs.). Such movements avoid contact with the stamen and pistils, so the value of ‘I‘iwi

6.12 The sickle-shaped bill of the ‘I‘iwi matches the curvature of many native flowers as in this very precise fit with flowers of Kaua‘i koli‘i, a native lobelioid; note contact of pistil with bird’s head. Photo © Jack Jeffrey.

6.13 ‘I‘iwi feeding in lehua blossoms (‘ohi‘a). Photo © Jack Jeffrey.

as ‘ohi‘a pollinators (Carpenter 1976) may be overstated. Spieth (1966) described the acrobatic movements of an ‘I‘iwi as it positioned its bill for insertion into a lobelioid flower. However, many recent observers (summarised by Fancy and Ralph 1998) report that ‘I‘iwi frequently pierce such corollas at the side of the base, as they do various introduced flowers whose corollas are not the proper ‘fit’.

Form and function 99

Short nectar-adapted bills Three members of the nectarivorous clade of Hawaiian honeycreepers have thin, short bills that are not suitable for feeding in deep tubular corollas. The ‘Apapane, Laysan Honeycreeper, and the ‘Akohekohe instead feed on flowers with open faces and exposed nectaries. Their bills have full nasal opercula and are slightly decurved, but not so much that they can be called sickle-bills, and the bill of the ‘Akohekohe is nearly straight.These bills have very close analogues among the honeyeaters (Meliphagidae) of the Indo-Australian region. In particular, the superficial resemblance between the two Himatione and members of the genus Myzomela is quite striking, both in bill shape and coloration. Why honeyeater-like honeycreepers should have evolved by convergence when genuine meliphagids (Moho and Chaetoptila) were present in Hawai‘i is an intriguing question that can probably never be answered definitively (but see Chapter 4 for some speculation). Both the ‘Apapane and the ‘Akohekohe feed primarily in flowers of ‘ohi‘a (Fig. 6.14), and the latter appears to be a Metrosideros specialist. Both are undoubtedly important pollinators (Carpenter 1976) and have foraging actions and postures that assure pollination of ‘ohi‘a flowers. Characteristically, a bird lands in a terminal clump of a flowering ‘ohi‘a, then moves to the outside of the inflorescences, completely exposed, and dips its bill straight through the brush-like stamens and pistils to reach the nectar at the base (Fig. 6.14). This approach behaviour is in contrast to the more surreptitious movements of the ‘I‘iwi. Birders have learned that, even though an ‘Akohekohe may skulk in dense leaves for a time, when it starts to feed it will pop into full view (Pratt 2002b). Whether that bird’s strategically placed, stiff, forward-projecting crest evolved as a pollen brush rather than an ornament is probably an unanswerable question, but it certainly functions in pollination, purposely or not. In a bird that is not sexually dimorphic, sexual selection was probably not a factor, yet we see no evidence of coevolution (Chapter 4) with ‘ohi‘a (Metrosideros is

6.14 Feeding by short-billed nectarivores: (a) ‘Apapane; (b) ‘Akohekohe behaving like a gymnast to reach lehua blossoms; note precise meeting of stamen tips and crest. Photos © Jack Jeffrey.

widely distributed in the Pacific with similar flower structure everywhere).The ‘Apapane has no such adaptation, but its activities, including almost

100 The Hawaiian Honeycreepers literally running through clumps of flowers, assures pollination.

Feeding apparatus: the tongue The tongue is just as important as the bill in food gathering, and the two work together very precisely.Thus we would not be surprised to find each bill type described above with its own peculiar tongue adaptation. But such is not the case. Honeycreeper bills are far more variable than their tongues (Fig. 6.15). Apparently a particular tongue structure will function with a variety of bill shapes. This relative uniformity of tongue morphology among divergent adaptive types was one of the earliest evidences that the Hawaiian honeycreepers are a natural group. Most drepanidine tongues are of two fundamental types, one resembling tongues of cardueline finches and the other being a complex and highly evolved structure unique to the Hawaiian honeycreepers. The only feature that reveals these very different tongues to be related is the presence in all of the obviously derived condition (see Chapter 4) in which the base of the tongue at the back of the throat is truncate or squared off, rather than notched or V-shaped. Early collectors often preserved the tongues of Hawaiian honeycreepers by drying them and attaching them to the study skins. Unfortunately, this practice caused subsequent workers to be badly misled. As the tongue dries, the fleshy part loses its turgor and the corneous underside pushes in, causing the tongue to curl longitudinally.Workers from Gadow (1899) to Gardner (1925) to Amadon (1950) believed that tongues of Telespiza, Psittirostra, and Pseudonestor were tubular or partly tubular, and cited this feature as evidence that they evolved from a tubular ancestor. Raikow (1977b), by studying fluid-preserved tongues, dispelled this myth (at least for Psittirostra and Telespiza), and showed that all drepanidine finch tongues were of the same fleshy, non-tubular type.

Seed-cup tongues Most drepanidine finches have tongues that, at the distal end, closely resemble those of

cardueline finches.They are sometimes called seedcup tongues because of a small concavity near the tip ‘that probably aids in scooping up small food items or holding them against the roof of the mouth during manipulation’ (Raikow 1977b: 106). Raikow describes such tongues as thick and fleshy with a shallow furrow down the middle of the dorsal surface. The fleshy part of the tongue tapers evenly toward the tip.The underside of the tongue has a corneous covering that is frayed at the anterior end so that a fringe sticks out beyond the fleshy part. The fringe may extend along the sides of the tongue as well.This resilient ventral covering curves up along the sides but never meets in the middle. This shaping helps to form the anterior cup. Interestingly, the tongue of Telespiza is closest in structure to those of some cardueline finches, and could easily have been the basic type from which all the others evolved.Variations on this theme are slight. The tongue of Loxioides is shorter and more rounded with a deeper cup; those of Rhodacanthis (Gadow 1899) and Chloridops (Amadon 1950) are very compact and deeply scoop-shaped, with very narrow distal fringes; that of Psittirostra is more attenuated with a pronounced terminal fringe (Raikow 1977b).

Warbler-like tongues Many small insectivorous passerines have tongues that are narrow, thin, non-tubular, and slightly bifid (divided in two) or frayed at the tip (Gardner 1925). Richards and Bock (1973) describe and illustrate similar tongues in Oreomystis mana and Paroreomyza montana.The tongue of the latter, however, is virtually identical to those seen in many passerines, but that of Oreomystis differs in lacking lingual wings, a feature that shows it to be derived from the drepanidine finch type just described and presumably convergent with the more generalised passerine type (Pratt 1992b). The tongue of O. bairdi as illustrated by Gadow (1899) appears identical to that of its congener, and because this type of tongue is unique to this genus among Hawaiian honeycreepers, it is one of the strongest pieces of evidence that the Hawai‘i Creeper belongs here and is not more closely related to the ‘amakihis or ‘akepas (Pratt

Form and function 101

6.15 Tongues of Hawaiian honeycreepers (from Pratt 1992b): (a) based on Gardner (1925) with alterations from Raikow (1977b); (b), (e), and (j) after Gadow (1899); (c), (g), (h), (i), and (n) after Richards and Bock (1973); (d) and (l) after Gardner (1925); (f ) after Frohawk (in Rothschild 1893–1900); (k) after Bock (1972); (m) after Bock (1978).

102 The Hawaiian Honeycreepers 2001b). As for the tongue of Paroreomyza, it once was viewed as indicating that the relationships of this genus lie outside the Drepanidinae (see Chapters 4 and 5). Even if we hypothesise that Paroreomyza is an early offshoot of the Hawaiian honeycreepers, before the loss of lingual wings occurred, the tongue’s similarity to those of wood warblers and other small passerines is not easily explained because it would have to have evolved secondarily from a cardueline finch ancestor.

Drepanidine tubular tongue The best detailed description of the characteristic tongue of nectarivorous Hawaiian honeycreepers is that of Raikow (1977b). Basically, such tongues look as if a secondary structure has been grafted onto the distal end of a finch tongue. This tubular part of the tongue is formed by narrow laciniae, thread-like projections along the lateral edges of the tongue that arch upward and inward to interlace dorsally forming a tube. Longer laciniae at the tip form a forward-pointing brush. The tube is formed mainly by cornified tissue and is not at all fleshy. Tongues that fit this description, varying only in the overall length and proportion of the tubular part (Fig. 6.16), are found in all of the ‘red and black’ honeycreepers including Drepanis, Himatione, Palmeria, and Ciridops (Bock 1972), and all of the hemignathine group including Loxops, Magumma, and all subgenera of Hemignathus. The lone exception would be the Hawai‘i Creeper if, as some have advocated (see Chapter 5), it were to be placed in Loxops or Hemignathus. This type of tongue is clearly an adaptation to nectarivory, which is accomplished at least partly by capillary action (Amadon 1950) possibly aided by muscular sucking at the back of the throat (Richards and Bock 1973), but no mechanism for that has been described. The yellowish brushy tip of the tongue often protrudes from the closed bill of birds that have been actively feeding. Perkins (1903: 401) said the tongue of a Black Mamo obtaining water from wet clumps of moss ‘darted in an out with great rapidity so as to appear like a liquid streak, the eye not being able to distinguish each separate movement.’ However, the drepanidine tubular tongue is

6.16 Tubular tip of drepanidine tubular tongue: (a) with fully developed brush-like tip in the ‘I‘iwi, photo © Jack Jeffrey; and (b) in a shorter, less brush-tipped version in the ‘Akiapola‘au, photo © Jaan K. Lepson.

not so specialised as to preclude feeding on arthropods and indeed seems to be well suited to that task (Richards and Bock 1970), and some of its possessors are almost totally insectivorous. The brush-like tip is especially useful for entangling prey. How they are extricated from the brush and swallowed is not known.

The parrotbill’s tongue The tongue of Pseudonestor is in a category by itself, although it is clearly drepanidine. Rothschild (1893–1900) illustrated a parrotbill tongue (Fig. 6.15f ) with its edges curled slightly upward. In contrast, Gadow (1899) described the tongue, as compared to that of Psittirostra, as ‘far less fleshy, more slender, more deeply split in the middle; the distal third of the horny sheath becomes gradually transparent towards the tip, is very slightly

Form and function 103 frayed out towards the tip and on the lateral edges, but shows no indication of curling up of the free margins’ (italics mine).Amadon (1950), based solely on the illustration and ignoring Gadow’s explicit statement to the contrary, described the tongue as ‘partially tubular’. In my first eclectic (i.e. noncladistic) classification (Pratt 1979a), I placed Pseudonestor in a position intermediate between the finchlike honeycreepers and the thin-billed ones. Previously, Bock (1970) had placed Pseudonestor in a similar intermediate position but in a totally different theoretical framework that envisioned derivation of seed-cup tongues from tubular tongues via this intermediate. Derivation of the Pseudonestor tongue from a seed-cup ancestor by lengthening the corneous portion and the terminal laciniae, or from a tubular ancestor by unrolling and shortening the tubular portion seem equally plausible, but Bryan and Seale (1900) note that tongues of nestlings of the ‘Anianiau are not yet rolled into a tube, so perhaps the parrotbill’s tongue is an example of neoteny. Because of a ‘unique character complex involving morphology of at least the bill, skull, and jaw muscles’, Zusi (1989) considered Pseudonestor unlikely to be ancestral to any other known honeycreeper. Thus it appears that, as has turned out to be the case many times, Perkins may have had it right all along in considering the parrotbill a highly derived relative of the heterobills (Pratt 2001b). As such it is the only member of the tubular-tongued clade known to possess a secondarily derived tongue structure.

blunt serrations along the medial groove and spoon-like tip to be vestiges of the laciniae that form the drepanidine tube. However, those serrations are in the fleshy part of the tongue, not in the cornified undersurface that actually forms the tube. In light of the other peculiarities, I doubt that the serrations have any connection whatsoever with the drepanidine tube. Indeed, the tongue of Melamprosops provides no evidence at all that it belongs among the Hawaiian honeycreepers.

The tongue of Melamprosops

Pterylosis

The tongue of the Po‘o-uli as described by Bock (1978) is unique among passerines. It is thick and fleshy throughout its length with a shallow groove down the dorsal midline that expands distally to form a spoon-like depression with a few blunt serrations along the sides. The tip of the tongue is rounded and smooth with no indication of the frayed tip or indentations seen in most passerine tongues, including those of drepanidines. At the rear, it has a very prominent pair of lingual wings armed with large tubercles. Bock considered the

Pterylosis is the distribution of feathers over a bird’s body. Although they may appear to be feathered all over, most birds’ feathers actually grow only in certain areas, called feather tracts, interspersed with patches of bare skin called apteria.These features are not visible externally because the overlapping feathers cover the bare places. Early researchers looked for phylogenetic significance in the variations in pterylosis, but found little among passerines. Feather distribution has been either illustrated or described for ‘O‘u, Palila, Lesser ‘Akialoa, Kaua‘i ‘Akialoa, and

Feeding apparatus: nostrils Many nectarivorous birds, including some of the Hawaiian honeycreepers, have a thin flap of tissue over their nostrils called a nasal operculum. Its exact function is unproven, but presumably it keeps the birds from getting pollen up their noses. In honeycreepers, the operculum grows from the dorsal margin and is shaped so that the opening of the nostril is a slit with a concave dorsal edge (Raikow 1977b). It is confined mostly to the species that also have a nectar-adapted tongue, the one exception being the ‘O‘u, which has a fairly large operculum but a simple finchlike tongue. Raikow (1977b) found that the nostrils of the other drepanidine finches and the ‘Akikiki do not differ in any important way from those of cardueline finches. The nasal operculum in fruit- and nectar-feeding honeycreepers is undoubtedly the result of convergence.

Plumage and coloration

104 The Hawaiian Honeycreepers ‘Akikiki (Gadow 1899); Laysan Finch (Clark 1912); Nihoa Finch, Hawai‘i ‘Amakihi, and ‘Apapane (van Tyne and Berger 1959; Berger 1982), a representative sample.All are rather similar in possessing a fully feathered head (capital tract) continuous with a longitudinal spinal tract.Two small lateral femoral tracts are found dorsally behind the legs. The legs themselves are feathered by the crural tract. The wings possess an inner humeral tract and a distal alar tract, plus two alular feathers, nine secondaries, nine functional primaries and a rudimentary tenth. The caudal tract has 12 rectrices (tail feathers), and 10 upper- and 10 undertail coverts. The ventral tract runs from the mandible to the base of the neck where it splits into branches that extend down the flanks leaving a large ventral apterion between. In these respects Hawaiian honeycreepers appear to be identical to cardueline finches (Newton 1973) and many other passerines. Areas between the feather tracts are not necessarily bald, and birds from colder places tend to have many small scattered contour feathers in them. In these same species the branches of the ventral tract extend further toward the tail (Berger 1982).

Moult In Hawaiian honeycreepers, Juvenal wing and tail feathers tend to be browner or paler than those of adults, allowing younger birds to be distinguished. The First Prebasic moult in Hawaiian honeycreepers is unusually long, extending on a population level from June to December (Baldwin 1953). Subsequent prebasic moults are about a month shorter, and individual birds are probably in moult for about 5 months. Honeycreepers and most cardueline finches have only a single annual prebasic moult.Thus honeycreepers do not have a ‘breeding plumage’ and a ‘non-breeding plumage’ as in the American Goldfinch, the only cardueline with a pre-alternate moult and a distinctive breeding plumage (Middleton 1977). Some Hawaiian honeycreepers in their First Basic plumage look much like adults (e.g. ‘Akeke‘e, Lepson and Pratt 1997) but Palila ( Jeffrey et al. 1993), Laysan Finch (Banks and Laybourne 1977), Hawai‘i ‘Amakihi (van Riper 1979),‘Akepa (Lepson and Freed 1997) and several

others exhibit delayed plumage maturation; all reach Definitive plumage by the third year. Some species have a distinctive Second Basic plumage, in which they are sometimes called subadults. For example, the second-year male ‘Akepa is duller orange than fully adult males (Lepson and Freed 1997). Closely related species may differ in the number of moults required to reach Definitive plumage. For example, Laysan Finches reach Definitive plumage in 3 years, whereas Nihoa Finches require only two (Banks and Laybourne 1977), and note the aforementioned difference between the two species of Loxops. Because feather replacement is a significant energy drain, birds usually cannot breed and moult at the same time. The aforementioned American Goldfinch, with two annual moults, is in moult throughout the year except for the months of breeding (Middleton 1978). In high latitudes, birds such as most cardueline finches have very sharply defined breeding and moulting seasons, but even within species those strictures become more lax toward the equator (Newton 1973). Ralph and Fancy (1994a) found that, like many tropical birds, Hawaiian honeycreepers have very extended breeding seasons, and consequently long periods of moult. Hawai‘i ‘Amakihi, for example, have been found in breeding condition in all months of the year. Breeding is not, however, evenly distributed within the season and all species studied have definite peaks. Although it varies among species (and from year to year), breeding by Hawaiian honeycreepers generally peaks between April and July (see Chapter 8). As a result, the percentage of distinctively plumaged juveniles in the population has a very noticeable peak at mid-year, with virtually none present by December. Obviously, the peak for moult of adults follows the peak for breeding. At a given time, a population may include birds in moult and in breeding condition. Individual honeycreepers, as in Palila (van Riper 1980a; Jeffrey et al. 1993), may do both at the same time, but most do not.

Plumage coloration As discussed briefly in Chapter 4, Hawaiian honeycreepers and cardueline finches have all the same

Form and function 105 pigments, and the drepanidine finches are remarkably similar in overall aspect to certain carduelines. (For colour illustrations of all cardueline species see Clement et al. (1993) and for the honeycreepers, of course, refer to the colour plates herein.) The genus Telespiza, in particular, could easily be merged with the cardueline Serinus without looking out of place. Other Hawaiian honeycreepers have diverged considerably in how the basic pigments are distributed in the plumage. Nearly all carduelines have longitudinal sparrow-like streaking, at least faintly, at some age or in at least one sex. Such streaking occurs only in immature and female Telespiza among drepanidines, and is one of the reasons for considering that genus the most primitive. Contrastingly coloured tips to the greater and median wing coverts are found in most carduelines. Such ‘wing-bars’ are also present in the Juvenal plumages of many Hawaiian honeycreepers, but they are less frequent among adults. Many carduelines have dark lores or facial feathers, as do many drepanidines. Many carduelines have brightly coloured body feathering with contrastingly dark wings, again a frequent pattern in Hawaiian honeycreepers. Nearly all cardueline finches are sexually dichromatic as adults, and most also have distinctive female-like immature plumages. That pattern is maintained in many Hawaiian honeycreepers, but in others adult males and females are alike in colour with distinctive immatures, and the Greater ‘Amakihi appears to have only one colour pattern for all sexes and ages. Species that lack sexual colour differences (dichromatism) or distinctive immature plumages can therefore be considered to have a derived condition, but one that has probably evolved several times independently in the family.

The ‘red and black’ group Even as early as Perkins (1903), observers recognised at least two groupings among the honeycreepers based on plumage. Perkins’s Division 1 corresponds to Amadon’s subfamily Drepaniinae. It comprises the so-called ‘red and black’ honeycreepers, but not all have red and within that designation few generalisations can be made. Richards and Bock’s (1973) comparison to rosy-finches notwithstanding, they are the least cardueline-like of all drepanidines.They

tend to have bold and sharply defined areas of bright colour, sexes alike as adults, and distinctive juveniles and immatures (except, as far as known, in the two mamos). The group is distinctive also in plumage texture. Their feathers are often described as ‘hard’ rather than ‘soft’. Indeed, with the exception of the Black Mamo, their feathers tend to have a glossy or shiny look rather than a matte surface. This change in the feather surface may be an adaptation to nectarivory. Each species (with the exception of the two Himatione) is distinctive in its own right and the group is believed to represent an old evolutionary line that has lost most of the intermediates. The ‘red and black’ group also includes some structural feather peculiarities that warrant closer consideration. Certain feathers may be stiffened in peculiar ways, others may be modified in shape. Throat feathers of the ‘I‘iwi, ‘Ula‘ai-hawane, and ‘Akohekohe are stiffened along the midrib and have sharp-pointed tips. The ‘Akohekohe has such feathers on other parts of the body as well, and its stiff, forward-curling crest is unique. One of the most intriguing feather adaptations in this group is an oblique truncation of the tips of the outer primaries that produces an audible note when the birds fly. Such unusual primaries are found in ‘I‘iwi and ‘Apapane, and, according to Perkins (1903), the mamos. Interestingly, the sound produced by these feathers in flight is also present in the ‘Akohekohe, whose primaries are not obviously modified. Drepanidine wing-flutter as a sound is discussed more fully in Chapter 7.

The ‘green’ group The rest of the honeycreepers, except for the Po‘ouli (Pratt 1992a), have long been known as the ‘green’ birds although, like ‘red and black’, that overstates the case. Not all are truly green, but it is a common enough colour in the group that it is a reasonable designation. Note that even the brightest green of Hawaiian honeycreepers is not like that of parrots, tanagers, parrotfinches (Erythrura) and other well-known green birds, but a dull olive green similar to that seen in such carduelines as greenfinches, citrils, and canaries. Perkins placed them in his Division 2, and Amadon named them the Psittirostrinae,

106 The Hawaiian Honeycreepers both of which included the drepanidine finches. More recent classifications (Pratt 1979a;AOU 1983) have separated the finches from the thin-billed ‘green’ birds if they recognised subtaxon groupings at all. Although Amadon (1950) denigrated coloration as an indicator of relationships, others (Pratt 1979a, 1992a; Conant et al. 1998) consider it one of the more useful characters, especially at the generic level. Within the great variational range of the family, one plumage pattern appears in more species than any other. I call this suite of feather characters ‘ ‘amakihi plumage’ (Pratt 2001b) from the most common species group to possess it. For a more detailed description, see Genus Hemignathus in Part II. All ‘amakihi plumages are dark above, pale below, with no sharp demarcation between. The overall colour is the typical drepanidine dull green, but the paler parts are more or less yellow and the lores are dark. Birds are sexually dichromatic, with males brighter than females. First-year birds resemble females but are even duller and have pale wingbars. Within these parameters there is some slight interspecific variation. Some species have more pronounced dorso-ventral contrast, but others are nearly uniformly coloured. Some have more pale coloration above the eye. Some have dark crowns with dark spots (Kaua‘i ‘Akialoa), and others have the top of the head nearly as pale as the underparts (nukupu‘us). Despite these variations on the theme, all species possessing this plumage pattern are very similar in all respects except bill length (Conant et al. 1998), and form a clearly definable cluster within the family. They are, in fact, the only such large species group that can be defined by coloration. The most aberrant possessor of ‘amakihi coloration is the Greater ‘Amakihi, which has almost no dorso-ventral contrast, sexual dichromatism, or maturational colour differences (but it may not actually belong to this group after all; see Chapter 5). No close approximation of this colour pattern occurs among cardueline finches, so it can reasonably be regarded as derived among the Hawaiian honeycreepers. It represents a complex suite of synapomorphies apparently evolving in tandem which, coupled with parallel synapomorphic clusters in bill shape and vocalisations (Chapter 7), defines ‘Greater Hemignathus’ (see Chapter 5).

Not all of the ‘little green honeycreepers’, however, exhibit ‘amakihi plumage. Although some of them, such as the Hawai‘i Creeper (Scott et al. 1979) and Maui and O‘ahu ‘Alauahios (Shallenberger and Pratt 1978), are similar enough in overall coloration to cause difficulty for field observers (Pratt et al. 1987), the details do not agree.The Hawai‘i Creeper is a slightly different, greyer shade of green, with virtually no yellow below even in adult males, no pale superciliary except in juvenile plumage, and a broad, triangular grey face mask instead of narrow black lores. Also, adult sexes are nearly alike but juveniles have a distinctive paler plumage without the mask. The ‘alauahios almost fit the pattern, but because coloration is the only feature in which they resemble Hemignathus and they are obviously closely related to scarlet Kakawahie, the resemblance can be regarded as coincidental or convergent (see Batesian mimicry below). Similarly, the ‘Akeke‘e, which has a general similarity to ‘amakihi plumage, differs in having a yellow crown and rump, broad black face mask, notched tail, non-distinctive immatures that lack wing-bars, and, like the Maui and O‘ahu ‘alauahios, is congeneric with a red bird. The ‘Anianiau has traditionally been considered closely related to the ‘amakihis, but close scrutiny of its plumage shows that it does not share the ‘amakihi plumage synapomorphies. Adult males show much less dorso-ventral contrast, both sexes lack the dark lores, no plumage ever exhibits wing-bars, and the bill colour, with pinkish tones, does not match. The closest approach to ‘amakihi plumage in a honeycreeper not usually grouped with Hemignathus is found in the Maui Parrotbill. In fact, Pseudonestor has ‘amakihi plumage except for a prominent, sharply demarcated superciliary and bold dark eye stripe that incorporates the black lores, and its bill coloration, like its thickness, is very different from that of other members of the group. On the other hand, Pseudonestor’s plumage is much more similar to that of Hemignathus than to any of the drepanidine finches with which it has usually been grouped. Thus, despite its deviations, plumage could be viewed as reinforcing the hypothesis (see Chapter 5) that the parrotbill is actually related to thin-billed insectivores rather than to drepanidine finches. The remaining species in the ‘green’ group is the ‘Akepa, and only the female is green (Scott et al.

Form and function 107 1979). The previously mentioned ‘Akeke‘e and the ‘Akepa differ from each other as much as any two honeycreeper species, yet they were long regarded as conspecific. Their distinctiveness is now well established (Pratt 1989b), and some have suggested that the nominal subspecies of the ‘Akepa might all be species (see Chapter 5; Pratt and Pratt 1999). The Maui form Loxops coccineus ochracea is of special interest in the discussion of plumage colour because it is the only honeycreeper that is colour polymorphic. Adult males can be either mustard yellow or orange in about equal numbers, with a small number of intermediates. Because the Maui ‘Akepa is nearly extinct and has been rare throughout historic times, the exact nature of this polymorphism may never be known, but an intriguing parallel may be a similar polymorphism in the cardueline Red Crossbill. Newton (1973: 203) describes it: The variation in colour of adult cock crossbills has attracted much comment. Most are predominantly reddish, but some are orange, bronze or yellowish-green, while others contain a mixture of feathers of different colours . . . but particular individuals may change from red to yellow or vice versa, according partly to when they moult. Most cocks which moult before the start of July grow yellowish feathers; most which moult after this date grow red ones; while many of those whose moults overlap this period end up part yellow and part red, sometimes with both colours on individual feathers. A similar process could explain the polymorphism in the Maui ‘Akepa, but the suggestion is purely speculative at present.

The creepers The two species of Oreomystis have long been considered to belong to the ‘green’ group of honeycreepers, but, as shown earlier they have a derived tongue type that places them in a group all their own. Now that they are no longer regarded as congeneric with the ‘alauahios (Pratt 1992b), their distinctive plumage characters have come into better focus. As discussed above, the Hawai‘i Creeper bears

only superficial resemblance to Hemignathus, but it does have some apparently synapomorphic plumage similarities to the ‘Akikiki. Both Oreomystis lack plumage dichromatism in adults, and both have distinctive juveniles. Both juvenile plumages have pale faces, unusual among Hawaiian honeycreepers, that extend above the eye. In O. bairdi the pale juvenile feathering takes the form of an almost Zosterops-like circle of feathers, but in O. mana it is more of a diffuse superciliary, so they are not identical. However, no other juvenile honeycreepers show a pattern anything like them. Finally, both creepers have bills that are pale essentially throughout, with only a trace of dusky pigmentation along the culmen.As with juvenile facial colours, the bills are not identically coloured (one pink, the other pale grey) but the only other thin-billed honeycreepers with pale bills are the ‘akepas.

The Hawaiian finches Coloration among the drepanidine finches is actually relatively uniform. All but one species whose plumages are known are sexually dichromatic, and younger birds look like adult females. Males tend to have brightly coloured, more or less contrasting heads, the colour of which may or may not extend onto the breast. Head colour is yellow in five species, golden-orange in one.The Kona Grosbeak stands apart from the others in exhibiting virtually no plumage variation; all ages and sexes look alike. Also, it is the dullest member of the group. Interestingly, its dull olive green plumage with black lores has a vague resemblance to ‘amakihi plumage, showing that the thin-billed birds’ plumage colours could easily have evolved from a finchlike ancestor. The Lana‘i Hookbill, possibly related to the ‘O‘u, is known from a single female specimen ( James et al. 1989), the coloration of which does not look out of place in the Hawaiian finch context. Based on the above generalisations, I would guess that the adult male had an all-yellow head and breast. Thus it would have looked like an ‘O‘u unless viewed from the side or front, and would not therefore have been easily distinguished in the forest. This may help explain why observers before Munro failed to notice the species on Lana‘i. Snetsinger et al. (1998) suggest that the adult male hookbill may have had

108 The Hawaiian Honeycreepers a brighter, whitish superciliary, but give no basis for such speculation. One could further speculate that the extinct members of the genus Telespiza might have been red or orange on the head and breast rather than the yellow seen in the two surviving species, both of which live in open, brightly lighted habitats. In fact, one might even suspect that when the living species inhabited the main islands ( James and Olson 1991), they might have had darker or more saturated pigmentation.

The Po‘o-uli One of the reasons for questioning the inclusion of Melamprosops phaeosoma in the Hawaiian honeycreepers (Pratt 1992a) was that it did not fit into any of the known plumage groupings. Indeed, its chocolate brown body coloration is unique in the family and has no counterpart among cardueline finches (although a few have other shades of brown). The black mask looks vaguely like the masks of a few honeycreepers, but is far more extensive than any, especially in the adult male. Adults are known only from field observations and photographs (Pratt et al. 1997; Baker 1998; see also species account) that reveal some sexual differences. Both of the two specimens in existence (Casey and Jacobi 1974) have turned out to be immatures, recognisable by a distinctive browner and buffier plumage. Certainly plumage colour provides no evidence as to the relationships of this enigmatic bird.

Aberrant coloration Many passerines display occasional ‘sports’ and oddities in their plumage colour. The only example of such freak coloration so far reported among Hawaiian honeycreepers was a local population of ‘Apapane in Hawaii Volcanoes National Park that produced several partly white individuals. The first of these ‘partial albinos’ had the ‘wings, lower chest, and areas of the back white. Other feathered regions appeared orange; tarsus and feet were salmoncoloured, while the beak was black’ (van Riper and van Riper 1978). The van Ripers reported similar birds, up to four at a time in one flock, between 1973 and 1978 in the same general area.

Factors influencing coloration Feather wear Colour patterns can be affected by feather wear as well as by moult. Dark coloured parts of feathers are more resistant to wear than pale areas, so pale edges and tips tend to wear away during the course of a year. Thus immatures with prominent wingbars late in the year may lack them by the following summer. As feathers wear, their colour may appear to change, usually becoming lighter. At certain seasons, for example, ‘Apapane may appear to have bright red heads contrasting with deep crimson bodies. This pattern results because the capital (head) feather tract is the last to complete moult (Baldwin 1953) and the worn feathers are brighter red than the fresh ones, some of which have grey tips that wear away quickly.The First Basic plumage of the ‘I‘iwi starts out greenish yellow, but wears to a more golden tone.

Natural selection One of the great mysteries in studies of Hawaiian honeycreepers was the adaptive significance of their colours. For no apparent reason, the honeycreepers, even the bright red and yellow ones, seemed to be cryptically coloured: red birds feed in red flowers, yellow birds in yellow, green birds feed among foliage, greyish birds climb tree trunks, dark birds live in the understorey, and pale birds live in open dry habitats. With only a single potential predator present, and it confined to one island only, naturalists from Perkins and Henshaw to Amadon and Baldwin were puzzled by honeycreeper coloration and grasped at such straws as ‘random consequences of genetic drift’ (Baldwin 1953: 383), which satisfied no one. Of course, we now know from prehistoric remains (Olson and James 1982b, 1991) that Hawai‘i had several avian predators that fed on small birds, so no further explanation is needed for the concealing coloration of Hawaiian honeycreepers (Ziegler 2002). Hawaiian honeycreepers’ cryptic colours also follow the predicted pattern for prey species (Stamps and Gon 1983) in that females are more cryptic than males, even in species lacking

Form and function 109 strong dichromatism. For example, the darker nape of female Palila helps conceal an incubating bird ( Jeffrey et al. 1993).

Batesian mimicry Named for Henry W. Bates, a nineteenth-century naturalist who discovered it in Amazonian butterflies, Batesian mimicry is an evolutionary development ‘in which the morphology and coloration of a certain species imitate those of another species that is less vulnerable to attack because of being dangerous, unpalatable, difficult to capture, and so on’ (Morris 1992: 229). Probably the most familiar example is the resemblance of the quite edible viceroy butterfly to the monarch, avoided by birds because it (apparently) tastes terrible. Few examples are known among birds (Barnard 1979), but the recent discovery of a possible example among Birds of Paradise (Frith and Beehler 1998) involving mimicry of certain species of Pitohui, whose skin and feathers have been found to be toxic to predators (Dumbacher et al. 1992), prompted me to consider whether drepanidine odour may have produced an example among Hawaiian birds. Because the predators that may have been the selective force are all long extinct, the case will be difficult to prove, and it seems a bit far-fetched, but I offer it at least as food for thought.We begin with the assumption that if an unpleasant odour makes some birds unpalatable to predators, a possibility suggested by Weldon and Rappole (1997), then those that lack the odour, such as the three species of Paroreomyza, could gain an advantage by looking like those that do, and may come to resemble them in this special case of convergence. Birders have long appreciated the close resemblance between the two ‘alauahios and ‘amakihis (Shallenberger and

Pratt 1978; Pratt et al. 1987), and the similarities misled Amadon (1950) to consider the ‘alauahios congeneric with them. One of the few things that recent systematists from a wide variety of disciplines (e.g. Johnson et al. 1989; Tarr and Fleischer 1995; Feldman 1997; James 1998; Pratt 2001b) agree on is that, regardless of the peripheral details, Paroreomyza is one of the most distinctive drepanidine genera and it diverged very early in honeycreeper evolution. Thus the plumage similarities may be the result of coincidence or convergence, but probably not phylogeny. But with so many ‘little green birds’ in the drepanidine radiation, how can we say that these are mimics? The key is in the details. Males of the two ‘alauahios simply resemble ‘amakihis in a general way, but the females parallel distinctive plumage features of the particular ‘amakihi female with which each is sympatric.The most striking convergence (or coincidence) involves females of the Oahu ‘Alauahio and O‘ahu ‘Amakihi, both of which have much-reduced yellow coloration so as to be more grey than green, and both of which have prominent pale wing-bars as adults, a feature not known in any other ‘green’ honeycreepers. Females of the Maui representatives present a less compelling case simply because they are so plain. But what about the striking flame-red male Kakawahie? As discussed above in the context of character displacement, P. flammea is likely the second invader of Maui-nui, and as such it may have had to develop a distinctive coloration to avoid interspecific hybridisation. But why red, when so many other Maui-nui birds were already that colour? Perhaps the reason is that the other red birds all possessed drepanidine odour. Of course, as Amadon (1950) appreciated, yellow/red colour shifts are easily accomplished in birds and Batesian mimicry may not be involved at all.

7 Behaviour

Locomotion Flight All Hawaiian honeycreepers are fully flighted, although the flight of the Laysan Honeycreeper was ‘weak and low’ (Munro 1960) and the Laysan Finch apparently prefers to walk (Fisher 1903). ‘O‘u (Rothschild 1893–1900; Perkins 1903; Conant et al. 1998), ‘Apapane (Fancy and Ralph 1997), ‘I‘iwi (Fancy and Ralph 1998), and occasionally even ‘Akiapola‘au (T. K. Pratt et al. 2001) are noteworthy for their long flights high over the forest, but most drepanidines only make relatively short flights between perches or feeding sites. Flight is direct in most species but ‘Apapane (Fancy and Ralph 1997) and ‘I‘iwi (Fancy and Ralph 1998) have the undulating, flap-and-glide pattern seen in many cardueline finches.

Walking, running, and hopping Most Hawaiian honeycreepers on the main islands rarely go to the ground except to gather nesting material, so running and hopping are not common means of locomotion. However, ‘I‘iwi (Fancy and Ralph 1998), ‘Apapane (Fancy and Ralph 1997), and ‘Akohekohe (Berlin and VanGelder 1999) often run over the nearly solid leafy surface of the ‘ohi‘a canopy. Both drepanidines on Laysan walk and hop along the ground more than they fly (Fisher 1903).

Creeping This term is rather imprecise, and has been applied to several bark-picking honeycreepers with different

locomotory movements. ‘Akikiki and Hawai‘i Creepers (Foster et al. 2000; pers. obs.) as well as Maui Parrotbills (Simon et al. 1997), creep over trunks and branches of trees in a manner reminiscent of nuthatches and holarctic creepers. None brace with the tail, however. They typically crouch low, with the legs hidden and the belly nearly pressed against the substrate. All three may proceed spirally or linearly, sometimes head downward. The parrotbill also uses its bill, parrot-like, to move from branch to branch (Simon et al. 1997). Heterobills and ‘akialoas (Rothschild 1893–1900; Perkins 1903; Lepson and Johnston 2000) and the Po‘o-uli (T. K. Pratt et al. 1997b) all ‘creep’ but include much more hopping in their progression.The creeping of the Maui ‘Alauahio also involves frequent hops along small branches, with stops to peer above and below branches, more like the movements of the Black-and-White Warbler than the more nuthatchlike motions of Oreomystis and Pseudonestor (H. and P. Baker 2000; pers. obs.).

Self-maintenance Bathing Hawaiian honeycreepers bathe primarily by rubbing against wet vegetation (Simon et al. 1997; Lindsey et al. 1998), or by simply sitting exposed in the rain (Eddinger 1970; Lepson 1997). A typical example is the ‘Anianiau, which Eddinger (1970: 137) described as hopping ‘from branch to branch shaking his body, wings, and tail in the wet leaves.’ Eddinger found similar bathing behaviour in ‘I‘iwi, ‘Apapane, and Kaua‘i ‘Amakihi, and others have

Behaviour 111 reported it in Hawai‘i ‘Amakihi (Lindsey et al. 1998), Maui Parrotbill (Simon et al. 1997), Maui ‘Alauahio (H. and P. Baker 2000), and Laysan Finch (Throp 1970). J. K. Lepson (in Lindsey et al. 1998) reported a Kaua‘i ‘Amakihi bathing at a vertical wall with dripping water, and Morin and Conant (2002) report bathing under freshwater seeps by Nihoa Finches. Bathing in pools is apparently quite rare. Eddinger (1970) never observed such bathing in any of his study species in the field, but found that captive ‘Anianiau bathed in pans of water. However, E. VanderWerf (in Lindsey et al. 1998) reported pool bathing by an O‘ahu ‘Amakihi and Nihoa Finches bathe in puddles in the wild (Morin and Conant 2002) and in pans in captivity (Berger 1981). Laysan Finches sometimes drown in deep containers that fill with water (Morin 1987), so pool-bathing is clearly not a customary behaviour for this species (Laysan normally has no standing fresh water).

Sunbathing Hawaiian honeycreepers also engage in sunbathing, and all that have been reported so far have a posture similar to that described by Eddinger (1970: 136) for an ‘Apapane in which the bird perches in a tree and ‘ruffs out its feathers, opens the bill, spreads the tail feathers, points the head and open bill upward, the eyes being open, stretches one wing and then the other. The bird may either face the sun or place one side toward the sun.’ Eddinger studied ‘I‘iwi, ‘Apapane, ‘Anianiau, and Kaua‘i ‘Amakihi, and his description would also fit Maui ‘Alauahio (H. and P. Baker 2000), Maui Parrotbill (Simon et al. 1997), Hawai‘i ‘Amakihi (Lindsey et al. 1998), and presumably other species.

Head-scratching All Hawaiian honeycreepers yet reported scratch their heads indirectly, i.e. by bringing the foot up and over the wing. Eddinger (1970) reported such head-scratching by ‘Apapane, ‘I‘iwi, ‘Anianiau, and Kaua‘i ‘Amakihi, and others have noted it in Maui ‘Alauahio (H. and P. Baker 2000), ‘Akepa (Lepson and Freed 1997), ‘Akiapola‘au (T. K. Pratt et al.

2001), Po‘o-uli (T. K. Pratt et al. 1997b), and O‘ahu and Hawai‘i ‘Amakihi (Lindsey et al. 1998).

Roosting Warner (1968) reported that, unlike most other passerines, Hawaiian honeycreepers sleep without tucking their bills into their back feathers and without crouching over their feet to cover them with feathers. This posture, according to Warner, makes them especially vulnerable to attack by mosquitoes (see Chapter 9) because it fails to protect the ‘soft parts’ where these insects mostly bite. The species he observed were Kaua‘i ‘Amakihi, ‘Anianiau, ‘Apapane, ‘I‘iwi, and Nihoa Finch, a representative sample with all branches of the family included. This sleeping posture is not immutable, however, and honeycreepers exposed to mosquito bites for a period of time apparently evolve more defensive rooting postures. Eighteen years after Warner’s study, van Riper et al. (1986) found that Kaua‘i ‘Amakihi and ‘Apapane slept with heads tucked and one foot raised into the feathers. Lindsey et al. (1998) report the same for Hawai‘i ‘Amakihi. Whether these observed behaviours are recent adaptations is problematical, but van Riper (1991) points out that ‘selection should favour modification of avian sleeping behaviour if the benefits outweigh the negative aspects, which in this case it certainly does.’ On the other hand, honeycreepers may simply have learned how to eliminate discomfort, particularly if sleeping posture is not genetically determined. Maui ‘Alauahio (H. and P. Baker 2000), ‘Akepa (Lepson and Freed 1997), and Po‘ouli (T. K. Pratt et al. 1997b) have been observed asleep, but their postures were not reported. Most honeycreepers roost in trees, but Nihoa Finches sleep in rock crevices (Morin and Conant 2002). MacMillen and Carpenter (1980) suggested that upslope evening flights of ‘Apapane and ‘I‘iwi (see page 148) indicated the possibility of communal roosting, as is characteristic of many cardueline finches (Newton 1973). Ralph and Fancy (1995) reported that birds moving downslope at dusk from high elevation mamane–naio forest on Mauna Kea scattered once reaching their destination (midelevation ‘ohi‘a forest).They also worked in the area

112 The Hawaiian Honeycreepers around Pu‘u Kulani, where MacMillen and Carpenter’s (1980) evening flights appeared to converge, and found no communal roosts. However, use of the Kulani forests as a ‘common roosting area’ (MacMillen and Carpenter 1980) too diffuse to qualify as a communal roost is not precluded.To date no true communal roosts of Hawaiian honeycreepers have been discovered.

Vocalisations Aside from external appearance, the most noticeable features of Hawaiian honeycreepers are the sounds they make. In fact, sounds are often a more reliable indicator of the species present and their numbers than visual cues (Scott et al. 1986). However, serious study of honeycreeper sounds did not begin until the final quarter of the twentieth century. Early naturalists certainly noticed and described honeycreeper vocalisations, but they were hampered by the lack of a technology for recording and visually displaying sounds. Word descriptions of bird sounds are useful, but cannot be used for critical comparisons, and are always subject to differing interpretations. Also, observers differ greatly in their ability to hear and transcribe bird voices. The mid-century development of small portable tape recorders and the development of the sound spectrograph solved both of these problems. Sound spectrograms (popularly known as sonagrams or sonograms) in particular have enabled much finer comparisons of sounds from different localities. Subtle differences that might not have been ‘remembered’ from island to island by early researchers can be obvious in these sound pictures. Also, vocalisations that sound like single notes to human ears are often revealed in sonograms to be much more complex. Hawai‘i got a late start in the modern study of bird sounds, and the pioneer stage of simply gathering and archiving representative sound specimens is still ongoing (Pratt 1996b). The biological study of honeycreeper vocalisations provides one of the richest sources of future research projects. Hawaiian honeycreepers have as rich a sound repertoire as any passerine group, and a representative sample of recordings has now been published

(Pratt 1996a). Even though most vocalisations are species-specific, some notes are widely shared among different species. Individual and geographic intraspecific variation is noticeable among honeycreepers, and some species also have age- and sexspecific vocalisations. Despite these variations, certain underlying patterns are discernible that provide useful phylogenetic inferences at all taxonomic levels. The groupings used here are arbitrary, but they correspond to customary vocal categories among other passerines. Detailed descriptions of each species’ vocalisations are presented with the species accounts (Part II).

Calls Calls, as distinguished from songs, are short vocalisations uttered throughout the year. They usually have little, if anything, to do with mating and territoriality. My experience is that Hawaiian honeycreepers have more varied call repertoires than most passerines.Very few species have a single ‘call note’ and in some cases (e.g. Hawai‘i ‘Amakihi) the calls seem almost infinitely varied. Sometimes the variation is age- or sex-related, but often Hawaiian honeycreepers just have many different calls that seem to be given at random. Perhaps that impression is only an artefact of our lack of knowledge.

Contact notes The most common calls can also be referred to as contact notes.They probably function to say simply ‘I am here’ to other members of the same species, or even other species with which the bird may associate. Some contact notes are quiet, apparently not intended for distant listeners. Others are loud and far-carrying. In the ‘red and black’ group, contact notes are not as strongly differentiated from other categories of vocalisations. In the ‘Apapane, call notes may often be incorporated into longer songs. In the ‘I‘iwi and ‘Akohekohe, the distinction between songs and calls is entirely blurred (pers. obs.; Berlin and VanGelder 1999). The Maui ‘Alauahio also incorporates its typical call note into the territorial song (pers. obs.), and the song of the Po‘o-uli is made up entirely of notes resembling the call (Engilis 1990).

Behaviour 113 While foraging, virtually all Hawaiian honeycreepers utter contact notes, the function of which is not always apparent. Some are obviously communication between members of a mated pair, especially when males have a distinctive call as in the case of the ‘Akiapola‘au and the Maui Parrotbill. My observations of Hawai‘i ‘Amakihi seem to indicate that the birds increase their call repertoire in response to population density. In most areas of Hawai‘i and Maui, the only ‘amakihi call note usually heard is a highly distinctive short buzzy note, but in the mamane–naio forest of Mauna Kea, where Hawai‘i ‘Amakihi are very abundant (Scott et al. 1987), they seem to have a wider variety of notes, including some that resemble calls of other Hawaiian honeycreepers. No detailed studies have been conducted to determine whether the denser population really has a more varied call repertoire. Perhaps their repertoire is simply more apparent in a locality where this one species vastly outnumbers all others. Some honeycreepers seem to use calls almost like songs. For example, male ‘O‘u utter a loud up-slurred whistle at regularly-paced intervals while perched high in emergent trees (Huber 1966; Conant et al. 1998).They may do so continuously for long periods without moving from the perch. Because the notes are very far-carrying, they may serve an advertising or even territorial function, although they are heard outside the nesting season. Palila also have a loud whistled call that may be given by a sedentary bird, but this note is uttered far less persistently than that of the ‘O‘u and does not seem to have any potential for territorial function. Berlin and VanGelder (1999) consider some ‘calls’ of ‘Akohekohe related to territoriality, but they may simply have not recognised a short song for what it was.

Social/alarm call As discussed in Chapter 4, calls of Hawaiian honeycreepers have a general resemblance to those of some cardueline finches, especially an up-slurred whistle that Mundinger (1979: 280) characterised as: associated with fear/attraction conflict, both in the social and in the alarm contexts. Socially, the call is typically given in courtship by a lone

individual (low fear) in a potential ‘mateattraction’ situation. Later, paired birds give it when separated from their mates. In the alarm context the call is given during low-intensity mobbing, e.g. when man approaches the general vicinity of the nest. Up-slurred whistles among Hawaiian honeycreepers have not been analysed to the same extent as those of carduelines, but their behavioural contexts appear to be similar.As in the Carduelinae, interspecific and intraspecific variation is obvious, but ‘crossspecies correspondence is clear’ (Mundinger 1979). Species that utter cardueline-like up-slurred whistles include ‘O‘u , ‘Akiapola‘au, Maui Parrotbill, Hawai‘i ‘Amakihi,‘Akepa, and ‘I‘iwi (pers. obs.), and possibly Po‘o-uli (T. K. Pratt et al. 1997).

Juvenile calls All drepanidine juveniles have begging notes given during feedings, and some of the insectivorous species have distinctive juvenile calls that continue past the fledging stage. Most of these are not strikingly different from adult calls, but several species have very noticeably distinct juvenile calls. These calls may be associated with unusually long periods of dependency, as is seen in the ‘Akiapola‘au and the Maui Parrotbill. In those species, juveniles may follow parents for up to a year and continue to be fed by them at least occasionally. The juvenile calls of both species are loud, regularly spaced chirps, quite different from any adult calls, and so continuous as to provide ‘a nearly constant vocal beacon’ (Simon et al. 1997). Both members of the genus Oreomystis also have distinctive juvenile calls (for sonograms, see Pratt 2001b), but they do not persist for as long a period after fledging and seem to be given by young birds following their parents closely. In both cases, these calls are rapid series of dry chips that have a syncopated rhythm with clusters of quicker notes (Scott et al. 1979; Pratt 2001b). Distinctive juvenile calls have been reported in Hawai‘i ‘Amakihi (Lindsey et al. 1998), ‘Akepa (Lepson and Freed 1997), and ‘Anianiau (Eddinger 1970), but not among any of the ‘red and black’ honeycreepers or drepanidine finches.

114 The Hawaiian Honeycreepers

Flock cohesion notes One of the most intriguing questions as yet unanswered about Hawaiian honeycreepers involves whether mixed-species foraging flocks (see Chapter 9) develop island-specific call notes shared by most or all member species. The most obvious possible example, and the one first noticed by observers, is on the island of Maui.There, the typical call (a sharp chick or cherk) of the Maui ‘Alauahio appears to have been adopted by several other species as a common note in mixed flocks (Engilis 1990).The note (or a very similar one) is given by Maui Parrotbill (Mountainspring 1987), Po‘o-uli (Engilis 1990), Kakawahie (Bryan 1908), and occasionally Hawai‘i ‘Amakihi (pers. obs.), all of which join (or may have joined in the case of the Kakawahie) flocks of ‘alauahio. A similar note was reported for the O‘ahu ‘Alauahio, but is not uttered by any other surviving birds there. On Kaua‘i, the ‘common note’ appears to be the typical call of the Kaua‘i ‘Amakihi. This may partly explain why the possible existence of such a note on that island was long overlooked. Until very recently, the call of the Kaua‘i ‘Amakihi was assumed to be a buzzy note similar to those of ‘amakihis on other islands. The bird does, indeed, utter such a note, but careful observation (Conant et al. 1998) has revealed that it is much less frequent than a piercing chirp that was long thought to be given only by other species, including ‘Akikiki,‘Anianiau, and ‘Akeke‘e. Could this be the ‘Kaua‘i flocking note?’ No such common flocking note has been reported on Hawai‘i, but a good candidate would be the Hawai‘i ‘Amakihi’s juvenile call, which is ‘a prominent part of the vocal environment during the flocking season’ ( J. K. Lepson, in Lindsey et al. 1998). Investigation of this question is made more difficult by the disappearance of so many species that might have participated in mixed flocks in earlier times. The designation of the Kaua‘i and Maui flock notes as those of ‘amakihi and ‘alauahio, respectively, is therefore rather arbitrary. On Kaua‘i, the ‘Akikiki is just as good a candidate as the original author of the note (because it has few others), as is the Po‘o-uli on Maui for a similar reason. Today’s relative population numbers, and therefore

percentage participation in mixed flocks, are clearly artificial.

Advertising songs Songs are loud vocalisations used to announce the singer’s presence to potential rivals for territory, food resources, or potential mates. They are much more seasonal than calls, but because Hawaiian honeycreepers have such long breeding seasons, songs can be heard through most of the year. The quietest months for singing are July–September, when most species have completed the breeding cycle for the year and not yet begun the pairing process for the next. Hawaiian honeycreepers also differ from mainland passerines in the distribution of song frequency during the day. Most observers are surprised to find that Hawaiian birds have no pronounced dawn chorus. Birds do sing vigorously at dawn, but no more so than later in the day and there is no obvious cessation of singing at any point.To the delight of birders, one need not rise at dawn to hear Hawaiian honeycreepers sing. Songs of most Hawaiian honeycreepers fall into one of three groupings that, perhaps not surprisingly, correspond to the traditional subdivisions of the family (Pratt 1992a). Indeed, one of those subdivisions, the ‘red and black’ group, was based in part on its distinctive vocal characteristics (Perkins 1903). Again not surprisingly, the two genera of questionable taxonomic position (Paroreomyza and Melamprosops) do not fit into any of the three large vocal categories (Pratt 1992a,b). Their vocalisations are described in the species accounts (Part II).

Canary-like songs The songs of Hawaiian finches are similar in a general way to those of many cardueline finches. Comparisons such as ‘linnetlike’ (Fisher 1906) and ‘canarylike’ (Berger 1981) are common in the literature (linnets and canaries are both carduelines). Such songs are usually loud, melodious, complex (containing whistles, warbles, trills, and chirps), unstereotyped, and lengthy. Drepanidine finches can be quite long-winded, with songs sometimes lasting several minutes (Fig. 7.1).

Behaviour 115 8 4

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7.1 Sonogram of song of Palila (LNS 06067), exemplary of complex canary-like songs of some Hawaiian finches.

Each species has its own variations on this theme, usually involving the degree of amplitude (e.g. the song of the Palila is much quieter than that of the ‘O‘u ) and the character of the individual elements included (some have more whistles, some

more trills, etc.). Unfortunately, many of these fine singers are either extinct, endangered, or confined to remote islands so that modern birders will never be able to share the experience of Henshaw (1901: 123):

116 The Hawaiian Honeycreepers the Ou possesses an even stronger claim to affection, for it is the most beautiful songster of the Hawaiian forest. The song is unmistakably fringilline in character, and so much resembles the Canary’s, that it is the generally received opinion among the settlers that the forest is full of escaped cage-birds; yet in purity, sweetness and power the song of the Ou far surpasses the Canary’s best efforts. Unfortunately the Ou, as a rule, is not very generous with its song, and too often the listener has to be content with snatches of melody in place of the finished performance. Yet I remember on one occasion to have heard more than a dozen males singing in a small patch of woods for at least an hour, and the chorus was worth going far to listen to. Perkins (1903), however, said that the song of the ‘O‘u ‘at times reminds one somewhat of a canary, but is much inferior to a first-class songster of that species.’ De gustibus non est disputandem; some like rock, some like country. We are fortunate that recordings of this song were obtained on Kaua‘i in the late 1960s (Gauthey et al. 1968; Pratt 1996b) so that we can make our own comparisons (Pratt 1996a).The finches of Laysan and Nihoa also have canary-like songs (Pratt 1996a; Morin and Conant 2002), the latter a bit sweeter than the former in my opinion, that resemble that of the Palila more closely than that of the ‘O‘u . The Kona Grosbeak was not highly vocal, but Munro (1960: 131) reported a ‘light sweet song . . . long with a variety of notes’ that may have been similar. The song of the Lesser Koa-Finch is unknown, but the Greater Koa-Finch had a distinctive song of humanlike whistles, some of which were ‘much prolonged’ (Perkins 1903) and ‘flutelike’ (Munro 1960). This song may have resembled the long whistles heard at the beginning of the ‘O‘u ’s song. Because it ‘frequently differs somewhat as whistled by different individuals, and also is sometimes distinctly varied, when repeated by the same bird’ (Perkins 1903: 438), it has at least some characteristics of the canary-like songs, but we will never know how close any resemblance may have been.

The ‘honeyeater mimics’ The ‘red and black’ honeycreepers have songs that resemble, sometimes strikingly, those of the honeyeaters (Meliphagidae) of the Indo-Australian region. Hawai‘i had its own meliphagids, the now-fabled ‘o‘os and the little-known Kioea. All were reportedly highly vocal, but the song of only one species, the Kaua‘i ‘O‘o , was ever preserved on tape (Pratt 1996b). That song does not closely resemble any Hawaiian honeycreeper song (pers. obs.). So why would I characterise the ‘red and black’ songs as resembling those of honeyeaters? When I was still in the early days of my research on honeycreeper songs, I heard a recording (Gunn and Gulledge 1977) of the song of the Tui, a honeyeater endemic to New Zealand.That song could be characterised as a jumble of highly varied elements, including bell-like, echoing, metallic, reedy, and flutelike notes. Portions of it are virtually identical to certain vocalisations of the ‘I‘iwi and ‘Apapane, and others closely resemble the song of the Kaua‘i ‘O‘o . In fact, one could approximate the song of the Tui by splicing together tapes of the songs of these three birds (Conant et al. 1998)! Of course, honeyeaters and Hawaiian honeycreepers are related only to the extent that both are passerines, so this vocal similarity must be the result of something other than shared ancestry. We can never know for sure, but the meliphagid resemblance may have developed when drepanidine nectar-feeders had to compete with meliphagids. Virtually all observers reported that the honeyeaters dominated the honeycreepers in feeding hierarchies (Chapter 8), so the smaller birds may have gained a feeding advantage by mimicking the sounds of the meliphagids.Thus, in the songs of ‘I‘iwi and ‘Apapane, we may be hearing ‘natural recordings’ of long-extinct Hawaiian honeyeaters (Pratt 1979a). If true, this situation would be a nearly exact parallel of the visual mimicry of large honeyeaters called friarbirds (Philemon spp.) by orioles (Oriolus spp.) in the Indo-Pacific (Diamond 1982), in which subordinate species gain acceptance into feeding flocks and immunity from attack by looking like their superiors. As with the honeycreeper vocalisations, the plumages of orioles that mimic honeyeaters differ strikingly from the usual coloration of that group.

Behaviour 117 Generalisations are difficult to make among the ‘red and black’ group because their songs are as individual as their plumages, but even observers of a century ago noticed a thread of similarity that ties them together. Perkins (1903: 394) said that, despite the variation among the species,‘most of the different forms frequently utter calls or notes very similar to one another’, and Baldwin (1953) also considered this vocal grouping valid. One characteristic of the group is the occurrence of almost incredibly wide intraspecific variation, from the individual to the population level. Derivation of the meliphagidlike vocalisations from a canary-like song is not as great a leap as it might at first seem.The pattern of sounds in the songs of the ‘Apapane and ‘Akohekohe is the same as in drepanidine finch songs, with whistles and trills interspersed with shorter notes. The main differences are in tonal quality. Except for the bell-like notes, reedy dissonances, and mechanical-sounding clicks and buzzes, the songs of the ‘Apapane are rather finchlike (Ward 1964; pers. obs.). An evolutionary step away from the highly varied ‘Apapane songs are those of the ‘Akohekohe. They are much lower pitched and slower, but have the same fundamental patterns of notes, as if an ‘Apapane recording were being played back at lower speed. The songs of the ‘I‘iwi are highly distinctive, even within the ‘red and black’ group, but maintain the same unusual tonal quality as the others. For one thing, ‘I‘iwi tend to vocalise for lengthy periods with short pauses between song elements, so that the decision as to where one song ends and another begins is purely arbitrary. In this pattern of delivery, it resembles the song of the North American Yellow-breasted Chat.The voice of the ‘Ula‘ai-hawane was never described and the two mamos became extinct before recording equipment was available, but Perkins (1903) and Bryan (1908) described their vocalisations as consisting of long penetrating whistles, much less complex than those of the other members of this group but with the same unusual tone. Note that among the drepanidine finches, the canary-like songs became reduced to long whistles in some species in a possible example of parallel evolution. One mysterious aspect of the songs of this group is why the birds sing so vigorously in situations

where they are obviously not defending territory or apparently seeking mates.When ‘ohi‘a and other flowering trees are in full bloom, ‘Apapane in particular gather in huge numbers. These feeding assemblages are characterised by an almost unbelievable chorus of song that produces an amazing cacophony. Early naturalists even suggested that the birds must be drunk on fermented nectar! Henshaw (1901: 122) referred to such scenes as ‘one mad revel all day long’, with the ‘I‘iwi and ‘Apapane singing ‘almost incessantly’.Why these (and probably related birds when they were more numerous) would put so much precious energy into vocalisation in such situations is difficult to surmise, but that they do so is without question. I have on occasion observed huge gatherings of these species, with dozens of individuals in a single large tree, often feeding within a metre of each other, each bird singing with reckless abandon and apparently oblivious of the others nearby. A variation on this behaviour was reported by Henshaw (1902: 56–7) who said of the ‘Apapane: this species and the iiwi rank as the most persistent songsters the writer has ever heard. The [‘Apapane] has a delightful habit of gathering together in loose companies in the tops of the leafy ohia trees about midday, when hunger is appeased and most of the other forest songsters are silent, when the males join in a subdued lullaby and literally sing themselves and their mates to sleep. These choruses are clearly distinct from territorial proclamations as described by Ward (1964), when advertising males are well spaced in the tops of separate trees. This behaviour resembles that of cardueline finches in winter, when ‘groups of cocks sometimes sit in the tree-tops and sing softly, in warm afternoons or in evenings prior to roosting’ (Newton 1973: 163). In both cases ‘song seems to result from the social stimulation of the group.’

Short simple songs The remaining Hawaiian honeycreepers whose vocalisations can be categorised sing much shorter

118 The Hawaiian Honeycreepers and relatively stereotyped songs, as compared with either of the above groups. Virtually all are either trills (a single element repeated rapidly) or warbles (a short series of frequency-modulated whistles), with some intermediacy.All are species-specific and easily identified in the field by an experienced observer (Scott et al. 1979), but the overall similarity caused difficulty for earlier naturalists. Henshaw (1901) claimed never to have heard the songs of the Lesser ‘Akialoa or the Hawai‘i Creeper, but in retrospect he probably simply failed to differentiate them from similar songs of the Hawai‘i ‘Amakihi, which are quintessential examples of the drepanidine short song: virtually always a simple short level pitch trill, the individual elements of which may be quite variable from song to song (Fig. 7.2). Vocal differences of the O‘ahu ‘Amakihi have not yet been adequately investigated, and it sounds much like its Hawaii/Maui relative. The Kaua‘i ‘Amakihi has a similar trill, but it has a tendency to drop in pitch, and sonograms (Part II, Fig. 11) have revealed a previously overlooked but clearly audible introductory note not present in the songs of other ‘amakihis. In a similar fashion, the song of the Greater ‘Amakihi, as described by Perkins (1903: 413), was identical to that of the Hawai‘i ‘Amakihi except for ‘two or three distinct additional notes’ following the trill. Their songs are poorly known, but the ‘akialoas apparently sang trills very similar to, but distinguishable from, those of ‘amakihis (Palmer in Rothschild 1893–1900; Perkins 1903). A canary-like song attributed to the Kaua‘i ‘Akialoa by Palmer may have actually been a whisper song (see below). The ‘Anianiau also sings a trill, but it characteristically includes much more complex elements that make it sound as if the elements are doubled or tripled (Part II, Fig. 10).The Hawai‘i Creeper’s trill is quieter and faster than that of an ‘amakihi, and has a distinctive rattling quality (Scott et al. 1979; Part II, Fig. 8). The ‘Akikiki sings a trill of short notes that descend in pitch (Part II, Fig. 8); it closely resembles some variations of the Kaua‘i ‘Amakihi song (Part II, Fig. 11). Trills of the ‘Akepa and ‘Akeke‘e tend to shift pitch and speed in midstream, and their individual elements may be more complex than those of ‘amakihi songs. The ‘Akepa

sometimes sings such slow, lackadaisical trills that they are really quick repetitions of short warbles (Part II, Fig. 9). Also tending toward a warble are the songs of the Maui Parrotbill which are trills in that the individual kHz

Hemignathus virens virens 05265

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7.2 Sonograms of songs of Hawai‘i ‘Amakihi, showing variation among individuals and within singing bouts of single individuals (songs with same number). Numbers from LNS catalogue.

Behaviour 119 kHz 8

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7.3 The warbled short song of ‘Akiapola‘au, LNS 05185.

elements are similar throughout, but the elements are short modulated whistles. They also virtually always drop in pitch.With a very similar sound quality is the song of the ‘Akiapola‘au, but it differs in that it is a short series of whistled notes, each element of which is different (Fig. 7.3); it is a warble rather than a trill, although Perkins used the latter term to characterise it. Henshaw (1901) described it as a quick version of the song of the Yellow-throated Vireo and that comparison is as good as any (pers. obs.). Perkins (1903) considered the songs of the Kaua‘i and Maui nukupu‘us very similar, but the Maui bird apparently had some distinctive variations. Unfortunately, none of the supposed observers of nukupu‘us in the twentieth century ever heard them sing (T. K. Pratt and Pyle 2000).

Whisper songs Those Hawaiian honeycreepers that belong to the ‘short song’ group have not abandoned long complex vocalisations entirely. Although their primary advertising songs are rather simple, they have an apparently unique class of vocalisation that is customarily called ‘whisper song’.Although they are similar in structure and amplitude to the subsongs sung by young birds in the course of learning their songs, drepanidine whisper songs are uttered by adults and their purpose is as yet a mystery.Although Perkins (1903) perfectly described a whisper song uttered by a female ‘Akepa, Baldwin (1953: 339) was the first to recognise such songs as a distinct class of vocalisation. He described the whisper song of the Hawai‘i ‘Amakihi as: a complex, extended song composed of quite a number of variations on rapid trills and warbles, frequently uttered in a soft tone. The phrases are strung together continuously and in any order.When the male is highly excited,

the slow trill [primary song] . . . may be interjected without interruption of the running song. Females may give snatches of soft notes resembling parts of the male song but lacking its variety and sustained, continuous character. Baldwin believed this song was primarily given by males but van Riper (1987) and Lepson and Freed (1997) attribute it also to females. Baldwin also considered the whisper song to be associated mainly with courtship and mating, but subsequent observations have revealed it to be uttered in a wider variety of situations including bathing (Eddinger 1970) and foraging (pers. obs.). In my own experience, it is just as likely to be given by a solitary foraging bird as by one engaged in any behaviour involving another individual. At times, a bird may perch quite still in concealment and sing the whisper song continuously for minutes at a time, as if just singing to entertain itself. These songs are indeed whispered and are usually audible from only a few metres away. I emphasise this point because listeners who study these songs only from recordings (Pratt 1996a) could easily be misled to think that they are louder in the field than they actually are. Nature has no volume control. Drepanidine whisper songs are, as Baldwin described, extremely complex with an almost infinite variety of elements that include chirps, chips, trills, and warbles, all of which resemble the primary song of the species. Possible examples of mimicry (see below) of other species are often included. In 1976, when whisper songs were first being widely noticed by field ornithologists, I recorded several minutes of such song by a male Kaua‘i ‘Amakihi from a distance of approximately 3 m (Pratt 1996b: Side C). I later played it back at high volume for other knowledgeable naturalists and not one guessed its author to be a Hawaiian honeycreeper (most thought it was a Northern Mockingbird!). Further complicating interpretation, drepanidine whisper songs exhibit surprisingly little specificity (Lepson and Pratt 1997), with many species sounding much the same (although studies of the details might reveal interspecific differences).They could be appropriately described as canary-like in pattern if not in

120 The Hawaiian Honeycreepers loudness, so can be viewed as a modified holdover of the ancestral finch song. Whether the whisper songs are phylogenetically informative is also problematical, partly because some species for which they have not yet been reported may have them. In many years of observation, the only example of whisper song by an ‘Akikiki ever recorded or reported was that given by a foraging single bird that I recorded in 1976 (Pratt 1996a), and the whisper song of the Hawai‘i Creeper was only reported recently (VanderWerf 1998). No whisper song has been reported for the O‘ahu ‘Amakihi, but because all other ‘amakihis have them, we can expect this little-studied species to have one also. Such ‘generic’ whisper songs are also known in the ‘Anianiau (Lepson 1997) and both species of Loxops (Lepson and Freed 1997; Lepson and Pratt 1997), that of the ‘Akeke‘e having been first recorded after publication of Pratt (1996a), and probably ‘akialoas. Palmer (Rothschild 1893–1900) described a canary-like song for Kaua‘i ‘Akialoa, and Munro (1960) refers to a ‘light sweet song’ given by both male and female of the same species. Both are likely to have been whisper songs. Whisper songs of ‘Akiapola‘au, recorded by T. Burr in 1979 (Pratt 1996b), and Maui Parrotbill, as described by Simon et al. (1997), may, in fact, be true subsongs because they comprise elements of primary song and are clearly species-specific. Whisper songs are not entirely restricted to the hemignathines. The Palila often gives its song very quietly (pers. obs.) but whether this qualifies as a true whisper song is questionable because the quiet performance does not differ from the normal advertising song. Berger (1981) reported a Palila whisper song but did not describe it. The Maui ‘Alauahio sings a whisper song much like that of the ‘amakihis. That song may be one of the few distinctively drepanidine vocal features possessed by that species. The only example yet reported of possible whisper song among the ‘red and black’ nectarivores is that of the ‘Akohekohe. Engilis (1990: 70) heard three different adults deliver ‘a soft warbling, gurgling subsong. In pattern it resembled the subsong of the ‘amakihi, but it was harsher and less musical. When delivering this subsong the birds sat motionless in the understory.’ This whisper song (or true subsong) is

possibly the same type that I previously (Pratt 1979a) thought was restricted to the short-song vocal group, but the very quiet vocalisations I have heard sung by ‘Akohekohe would not qualify because they were simply quiet versions of species-specific primary songs and calls.We may simply be drawing arbitrary lines in a continuum.Whether drepanidine whisper songs are homologous to the quiet songs given by cardueline finches in winter (Newton 1973) is an intriguing question, unanswerable for the present.

Vocal mimicry According to Munro (1960: 130), ‘Freeth thought [the song of the Laysan Finch] was a musical imitation of the various bird cries of the island.’ Two decades ago I stated (Pratt 1979a), that whether any Hawaiian honeycreepers engage in true vocal mimicry was problematical and required further study. Drepanidine whisper songs (see above) are so varied and variable that some of the included notes are likely, by simple coincidence, to resemble those of other species. That may be the case with the most frequent apparent imitation, a note that sounds like the call of the introduced Nutmeg Mannikin. Why Hawaiian honeycreepers would mimic this exotic seedeater is not easily explained, but this species was among the earliest introductions to Hawai‘i and is widespread today even in forest openings. Since 1979, so many authors have cited examples of drepanidine mimicry that its existence now seems undeniable.VanderWerf (1998) reported mimicry of Hawai‘i ‘Elepaio and ‘I‘iwi in the whisper song of the Hawai‘i Creeper. T. K. Pratt et al. (2001) reported imitations of ‘virtually all syntopic passerines’ by whisper-singing ‘Akiapola‘au, whose whistle call is in turn mimicked by Palila, whose contact call is imitated by Hawai‘i ‘Amakihi (Pratt 2002b)! Careful comparative analysis of sonograms of whisper songs and their supposed sound models would be helpful in proving whether true mimicry is involved. A few honeycreepers present more compelling possible examples of vocal mimicry. For example, modern birders have noted, often with considerable annoyance, that Hawai‘i ‘Amakihi within the range of the Palila produce a short call that is very close to that of the larger bird. This note is not heard

Behaviour 121 elsewhere, but why ‘amakihi would imitate a bird with which they presumably do not compete is an intriguing question. Likewise, one population of ‘I‘iwi, on the kona slope of Hualalai, seem to produce a nearly perfect imitation of the loud whistled song of the Hawai‘i ‘Elepaio. Although the two species are broadly sympatric and syntopic elsewhere, this is the only locality where a human listener could be confused as to which species was vocalising. I learned of this situation during a study of intra-island variation in the Hawai‘i ‘Elepaio (Pratt 1980), when virtually every ‘ ‘elepaio’ on Hualalai turned out to be an ‘I‘iwi. Obviously, much remains to be learned about the voices of Hawaiian honeycreepers.

Non-vocal sounds Wing-flutter The most intriguing non-vocal sound produced by any Hawaiian honeycreepers may be the peculiar audible wing-flutter characteristic of the ‘red and black’ nectarivores.This sound is quite different from the usual wing-flutter produced by many birds, including other honeycreepers, and sounds more like a vocalisation. Henshaw (1902) called the notes ‘rhythmic pulsations’. One of the most frequent questions asked by novice birders in Hawai‘i is ‘What dove is that calling?’ when the cooing notes are actually the wing-flutter of honeycreepers. Perkins (1903: 404) noted that ‘I‘iwi wing beats were ‘very audible even when [the birds] are high up and at a distance.’ The function of this ‘wing-note’ is not known, although some authors have speculated that it is associated with flocking or territorial behaviour. Unlike vocalisations, of course, it is present year round and in all contexts.Whether this sound is analogous to wing noise produced by the North American Broad-tailed Hummingbird, also a nectarivore, is likewise unknown. Wing-notes are produced by ‘Apapane, ‘I‘iwi, ‘Akohekohe, and the two mamos (Perkins 1903; pers. obs.). Whether the ‘Ula‘aihawane produced wing-notes is not known.The first two species have primary feathers with obvious modifications (Chapter 7) that supposedly enhance this sound (Amadon 1950), and Perkins (1903)

reported similar modifications, not apparent to me, in the mamos. Henshaw (1902) states that the ‘Akohekohe lacks the wing-note characteristic of the ‘I‘iwi and ‘Apapane, but I must disagree. I have not noticed it in sustained flight, but the note is clearly audible during short flights between feeding perches. I agree with Perkins (1903) that the ‘Akohekohe lacks the truncate primaries, but they produce a wing-note (audible on Pratt 1996a) nevertheless. Thus Perkins’s statement that the note is ‘almost confined’ to species with modified primaries allows for its presence in ‘Akohekohe. Apparently selection favoured the sound itself, and the secondary feather modifications evolved to enhance it. Charles van Riper (in Lindsey et al. 1998) reports that Hawai‘i ‘Amakihi use ‘wing-whirring’ to deter intruders at their nests, but whether this sound is homologous or analogous to that produced by the ‘red and black’ birds is not known.

Bill clapping In aggressive encounters with conspecifics and other birds, Maui ‘Alauahio frequently make ‘a loud, sharp, snapping noise’ (H. and P. Baker 2000) with their bills. This sound is very similar to bill sounds made by birds that capture insects on the wing, such as ‘elepaios. The Maui ‘Alauahio is the only honeycreeper reported to feed in this manner, and it indeed produces bill claps in that context, but also uses them apparently for aggressive signalling. Bill clapping associated with aggressive chases has also been observed in the Maui Parrotbill (Simon et al. 1997).

Feeding noises In addition to the just-mentioned bill clapping associated with aerial sallies for insects, Hawaiian honeycreepers produce other sounds while feeding. These sounds are produced by the bill making contact with a feeding substrate, such as a tree trunk, or with a food item. Early naturalists and modern birders alike have noted the woodpeckerlike pecking sounds produced by the ‘Akiapola‘au, but the honeycreeper never pecks as loudly as its mainland counterparts. Similar, but quieter, pecking

122 The Hawaiian Honeycreepers noise is produced by the closely related nukupu‘us (Perkins 1903), and still quieter tapping was reported for ‘akialoas. Other bark-picking species, such as Hawai‘i Creeper,‘Akikiki, and Maui Parrotbill, produce cracking or snapping sounds in the course of foraging.The Kona Grosbeak was said to be most easily located by the loud noises it made as it crushed hard, dry naio fruits (Munro 1960).

Displays Display flights Displays involving flight are known in several, but by no means all, Hawaiian honeycreepers. The Palila has a ‘low advertising flight’ described as ‘a slow fluttering of wings followed by a short glide’ (van Riper 1987), similar to the cardueline ‘butterfly flight’ in which ‘a series of slow deliberate wingstrokes are interspersed with long glides’ (Newton 1973: 164).A similar flight has been reported in the Hawai‘i ‘Amakihi,‘usually given in a gentle arc over an established territory, either in silence or accompanied by primary song’ (van Riper 1987: 88).The ‘Akiapola‘au has a ‘Helicopter Display’ directed at territorial rivals that is apparently homologous (T. K. Pratt et al. 2001). When such flights involve vocalisation, as they frequently do, they are often called ‘song flights’. The typical drepanidine song flight involves the bird rising high above the canopy with very deliberate wing-beats, hovering and singing for a while, then gliding back into the foliage. Such flights are performed by ‘Apapane (Eddinger 1970; Berger 1981), Maui ‘Alauahio (H. and P. Baker 2000), Hawai‘i ‘Amakihi (van Riper 1987), ‘Akiapola‘au (T. K. Pratt et al. 2001), and ‘Akohekohe (Engilis 1990), and possibly Nihoa Finch (Morin and Conant 2002). In ‘Akohekohe it may involve 2–6 birds (Berlin and VanGelder 1999; pers. obs.), but whether both sexes are involved is not known. Such song flights may be homologous to the ‘distinctive fluttering or ‘whirring’ flight with rapid, silent wing-beats’ accompanied by song of groups of male ‘Akepa (Lepson and Freed 1997: 7). Following an aggressive encounter, Maui ‘Alauahio will ‘rise spirally upwards to a height of twenty or thirty feet pouring out its little song while on the

wing’ before ‘suddenly darting down again to cover in the underbrush’ (Perkins 1903: 414; pers. obs.). Song flights are not to be confused with simple singing on the wing, which has been reported in several species. Eddinger (1970) found that ‘I‘iwi, ‘Anianiau, and Kaua‘i ‘Amakihi all sang in flight occasionally, but had no special song-flight display; Berger (1981) reported singing in flight by Palila; and Perkins (1903) reported similar behaviour by Maui Parrotbill and ‘O‘u .The latter may have actually been a display flight, but the information is insufficient to say for sure. The same is true for Rothschild’s (1893–1900) report of Palmer’s observation of a Kona Grosbeak and Munro’s of a Kaua‘i ‘Akialoa vocalising in flight.

Mating displays Mating displays involve one or both members of a prospective pair, and are used in an effort to form a pair-bond.They are not advertising displays such as those just described, and are therefore much more difficult to observe. Probably all species of Hawaiian honeycreepers have mating displays, but only a limited number have been described at all, and even fewer in detail. Like songs, which usually accompany these displays, each species has its own characteristic choreography, but some underlying patterns can be discerned. For example, both Laysan and Nihoa Finches have displays that involve spreading the wings, but they differ in details and associated movements. Morin (1992: 650–1) reports that both sexes of Laysan Finches spread their wings ‘rapidly in and out and the tail [is] rapidly flipped up and down.’ In contrast, the Nihoa Finch male ‘holds his wings horizontally away from the body; sometimes he also sways slowly to the right and the left, and he may move short distances along the perch’ (Berger 1981: 116). Virtually all species yet reported have two mating displays in common: sexual chasing and courtship feeding. Wildly cavorting birds engaged in sexual chases are a conspicuous part of the Hawaiian environment late in the winter months when pair-bonds are forming. Some species, such as the Hawai‘i ‘Amakihi (van Riper 1987) and ‘Akepa (Lepson and Freed 1995), have specific subtypes of sexual chases

Behaviour 123 (see species accounts for descriptions). Some chases involve males competing for females (Lepson and Freed 1997), others involve potential mates. Courtship feeding is apparently very similar in all species. The female may solicit feeding by crouching and quivering her wings, often with a vocalisation that resembles begging notes of nestlings or fledglings. The same posture is used when she submits to copulation. Among the hemignathine group, copulation is usually solicited by what has been variously called a ‘hopping’ or ‘flitting’ display in which the male flits or hops rapidly back and forth near the female, usually giving either the advertising song or the whisper song. Displays of this type have been reported for Hawai‘i ‘Amakihi (van Riper 1987), O‘ahu ‘Amakihi (Russell and Ralph 1981), and Kaua‘i ‘Amakihi (Eddinger 1970), but each apparently has distinctive versions.The ‘Akiapola‘au also has a flitting display that has not been described in detail (van Riper 1978). This display in ‘Akeke‘e (Lepson and Pratt 1997) and ‘Akepa (Lepson and Freed 1995, 1997) involves more hopping than flitting, although the difference may be more semantic than real. The ‘Anianiau has a variation, reminiscent of the spread-wing displays of the Telespiza finches, in which the male hops and bobs along near the female with wings extended and held horizontally (Eddinger 1970). ‘Akepa are apparently unique among Hawaiian honeycreepers in employing group displays (Lepson and Freed 1995). These displays involve up to six males displaying for one female, and somewhat resemble lek behaviour.The only mating display reported among the ‘red and black’ honeycreepers is that of the ‘I‘iwi, which seems to combine elements of the Nihoa Finch display and the flitting display (for detailed description see species account). Apparently in species such as ‘Apapane and ‘Akohekohe, song is more important than physical actions in forming pair-bonds.

Agonistic displays Agonistic displays are those that involve aggressive confrontations, most often between two males competing for territory or a female, as in the ‘Helicopter

Display’ (see species account) of the ‘Akiapola‘au (T. K. Pratt et al. 2001).The most frequent examples among Hawaiian honeycreepers are chases that are very similar to sexual chasing, except that they often involve actual contact with foot grappling and pecking. Sometimes a pair of grappling males may plunge all the way to the ground (Lepson and Freed 1997; H. and P. Baker 2000). Maui ‘Alauahio use bill claps and may actually pull tail feathers of a rival (H. and P. Baker 2000). Maui Parrotbills use bill clapping (Simon et al. 1997) and pecking with the upper mandible (Mountainspring 1987). ‘Akepa males have an aerial display in which up to eight males engage in ‘dogfights’ high above the ground (Lepson and Freed 1995, 1997). Birds fly in from as much as 200 m away to join these battles, which eventually break up into individual chases back into the treetops. Morin (1992) reported a stationary threat display by Laysan Finches (see species account for details) virtually identical to the ‘head-forward threat’ of cardueline finches (Newton 1973). A similar display by the Maui Parrotbill (Baker and Baker 1997) and Po‘o-uli (Kepler et al. 1996) has also been observed, and the ‘Akiapola‘au’s ‘lion head display’ (T. K. Pratt et al. 2001) may be homologous. No perched agonistic displays have been reported for other honeycreepers.

Flocking Any time more than two birds are present together and interacting in a co-ordinated manner, they constitute a flock. Some authors regard any group of birds as a flock, others exclude from the definition gatherings of individual birds at a food source, where no co-ordinated activity occurs. Gatherings of several species of Hawaiian honeycreepers at an abundant food source are common (Mountainspring and Scott 1985; VanderWerf and Rohrer 1996; Lindsey et al. 1998; pers. obs.), but are not really flocks in the restricted sense meant by most authors because individual birds come and go independently. Most true flocks involve a single species (‘Birds of a feather. . .’), most often family groups. Cardueline finches, especially such species as goldfinches and crossbills, commonly move about in flocks (Newton 1973), usually in response to seasonally available food

124 The Hawaiian Honeycreepers supplies (Nethersole-Thompson 1975; Kaufman 1999). Insectivorous birds of tropical rainforests and of winter woodlands in temperate climates often form flocks involving several species. Both mixedspecies and conspecific flocks are found among Hawaiian honeycreepers.

Mixed-species flocks E. O. Willis (1972) posed the question ‘Do birds flock in Hawaii, a land without predators?’ He was, of course, like everyone else at the time, labouring under an assumption (lack of avian predators) that we now know to be false (Olson and James 1991). But his question still has merit because it goes to the heart of one of ornithology’s current controversies: do the mixed-species flocks so characteristic of tropical forests worldwide form as a response to predation or to gain trophic (feeding) advantages. If the former, then flocks should not form in a tropical avifauna that lacks predators. But Hawai‘i was not the place to conduct a test of this hypothesis,1 not only because avian predation was certainly an important element in the evolution of its birds, but also because so many species that potentially would have joined flocks under pristine conditions are now extinct or very rare. Modern observations of flocking behaviour in Hawai‘i must take these factors into account. Contrary to the artificial situation that led Willis (1972) to conclude that Hawai‘i lacks mixed-species flocks (despite reports of them by early naturalists), recent observations in less disturbed habitats show that such flocks remain a feature of the Hawaiian environment, even though the predators that probably instigated the behaviour are long extinct. O‘ahu lost its native avifauna too early to provide much useful information about mixed-species flocks, but the other large islands provided considerable information to naturalists at the end of the nineteenth century and continue to do so today. On all of them flocking behaviour is seasonal, following the breeding season when many juvenile birds are present in the population. In most places, flocks are most evident in the second half of the year. Thus Hawaiian mixed flocks differ from the highly organised, closed social systems maintained throughout

the year elsewhere in the tropics (Gill 1990). Another fundamental difference is that flock size increases by recruitment of additional members of the same species already present, rather than by adding new species to the mix as is the case with flocks in the American tropics (Powell 1979). Some inter-island differences are apparent, not only in the species composition of these flocks, but in their size and possibly even their purpose. On Kaua‘i flocks are not as tightly organised as those I have observed elsewhere but are clearly present nevertheless. In Fiji and Panama, one may go for long periods without seeing any birds, then be overwhelmed by a horde of varied species at every hand, with many individuals foraging quite close together, only to have them melt away into the forest just as suddenly. Following such flocks is very difficult. Modern flocks on Kaua‘i usually include several species, but they are so loosely associated that one can easily fail to notice the flocking behaviour. For example, when birding along a trail, one may have fairly long periods with no sightings, then see one bird. Then, at some distance, another appears, then another.The process is quite slow and deliberate, and unless one remains stationary, the sense of flock movement is not apparent. However, by following call notes, one gets a definite impression that a loose, multi-species party is moving through the forest in concert. The species involved today almost always include the non-drepanidine Kaua‘i ‘Elepaio and such honeycreepers as ‘Anianiau, Kaua‘i ‘Amakihi, ‘Akeke‘e, and ‘Akikiki (pers. obs.; Mountainspring and Scott 1985). Occasionally the nectarivorous ‘Apapane (Fancy and Ralph 1997) and ‘I‘iwi (pers. obs.) follow such groups for a short distance. Present-day flocks on Kaua‘i rarely comprise more than 10 individuals. Historically the situation was quite different, and Kaua‘i ‘Akialoa and Kaua‘i Nukupu‘u were most often seen associated with mixed flocks (Perkins 1903). Indeed, Perkins (1903: 429) stated that at times ‘nearly all the known forest birds of Kauai may be found in a single assemblage.’ At that time, the ‘Akikiki appeared to be the nuclear species that was then joined by ‘a pair or two of Elepaio and Akialoa, several Chlorodrepanis, and possibly several Oo overhead’ and a few Kaua‘i Nukupu‘u in the understorey below. Perhaps the

Behaviour 125 lack of tighter organisation in Kaua‘i flocks today results from the scarcity of ‘Akikiki, which is by no means now present in every flock. Mountainspring and Scott (1985) considered mixed-species flocks more common on Maui than on other islands. Present-day flocks on that island seem somewhat more tightly organised than those on Kaua‘i, but that may be because they most often involve a single species, the Maui ‘Alauahio, with some participation by Hawai‘i ‘Amakihi.That biased species mix is clearly an artefact, because in forests that still have a representative sample of their original species, others are regularly involved (Engilis 1990; pers. obs.). In fact, naturalists searching for rare birds routinely look for them among the conspicuous, loudly calling flocks of ‘alauahio.A fully formed mixed-species flock on Maui would include Maui Parrotbill, Maui Nukupu‘u, and Po‘o-uli. As on Kaua‘i,‘I‘iwi may join the group, but whether their participation is purposeful or coincidental is not known. Maui mixed flocks may be twice as large as those seen today on Kaua‘i, probably because the ‘nuclear’ species is still common. Remarkably, careful reading of the early naturalists reveals no specific reference to mixed-species flocks on Hawai‘i, even though it was the only island to retain a native bird predator. Baldwin (1953: 342) was the first modern writer to describe a mixed-species flock on The Big Island: On October 24, 1948, I encountered a loose grouping of six species of drepaniids slowly moving along together near Solomon’s Waterhole, Keauhou. Many other instances could be cited, but it will suffice to add that the groups are seldom well enough knit to look like flocks moving in unison, and often they appear to be merely individuals acting independently, perhaps together only by coincidence. It is not known how far these small groups go or whether the same individuals remain together for any length of time. In summer one may occasionally hear male trills given by [Hawai‘i ‘Amakihi] feeding with others, indicating that territorial ties in such individuals are not binding at this time. Later in the fall, when older males are vigorously

defending territories, it is evident from banding work that a large number of individuals, especially females and young males, are still wandering at large. Flocking of this type reaches its peak after the breeding season and continues into fall and winter. Baldwin’s report was largely overlooked, as witness Willis’s (1972) paper and the amazement with which I and my co-workers viewed a similar phenomenon in the same general area on 19 August 1975 (Pratt et al. 1977). During a light, warm rain we observed incredibly high numbers of endangered species in an area well studied by other observers.The most numerous were Hawai‘i Creeper, but ‘Akiapola‘au were also unusually abundant and conspicuous. During 5 hours of observation at midday, we saw 50–75 creepers and 50 ‘Akiapola‘au. Other birds in the area were present in their usual numbers, including such honeycreepers as ‘Apapane, ‘I‘iwi, ‘Akepa, and Hawai‘i ‘Amakihi. The following day the weather had cleared and the huge numbers of creepers and ‘Akiapola‘au were no longer present. At the time, we did not realise that we were seeing an example of flocking behaviour, although we speculated (Pratt et al. 1977) that it may have been ‘a wave of postbreeding wanderers’. On 11 August 1992, I observed a similar phenomenon at Hakalau Forest National Wildlife Refuge. After a period of seeing very few birds, we came upon a large flock of ca. 50 ‘Akepa, 10 Hawai’i Creeper, and 10 ‘I‘iwi, moving together from tree to tree in a semi-open area of the refuge. As before, a light, warm rain was falling, not hard enough to prevent observation. At times, the entire assemblage occupied only two or three mediumsized ‘ohi‘a trees. According to the careful 1988–93 observations by Lepson and Freed (1995) in the same area, ‘Akepa-centred flocks of up to 40 individuals form around family groups, with nonbreeders and unsuccessful breeders joining as well as large numbers of Hawai‘i Creeper, a few Hawai‘i ‘Amakihi, occasional ‘Akiapola‘au, and Japanese White-eyes, an introduced species. Fancy and Ralph (1997) report that ‘Apapane also join such flocks. Flocks are present every day in this area

126 The Hawaiian Honeycreepers between July and September, although the amount of flocking varies from year to year. These groups disperse after September, when birds begin courtship. Mixed-species flocks observed on The Big Island, especially those led by ‘Akepas, give an entirely different impression from those on Kaua‘i and Maui, with much larger numbers and much more closely associated participants. They also differ in being confined to a relatively short period in late summer.

Conspecific flocks Before honeycreepers gather into mixed flocks, they travel about in small post-breeding parties, usually assumed to be made up of several family groups, but sometimes including juveniles only (Berlin and VanGelder 1999). Such small conspecific flocks have been reported in Palila (van Riper 1980), Kona Grosbeak (Perkins 1903), Hawai‘i ‘Amakihi (Baldwin 1953; van Riper 1987), ‘Akepa (Lepson and Freed 1995, 1997), ‘Akeke‘e (Bryan and Seale 1900; Lepson and Pratt 1997), Maui ‘Alauahio (H. and P. Baker 2000), ‘Anianiau (Lepson 1997), ‘Apapane (Fancy and Ralph 1997), ‘Akohekohe (Perkins 1903; Berlin and VanGelder 1999), and ‘I‘iwi (Baldwin 1953; Fancy and Ralph 1998). Other species maintain single-family parties after the young have fledged and do not gather into larger parties unless and until they join mixed flocks. Both the ‘Akikiki and the Hawai‘i Creeper have conspicuous family groups that forage as tight parties of a pair and one or more offspring (Pratt 2001b). These groups may form the nucleus of mixed flocks on Kaua‘i and Hawai‘i.Young of the ‘Akiapola‘au (T. K. Pratt et al. 2001) and Maui Parrotbill (Simon et al. 1997) keep company with their parents for much longer periods than other Hawaiian honeycreepers, perhaps because of a prolonged learning period associated with their unusual bills. An entirely different kind of conspecific flock is formed by ‘Apapane (Carothers 1986a). These could be called feeding flocks, because they enable a subordinate species to gain advantage over a dominant species by sheer force of numbers (see discussion of nectarivory, Chapter 8).The larger the flock, the greater the advantage up to a point.

Predator response Hawaiian honeycreepers are peculiar among passerine birds in the ways they deal with predators and other intruders into their habitat. Mainland birders have long noticed that Hawaiian honeycreepers very rarely respond to ‘spishing’ or ‘squeaking’ (kissing the back of the hand to produce a high-pitched note), auditory lures often used to attract birds to an observer (Homel 1991; Pratt 1992a,b, 2002b). A popular birder’s joke is that spishing helps to pick the honeycreepers out of an assemblage because it attracts Japanese White-eyes, leaving the native birds behind! Other Hawaiian passerines, such as the ‘Elepaio,‘Oma‘o, and Hawaiian Crow, readily ‘spish up’ (Pratt 1992a), as do many continental cardueline finches (pers. obs.). Most honeycreepers, including Palila, ‘Akiapola‘au, Maui Parrotbill, ‘Akepa,‘Akeke‘e, and notably Hawai‘i Creeper and ‘Akikiki, apparently never respond to such lures (Pratt 1992a,b). Members of the ‘red and black’ group, however, do sometimes respond.‘I‘iwi,‘Apapane, ‘Akohekohe (Engilis 1990), and formerly the two mamos (Perkins 1903), are all attracted by man-made sounds.The ‘amakihis and the ‘Anianiau usually ignore auditory lures but occasionally exhibit a very strong response (Willis 1970; Engilis 1990; pers. obs.). Interestingly, the members of the two genera that stand apart in so many other ways, Paroreomyza and Melamprosops, readily respond to spishing (Engilis 1990; Pratt 1992a,b). Some of the same honeycreepers that respond to auditory lures also engage in ‘approach-and-follow’ behaviour. The Hawaiian bird most noticeable for this behaviour is the non-drepanidine Kaua‘i ‘Elepaio (Conant et al. 1998), and it is also characteristic of the Hawaiian Crow (Perkins 1903). Small flocks of Maui ‘Alauahio often approach an observer, seemingly out of simple curiosity. Bryan (1908) reported similar behaviour for the Kakawahie. Perkins (1903: 406) gave a particularly good account of this behaviour in the ‘Akohekohe: The adults, though easily called and curious, are less tame than the young, as is shown by their restless movements when they are attracted, but in dense untrodden forests they

Behaviour 127 will not infrequently approach the collector from mere curiosity, without any imitation of the call note being necessary. On one occasion I assembled no less than nine adult birds at the same time in one small Ohia tree not more than twenty feet high, and a pair of adults and several young were quite an ordinary gathering. One of the best known avian responses to predators is the phenomenon known as mobbing (Curio 1978), which is related to the behaviours just discussed (Emlen 1969). Mobbing has been documented for many cardueline finches and is believed to be innate rather than learned (Hinde 1954; Altmann 1956). Curio (1978: 176) described this specific behaviour pattern: While mobbing, birds of one or more species assemble around a stationary or moving predator (potentially dangerous animal), change locations frequently, perform (mostly) stereotyped wing and/or tail movements and emit loud calls usually with a broad frequency spectrum and transients . . . As a rule, a mobbing assembly builds up within less than a min after the first individual has discovered the predator.Thus mobbing has a dual nature:The mere presence of a predator suffices to initiate it in the first individual, whereas later members of the chorus are attracted by the mobbing behaviour of the initiator (and by the enemy); vocalisations alone are sufficient as can be demonstrated by mere playbacks of mobbing calls luring conspecifics and other birds. Mobbing behaviour that fits this description has been reported in only one Hawaiian honeycreeper, the Maui ‘Alauahio. Perkins (in Wilson and Evans 1890–99: 41) reported that when Short-eared Owls fly overhead, ‘alauahio ‘become greatly excited, all those in the neighbourhood joining in the clamour.’ Jack Jeffrey and I observed an example of this behaviour in October 1994 in the Hanawi Natural Area Reserve. In response to the overhead passage of an owl, seemingly all the Maui ‘Alauahio in the

vicinity left their usual haunts in the forest understorey and rose to the upper branches, chipping loudly all the while and producing a very noticeable commotion.As they did so, other honeycreepers nearby (‘I‘iwi, ‘Akohekohe, Hawai‘i ‘Amakihi, and ‘Apapane) had just the opposite reaction, leaving the canopy for concealment and falling silent. Perkins reported ‘twenty or thirty’ ‘alauahio gathered about a sleeping owl. They also mob other kinds of predators, including feral cats. In the Hanawi area in July 1975, I watched a mobbing incident involving a cat (Pratt 1992b). As the animal crept slowly along the forest floor, Maui ‘Alauahio began to follow and set up a loud clamour with their chipping calls. Others joined the group until about a dozen birds became involved. ‘Apapane, Hawai‘i ‘Amakihi, ‘I‘iwi, ‘Akohekohe, and Maui Parrotbill were within auditory range of the mobbing sounds, but none joined the fray. Kepler et al. (1996) report what may have been an example of mobbing, directed at something on the ground (mammal?), by a male Po‘o-uli. Because small bird bones are prominent components of pellets from prehistoric stilt-owls (Olson and James 1991), we know that predation was a force in the evolution of Hawaiian honeycreepers, so the lack of mobbing behaviour in most species is quite puzzling until the presence of drepanidine odour is considered. The odour (Chapter 4) is apparently confined to those species that do not mob. The Hawaiian honeycreepers’ loss of mobbing may have resulted from the development of a more energyefficient way of fending off predators. Nestlings of at least some Hawaiian honeycreepers have a response to predation threats that Eddinger (1970: 151), who discovered it, could not explain. He stated: Weight changes [of nestlings] could not be taken late in the nestling period because the nestlings developed a fear reaction and jumped from the nest if disturbed. This fear reaction was evident on day 11 in the Anianiau and Apapane, on day 14 in the Iiwi, and on day 17 in the [Kaua‘i] Amakihi. After the fear reaction developed the nestlings would often jump if I shook the tree while I was

128 The Hawaiian Honeycreepers climbing and even if the nestlings could see me in a neighboring tree. The adaptive value of jumping from the nest is difficult to understand. Presumably the birds had no predators when this behavior pattern evolved. Today it makes them an easier prey to rats. The nestlings jump from the nest when their feathers are just beginning to unsheathe and they are thus exposed to rain and cold temperatures. I have observed nestlings that jumped from the nest giving food calls from the ground, but I have never seen an adult [honeycreeper] fly to the ground to feed a nestling. Berger (1981) reported similar behaviour in Hawai‘i ‘Amakihi and was equally at a loss as to its adaptive value. Subsequent researchers have found this trait to be characteristic of honeycreeper nestlings, and have even coined the term ‘jumpling’ for the nest deserters (Lepson 1997). Despite the apparent lack of parental care for nestlings that leave the nest, jumping to avoid predation from overhead makes sense, now that we know that prehistoric predators hunted aerially, and that in pre-human times no ground predators waited below for the hapless chicks. This is an example, among others (Conant et al. 1998), of an originally adaptive trait

that became maladaptive in the presence of modern changes. In fact, the trait may not have been restricted to honeycreepers. When they first leave the nest, Hawaiian Crow fledglings are now known to spend several days on the ground ( Johnston and Banko 1992), where they are vulnerable to introduced ground predators, but which was a safer place than the treetops when all attacks came from above.This behaviour and the odd development of drepanidine odour (Chapter 4) show how important predation by owls, harriers, and eagles (Olson and James 1991) may have been in the evolution of Hawaiian birds. 1. Other Pacific islands are much better natural laboratories for the study of flocking, and my own studies support the predation-avoidance hypothesis. Mixed-species flocks are a prominent feature of the avifauna of Fiji (Watling 1982; Pratt et al. 1987), which has three avian predators (a harrier, a falcon, and an Accipiter ). However, in Samoa and the high islands of Micronesia, whose avifaunas include many of the same elements as that of Fiji but which have no avian predators, mixed-species flocks have never been observed. Further evidence is that the Fiji Peregrine Falcon Falco peregrinus nesiotes feeds disproportionately (Clunie 1972) on nectarivorous canopy birds that do not usually join mixed flocks (pers. obs.).

8 Ecology and breeding biology

The interactions of Hawaiian honeycreepers and their environment are a pervasive underlying theme in all discussions from evolution to distribution and status. The Hawaiian environment in general was discussed in detail in Chapter 2. Ecological observations appear throughout the discussion of evolutionary phenomena (Chapter 4), and are intimately intertwined with anatomical and functional analyses and natural selection of coloration (Chapter 6), and even to some extent in behaviours such as locomotion and self-maintenance (Chapter 7).This chapter focuses on several aspects of honeycreeper ecology that can be reasonably separated from other subjects, but a large amount of crossreferencing was necessary nevertheless. Likewise, breeding biology is entangled with both behaviour (displays) and ecology, and again I have resorted liberally to cross-referencing to avoid repetition wherever possible.

Feeding guilds The foods of adult Hawaiian honeycreepers cover the spectrum of those reported for passerine birds, but most species choose from a menu with only four categories: seeds, fruit, nectar, and invertebrates. Some species specialise in only one or two, but most of them at least occasionally take foods from all categories and a few are true generalists. Conversely, each category can be subdivided and some species have very exacting tastes. A few unconventional items (leaves, eggs, etc.) appear occasionally in honeycreeper diets, and all species feed invertebrates to their nestlings. Thus any classification of honeycreepers on the basis of diet

alone is very imprecise, with broad overlap between the categories. I have used the foods themselves as a basis for categorising the honeycreepers, and I use the term ‘guild’ for associations of species that exploit one type of food. Any given species may belong to more than one guild.

Seed-eaters The founder of the honeycreeper dynasty was primarily a seed predator. But seeds come in a wide variety of sizes and packaging and we do not know exactly what kind it preferred.We tend to think of finches as granivores, feeding on small seeds such as those of grasses (Poaceae), but although Hawai‘i has many native grasses (Wagner et al. 1990), few historically known honeycreepers exploited this food source. The only ones to do so commonly are the Telespiza finches, but grass seeds form only a small proportion of foods taken by these generalists (Morin and Conant 2002; see ‘Eclectic diets’ below). However, among the numerous prehistoric drepanidine finches must have been some, perhaps the smaller Telespiza, that specialised on small seeds, particularly considering the absence of mammalian competitors such as rodents. All other known drepanidine seed-eaters eat larger seeds removed from sizeable fruits, and all are specialists on one kind of plant with a few others as backup food sources. The Palila is a mamane seed-pod specialist (Fig. 6.2) with secondary feeding on naio (Berger 1981); the well-named koa-finches fed almost exclusively on seeds of koa, and secondarily on ‘a‘ali‘i (Part II, Fig. 11.5; Munro 1960); and the Kona Grosbeak was specialised to crack the extremely

130 The Hawaiian Honeycreepers hard endocarps of naio to extract the seeds, nearly its only food (Olson 1999).

Frugivores Perhaps because of competition from other frugivores such as corvids (Sakai et al. 1986) and solitaires (Snetsinger et al. 1999; Wakelee and Fancy 1999), few Hawaiian honeycreepers can be considered mainly fruit-eaters, although many take fruit opportunistically. Only the ‘O’u can certainly be classed as a frugivore. It eats small, mostly soft fruits, favouring those of ‘ie‘ie (Fig. 8.1), but can hardly be considered a specialist because of the wide variety of foods, including introduced fruits and insects, it is known to have eaten (Henshaw 1902; Perkins 1903; Snetsinger et al. 1998). ‘O’u ate fruits that

8.1 Fruit of ‘ie‘ie, a composite fruit the small components of which are eaten whole by the ‘O‘u, whose bill is apparently adapted for the task.

could be swallowed whole (Fig. 8.2) as well as larger soft fruits of native lobelioids and introduced fruit trees (Fig. 8.3). Other possible true drepanidine frugivores include the Lana‘i Hookbill, although James et al. (1989) thought not. When collected, the single specimen had been eating small ripe berries of opuhe, and Munro (1960), who collected it, speculated that the species might also formerly have fed on a then-destroyed forest of ‘akoko, which has similar-sized fruits. Likewise we can never know for certain the dietary habits of the ‘Ula-‘ai-hawane, whose name implies that it fed on fruits of native Pritchardia palms (Part II, Fig. 11.14). Most Pritchardia have fruits much too large (Wagner et al. 1990) to have been eaten whole by Ciridops, but the bird could have devoured them piecemeal in the same way that the ‘O‘u eats large fruits. The ‘Ula‘ai-hawane has a short version of the drepanidine nectar-adapted tongue (Bock 1972), and may have used it to extract juices from soft palm fruits (Pratt 2002a) in the same way other nectarivores do (Fig. 8.4). Hawaiian honeycreepers known to occasionally eat fruit comprise a disparate array that includes the aforementioned generalist Telespiza finches (many plant species, Morin and Conant 2002), Po‘o-uli (olapa, Mountainspring et al. 1990; kanawao, T. K. Pratt et al. 1997b), Kaua‘i ‘Amakihi (‘akia, Perkins 1903), Hawai‘i ‘Amakihi (‘ohelo kau la‘au, Fig. 8.4), ‘Akohekohe (olapa, Berlin and VanGelder 1999), and Maui Parrotbill. For the latter, Simon et al. (1997) state that the bird does not actually eat fruits (pilo, kolea, and kanawao) but ‘may test berries for larvae while still on the plant or, after plucking them, steadying them between toes and perch.’ However, my own observations in the Upper Hanawi area in the fall of 1995 seem to indicate otherwise. I observed parrotbills frequently plucking fruit of pilo (Fig. 8.5) and apparently eating the seeds and discarding the orange pericarp. I examined many of these and could see no evidence of the presence of insect larvae, nor did the birds appear to be selecting some fruits and leaving others. Birds worked along a heavily fruiting branch, eating essentially every ripe fruit. The ground below was littered with discarded pericarps. The Kona Grosbeak was never seen to feed in ‘ie‘ie, but

Ecology and breeding biology 131

8.2 Representative small fruits eaten whole by ‘O‘u and other honeycreepers: (a) pilo; (b) ‘olapa; (c) kanawao; (d) kolea lau nui.

scanning electron micrography revealed traces of ‘ie‘ie pollen on the birds’ feathers (Cox 1983); we do not know whether it ate the fruit.

Invertebrate predators All Hawaiian honeycreepers eat insects or other invertebrate prey. The fact that growing nestlings need animal food has long been appreciated, but some high-protein items are necessary to ‘balance’ the diet even of adult frugivores and nectarivores. In many cases, these birds prefer soft-bodied prey such as caterpillars over hard-shelled insects. For example, during outbreaks, looper caterpillars (Geometridae) are heavily exploited by ‘O‘u and ‘I‘iwi (Perkins 1903).A few drepanidines eat almost no vegetable matter as adults. We usually refer to such birds as insectivores, although their diets may include spiders, mites, centipedes, millipedes, snails,

and other non-insect prey. Hawaiian honeycreepers that are more or less predominantly ‘insectivorous’ include the ‘alauahios, Kakawahie,‘Anianiau,‘amakihis, ‘Akepa, ‘Akeke‘e, ‘Akikiki, Hawai‘i Creeper, Po‘o-uli, heterobills, ‘akialoas, and Maui Parrotbill. (The ‘amakihis, ‘akialoas, and ‘Anianiau also take large amounts of nectar.) With so many species exploiting the same general food resource, specialisations are inevitable. Niche differentiation is accomplished by an interplay of feeding techniques (Chapter 6), substrates, and prey choices.The spectrum of insectivory includes everything from very narrow specialists to broad generalists. The insectivore community on Maui provides an excellent example of fine-tuned niche partitioning. The Po‘o-uli may be the most specialised of the invertebrate predators as it is the only honeycreeper, as far as we know (but see James et al. 1989), that feeds primarily on land snails (Baldwin and Casey

132 The Hawaiian Honeycreepers

8.4 Hawai‘i ‘Amakihi piercing soft fruit of ‘ohelo kau la‘au to extract juices. Photo © Jack Jeffrey.

8.5 Maui Parrotbill plucking fruits of pilo. Photo © Jack Jeffrey.

8.3 Larger fruits eaten piecemeal by ‘O‘u and other honeycreepers: (a) haha‘aiakamanu; (b) guava, an introduced fruit; (c) ‘akala, the native raspberry.

1983; T. K. Pratt et al. 1997b). It also takes insects from bark surfaces and crevices but does not excavate tunnelling insect larvae. In the same habitat

and foraging sites, subsurface insects are preyed upon by the Maui Parrotbill, another specialist whose diet consists almost entirely of cerambycid and other wood-boring beetle larvae (Simon et al. 1997). On the same trunks and branches the Maui Nukupu‘u has foraging motions similar to those of the parrotbill (T. K. Pratt et al. 2001) but its weaker bill is unsuited for extracting cerambycid larvae and Perkins (1903) believed this prevented competition between the birds. Palmer (in Rothschild 1893–1900) found remains of adult beetles and other insects as well as ‘large grubs’ in stomachs of Maui Nukupu‘u. Maui ‘Alauahio join flocks (see below) with the above specialists and apparently glean the leftovers from their excavations, rather like

Ecology and breeding biology 133 jackals following lions. They also exploit a much wider variety of small insects gleaned from bark and leaf surfaces or even captured in flight (H. and P. Baker 2000). The Hawai‘i ‘Amakihi takes some of the same foliage and bark insects, but spends much more time in leaves (and feeding on nectar) than the ‘alauahio (Lindsey et al. 1998; pers. obs.). In prehuman times, at least one ‘akialoa ( James and Olson 1991) also belonged to the Maui bark-picking guild. Its long but weak bill would enable it to reach insects deep in crevices that the other birds could not exploit. Finally, the Maui ‘Akepa avoids competition with the other insectivores by exploiting larvae that live inside leaf buds (Lepson and Freed 1997). The other islands have similar niche partitioning among insectivores, but because each has a different complement of species, they are not directly comparable.Thus on The Big Island, which has no parrotbill equivalent, the wood-borer specialist is the ‘Akiapola‘au (T. K. Pratt et al. 2001), while on Kaua‘i the local ‘amakihi is a more dedicated bark forager and the ‘alauahio niche is essentially divided and expanded by the ‘Anianiau and ‘Akikiki (Lepson 1997, Foster et al. 2000). The Big Island had a smaller and a larger ‘akialoa and a much larger nukupu‘u ( James and Olson 2003.), plus the Hawai‘i Creeper, but had no known ecological equivalent of the Po‘o-uli. Kaua‘i and O‘ahu each had two ‘akialoas. These inter-island differences in the way feeding niches are partitioned obviously played a role in driving adaptive radiation by means of character displacement. A new coloniser could not simply slip into the niche it occupied on its parent island because that imaginary ‘ecological space’ had a different ‘shape’ on the new one.Variations on all themes were inevitable.

Leaf-eaters Many birds habitually forage among leaves but actual folivory (the eating of leaves) is very rare in passerines (Munson and Robinson 1992). Leaves are very tough and difficult to digest, and more often eaten by mammals than small birds. A few scattered observations indicate that some Hawaiian honeycreepers may occasionally eat leaves, but none do so predominantly. Perkins (1903: 434)

reported some ‘O’u stomachs were filled with ‘finely-divided green vegetable matter’, which Snetsinger et al. (1998) considered to be young leaves. Laysan finches occasionally eat leaves of several succulent shrubs as well as tree heliotrope (Newman 1988), and Nihoa Finches eat leaves of ‘aheahea and leaf buds of ‘ilima (Morin and Conant 2002), but these represent very small percentages of their omnivorous diets. Munro (in Olson 1999) reported green leaf bits in some stomachs of Kona Grosbeak, and van Riper (1980a) reported a small percentage of mamane leaves in the diet of the Palila. Another possible example is my observation of an ‘Akeke‘e nipping two small pieces from the edge of an ‘ohi‘a leaf, but I could not determine whether it ate them (Lepson and Pratt 1997). Interestingly, Darwin’s finches include one folivore, Platyspiza crassirostris (Grant 1986).

Sap suckers Several nectarivorous honeycreepers have been observed taking tree sap at natural fluxes, as does the endemic Kamehameha butterfly Vanessa tameamea (pers. obs.). No study has been done to determine the nutrient value of sap as compared to nectar. Uniquely, the ‘Akiapola‘au purposely exploits this resource even though it only rarely takes nectar (T. K. Pratt et al. 2001).Although it rarely forages for insects in ‘ohi‘a trees, these are the only ones exploited for sap. According to Jack Jeffrey (in T. K. Pratt et al. 2001), the bird hammers a 2–4mm deep hole in the cambium, then repeats about 1 cm away, producing a horizontal line of holes (Fig. 8.6b) that look very much like sap wells of ‘true’ sapsuckers (pers. obs.). Each well produces only a few drops of sap before flow stops, so the birds must reopen them on repeat visits. Certain trees are favoured as sap sources, and may be thoroughly riddled with holes (Fig. 8.6a). Other honeycreepers, especially Hawai‘i ‘Amakihi, often feed at the sap wells, but do not create them. ( J. Jeffrey, pers. comm.).

Eclectic diets Quite a few honeycreepers divide their attentions among several food categories so that none predominates. Such birds are often thought of as

134 The Hawaiian Honeycreepers

8.7 A park-tame panhandling Hawai’i ‘Amakihi attempting to eat “inappropriate” food, in this case dried coconut from trail mix.

8.6 ‘Ohia trees scarred with sap wells dug by ‘Akiapola‘au: (a) heavily riddled large tree, photo © Jaan K. Lepson; (b) close-up of rows of sap wells, photo © Jack Jeffrey.

generalists or omnivores, but that does not mean that their diets are not species-specific. The percentages of foods vary widely, and some are probably better thought of as dualists. For example, Perkins (1903) considered ‘akialoas to be nectarivorous and insectivorous in about equal proportions, and the ‘Anianiau likewise has a divided diet of insects and nectar (Lepson 1997). In addition to having a warbler-like bill (Fig. 6.7c), the Maui ‘Alauahio resembles some wood warblers (Parulidae), such as the Cape May Warbler, in their diets of foliage and bark insects supplemented by nectar (pers. obs.). But true omnivores have even more catholic tastes. The ‘amakihis are quintessential generalists, so much so that I have jokingly called them universalists (Pratt 1999b). They all eat a combination of arthropods, fruit, tree sap, and nectar. They apparently have no particular preferences when it comes

to nectar, and do not disdain introduced flowers; O‘ahu ‘Amakihi commonly feed in eucalyptus and other alien trees (Lindsey et al. 1998; Pratt 2002b). ‘Amakihi also exploit a variety of native and introduced fruits by piercing them and sucking out the juices and pulp rather than by swallowing them whole (Baldwin 1953).As insectivores, they are frequent leaf-gleaners and bark-pickers (Lindsey et al. 1998). No broad generalisations are possible because even within species, feeding methods vary with habitat. For example, Ralph and Noon (1988) studied a population of Hawai‘i ‘Amakihi in ‘ohi‘a forest on Hawai‘i and found that the birds fed by 33% foliage-gleaning, 31% flower-probing, 15% foliage-probing, 6% bark-picking, and 3% fruitpiercing. In a mixed ‘ohi‘a and koa forest on Maui, Lindsey et al. (1998) report 50% nectar-feeding, 39% gleaning of all types, and 11% fruit-eating. In subalpine shrubland on Maui, the same authors report 67% gleaning, 31% nectar-feeding, and 1% fruit-eating. Some ‘amakihi even attempt to eat foods that are clearly unsuitable for them (Fig. 8.7). This bird and its apparent family group also consumed potato chips, bread crumbs, soft drinks, and other garbage obtained by panhandling from picnickers, and even entered parked cars to search for titbits (Pratt 1999b). Apparently some ‘amakihi like junk food as much as people do. The Laysan and Nihoa finches are the champion omnivores among Hawaiian honeycreepers (see species accounts for details). The fact that they can

Ecology and breeding biology 135 quite literally eat whatever is available enabled the Laysan Finch to survive the early twentieth-century rabbit plague (Chapter 2; Pratt 1994a) when all vegetation on the island disappeared for a time. As summarised by Morin and Conant 2002, Laysan Finches consume ‘seeds, fruits, leaves, flowers, stems, seedlings, roots, carrion, invertebrates, and eggs’, probably the broadest menu of any Hawaiian honeycreeper. Nihoa Finches are less well known and inhabit a smaller island with a different plant community, but their diet is very similar and includes insects, flowers, fruits, seeds, leaf buds, leaves, and eggs (Morin and Conant 2002). None of these items (i.e. insects, seeds, fruits) predominate. Interestingly, the one food missing from these lists is nectar!

Nectarivores1 The honeycreepers’ namesake nectarivores, like the insectivores, include many variations on the theme, from broad generalists that feed on any available nectar to birds that specialise in particular flowers. However, the community of nectar-feeders differs strikingly from all others among Hawaiian honeycreepers in the degree of overlap of feeding niches, apparently because there are only so many ways to eat nectar and it is a super-abundant, though often patchy or irregular, resource (Carpenter and MacMillen 1976b; Carpenter 1978).The most important source of nectar in Hawaiian forests are lehua blossoms, the flowers of the ‘ohi‘a tree. They are open flowers lacking petals, but with the calyx forming a cup, in which nectar collects, at the base of the red (sometimes yellow or orange) stamens and pistils (Fig. 8.8). Pollination is mostly by Hawaiian honeycreepers (Carpenter 1976). ‘Ohi‘alehua blooms year-round, but blooming is periodic and irregular, depending on climatic and other factors, in any given locality (Scott et al. 1986). Almost all Hawaiian honeycreepers, including nectarivores with bills adapted to other flower types and even such dedicated insectivores as ‘Akikiki (Conant et al. 1998) that lack a nectar-adapted tongue, sample the sweetness of lehua at least occasionally, and some depend heavily on it. The ‘Apapane and ‘I‘iwi spend 80–90% of their foraging time in ‘ohi‘a in some localities (Fancy and

8.8 ‘Ohi‘a-lehua, the staff of life for Hawaiian nectarivores: (a) brush-like stamens and pistils and calyx cups (seen in disintegrated flower, lower right) in which nectar collects; (b) ‘Akohekohe sipping from the chalice. Photo (b) © Jack Jeffrey.

Ralph 1997, 1998) and the diet of the ‘Akohekohe is as much as 75% ‘ohi‘a nectar (VanGelder 1996; Berlin and VanGelder 1999; Berlin et al. 2001b). These three species also forage seasonally in other native flowers (Fig. 8.9) including koa, naio, mamane, akala, kanawao, tree ‘ohelo, native Hibiscus, and alani (Fancy and Ralph 1997, 1998; Berlin and VanGelder 1999; Berlin et al. 2001b).They are also attracted to several introduced plants including tagasaste (tree alfalfa) (Waring et al. 1994) and the invasive banana poka (Fancy and Ralph 1998; pers. obs.). ‘Apapane were even reported to feed in coconut flowers along the coast in earlier times (Perkins 1903). The closely related Laysan Honeycreeper had no lehua blossoms on which to feed, but early naturalists reported it feeding on maiapilo (Fig. 8.10), whose white flowers bear a striking

136 The Hawaiian Honeycreepers

8.10 White flower of maiapilo, whose shaving-brush form somewhat resembles that of lehua blossoms (Figure 8.8).

8.9 Open-faced or short tubular flowers exploited for nectar by Hawaiian honeycreepers: (a) ‘akala, a favourite of ‘I‘iwi; (b) kanawao being visited by Kaua‘i ‘Amakihi; (c) Hawai‘i ‘Amakihi feeding on inconspicuous dangling flowers of ‘ohelo; (d) ‘Anianiau on full flower cluster of kanawao. Photos © Jack Jeffrey.

morphological resemblance to those of ‘ohi‘a, but it did not specialise on such flowers.All authors mention the honeycreeper’s feeding on other open flowers such as ihi and sea purslane, and the bird survived for several years after maiapilo was eliminated from Laysan by rabbits (Ely and Clapp 1973). Other honeycreepers with a substantial, though not dominant, proportion of nectar from a wide range of flowers in their diets include the ‘amakihis (Lindsey et al. 1998) and ‘Anianiau (Lepson 1997), discussed above under ‘Eclectic diets’. Considering the variety of native and introduced flowers at which the unspecialised drepanidine nectarivores have been observed to feed, we can reasonably predict that they will take nectar wherever it is available. The Hawai‘i ‘Amakihi will even accept artificial nectar feeders in the wild (van Riper 1984) and many species do so in captivity (A. Lieberman, pers. comm.). The two mamos, in contrast to the broad tastes of other nectarivores, fed preferentially on ornithophilous flowers of shrubs belonging to the genera Cyanea and Clermontia (Fig. 8.11) and possibly Delissea (Pratt 2002a). So attracted was the Hawai‘i Mamo to such flowers that one method of capture used by early bird-catchers was to lie concealed beneath the plant with a hand-held flower, then seize the bird when it inserted its bill (Townsend 1839; Perkins 1903). The relatively few specimens taken of the Black Mamo had no remains of insects

Ecology and breeding biology 137

8.11 Ornithophilous flowers of Hawaiian lobelioids: (a) haha‘aiakamanu; (b) haha; (c) ‘oha; (d) ‘oha kepau.

in their stomachs (Perkins 1903), indicating their near total devotion to nectar. The ‘Ula-‘ai-hawane may also have been a specialised nectarivore (Pratt 2002a) rather than a frugivore as hypothesised above. Because its tongue is nectar-adapted (Bock 1972), it may have fed on the nectar of loulu palm flowers (Part II, Fig. 11.14), perhaps as a Hawaiian analogue of small-billed, brush-tongued lorikeets of other Pacific islands (Pratt 2002a). The ‘akialoas are most often thought of as insectivores, but Perkins mentioned particularly their fondness for nectar of certain species of arborescent lobelioids called haha, especially ‘those with large corollas’. He wrote (Perkins 1903: 422–3): In the heart of the forest on Mauna Kea, during three weeks of continuous rain in the winter months I daily observed the Akialoa visiting these flowers; as also on the rough lava-flows in the Kona district on the other

side of the island. H. procerus on Kauai likewise obtains nectar from the same remarkable flowers, and is likewise fastidious in the choice of species that it frequents, some of the most abundant Lobeliaceae (even when well-liked by other honey-suckers) being rarely, if ever, visited. At times, even when the Akialoa was feeding on insects, and in places where I noticed no flowering lobelias, I found the forehead to bear more or less evident traces of the pollen of these flowers, showing that they must have lately visited these. Wolstenholme himself informed me that he shot the Lanai Akialoa, while they were seeking the deepseated nectar of the Haha. At other times both the common species resort to the flowers of the Ohia with their exposed nectaries. In Kona H. obscurus was common in the very tallest flowering trees, frequently two to eight in a single tree, but

138 The Hawaiian Honeycreepers requiring to be looked for amongst the extraordinary numbers of commoner Drepanids. In some localities and at certain seasons the Kauai bird is an eager visitant to the Ohia flowers. One of the best localities known to me for entomological work was an open dry slope . . . In the middle of the open space grew several low and bushy Ohia trees, at that season covered with flowers, although those in the surrounding forest bore few or none. To these isolated trees the Akialoa would come at intervals throughout the day, sometimes a number of individuals being present at one and the same time.Yet in the surrounding forest the birds appeared to be always hunting for insects on the branches of the trees, and were not seen to pay any attention to such flowers as were to be found . . . I am quite satisfied that the different kinds of Akialoa are all still largely nectar-eaters, although possibly on the way to become entirely insectivorous. Nectar-feeding is very rare but not altogether unknown among the heterobills (see below), especially the nukupu‘us. Perkins (1903) reported that the Maui Nukupu‘u takes nectar from ‘ohi‘a flowers, and Wilson’s Hawaiian assistant assured him that the Kaua‘i Nukupu‘u ate bananas and oranges, which Perkins, probably correctly, assumed meant the nectar of their flowers inasmuch as similar feeding on banana flowers had been reported by Deppe and Townsend for the O‘ahu Nukupu‘u (Perkins 1903). Despite these reports, most authors have regarded the subgenus Hemignathus as almost entirely insectivorous. Even the most devoted vegetarian honeycreepers feed invertebrates to their nestlings and begging young, presumably because the protein is necessary for rapid growth and development. Nectar is particularly deficient in amino acids (Baker and Baker 1973, 1975). Some juvenile nectarivores apparently have specialised intermediate diets before adopting that of their parents. Carothers (2001) found that 3-month-old ‘Apapane and ‘Akohekohe on Maui had a significantly higher proportion of invertebrates in their diet than adults, but converted to adult proportions by the age of 9 months. These

differences are related to differing metabolic requirements rather than to interference competition from adults.

Special aspects of nectarivory Feeding territories and competition The nectar-feeding guild presents a very different ecological picture when compared to insectivorous honeycreepers. In most years, nectar is so abundant that its members can co-exist amicably at food sources with both conspecifics and other nectarivores (Carpenter 1978). Perkins (1903) reported a ‘birder’s dream’ of a single ‘ohi‘a tree on Moloka‘i in which Bishop’s ‘O‘o, Black Mamo,‘Akohekohe, and ‘Apapane fed simultaneously. But nectar resources are more subject to vagaries of weather and climatological events (Lindsey et al. 1997) than are insects (Ralph and Noon 1988), and in seasons of scarcity, the nectarivores engage in what amounts to niche partitioning, but it is a rather different phenomenon from that exhibited by the insectivores (Pimm 1991). It is in ‘nectar famines’ that the specialised members of the guild gain their evolutionary advantage because they then have a small but consistently available resource that others cannot exploit (Wolf and Gill 1980; Pimm and Pimm 1982). Others establish feeding territories that lead to a size-correlated dominance hierarchy (Pimm and Pimm 1982; Carothers 1986a). We know that the now extinct Hawaiian meliphagids (all of which were larger than the nectarivorous dreps) were at the top of this hierarchy (Conant et al. 1998; Sykes et al. 2000), and that on Moloka‘i the Black Mamo was the dominant (and largest) honeycreeper (Perkins 1903). We do not know where the ‘akialoas fit into this scheme, but presumably it varied from island to island depending on where the island’s ‘akialoa fell within the size array.Today, only the ‘I‘iwi (Fancy and Ralph 1998) and ‘Akohekohe (Berlin and VanGelder 1999) survive to establish feeding territories.The lower members of the hierarchy have behavioural ways of dealing with territorial dominants: the ‘Apapane simply overwhelms a defender by foraging in flocks and the Hawai‘i ‘Amakihi employs stealth (Carothers 1986b). The subordinates are nevertheless relegated

Ecology and breeding biology 139 to areas of reduced resource availability (Carpenter 1978) and ‘amakihi may also shift to the insectivorous part of their varied feeding repertoire during times of competition for nectar (Carothers 1986a, b; Carpenter 1987). Carothers (1986a) found that ‘Akohekohe and ‘I‘iwi were much more aggressive toward members of their own species than toward other species. Both maintain strongly defended feeding territories outside the breeding season from which both conspecifics and other species are chased (Carothers 1986b). As predicted by Wolf and Gill (1980), the structure of the Hawaiian nectarivorous community is qualitatively very similar to that exhibited by others such as hummingbirds and sunbirds.

Trap-lining Birders have long been aware that both the ‘Akohekohe and ‘I‘iwi sometimes engage in a foraging method called ‘trap-line feeding’ because of its resemblance to a fur trapper’s running of a line of traps. It occurs when nectar resources are scarce and scattered in an area.An individual bird sets up a ‘trapline’ of foraging sites that it visits on a regular cycle. These birds presumably are not defending feeding territories because the resources individually are not worth the effort. In Waikamoi Preserve on Maui, a knowledge of this behaviour is often the key to repeated observations (Pratt 1993, 2002b). Based on numerous visits to the area with groups of birders, I have found that the ‘periodicity’ for ‘Akohekohe is between 20 and 30 minutes. When a feeding bird departs, an observer simply has to wait a half hour at the same blooming ‘ohi‘a tree and the bird will return to the spot. I have seen ‘Akohekohe feeding only in ‘ohi‘a in Waikamoi Preserve. Apparently, the tree requires that amount of time to replenish its nectaries, but I know of no direct data on the subject.‘I‘iwi may also engage in trap-lining under similar circumstances (Carpenter and MacMillen 1975, 1980), as in a narrow tongue of habitat such as Waikamoi Gulch behind Hosmer Grove in Haleakala NP (Pratt 2002), but birders have paid less attention to its patterns and no periodicity has been established. It may well be different from that of the ‘Akohekohe, because ‘I‘iwi feed on a wider variety of flowers that may have varied ‘recycling’ times.

Some of the observations of Perkins (1903) suggest that the mamos may have also been trap-liners of widely scattered Clermontia trees.

Nectar-stealing 2 The behaviour pattern in which a pollinator takes nectar from a flower in the ‘wrong’ way and therefore avoids pollination is called nectar-stealing. Taking nectar from a flower adapted for bird pollination (see below) in such a way as to avoid pollination constitutes nectar-stealing. The earliest report of such feeding by any Hawaiian honeycreeper is that of Henshaw (1902: 53) who said that the ‘I‘iwi takes nectar mainly from ‘ohi‘a but: does not disdain to rifle it from other flowers, both wild and cultivated. Nasturtiums and canna are especial favorites with it, and any one who is fortunate enough to live where the bird is common may attract numbers close to the house by planting the above named flowers. The birds will soon learn to visit the flower beds daily, especially in early morning and at eventide. Unable to reach the nectaries from the mouth of the nasturtium flower, even with their long bills, the iiwi has learned to pierce the spur of the blossom just above the coveted honey, and the brush tipped tongue enables them to scoop out the sweet drops and leave not a trace behind. ‘I‘iwi obviously are quick studies, and their recent seeming addiction to banana poka flowers (Fancy and Ralph 1998; pers. obs.), which have long straight corolla tubes (Fig. 8.12), may be spoiling them for their role as pollinators. Conant et al. (1998) speculate that such nectar-stealing is recently learned and is a danger to native plants that depend on the ‘I‘iwi for pollination (see also Chapter 10). On the other hand, banana poka need not be the culprit. ‘Akohekohe steal nectar from native lobelioids and mints Stenogyne spp. (VanGelder 1996; Berlin and VanGelder 1999) in the same manner, and VanderWerf and Rohrer (1996) report that ‘I‘iwi steal nectar from open-faced flowers of the native hibiscus kokio ke‘oke‘o (Fig. 8.14), so at least in

140 The Hawaiian Honeycreepers

8.13 Kaua‘i ‘Amakihi stealing nectar from ornithophilous Kaua‘i koli‘i. Compare with ‘correct’ behaviour of ‘I‘iwi in Figure 6.12. Photo © Jack Jeffrey.

8.12 Flower of banana poka, an invasive introduced passion flower with straight, tubular corolla.

those cases the behaviour may not be a recent development. However, the latter example may be related to the recent ecological shift to non-tubular flowers by ‘I‘iwi (see below). Kaua‘i ‘Amakihi (Conant et al. 1998) and ‘Anianiau (Lepson 1997) may also pierce long corollas at the base (Fig. 8.13), or possibly use holes already made by ‘I‘iwi to steal nectar. I have observed apparent nectar-stealing by ‘Apapane from banana poka flowers, but the birds may have also been taking advantage of ‘I‘iwi piercings rather than making their own holes. I find that most flowers of ohe naupaka (Fig. 8.15c) are pierced at the base.This phenomenon has also been reported among supposedly co-evolved honeyeaters in Australia, and there, too, the malevolent influence of introduced plants on the birds’ behaviour is suspected (McCulloch 1977). Inouye (2001) sees nothing sinister in

8.14 Flowers of kokio ke‘oke‘o, a Hawaiian hibiscus: (a) typical form adapted for insect pollination; (b) early-stage development of ornithophily in variety punaluuensis.

Ecology and breeding biology 141 nectar-stealing, however, and believes such behaviour may in fact benefit the plants by forcing the co-evolved pollinators (see below) to make more frequent visits to the flowers. In the case of nectarstealing by ‘I‘iwi from native Hibiscus, it is difficult to see how the bird could take nectar from these flowers, which, unlike close relatives Kokio and Hibiscadelphus (Fig. 8.17), have no ornithophilous adaptations, any other way than by piercing the corolla tube at the base. These flowers appear adapted for insect pollinators such as hover-feeding hawk moths, but one variety, at least, has the beginnings of adaptations to force ‘I‘iwi to pollinate when it takes nectar (Fig. 8.14).

Plant/pollinator co-evolution Numerous popular writers on Hawaiian natural history have pointed to the apparent connection between the curved bills of some Hawaiian honeycreepers and the curved corollas of certain endemic plants, particularly the lobelioids (Campanulaceae). Both the birds and the plants involved belong to families in which these supposedly coevolved features are novel, i.e. ancestral finches did not have curved bills and mainland lobelioids do not have curved flowers. In many (but by no means all) cases the flower and bill fit with keyin-lock precision, with the pistils positioned strategically for bird pollination (Spieth 1966). Superficially, this bird/plant evolutionary interplay seems so obvious and compelling that it has become a cliche of biological folklore without anyone really questioning whether it actually happened the way it appears, or analysed how the apparent co-evolution occurred.Thus, the claim of Fleischer and McIntosh (2001) that the age of evolutionary divergence of the nectarivorous honeycreepers (2–3 Ma) is so incompatible with the 8–17 Ma age of the ornithophilous lobelioids (Givnish 1998; Givnish et al. 1995) that ‘it is highly unlikely that drepanidines “coevolved” with these plants in the islands’ comes as something of a surprise. They suggested instead that Hawaiian meliphagids ‘could be the coevolved taxon’, a statement that seems to contradict the obvious. (None of the five known meliphagid species have

sickle-shaped bills, nor are any known from subfossils, so how could they be co-evolved with sickle-shaped flowers?) In a study of two lobelioids on O‘ahu, Cory (1984; as reported in Buchmann and Nabhan 1996) observed no feeding at the flowers by honeycreepers and thus concluded that no co-evolution could have occurred. Of course, this conclusion ignores the important fact that all of the sickle-billed honeycreepers on O‘ahu were by that time either extinct or very rare. Other objections to possible co-evolution of lobelioids and honeycreepers were raised by Lammers and Freeman (1986), who considered the question unresolved. Again, their study considered only the lobelioids. The following review shows that the circumstantial evidence that Hawaiian honeycreepers and Hawaiian plants do, indeed, present a spectacular example of co-evolution is overwhelming, though the exact mechanisms involved remain controversial. The honeycreepers involved would obviously be those with sickle-shaped or curved bills: mamos, ‘I‘iwi,‘amakihis,‘akialoas, and possibly one or two of the prehistoric gapers (nectar-feeders are enigmatically under-represented among subfossil remains; James and Olson 1991). Possibly because early writers emphasised so strongly the relationship between the Hawai‘i Mamo and certain Clermontia flowers, other possibly co-evolved plant taxa were largely overlooked as in the above cited studies. Certainly from the semipopular literature (e.g. Carlquist 1970, 1974; Frierson 1991; Buchmann and Nabhan 1996) one might gather that Hawaiian lobelioids are the only plants co-evolved with bird pollinators. However, even a cursory survey reveals that ornithophily of flowers in groups that are not normally birdpollinated is a widespread and obvious feature of the Hawaiian flora (Carlquist 1970; Pratt 1999a). Members of the genus Scaevola, such as the widespread Pacific beach plant known in Hawai‘i as beach naupaka, typically have small, rather open, asymmetrical white flowers. Most endemic upland Hawaiian species show an increasing tendency to tubularity but are not bird-adapted. However, one species, ohe naupaka, is strikingly different from its congeners in having large, waxy, yellow, curved corollas almost exactly the size and shape of the

142 The Hawaiian Honeycreepers bill of the ‘I‘iwi (Patterson 1995; Fig. 8.15) and superficially closely resembling flowers of Clermontia (Fig. 8.11). Likewise, Hawaiian geraniums typically

8.15 Varying degrees of ornithophily in flowers of Scaevola: (a) short, open flowers of coastal naupaka kahakai showing ‘half-flower’ configuration characteristic of the genus; (b) flowers of montane huahekili ‘uka with longer, but straight, corolla tube; (c) flowers of ‘ohe naupaka with falcate ornithophilous flowers which nevertheless maintain generic pattern at distal end.

have broadly open white flowers pollinated by insects, but one species, nohoanu, has partly tubular, curved red corollas designed, apparently, for pollination by honeycreepers (Carlquist 1970;Wagner et al. 1990). Typically, mints (Lamiaceae), including several Hawaiian species, have straight corolla tubes and no particularly ornithophilous features. However, the endemic genus Stenogyne (Fig. 8.16) includes several species with adaptations (small lower lip, exserted stamens, abundant odourless nectar, and sickle-shaped corolla tubes) for bird pollination (Wagner et al. 1990). Both ‘I‘iwi and Hawai‘i ‘Amakihi feed on them, with an excellent ‘fit’ of flower to bill (Fig. 8.16c). Members of the hibiscus family (Malvaceae) are generally characterised by open flowers with broadly spreading, often showy, petals. Hawaiian hibiscus have undergone a minor radiation that has produced two endemic genera with very distinctive characteristics. In hau hele ‘ula (Fig. 8.17a), the flowers are not only bright red like many bird-pollinated flowers (Raven 1972), but also the petals are rolled into a short, curved tube and the staminal column is likewise curved. In Hibiscadelphus, the adaptations for pollination by curve-billed birds are even more pronounced, although the flowers are less colourful (Fig. 8.17b). Finally, several examples come from the pea family (Fabaceae), whose familiar asymmetrical flowers (adapted for bee pollination) can hardly be thought of as long and curved. The rare Hawaiian vetch has unusually large, showy flowers with a distinct curve that are clearly adapted for bird pollination (possibly by ‘I‘iwi or Hawai‘i ‘Amakihi;Wagner et al. 1990). A vine called nuku ‘i‘iwi (‘‘I‘iwi beak’) was figured in Frohawk’s illustration of the ‘I‘iwi for Wilson and Evans (1890–99; see also Wilson and Buff 1989), perhaps in an artistic effort to make a point about co-evolution. A common dry-forest tree, the native coralbean called wiliwili, has those petals known as standards modified and curved (Fig. 8.18a) so that they guide the bills of such birds as ‘amakihis to promote contact with the prominently exserted stamens. Even the more typical pealike flowers of mamane (Fig. 8.18b, c) exhibit a short curvature that ‘fits’ the bill of pollinating ‘amakihi and ‘I‘iwi. Unfortunately, lineage ages for the non-lobelioid co-evolved plants are unavailable, but even if the ages

Ecology and breeding biology 143 for lobelioids and honeycreepers do not coincide precisely, apparent examples of co-evolution leading

8.16 Flowers of Stenogyne, a Hawaiian genus of the mint family (Lamiaceae): (a) S. microphylla, a dry forest species with no ornithophilous adaptations (corolla straight); (b) S. scrophularioides, a larger-flowered species with falcate corollas; (c) ‘I‘iwi showing acrobatic movements to access nectar and pollinate flower of S. calaminthoides. Photo © Jack Jeffrey.

to falcate flowers are so common in the Hawaiian Islands that surely some of them must have involved honeycreepers because no other viable birdpollinator candidates exist. Pimm (1991) and Cox and Elmqvist (2000) believe the reason a disproportionate number of ornithophilous plants are rare in Hawaii is the extinction of co-adapted avian pollinators, and Diamond (1984b) directly attributes the near extinction of all species of Hibiscadelphus to the loss of honeycreeper pollinators. Other birdpollinated but not co-evolved plants, such as the indigenous ‘ie‘ie, widely distributed in southeastern Polynesia (Wagner et al. 1990), now depend on introduced species such as the Japanese White-eye for pollination originally performed by honeycreepers and other now-absent native birds (Cox 1983; Cox and Elmqvist 2000).

8.17 Varying ornithophily in flowers of Hawaiian Malvaceae (hibiscus family): (a) hau hele ‘ula with splayed petals, shorter corolla tube, and curved staminal column; (b) more tightly tubular and curved corollas of hau kuahiwi.

144 The Hawaiian Honeycreepers

8.18 Ornithophily in flowers of Hawaiian Fabaceae (pea family): (a) ornithophilous flowers of wiliwili, the native coralbean; (b), (c) short but curved flowers of mamane pollinated by Hawai‘i ‘Amakihi and ‘I‘iwi, respectively. Photos (b) and (c) © Jack Jeffrey.

But can we dismiss the drepanidine–lobelioid association? The estimates of age for the lobelioid lineage ‘are admittedly crude but are the first such estimates for a Hawaiian plant group and among

the very few for any plant group’ (Givnish et al. 1995). I am not ready to make such a sweeping statement as Fleischer and McIntosh (2001) on the basis of these pioneering estimates. But even if Cyanea did have ‘8.7 to 17.4 million years to coevolve with its avian pollinators, frugivores, and . . . herbivores’ (Givnish et al. 1995), nothing in these studies can tell us when its ornithophilous adaptations first appeared. None of the present-day birdadapted species are older than the islands they inhabit, and they exhibit a conveyor-belt pattern of speciation from older islands to younger ones (Givnish et al. 1995) that parallels those of the honeycreepers (Fleischer et al. 1998). The possibility that the lineage split from its ancestor earlier than the drepanidines separated from the carduelines has nothing to do with whether it may have co-evolved with the honeycreepers following the birds’ arrival. Other examples of co-evolved flowers, such as ‘ohe naupaka and nohoanu mentioned above, are the only members of their genera to be so adapted. Obviously, ornithophily can develop later in a lineage once appropriate birds are present, and need not be present at the start. The lobelioid genus Clermontia (more often cited as an example of bird/plant co-evolution) has no problems of timing. It is a relatively young lineage that appears to have evolved first on The Big Island and colonised the older ones in a rare reversal of the usual pattern (Lammers 1995). The genus is clearly much younger than the honeycreepers and it has all of the expected adaptations for ornithophily, including nectar with the distinctive sugar composition (Lammers and Freeman 1986), characteristic of bird-pollinated flowers (Baker and Baker 1983). Furthermore, Clermontia pollen has been found on the foreheads of century-old skins of the ‘I‘iwi (Lammers 1995) and was reported on collected specimens of ‘akialoa (Perkins 1903, quoted above). Even more telling is the recent discovery that I‘iwi bills are shorter today than they were a century ago, apparently as a result of a shift to feeding primarily on ‘ohi‘a and away from the increasingly rare lobelioids and other ornithophilous flowers (Smith et al. 1995;Thompson 1996; but see Freed et al. 1996 and Winker 1996). Although this shift certainly seems to have occurred, the reasons for it may have

Ecology and breeding biology 145 been exactly the opposite: the bird-adapted flowers declined because they were no longer being pollinated by sickle-billed birds (Cox 1983; Pimm 1991; Hopper et al. 1996; Cox and Elmqvist 2000). Parker (1977) suggests a similar connection between bird extinctions and scarcity of certain plants in New Zealand. Despite extinctions of mamos and ‘akialoas, the ‘I‘iwi is still common enough on most islands to do the job of pollination. However, possibly as a result of learning to steal nectar (as discussed above) from introduced flowers such as banana poka with long straight corollas (Conant et al. 1998), the birds have in many cases forsaken their oftenreported (Perkins 1903; Spieth 1966) co-evolved feeding methods and begun to pierce the falcate corollas at their base (Engilis 1990; Conant et al. 1998; Fancy and Ralph 1998; pers. obs.), circumventing the ‘return favour’ of pollination. Thus, though the birds still feed at the flowers, the chain of mutualism may be broken. Determining the processes by which such coevolution proceeds begins with a classic chickenand-egg question:Was it the birds or the plants that started the trend toward falcate shapes of bills and flowers? We may never know the answer, but some recent findings from research on hummingbirds (Trochilidae) suggests that the flowers actually issued the invitation (Temeles and Ewald 1999) to the ‘intricate dance of flower and bird adapting together’ (Frierson 1991: 73). However begun, the synergism led inexorably to the outer limits of adaptability in such species as the mamos and ‘akialoas (bills and corollas cannot increase in length indefinitely).The selective forces driving this adaptational interplay have not been investigated in Hawaiian communities, but Ziegler (2002) offers a hypothesis built upon comparisons with bird/plant co-evolution in hummingbirds (Grant and Temeles 1992;Temeles and Roberts 1993;Temeles 1996). In this scenario, co-evolution is driven by competition within both the bird and plant communities. Suppose that two morphologically similar nectarivorous birds find themselves in competition on an island. By chance alone, one species will inevitably have a slightly longer average bill length than the other, and the individuals with the longest bills will have a feeding advantage at flowers that are slightly

longer. Selection would then favour individuals in both species that select flowers with characteristics that promote optimal foraging efficiency, and that in turn would progressively emphasise their differences. At the same time, two morphologically similar plant species on the same island are in competition for pollinators, with an important advantage being the possession of a fail-safe pollinator, i.e. one that might visit other flowers but which would always visit those of the plant in question. This example is for bill and corolla length, but could apply equally well to shape (i.e. curvature) or flower colour. Eventually, an equilibrium is reached in which further changes do not occur. Recently, a new twist on the paradigm has been suggested by Temeles and Ewald (1999).Their hummingbird research involving bill and corolla curvature suggests that, in contrast to the mutualistically beneficial model proposed above, birds and plants ‘have somewhat conflicting interests’. They found that the optimal condition for the plant is a slight mismatch with the bird’s bill (which promotes pollination by physically disturbing the flower), whereas the birds gain most from a precise fit that promotes feeding efficiency.They refer to the result as an ‘evolutionary arms race’, an idea that originated with Charles Darwin and that may apply equally well to the drepanidine example. Selection would first favour flower curvature to promote pollination, then bill curvature would be selected to increase efficiency, which in turn would lead to selection for greater curvature (to regain the pollination advantage), and so on. Ultimately, the most curved corolla in the community should be more curved than the most falcate bill (Temeles and Ewald 1999). Whether this prediction holds for Hawaiian communities is fertile ground for investigation.The most obvious first questions are whether Hawaiian flowers gain the same pollination advantage from a slight mismatch of shape, and whether foraging methods of honeycreepers are really comparable to the hover-feeding of hummingbirds. Such studies could help answer the chicken-andegg dilemma posed earlier. One might suppose that the plants in the above model might be leading the birds down the garden path to overspecialisation. In fact, just the opposite is true.The longer-billed birds

146 The Hawaiian Honeycreepers can always feed on shorter flowers, and surprisingly may even do so more efficiently than birds whose bills are of a more appropriate length (Carothers 1982), but they will always have a certain number of flowers that only they can exploit.

Interspecific competition As the above discussions show, natural communities tend to structure themselves to avoid competition between species. The debate over how this happens is still ongoing (Pimm 1991), but Hawaiian honeycreepers seem to confirm the hypothesis, first articulated by Lack (1947) in reference to Darwin’s finches, that character displacement in morphological features underlies the observed community structure. Even though a particular guild may have many members on a given island, they differentiate behaviourally and trophically so that niches do not overlap significantly (Conant 1981a).This differentiation among Hawaiian honeycreepers has led to an unusually large proportion of specialists (Ralph 1990a), but which species are considered specialists and which generalists depends on one’s definition. In a study of Big Island communities that included six honeycreepers, Ralph and Noon (1988) found no ‘common theme connecting the various species and their adaptations to their environment.’ As stated above, the nectarivores show the greatest degree of overlap based on food resource, but use competition in the form of dominance and territoriality to subdivide the niche (Pimm and Pimm 1982). However, that sort of competition does not produce negative distributional correlations (Mountainspring and Scott 1985). Bock (1970) suggested that morphological differentiation among the insectivores might have been driven by past competition during resource shortages, but no such direct competition is evident today (Mountainspring and Scott 1985). Freed (1999) defines niches somewhat differently by recognising eight ‘ecomorphs’ among Hawaiian honeycreepers: frugivore (‘O‘u), creeper (Paroreomyza/Oreomystis), ‘akepa, ‘amakihi, heterobill, ‘akialoa, short-billed nectarivore (‘Apapane), and long-billed nectarivore (‘I‘iwi). Each large island has at least one species of each ecomorph,

but the communities of each island are unique not only because the species may differ within ecomorphs (as in creepers, ‘akepas, ‘akialoas, and heterobills), but each island has its own specialities that may or may not fit into one of these categories (e.g. Maui Parrotbill).This differentiation allows all ecomorphs to co-exist in the forest community. Whether Hawai‘i’s many introduced bird species compete directly with honeycreepers has been the subject of considerable speculation. Mountainspring and Scott (1985) suggest that the unusually high success rate for alien species in Hawai‘i as compared to other islands, where aliens are often unsuccessful because of competition with residents (Lack 1976), is the result of the decimation of the native lowland avifauna by Polynesians (Olson and James 1982a, 1984; Olson 1989a) and the subsequent exclusion of native birds from the lowlands by avian diseases (see Chapter 9). Mountainspring and Scott (1985) found that most of the introduced species in the community fill niches unoccupied by honeycreepers, and do not compete directly with the dreps.The Japanese Whiteeye, however, exhibits limited and sporadic negative distributional correlations with both ‘I‘iwi and ‘amakihis, and plays a ‘potent’ role in reducing native bird populations.When the native/alien community is viewed as a whole, a significant amount of diffuse competition is apparent that does not show up in species-by-species analyses.As a result of a ‘combination of direct and diffuse competition’, Mountainspring and Scott (1985: 233, 235) conclude that ‘competition with exotic species must be included among the many factors causing the decline and extinction of native Hawaiian birds.’

Daily and seasonal movements Hawaiian honeycreepers do not migrate, but early naturalists used the term to refer to altitudinal movements. Though none are apparent today, whether any of these elevational shifts represented regular seasonal movements in earlier times cannot be determined. Observers such as Perkins (1903), Henshaw (1902), and Munro (1960) refer to such movements, which ‘were especially prominent during the early historical period when the native bird populations were still more or less intact’ (Warner

Ecology and breeding biology 147 1968).What is certain is that birds of the mountains could be seen in the lowlands during severe storms. Perkins (1903: 393) states: Even now, in winter storms, large numbers of birds resort to the lowest skirts of the existing forest, generally at an elevation of 1200–1500 feet; and it is well known that in Cook’s time such forms as [‘O‘u, ‘Apapane, and ‘amakihi] actually came down to the coast in Kealakeakua [sic] Bay, though now such flights would mean death to the visitants. Warner (1968), in reporting a 1958 interview with long-time Kaua‘i resident James Clapper, described such an event at Waimea, Kaua‘i, during winter storms in 1920: A flock of [honeycreepers] including several species was observed congregating in an introduced Hibiscus hedge that was blooming at the side of the road.The birds were feeding on the flowers. Some of the birds were red, others yellow.The flock numbered at least 30. The bills of some were . . . ‘long and decurved, like that of the curlew, and at least two inches long . . . ,’ certainly the now rare Kauai Akialoa, Hemignathus procerus. First observed in the afternoon, the birds stayed in the area only a few hours and were gone the following morning. Henshaw (1902: 20) believed that ‘sudden and marked changes of temperature’ caused the forest birds to descend to the lowlands. He reported that many ‘I‘iwi and ‘Apapane ‘are driven into sheltered valleys and even along the sea-shore far from their woodland haunts’ after heavy and prolonged storms. Although less noticeable because there are fewer native birds present now, this phenomenon continues to the present day, at least during extremely severe storms. ‘Apapane and ‘I‘iwi were seen at unusually low elevations on Kaua‘i after Hurricane Iniki in 1992 (Conant et al. 1998). Circumstantial evidence suggests that it was just such formerly adaptive movement to the lowlands that proved the undoing of the last ‘O‘u on Kaua‘i.

Their numbers were dramatically reduced after Hurricane Iwa in 1983, and none have been seen for certain since Iniki. Although both occurred within a single decade, these were the only severe hurricanes to impact Kaua‘i in the twentieth century, so the last time upland birds retreated to the lowlands there, it was a good strategy.As with ‘nestjumping’, a once adaptive trait has gone awry because of anthropogenic ecological changes. Early naturalists also used the term ‘migration’ to describe seasonal movements of some species in pursuit of food resources. None of the exclusively insectivorous or seed-eating species that survive today exhibit seasonal movements, but the nectarivores and generalists do. Birders are well aware of the fact that ‘I‘iwi and ‘Apapane invade mamane– naio forests, where they do not breed, when the mamane is in peak flower both on Maui (Pratt 1995, 2002b) and Hawai‘i (Hess et al. 2001), but on the latter, at least, the birds may return to their more familiar koa/‘ohi‘a forests to roost (Ralph and Fancy 1995; see below). On O‘ahu, ‘Apapane may appear at unusually low elevations when blue gum and other introduced trees are in bloom (Pratt 1995, 2002b), but whether they return to higher elevations at night is not known. Even within habitats, these species and the ‘Akohekohe (VanGelder and Smith 2001) move about in search of patches of heaviest bloom. Lamoureaux et al. (1981) found that the flowering of Hawaiian trees followed an elevational gradient, with lower elevations blooming in the summer and higher ones in the winter. My own observations indicate that honeycreeper movements are less predictable than might be supposed because blooming cycles are partly dependent on rainfall patterns that in recent years have been rather irregular. Hawai‘i ‘Amakihi on Mauna Loa forsake high-elevation scrub for wetter forests below from August to January (Baldwin 1953), probably because of food scarcity at higher elevations at the time (Lindsey et al. 1998), although they do not seasonally forsake subalpine mamane forest on Mauna Kea (Hess et al. 2001). O‘ahu ‘Amakihi often congregate at flowering eucalyptus and wiliwili haole trees (pers. obs.; Lindsey et al. 1998), but whether they visit the trees daily then return to a home range is not known.

148 The Hawaiian Honeycreepers MacMillen and Carpenter (1980) reported movements of huge numbers of ‘Apapane and ‘I‘iwi upslope at dusk on the south-eastern slope of Mauna Loa. Berger (1981: 163) described such a flight on 20 July 1972: Beginning at 4:00 P.M., loose flocks of birds (primarily Apapane but with occasional Iiwi) were noted flying upslope from Volcanoes National Park toward the direction of the Kulani Cone. Many flocks contained 20 or more birds, and 300 birds were counted during the first 20 minutes of observation; we estimated that more than 600 birds flew up the slope of the mountain during one hour. Some birds flew at treetop level but others, 100 or more feet above the crown of the forest. MacMillen and Carpenter’s (1980) data showed a pronounced seasonality in these mass movements, which begin during spring and continue through the summer, but ‘likely’ cease during the rest of the year. They believed that these energetically costly flights were nevertheless advantageous because of the thermal protection afforded by the forests at 1580 m elevation. Although higher than the forests the birds leave daily, the Kulani area is ‘above the level of heavy fog that occurs almost nightly on the lower slopes during the summer months’ (Macmillen and Carpenter 1980: 35). Ralph and Fancy (1995) reported evening movements downslope from high elevation mamane–naio forest to mid-elevation ‘ohi‘a forest on Mauna Kea. In this case the lower forests would obviously be warmer; night-time temperatures approach freezing in the mamane–naio forest (Weathers and van Riper 1982; pers. obs.). Ralph and Fancy (1995) believed the daily flights were primarily resourcerelated. MacMillen and Carpenter (1980) also acknowledge that the upslope movements would have ‘the additional advantage of reducing encounter (sic) with night-flying Culex mosquitoes, carriers of avian malaria’ to which the species in question are highly vulnerable (see Chapter 9). Van Riper et al. (1986) believed that these daily movements resulted entirely from selection pressures resulting from the presence of malaria at lower elevations. In this hypothesis, summarised by van

Riper (1991a), birds leave the roost early when Culex activity is waning, work downslope to reach the lower elevations when mosquitoes are least active, then return to their mosquito-free haven well before the start of mosquito activity at 2000 hr (8 PM). This behaviour could have evolved from the gradual downslope movements following nectar availability discussed above. As explained by van Riper (1991a: 77–8): those birds that either remained in the upper elevation breeding areas or that moved laterally across the mountain, would not have been appreciably affected. Neither would have the small segment of the V. coccinea and Himatione sanguinea populations that returned up slope each evening. However, those individuals that gradually moved down slope would eventually enter malaria zones. Owing to the presumed extreme susceptibility of the native passerine species, the great majority of those individuals most likely succumbed to malaria. Because of the selective pressure exerted by the malarial parasite at lower elevations, we are, therefore, left today with an altered host behavioral pattern in which a large majority of the V. coccinea and Himatione sanguinea undergo daily altitudinal migrations. Ralph and Fancy (1995), however, believe the movements observed today are little changed from those that evolved before the introduction of mosquitoes. Probably the truth is that thermal advantage, patchy resource availability, and escape from mosquitoes all play a part in the daily movements of ‘I‘iwi and ‘Apapane observed today.

Bioenergetics Relatively little has been published about the physiological responses of Hawaiian honeycreepers, but a few generalisations can nevertheless be made.Those species that have been studied tend to have unexpectedly slow metabolisms as compared to passerines of similar size elsewhere, and the difference is inversely correlated with body mass, i.e. the smaller species have lower rates. Also, homeothermy is

Ecology and breeding biology 149 apparently weak in these birds. Body temperature of the ‘Anianiau becomes lower the colder the ambient temperature and both it and ‘amakihis become hyperthermic at high temperatures (MacMillen 1974).The ‘amakihis in this study died after exposure to temperatures that would be easily tolerated by most passerines. Palila also are sensitive to high ambient temperatures, although Laysan Finches do not differ from other passerines in this respect (Weathers and van Riper 1982).Weathers and van Riper (1982) believed the Palila’s ‘upper critical temperature (31°C), one of the lowest among birds (see Weathers 1981)’ restricted it to cooler high elevations, an interesting observation in light of recent discoveries of Palila remains at sea level on Kaua‘i (S. L. Olson, pers. comm.). The Laysan Finch has several metabolic adaptations to its low, hot environment (Weathers and van Riper 1982) that might not have been present in prehistoric populations on the main islands ( James and Olson 1991).Also,‘amakihi are known to have occurred historically at much lower elevations, where they would encounter much higher temperatures than would MacMillen’s (1974) mamaneforest test subjects. Lepson (1997) speculated that low-elevation ‘Anianiau must have had higher heat tolerance than that reported by MacMillen (1974). Apparently, metabolic adaptations are flexible within honeycreeper species (but perhaps not within local populations). Ability to thermoregulate increases with age in nestling Maui ‘Alauahio (H. and P. Baker 2000), the only species for which data are available. Interestingly, metabolic rates for honeycreepers tend to be positively correlated with body size, just the opposite of what is ‘predicted by conventional endothermic allometry’ (MacMillen 1981). MacMillen (1974) found that the standard metabolic rate for ‘amakihi (Hawai‘i and Kaua‘i species combined) was 96% of the expected value, but that of the ‘Anianiau, at about half the mass, had a rate 76–77% of expected. Metabolic rates for ‘I‘iwi, ‘Apapane, Hawai‘i ‘Amakihi, and ‘Anianiau conformed to the pattern (MacMillen 1981). However, generalisations are difficult to make on the basis of these relatively few studies. Weathers et al. (1983) found a much higher body temperature in ‘Apapane than that found by MacMillen (1981); they attributed the difference to differing times of tak-

ing the measurement (i.e. just after capture as opposed to some hours later when birds had fasted). MacMillen (1981) believed the reduced metabolic rate in the smaller species of nectarivores he studied to be related to the size-determined dominance hierarchy discussed above, and that the lower rates were an energy-conserving adaptation. Evaporative water loss parallels metabolic rate in being higher relative to body size for smaller species (MacMillen 1974). The Laysan Finch, however, is an exception in having a reduced rate of water loss, undoubtedly an adaptation to its sunny atoll environment.

Breeding biology In striking contrast to their wide variation in morphology, feeding techniques, behaviour, and ecology, the reproductive habits of the Hawaiian honeycreepers are remarkably uniform. The details for individual species are included in the species accounts (Part II), so this discussion will be a general overview. Unfortunately, the information we have on drepanidine breeding biology is based on very few detailed life history studies, and only a small subset of species. Just two decades ago (Scott et al. 1980), no breeding information at all was available for about half of the 37 historically known species recognised herein. The two koa-finches, Kona Grosbeak, ‘Ula-‘ai-hawane, both mamos, Greater ‘Amakihi, at least one (and probably all three) nukupu‘u, three of the ‘akialoas, and the ‘O‘u all disappeared before anything at all could be learned about their breeding habits. As of 1980, nests had been described for 18 species, eggs for 14, and nestlings for 14 (not the same list as for eggs). Scott et al. (1980) made a plea for study of those extant species whose breeding was still unknown, and much progress has been made since. Nests of O‘ahu ‘Amakihi (Russell and Ralph 1981; Eddinger 1984;VanderWerf 1997), eggs of ‘Akiapola‘au (Banko and Williams 1993), eggs and young of Hawai‘i Creeper (Sakai and Johanos 1983), nest, eggshells, and chicks of Po‘o-uli (Engilis et al. 1996; Kepler et al. 1996), and nests, eggs, and chicks of ‘Akohekohe (VanGelder 1992, 1996; Berlin and VanGelder 1999; Berlin et al. 2001b;VanGelder and

150 The Hawaiian Honeycreepers Smith 2001) and Maui Parrotbill (VanGelder 1993; Lockwood et al. 1994), have all been added to Scott et al.’s (1980) tally, leaving very few gaps with any potential of being filled. However, discovery of a single (or a few) nests does not mean we can accurately characterise a species’ nesting behaviour, as shown by the wide variations demonstrated by some species (Sakai 1983; Kern and van Riper 1984; H. and P. Baker 2000).Thus, only a few honeycreepers’ breeding is well known, and much remains to be learned despite recent discoveries. Particularly pressing needs, and quite realistic goals, are life history studies of ‘Akeke’e, ‘Akikiki, and O‘ahu ‘Amakihi.

Seasonality Hawaiian honeycreepers, like many tropical birds, have extremely lengthy breeding seasons. Seasonality is more protracted in insectivorous species, whose food supply is relatively constant year-round, but even the nectarivores have longer nesting periods than most continental birds (Ralph and Fancy 1994a). In a given year, the breeding season will be shorter than that established for the species as a whole because nesting may shift from year to year in response to various environmental factors. Also, seasonality can vary within a single island because of local climatic conditions. Most honeycreepers that have been studied begin the breeding cycle during the winter months, some as early as October (Berger 1969). Activity builds over a period of months and peaks during spring and summer, although different species will peak earlier or later within this period (Ralph and Fancy 1994a). Nesting then tapers off into the fall months.Very little nesting activity occurs from August through October, but active nests of one species or another have been found in every month of the year. This cycle coincides with periods of heaviest rainfall in the Hawaiian Islands and partly accounts for our poor knowledge of the birds’ breeding biology (Scott et al. 1980). A few species appear to breed year-round. Active ‘Akiapola‘au nests have been reported in January, February, July, and October (Banko and Williams 1993), and Ralph and Fancy (1994a) captured individuals in breeding condition in March, May, July, August,

October, and November. Such a cycle, with no obvious peak, would be exceptional and additional observations seem to establish at least a predominance of breeding in the first half of the year (T. K. Pratt et al. 2001).

Courtship and pair-bonding Courtship displays were discussed in Chapter 7. Hawaiian honeycreepers tend to be monogamous and the same individuals may remain paired for several years. Maui Parrotbill (Simon et al. 1997) and ‘Akiapola‘au (T. K. Pratt et al. 2001) pairs may associate throughout the year, but most other species remain together only through the breeding cycle, then may join flocks of other unpaired birds or simply disperse. ‘Akepa begin pairing while still following postbreeding flocks in July and August (Lepson and Freed 1997), but that is exceptionally early. The nectar-feeding ‘I‘iwi (Fancy and Ralph 1998), ‘Apapane (Fancy and Ralph 1997), and ‘Akohekohe (Berlin and VanGelder 1999) begin pairing in the fall (September–October), but ‘I‘iwi on Kaua‘i may not do so until January (Eddinger 1970). Kaua‘i ‘Amakihi, Hawai‘i ‘Amakihi (Lindsey et al. 1998), ‘Anianiau (Lepson 1997), Maui ‘Alauahio (H. and P. Baker 2000), and Laysan Finch (Morin 1992b) start pairing in the winter (November–January).

Territoriality As discussed briefly in Chapter 4, Hawaiian honeycreepers share many features of their territoriality with cardueline finches, although a few deviate from the ancestral pattern. Early in the breeding cycle males may defend a moving territory centred on the female, then establish a very small territory, often only a few metres in diameter, around the nest. Such territoriality has been documented for Palila (van Riper 1980a), Laysan Finch (Morin 1992b), and ‘Akepa (Lepson and Freed 1995, 1997). Small breeding territories (but not matecentred ones) have been reported in ‘Anianiau (Lepson 1997), Kaua‘i ‘Amakihi (Eddinger 1970), ‘Apapane (Ralph and Fancy 1997), ‘I‘iwi (Fancy and Ralph 1998), and ‘Akeke‘e (Eddinger 1972a),

Ecology and breeding biology 151 although the latter may exhibit roving mate defence (Lepson and Pratt 1997). A different territorial pattern occurs in species that tend to remain in pairs or family groups throughout the year.They defend a fairly large home range (or feeding territory) that either contracts to a smaller territory around the nest or remains the same size all year. Such a pattern has been described for Maui ‘Alauahio (H. and P. Baker 2000), dry-forest Hawai‘i ‘Amakihi (van Riper 1987), probably Greater ‘Amakihi (Perkins 1903), Maui Parrotbill (Simon et al. 1997), ‘Akiapola‘au (T. K. Pratt et al. 2001), and ‘Akohekohe (Berlin and VanGelder 1999). The Hawai‘i ‘Amakihi varies the size of its territory in different habitats, smaller in mamane forest (van Riper 1987), larger in ‘ohi‘a forest (Baldwin 1953). No territoriality of any kind has been observed in Po‘o-uli (T. K. Pratt et al. 1997) or ‘Akikiki (Foster et al. 2000). Interestingly, all variations of honeycreeper territoriality are found within each major taxonomic grouping.

Nest sites The typical nest of a Hawaiian honeycreeper is located high in a terminal leaf cluster of a nonblooming ‘ohi‘a tree, which seems to be preferred because of the density of its foliage (Fig. 8.19). Such nest sites have been reported for nearly every species that nests in ‘ohi‘a forest, but some have a wider range of site choices and a few deviate strikingly from typical nest placement. Of course, those on

Laysan and Nihoa must use other sites. Laysan Honeycreepers (Schauinsland 1899; Bailey 1956) and Laysan Finches (Morin 1992a) use grass clumps, whereas Nihoa Finches nest in rock crevices (Sincock and Kridler 1977; Morin and Conant 2002). In dry forests, Hawai‘i ‘Amakihi (van Riper 1987) and Palila (van Riper 1980a) nest both in mamane and naio trees but more often in the former. Maui ‘Alauahio tend to build lower in the forest than other honeycreepers and often choose a variety of understorey trees other than ‘ohi‘a.They also do not disdain introduced trees, including conifers such as pines and sugi, but never nest in eucalyptus (H. and P. Baker 2000), which is readily available.The ‘Apapane, although it usually builds in a typical site, has perhaps the widest range of nest sites reported for any honeycreeper (van Riper 1973a; Sakai 1983), including lava tubes, koa trees, low shrubs such as kawau and hapu‘u (tree ferns), and tree cavities. Cavity nesting has also been found as a variation in ‘Akiapola‘au (Ralph and Fancy 1996) and Hawai‘i Creeper (Freed et al. 1987b;Woodworth et al. 2001), and the ‘Akepa is an obligate cavity nester at least on Hawai‘i (Lepson and Freed 1997). It depends entirely on natural cavities in huge old ‘ohi‘a trees and consequently is probably dependent on oldgrowth forests for long-term survival (Hart 2001). Where the nest is placed in a tree is, at least in part, determined by elevation and rainfall, so that the most advantageous thermal environment is obtained (Kern and van Riper 1984). Hawai‘i ‘Amakihi vary nest placement seasonally, and even a single pair that renests may choose a different site later in the season (van Riper et al. 1993).

Nests

8.19 Location of ‘Apapane nest (arrow) in dense ‘ohi‘a tree, a typical honeycreeper nest site.

Nests are always built by the female, with or without help from the male. The typical pattern is for the female to carry on most of the building chores, with relatively little help from the male who may bring nest materials that the female then places in the nest. Species in which only the female builds include Hawai‘i ‘Amakihi (Lindsey et al. 1998), ‘Akepa (Lepson and Freed 1997), Maui Parrotbill (Simon et al. 1997), Maui ‘Alauahio (H. and P. Baker 2000), and Laysan Finch (Morin 1992a;

152 The Hawaiian Honeycreepers Morin and Conant 2002). Both sexes contribute about equally to nest construction in the ‘Apapane (Fancy and Ralph 1997), and may do so in ‘Akikiki (Foster et al. 2000) and Po‘o-uli (T. K. Pratt et al. 1997b), but observations are too few to say for sure. Virtually all drepanidine nests are open cups (Fig. 8.20) often referred to as ‘statant’.3 All have a coarser outer wall and a central cup very neatly lined with fine materials. The actual materials used as well as the ‘style’ of construction vary widely even within species. For example, Kern and van Riper (1984) found that Hawai‘i ‘Amakihi, whose nesting is arguably the best known, build five types of nest with rather striking differences in appearance, and H. and P. Baker (2000) describe seven variations of Maui ‘Alauahio nests. By far the most common materials in honeycreeper nests are, not surprisingly, twigs and bark of ‘ohi‘a, but the ‘Akohekohe even uses the flowers (stamens) for lining the nest cup (Berlin and VanGelder 1999). ‘Olapa and pukiawe are also frequent sources of twigs and bark strips. Soft materials, probably included for insulation, include Usnea lichen (Fig. 8.21), mosses, and pulu, a cottony material produced by hapu‘u tree ferns (Fig. 8.22). Linings of the central cup are always made of very fine material, the most frequent being rootlets, especially those of ferns, and grass fibres. Although the outer nest walls vary in the amount of soft material vs. ‘hard’ twigs and stems, even intraspecifically depending on their environment (Kern and van Riper 1974), the basic pattern described applies to all but two. The ‘Akiapola‘au builds a unique parapet of ‘ohi‘a bark strips that stick up above the rim of the nest (Banko and Williams 1993), and the three Paroreomyza are the only Hawaiian honeycreepers that routinely use spider silk, especially egg cases, to bind their nests (H. and P. Baker 2000; P. and H. Baker 2000), although Hawai‘i Creepers have been observed apparently gathering spider silk as nest material (Sakai and Ralph 1980). Spider silk is an important component in nests of monarchine ‘elepaios (Berger 1981) and they may compete with ‘alauahios for this material (see discussion of competitive exclusion, Chapter 4). Hawaiian honeycreepers are opportunistic and choose whatever material is available that will do the job. For example, one Maui ‘Alauahio nest

8.20 Collected nest of Maui Parrotbill; typical honeycreeper nest comprised largely of lichens and twigs and situated in terminal leaf cluster of ‘ohi‘a.

8.21 Usnea lichen, probably the second most frequent component (after ‘ohi‘a bark) of Hawaiian honeycreeper nests.

used the sticky scales from pine buds instead of spider silk to bind the nest (H. and P. Baker 2000). Apparently anything that has the proper physical properties may be used, including man-made fibres (H. and P. Baker 2000).When they can find it, both Palila and Hawai‘i ‘Amakihi use sheep wool in the outer walls (but not the lining) of their nests (van Riper 1977); other animal hair, such as that of horses and feral pigs, has also been reported in nests of several species. Maui ‘Alauahio, Hawai‘i ‘Amakihi, and ‘Akepa may use exotic pine needles or grasses (Lepson and Freed 1997; Lindsey et al. 1998; H. and P. Baker 2000). Many honeycreepers dismantle old nests of their own and other species and reuse the

Ecology and breeding biology 153 to orange (‘I‘iwi,‘Apapane).A typical clutch is three, but clutches of from one to five have been reported and some species never have more than two eggs. Incubation periods vary among species from 14 to 18 days, with 16 being typical. Intraspecific variation is no more than 2 days. Only females develop a brood patch and incubate. Males feed females by regurgitation during incubation, occasionally on the nest but more often away from it.The female is usually silent while incubating, but a few will answer the calls of a mate approaching to feed.

Nestlings and fledglings

8.22 Furry-looking ‘pulu’, soft, fibrous scales at base of hapu‘u tree fern fronds, often used as a nest lining by Hawaiian honeycreepers.

materials, and stealing of materials from nests under construction is common (Lepson 1997). A few species at least occasionally place a new nest on top of an old one, not necessarily their own (van Riper 1973a, 1976;VanGelder and Smith 2001).

Eggs Honeycreeper eggs are quite uniform in external appearance. All are nearly white, spotted and blotched with various earth tones, brown, and purple.The spots tend to concentrate in a ring around the larger end, and the amount of spotting varies widely within both species and individuals.The egg surface is smooth but not glossy.Yolk colour varies (Eddinger 1970; Berger 1981) from yellow (Nihoa Finch, Hawai‘i ‘Amakihi, ‘Anianiau, Kaua‘i ‘Akepa)

The eggs usually hatch in the morning, with most of the clutch hatching by noon or at least within a day. Honeycreeper chicks are altricial and nidicolous (i.e. helpless nestlings with eyes closed).They are nearly naked with sparse grey down in various shades, mostly on the head and back. They are brooded only by the female, especially during rainy weather and at night, and late in the nestling period brooding may be restricted to those times. Both parents feed the young, although the female takes the larger role in most species. Sometimes food is passed from the male to the female and then fed to the nestlings. The young spend between 15 and 27 days in the nest before fledging. The length of time varies among species, with no discernible pattern. The shortest nestling periods (15 days) reported are for Hawai‘i ‘Amakihi (Lindsey et al. 1998) and Maui Parrotbill (Simon et al. 1997), and the longest (26 and 27 days, respectively) in Laysan Finch (Morin 1992a; Morin and Conant 2002) and ‘Akohekohe (VanGelder and Smith 2001). Once the chicks fledge, they remain dependent on their parents for periods varying from 1 to 4 months in most species, to 5 to 8 months in the Maui Parrotbill (Simon et al. 1997), to sometimes over a year in the ‘Akiapola‘au (T. K. Pratt et al. 2001). Long dependency periods are probably associated with difficult-to-master feeding apparatus.

Nest sanitation As discussed in Chapter 4, some cardueline finches and some Hawaiian honeycreepers are the only

154 The Hawaiian Honeycreepers passerines that allow faecal sacs to accumulate on the rim of their nests during the last days of the nestling period. When the chicks first hatch, the parents diligently carry away the faecal sacs. When the nestlings become more mobile, they present the sacs to the parent for disposal. Later, the nestlings perch on the nest rim to defecate over the side, and in some species faecal sacs become caked around the outside of the nest late in the breeding cycle. This behaviour is in stark contrast to that of the Maui ‘Alauahio (H. and P. Baker 2000) and the nectarivorous honeycreepers that fastidiously remove any faeces that stick to the nest. In honeycreepers it is a difficult trait to assay in species for which we have limited knowledge because many collected nests were abandoned before the breeding cycle was complete. Thus it was not until recently that this trait was confirmed in the Maui Parrotbill (Simon et al. 1997), and it appears to be much less pronounced in that species than in better known examples such as Palila (van Riper 1980a) and Laysan Finch (Berger 1981; Morin 1992a).The trait is also seen in the ‘Akepa (Lepson and Freed 1997) and the Hawai‘i ‘Amakihi, but in the latter it is variable: dry-forest birds have complete nest sanitation (van Riper 1978b) but elsewhere on Hawai‘i and on Maui, their faeces sometimes accumulate on the outside of the nest (Lindsey et al. 1998).

Co-operative breeding Nest helpers are rare among Hawaiian honeycreepers. Reports of a female helper at a Hawai‘i ‘Amakihi nest on Maui (Lindsey et al. 1998) and of two helpers at one Palila nest (van Riper 1979) are clearly exceptional.The only species that routinely has nest helpers is the Maui ‘Alauahio (H. and P. Baker 2000), a species that stands apart from other honeycreepers in many other fundamental aspects (see Chapter 4). One or two offspring from the previous year may help feed the female, nestlings, and fledglings. Virtually all pairs have helpers in optimal habitats, but less than a third may have them in disturbed areas, undoubtedly a reflection of differential nesting success (H. and P. Baker 2000). Apparently, if young of the previous year are available to help, they do so. Whether the con-

generic O‘ahu ‘Alauahio and Kakawahie had nest helpers is not known.

Demography Reproductive age Most Hawaiian honeycreepers can breed at 1 year of age, but only ‘Apapane (Ralph and Fancy 1997), ‘I‘iwi (Ralph and Fancy 1998), and Hawai‘i ‘Amakihi (van Riper 1987) are known to do so routinely. Most species breed at the age of two, but ‘Akohekohe (Berlin and VanGelder 1999), ‘Akepa (Lepson and Freed 1995), and some Laysan Finches (Morin and Conant 2002) may wait until 3 years of age.

Reproductive success Most drepanidines raise a single brood each year, but ‘Akohekohe usually raise at least two and sometimes three (Berlin and VanGelder 1999). Others raise a second brood if they can, and most species that have been studied renest if the first attempt fails. Published data on nesting success are difficult to summarise because of variability in what is reported, variability from locality to locality within the range of a species, and variability from one year to the next. For example, van Riper (1980a) found a Palila success rate of 50%, but Pletschet and Kelly (1990) found half that. H. and P. Baker (2000) found nesting success of Maui ‘Alauahio varied from 20% to 30% from year to year in one ecologically disturbed locality, but approached 100% in a presumably better habitat.‘Akohekohe in Waikamoi Preserve had rates of 80% and 36% in succeeding years, and in Hanawi Natural Area Reserve had successive rates of 78%, 87%, and 62% (Berlin and VanGelder 1999). Probably all Hawaiian honeycreepers exhibit fairly strong annual and locality variation. Nevertheless, the rate of young fledged per nest remains relatively constant (1.4–1.6) over these same variables. Based on a limited sample of species and limited number of nests for the rarer birds on Kaua‘i, Freed (1999) suggested that smaller clutch sizes and single broods per year make the endangered species less adaptable than the others that have larger clutches and multiple broods.These generalisations do not appear to hold up when

Ecology and breeding biology 155 more species and other islands are considered (and one of Freed’s supposedly ‘listed’ species, ‘akepa = ‘Akeke‘e, is not). Some endangered honeycreepers have relatively large clutch sizes and more than one brood per season (Berlin and VanGelder 1999; Morin and Conant 2002).

Survival and longevity Among the Hawaiian honeycreepers for which it is known, annual survival rates for adults are remarkably high, especially for small insectivores (Woodworth et al. 2001), although Lindsey et al. (1995) considered them ‘comparable to those . . . of birds in temperate and tropical forests’.Their conclusion was probably valid at the time, but more recent data reveal that two (Palila and ‘I‘iwi) of the five honeycreepers they compared have the lowest survival rates among honeycreepers by a wide margin, which biased their generalisations. Most Hawaiian honeycreepers do, indeed, have much higher survival rates than continental birds (Table 8.1).

‘Conventional wisdom’ among ecologists has long held that tropical birds in general have higher survival rates and smaller clutches than those of temperate climates, but Karr et al. (1990) challenged that belief by showing that rates and clutch sizes in Maryland (54%) and Panama (56%) did not differ. However, Faaborg and Arendt (1995) found a higher rate (68%), but not smaller clutches, in Puerto Rican birds, and Johnston et al. (1997) found a similar rate on the island of Trinidad. Faaborg and Arendt (1995: 506) asked ‘is it possible that there is something distinctively different about island populations that we do not yet understand?’ Clutch sizes of Hawaiian honeycreepers are not smaller than those reported in most of these studies, but survival rates are higher even when compared to those on Caribbean islands. Survival rates for dreps in their first year, however, are much lower across the board. Apparently, if a honeycreeper manages to reach adulthood its chances for a long life are quite good, barring outside perturbations. Lifespans of up to 12 years (Table 8.1) are substantial

Table 8.1 Longevity and survival rates of Hawaiian honeycreepers. Species

Longest known lifespan (years)

Survival rate (adults %)

References

‘Akepa ‘Akiapola‘au ‘Akohekohe ‘Anianiau ‘Apapane Hawai‘i ‘Amakihi Hawai‘i Creeper ‘I‘iwi Kaua‘i ‘Amakihi Laysan Finch Maui ‘Alauahio Maui Parrotbill Nihoa Finch Palila

⬎9 ⬎7 ⬎5b ⬎9.5 (captive) ND ⬎12 ND ND ⬎9 (captive) ⬎12 ⬎8b ⬎5 ⬎12 ⬎9

0.70, 0.82 NDa ‘high’ ND 0.72 0.85 0.88 0.55 ND ND 0.87 ND ND 0.63c

Lepson and Freed 1995, 1997; Ralph and Fancy 1994a Snetsinger et al. 1998 Berlin and VanGelder 1999 Berger 1980 Ralph and Fancy 1995 Freed 1988 Woodworth et al. 2001 Fancy and Ralph 1998 Berger 1980 Morin 1992b; Morin and Conant 2002 H. and P. Baker 2000 Simon et al. 1997 Conant 1983 Lindsey et al. 1995

aND = No

data. on short-term studies; probably higher. cActual figures for males (0.65) and females (0.62) not significantly different. bBased

156 The Hawaiian Honeycreepers for passerines. Apparently island birds do differ in survival in some as yet unexplained way, and the honeycreepers’ very high rates are simply a function of their extreme insularity. Nevertheless, both Palila (Lindsey et al. 1995) and ‘I‘iwi (Fancy and Ralph 1998; Tummons 2001b) are struggling to maintain their populations in the presence of threats such as disease and predation (see Chapters 9 and 10).

Philopatry Philopatry (site tenacity) varies widely among Hawaiian honeycreepers. Some species such as Hawai‘i Creeper (Woodworth et al. 2001) and Palila (Fancy et al. 1993b) are so strongly philopatric that they are less able to recolonise suitable habitats in their former ranges. All insectivorous honeycreepers that have been reported tend to occupy the same home ranges for years, and may even nest in the same tree in succeeding years, with relatively little dispersal even of young. In contrast, ‘Apapane and ‘I‘iwi, both of which wander widely following patches of bloom, may undergo large population

shifts. ‘I‘iwi disperse so readily that measures of survivorship may be biased (Fancy and Ralph 1997, 1998). However, the ‘Akohekohe, though it wanders widely after the breeding season, appears to be much more philopatric than its relatives and may, like the insectivores, even nest in the same tree year to year (Berlin and VanGelder 1999). 1. For some reason the word ‘nectarivory’ came into widespread use only in the past two decades. Some authors (e.g. Freed et al. 1987a) have used ‘nectivory’ but there is no such word and if there were, it would mean feeding on dead matter (Gk. nekton). 2. In much of the literature the term ‘nectar-robbing’ is used for this phenomenon, but it is the flower that is being robbed; the nectar is being stolen. 3. I do not know how the heraldic term ‘statant’, which means ‘standing on all four feet’, came to be used in this context. I am not aware that bird nests have feet, let alone stand on them, so I assume what is meant is a nest that will stand alone without collapsing. It cannot mean ‘self-supporting’ because many nests so described are supported by tree branches.

9 Diseases and parasites

Native birds have now experienced over a hundred generations of life with a pathogen that kills most of them. How they are marshaling the few defenses still present and, hopefully, evolving new ones, will demonstrate to evolutionary biologists the creative aspects of extreme natural selection in changing ecosystems. Cann and Douglas (1999: 134)

In a typical life history account of a bird species, or in books like this one, a few perfunctory lines are usually included to indicate that the species or group in question is known to have certain diseases and parasites. Such information is included for completeness, but is frequently ignored by readers who prefer to concentrate on more important or more interesting subjects such as ecology and behaviour. Unfortunately, no one can ignore disease in any discussion of Hawaiian honeycreepers. Indeed, for the past two centuries, disease has probably been the single most important factor in drepanidine natural history, directly causing the extinction of many species, contributing to the demise of others, and determining the present-day distributions of survivors. No other group of birds have been so strikingly affected by disease, and those who have studied it among honeycreepers have become world leaders in research on disease in wild bird populations. In fact, much of our knowledge of the workings of disease in wild birds comes from studies of Hawaiian honeycreepers (van Riper et al. 1994), most studies of avian diseases having involved domestic poultry or birds in captivity (Atkinson and van Riper 1991).The sad history of bird diseases in Hawai‘i reads much like a whodunit. Because many of the key clues have emerged only in the past few

years, this book will be the first to offer a hypothetical solution that addresses all outstanding questions.

What is killing Hawai‘i’s birds? Evidence of epizootic Beginning about 1895 (earlier on O‘ahu), entire species of birds, many of which had until then been common, suddenly and mysteriously disappeared from the Hawaiian Islands. The story has been recounted in varying detail by Warner (1968, 1973), Atkinson (1977), and Pratt (1994a). The unprecedented rapidity of this catastrophe might have gone undetected except for the fortuitous presence of several competent naturalists in the islands at the time. Except on O‘ahu, the Hawaiian avifauna in 1890 did not look very different from that seen by Captain Cook a century earlier. A few Hawaiian honeycreepers had become extremely rare but none were known to be extinct (see Table 10.1 for a complete list of drepanidine extinctions). It was a time of intense interest in Hawaiian birds, with naturalists such as Scott B. Wilson, Henry Palmer, George C. Munro, R. C. L. Perkins, Henry W. Henshaw, and William Alanson Bryan actively collecting both specimens and information (see Chapter 3). They found nearly all of the drepanidine species known previously and discovered many ‘new’ ones. The earlier visitors, such as Wilson and Palmer, noted nothing amiss except on O‘ahu, where all agreed the dearth of native birds was obvious and puzzling. Perkins (1903: 403) stated: The general scarcity of birds on Oahu as compared with the other islands is noticeable,

158 The Hawaiian Honeycreepers both as to the number of species and individuals, but this scarcity applies only to recent years, for the well-known ornithologist Townsend expressly states that he found birds more abundant on Oahu than on Kauai, some 65 years ago. On the other islands, Perkins (1903), and especially Henshaw (1901, 1902) and Munro (1960) who worked in the islands through the turn of the century, would bear witness to a sudden and mysterious calamity. Birds disappeared from seemingly pristine habitats almost overnight in what Bryan (1912c) called ‘one of the wonder tales of ornithology’. Although O‘ahu had lost several honeycreepers and other birds before 1850, the O‘ahu Nukupu‘u (based on anecdotal evidence from Perkins reported by Munro 1960) still ‘abounded’ in 1860, but thirty years later, none could be found.The O‘ahu ‘Akepa, ‘Akialoa, and ‘O‘u were still present, though very rare, in 1890, but were gone by 1900 (Warner 1968; Scott et al. 1986). Indeed, the naturalists of the 1890s spent little time on O‘ahu because bird life was so scarce there. Henshaw (1902: 66) was mystified: The cause of the extinction of the ou upon Oahu seems to be very obscure. The fruit of the ieie vine is the particular food of the bird, and there are considerable tracts of timber on the mountains of the island where this vine still abounds. So too, there are sections where the introduced guava and the mamaki are still plentiful, and the ou is very fond of the fruit and berries. There apparently being no scarcity of food and shelter, why should the ou have disappeared from Oahu, and yet persist upon other islands . . .? The O‘ahu experience was a foretaste of things to come on other islands. Although he used the meliphagid Hawai‘i ‘O‘o as his main example, Perkins (1903: 393) was as puzzled as Henshaw by similar events on Hawai‘i when he reported that: on a very rough lava-flow on Hawaii in 1892, the ‘Oo’ . . . was very numerous, and as

many as a dozen of these birds could be seen in a single tree, making, with hosts of the scarlet ‘Iiwi,’ the crimson ‘Apapane,’ and other birds, a picture never to be forgotten. A few years afterwards, on revisiting the spot at the same season, although the trees were, as before, one mass of flowers, hardly a single ‘Oo’ was to be seen. Henshaw, arriving in the islands essentially on the heels of Palmer and Perkins, who had observed a nearly intact avifauna in the early 1890s, provides some of the best information on these events, at least on The Big Island. Although he arrived in Hawai‘i in 1894, poor health kept him from serious field work until several years later (Nelson 1932). In an 1896 letter (reported by Nelson 1932), a perplexed Henshaw wrote that ‘they are a curious lot, these Hawaiian birds and any change whatever in environment seems fatal to them. It is simply astounding the way they disappear’, and later observed (Henshaw 1901: 158–9): That extensive deforestation should have a marked effect upon Hawaiian birds, wholly unused as they are to competition of any kind, is what we might expect; but there remains to be recorded a still more remarkable fact indicative of the singular sensitiveness of Hawaiian birds to change. Large sections of forest land on Hawaii that have been but slightly interfered with by man, and that are nearly as dense and impenetrable as they ever were, have been almost wholly abandoned by birds within the last ten years. For this abandonment, no reasonable explanation suggests itself. Henshaw was the last to see the Hawai‘i Mamo (1898), but the bird had been common enough in the Hilo District for feather-hunters to have supposedly shot up to 12 individuals in a day in 1880 (Perkins 1903: 399). Henshaw was also the last naturalist to observe the Greater ‘Amakihi and Hawai‘i ‘Akialoa, neither of which was reliably reported in the twentieth century, but which were not rare (although the former was very local) in the

Diseases and parasites 159 early 1890s (Wilson and Evans 1890–99; Rothschild 1893–1900; Henshaw 1902; Perkins 1903; Greenway 1967). He was a bit too late, apparently, for the Greater Koa-Finch and Kona Grosbeak, which he failed to find even though he visited the localities where Palmer and Perkins had found them only a few years earlier. Palmer’s assistant Munro (1960: 131) lamented: Henshaw failed to find [the Kona Grosbeak] and considered our wonderful collecting area, which we thought a collector’s paradise, one of the poorest over which he had hunted. It can be seen from some of Perkins’ experiences how quickly the birds disappeared from some localities. Perkins (1903) found both the Black Mamo and ‘Akohekohe in good numbers on Moloka‘i in 1893, but by 1907 Bryan (1908) found the former absent from the areas where Perkins had found it and greatly reduced in numbers, and the latter nearly extirpated. Neither were ever seen again on the island, although no one was present to record exactly when they finally disappeared. The Lana‘i ‘Akialoa, rare in 1893, was gone before 1911 and the Lana‘i Hookbill was last seen in 1918 (Munro 1960). Throughout the islands, species that did not die out completely withdrew from parts of their ranges where they had formerly been common. On Maui, the parrotbill, nukupu‘u, and ‘Akohekohe became so rare that for a long time all were believed extinct (Richards and Baldwin 1953; Banko 1968), and the ‘O‘u, common in the 1890s (Henshaw 1902; Perkins 1903), was never reported from Maui in the twentieth century. The Kaua‘i ‘Akialoa, abundant in the 1890s (Henshaw 1902; Perkins 1903; Munro 1960), was not reported again until 1960 (Richardson and Bowles 1964), and the Kaua‘i Nukupu‘u may not have survived the turn of the century despite subsequent reports (Pratt and Pyle 2000). Aftershocks of this cataclysm continued through the twentieth century, although, except on Lana‘i, they were less well documented. For the story on that island, we are indebted to Munro (1960), who went there as ranch foreman in 1911 and

remained for 20 years.Although the Lana‘i ‘Akialoa was last seen by Perkins some time in the early 1890s, most of the other birds were benefiting from reforestation efforts to such an extent that Munro could report to the Bishop Museum in 1923: the forest birds of Lanai are holding their own and some species probably increasing, in the following order, ou, olomao, amakihi, apapane, [Maui ‘Alauahio] and iiwi and there is every reason to believe that these species, with the proposed extension of the forest area planned by the Hawaiian Pineapple Company, will flourish indefinitely. Shortly thereafter, Lana‘i City was built right at the base of the forested area, and the situation changed so dramatically that Munro’s 1932 report stated that native birds ‘are getting scarcer; the ou has not been seen for some time.’ By 1940, only the ‘Apapane and Hawai‘i ‘Amakihi survived on Lana‘i, and the latter disappeared by the 1970s (Hirai 1978; Scott et al. 1986). On other islands, some populations and species that were still common after the holocaust of the 1890s became vanishingly rare or extinct in a second wave of declines that may have paralleled the events on Lana‘i, but we can only speculate about the timing because so few observers were active in Hawai‘i, especially on the outer islands, during the first half of the twentieth century. On Moloka‘i, the ‘O‘u and Kakawahie were both still common in 1907 (Bryan 1908), but could not be found in surveys in 1937 (Munro 1960) and 1948 (Richardson 1949), although the latter apparently survived in very low numbers into the early 1960s (Pekelo 1963, 1964). On Hawai‘i, Baldwin’s (1941, 1953; Richards and Baldwin 1953) surveys in the late 1930s located ‘O‘u, Hawai‘i Creeper, ‘Akepa, and ‘I‘iwi in a mid-elevation area of Hawaii Volcanoes NP, from which all subsequently disappeared (Conant 1975; van Riper et al. 1986). Kaua‘i lost no species (except perhaps the Kaua‘i Nukupu‘u, see Pratt and Pyle 2000) in the 1890s catastrophe, but by the time they were found again in 1960 (Richardson and Bowles 1964), several had become vanishingly rare and the Kaua‘i ‘Akialoa apparently

160 The Hawaiian Honeycreepers

9.1 Understorey of nearly pristine native rainforest virtually devoid of native birds because of its low elevation; Kamakou Preserve, Moloka‘i.

did not survive the decade of its rediscovery. The legacy of these disasters is that, even today, Hawaiian honeycreepers are virtually absent from some native rainforests that still appear otherwise ecologically intact. Such places include the upper elevations of eastern Moloka‘i (Scott et al. 1977), Kohala Mountain (van Riper 1973b) and the east flank of Mauna Loa (Conant 1975) on Hawai‘i, and West Maui (Scott et al. 1986). On a visit to the upper reaches of eastern Moloka‘i in 1975, I was struck by the eerie silence of a beautiful native rainforest (Fig. 9.1) that showed almost no evidence of human influences (Scott et al. 1977).The one thing all these places have in common is that they are at relatively lower elevations than other areas that still have native bird populations. Henshaw (1902) searched for causes for the population crashes and abandonment of seemingly good habitat, but could only hypothesise a hypersensitivity to disturbance or the effects of inbreeding as causes. We now know that the explanation was, quite literally, right under Henshaw’s (and his contemporaries’) nose. Henshaw (1902) reported

that when birds came to the lowlands during storms (see Chapter 8), ‘scores . . . are picked up dead or dying, and the mortality . . . is unusually great’, and that ‘dead birds are . . . found rather frequently in the woods on the island of Hawaii.’The Hawaiian lowlands had become, for some reason, a honeycreeper death trap. Furthermore, nearly all bird collectors in the 1890s remarked about the frequency of birds found with swellings and lesions on their bare parts. According to Perkins (1903) these lesions affected many species of Hawaiian honeycreepers on all islands. He told of picking up a sick Kaua‘i ‘Akialoa that was unable to fly, and of finding two Kakawahie hiding in a dense shrub that would not move until actually touched. Both Perkins and Henshaw found diseased birds more often in wetter parts of the islands, and Henshaw thought the pathogen might be picked up from the wet bark of trees. But neither considered disease the main culprit in the precipitous bird declines they were witnessing. Munro (1960), however, was convinced that circumstantial evidence clearly implicated an epizootic as the cause in the same way that European diseases such as smallpox, syphilis, and measles had decimated the native Hawaiian population earlier in the nineteenth century. But what were these diseases and how were they transmitted? And why were their effects seen primarily at low elevations?

The vectors and the pathogens The two diseases most often hypothesised as being causative agents in extinctions and declines of Hawaiian honeycreepers are avian pox and avian malaria (Warner 1968; Ralph and van Riper 1985; van Riper and van Riper 1985; van Riper 1991a). They are among the relatively few avian diseases capable of infecting a wide variety of hosts (most others are host-specific; van Riper and van Riper 1985), and both are transmitted by mosquitoes (although pox can spread by other means as well). We cannot fault scientists of the 1890s for failing to appreciate the importance of vectors; the role of mosquitoes in transmission of such human diseases as malaria and yellow fever was just then being worked out (Harrison 1978), and avian malaria had

Diseases and parasites 161 only recently been discovered (for a thorough review of the history of avian malaria, see van Riper et al. 1994). But by 1944, when Munro (1960) published his first edition, he could state almost dogmatically: ‘The ou’s habit of coming down to the lowest levels where food could be found, exposed it to introduced diseases, probably mosquito-borne.’ But his evidence was all circumstantial; the actual link between diseases, mosquitoes, and bird die-offs was not demonstrated until Warner’s (1968) pioneering studies. Warner (1968) examined the history of mosquitoes and mosquito-borne parasites in Hawai‘i, and, unless otherwise stated, the following summary is based on his report (with taxonomy updated). Originally, the Hawaiian Islands had no mosquitoes. They were first noticed at Lahaina, Maui, in 1827, and traced to the ship Wellington that had, the year before, drained containers filled with water from the west coast of Mexico into a freshwater stream in West Maui. The waste water contained larvae of the night-flying mosquito Culex quinquefasciatus (Fig. 9.2), which became established and presumably spread widely (although no records were kept). This mosquito has a flight range of at least 23 km, and can breed in brackish coastal pools, so it was ideally suited for rapid spread in the islands. Importantly, the insects were of the subspecies C. q. fatigans, which is adapted to warmer climates (it is replaced by another subspecies north of Stockton, California), and thus were not suited

9.2 Mosquito biting an ‘Apapane in the vulnerable bare skin of eyelid. Photo © Jack Jeffrey.

to colder higher elevations, a very important point (see below). A second species of introduced mosquito, Aedes albopictus, has been implicated as a minor vector elsewhere, but in Hawai‘i has not been shown to carry avian malaria and is restricted to even lower elevations than C. q. fatigans (van Riper et al. 1986). Blood-sucking hippoboscid flies, ectoparasites that were undoubtedly introduced with the first European domestic fowl, are also potential vectors of bird diseases. Perkins (1893) found hippoboscids on an ‘I‘iwi in Kona, and Warner regarded them as widely established, but nothing has been published on their elevational distribution, and their role, if any, in transmission of avian diseases in Hawai‘i has not been well researched. With vectors established, pathogenic organisms previously unknown among Hawaiian honeycreepers could begin to spread. Warner and Atkinson both believed that a reservoir of bird diseases that could be spread by mosquitoes had been present in the Hawaiian Islands for a long time, carried by migratory waterfowl and shorebirds, seabirds, domestic fowl, and introduced game birds. Poxvirus avium, the cause of avian pox, probably did arrive with the first domestic chickens, but avian malaria is another story. In an exhaustive literature review of the distribution of malaria haematozoa in suspect reservoir groups, van Riper et al. (1986) revealed that seabirds and shorebirds almost never harbour avian malaria, and that although ducks can carry malaria, the specific organism that is the infective agent in Hawai‘i, Plasmodium relictum capistranoae, has never been reported from waterfowl or any other migratory birds in Hawaii (van Riper and van Riper 1985). Also, the habitats of these groups are rarely, if ever, frequented by Hawaiian honeycreepers and other forest birds. Consequently, van Riper et al. (1986) concluded that migratory birds could not have been a significant malaria reservoir. However, by the last quarter of the nineteenth century, several introduced birds that are definitely known to carry avian malaria (van Riper et al. 1986; Cann and Douglas 1999), including Spotted Dove, Nutmeg Mannikin, and House Finch, had penetrated deeply into native forests (Henshaw 1902).

162 The Hawaiian Honeycreepers During Warner’s study, the approximate upper limit for the mosquitoes was 600 m, but that has moved upward since (Scott et al. 1986; Conant et al. 1998). Scott et al. (1986) found the functional upper limit of mosquitoes, above which their epidemiological significance is minimal, to be as high as 1500 m. When I first visited the Koke‘e area at ca. 1100 m on Kaua‘i in 1974, I did not notice any mosquitoes, but by the 1990s they had become a serious annoyance. Their presence by 1996 near Koai‘e Stream in Kaua‘i’s Alaka‘i Plateau, and the apparent first appearance of avian pox in the area (Herrmann and Snetsinger 1997), demonstrates their continued assault on the higher ramparts. One would expect Culex quinquefasciatus fatigans to adapt slowly to cooler regions because other subspecies have done so. However, evidence is accumulating that a general warming trend in recent years has assisted them in their upward advance (Tummons 2001a). Warner noted that Hawaiian honeycreepers (and most other native birds) generally occurred only in areas relatively free of mosquitoes, but he had no hard data to support the claim and Atkinson (1977) considered it unconvincing. However, the HFBS (Scott et al. 1986) found precise mathematical correlations that showed unequivocally that Warner was essentially correct: mosquitoes and honeycreepers inhabit mutually exclusive areas. Not only were honeycreepers rare in areas with substantial mosquito populations, but ‘staggering drop-offs of densities in mosquito-infested areas’ (Scott et al. 1986: 367) were documented for several species. With a few scattered exceptions, the surveys showed that mosquito-borne disease currently restricts the ranges of virtually all surviving Hawaiian honeycreepers. Furthermore, they showed that species such as the nukupu‘us and ‘O‘u, whose densities were greatest at the lowest portions of the mosquito-free refugia, were the most vulnerable to extinction by disease, which, in the case of the ‘O‘u at least, seems clearly shown by the pattern of its disappearance in the twentieth century.

Epidemiology Warner conducted several experiments to demonstrate the vulnerability of Hawaiian honeycreepers

to avian pox and malaria. Laysan remains today free of mosquitoes, so its birds had never encountered mosquito-borne diseases. Attempts had been made in the past to keep Laysan Finches at the Honolulu Zoo, but they ‘lived only a short time and then died of unknown causes’ (Warner 1968: 108). In 1958, Warner brought some of the finches to Honolulu, where they were maintained in apparent good health for 2 months in a mosquito-proof enclosure, and then were exposed to local mosquitoes. Because of control measures, the mosquito population at the site of the experiment was unusually low. Nevertheless, within 2 weeks of exposure to them, six of the finches had obvious pox lesions, and by the end of a month virtually all were infected. Some of these birds had possible symptoms of avian malaria, but that could not be confirmed and the low mosquito population made this study group unsuitable for detecting that disease. So a year later, Warner brought 36 Laysan Finches to Lihu‘e, Kaua‘i, where the mosquito population was relatively high.As before, the birds were maintained in apparently good health (one bird died but had no disease symptoms) in mosquito-proof enclosures for one month.Then they were divided into experimental and control groups, and the former were exposed at night to mosquitoes. None of the birds in the two groups were diseased at the start of the experiment, but after five nights of exposure the first experimental bird died, and all were dead by the 16th night. Blood smears from the dead birds revealed massive infections with malaria parasites. Meanwhile, the control group remained healthy. In a follow-up, Warner determined that only three nights’ exposure to mosquitoes was sufficient to induce lethal avian malaria in Laysan Finches. In an experiment of similar, though not identical, design,Warner captured a small group of ‘Apapane, Kaua‘i ‘Amakihi, and ‘Anianiau at 1200 m and brought them to Lihu‘e along with a group of introduced Japanese White-eyes from the same elevation. After a week at low elevations in mosquitoproof cages, none of the birds had blood parasites. Five days after exposure to mosquitoes, two of the honeycreepers had malaria and by the 10th day, all had massive infections. In contrast, only one of the exposed white-eyes developed malaria, and it was

Diseases and parasites 163 a mild case. Lowland introduced passerines from the Lihu‘e area (House Finches and white-eyes) all had light infections of Plasmodium, as well as evidence of avian pox such as healed lesions, but were apparently not debilitated by either. Recent studies have confirmed and refined Warner’s findings with regard to the extreme susceptibility of Hawaiian honeycreepers to avian malaria, as well as the striking differential between them and introduced birds in their responses to the disease. Using artificial infection techniques, van Riper et al. (1986) demonstrated that Laysan Finch, Hawai‘i ‘Amakihi,‘I‘iwi, and ‘Apapane are highly susceptible to avian malaria. Mortality rates from the challenges ranged from 100% in the Laysan Finch to 20% in Hawai‘i ‘Amakihi from dry habitats (the same species from wetter habitats had a rate about twice as high). Control birds (Island Canary, Red-billed Leiothrix, and Japanese White-eye) not only survived the challenge but the latter two failed to become infected. Warner had been unable to demonstrate the existence of malaria in wild honeycreepers, but van Riper et al. (1986) found it in Hawai‘i ‘Amakihi, ‘I‘iwi, and ‘Apapane. Using mosquitoes carrying Plasmodium relictum,Atkinson et al. (1995) challenged 22 ‘I‘iwi from essentially mosquito-free zones. The birds were divided into low-dose (one mosquito bite only) and high-dose (4–10 bites) groups. Within 4 days, all had developed malaria, and by the eighth day the first death occurred. By day 37, all but one bird, the largest adult male in the low-dose group, had died. On average the low-dose birds survived longer, but interestingly the first bird to die came from that group.The lone survivor developed a chronic, barely detectable, low-level infection with P. relictum.When challenged again at day 167, that bird exhibited resistance to reinfection. Nutmeg Mannikins from lower elevations were used as controls in this study, and none developed malaria although the species is known to be susceptible (van Riper and van Riper 1985; Cann and Douglas 1999). Cann and Douglas (1999) reported that as few as two mosquito bites could produce 100% mortality in some populations of ‘Apapane and Hawai‘i ‘Amakihi, and that the numbers of parasites carried by individual mosquitoes in Hawai‘i are among the most extreme ever observed.

Anomalies Although Warner’s study was fairly convincing to most scientists, acceptance was not universal. Ian Atkinson (not to be confused with Carter Atkinson, a leader in ongoing research on avian disease in Hawai‘i) cited several anomalies that he thought could not be explained by Warner’s hypothesis (Atkinson 1977), and van Riper et al. (1986), who confirmed most of Warner’s findings, nevertheless did not believe avian malaria was present in Hawai‘i until long after the first wave of extinctions and declines. They viewed the extinction event as bimodal, with the declines on Lana‘i part of a second wave that was caused by avian malaria. However, recent discoveries and resultant reinterpretation of data cited by both Atkinson and the van Riper team show that these anomalies are easily explainable, and that avian malaria and avian pox, probably working synergistically, were, in fact, the cause of the holocaust as well as the ongoing threat to surviving honeycreepers that they are known to be. Both Atkinson (1977) and van Riper et al. (1986) believed, for different reasons, that the timing of various events was wrong for the ‘disease hypothesis’, at least for the 1890s. As Atkinson (1977: 120) states, ‘any hypothesis that relates the post-1892 accelerated decline of forest birds to disease requires evidence that either a new disease . . . or a new disease vector reached the Hawaiian Islands in the latter third of the last century.’ Now that van Riper et al. (1986) have shown that no reservoir of avian malaria had been lurking for centuries in migratory birds in Hawai‘i, Atkinson’s premise that avian malaria and pox must have been widespread in the islands long before 1892 does not hold up. Atkinson (1977) believed that Warner’s inability to document the spread of mosquitoes through the archipelago partly discredited the ‘disease hypothesis’ of Hawaiian bird disasters. But Atkinson’s supposition that mosquitoes were widespread in the islands long before the bird catastrophes is likewise undocumented, and his hypothesis that the roof rat Rattus rattus (Fig. 9.3) was the major causative factor suffers from the same flaws of timing that he imputes to Warner’s scenario, and has several other

164 The Hawaiian Honeycreepers

9.3 Roof rat caught in the act of depredation at a nest. Photo © Jack Jeffrey.

fatal conceptual flaws as well. For one thing, no one at the time reported an unusual rat plague, whereas in all other examples of rat-caused bird population crashes on islands (e.g. McCulloch 1921; Fisher and Baldwin 1946a), the cause was obvious at the time (Pratt 1994a). Secondly, Atkinson’s (1977, 1985) claim that R. rattus was not present in the Pacific before the late nineteenth century is demonstrably false: the species was established on at least one island in the Galapagos when Darwin visited in 1835 (Orr 1966); and it is the only rat present on Kosrae, Micronesia (Strecker et al. 1962), where the extinction of several birds before 1850 has been attributed to rats (Engbring and Pratt 1985). The van Riper team believed that avian malaria did not reach epizootic proportions until as late as the 1920s ‘following the numerous releases of introduced birds, particularly those from Asia’ (van Riper et al. 1986). However, two alien birds of Asian origin were well established long before that, and their introduction and spread coincides with the bird declines. The Nutmeg Mannikin was introduced by Hillebrand ‘about 1865’ (Caum 1933) on O‘ahu and soon became a pest in ricefields. The Spotted Dove appeared on O‘ahu at about the same time. Caum (1933) states that Chinese immigrants kept them for food purposes and escapes or purposeful releases were the source of the naturalised population. Both species (Fig. 9.4) became common on O‘ahu by 1879, at just the time when the island’s honeycreepers were disappearing. By

9.4 Alien birds implicated in the spread of avian diseases in Hawai‘i in the late nineteenth century. (a) Nutmeg Mannikin; (b) Spotted Dove.

the end of the century, both species had spread throughout the islands, and both species, then (Henshaw 1902) as now (Scott et al. 1986), commonly occur in forest openings and along roads and trails even in upland rainforests. A surprising recent discovery is that both species are among the five introduced birds with the highest prevalence rates for avian malaria on O‘ahu (Cann and Douglas 1999; Shehata et al. 2001). (The other three are two bulbuls found only on O‘ahu and the Whiterumped Shama found elsewhere only on Kaua‘i; all relatively recent introductions (Berger 1981).) We have no contemporaneous data on the spread of these two species, but comparisons with related birds can provide some inferences. The African Silverbill was released near Pu‘u Wa‘a Wa‘a on Hawai‘i, presumably in the early 1960s because by the time they were reported (Berger 1975a) the population was quite large. They colonised the Maui-nui group by the late 1970s and O‘ahu by

Diseases and parasites 165 1984 (Scott et al. 1986).A similar rate of inter-island spread for the Nutmeg Mannikin would have placed it on Hawai‘i at almost precisely the start of the bird declines, with population build-up tracking the event. Likewise, the Zebra Dove was introduced apparently around 1920, but did not reach The Big Island until 1937 (Schwartz and Schwartz 1949), where it first became established in Kona and then spread throughout the island. Thus the Spotted Dove could well have appeared in Kona coincidentally with a large increase in human population that Munro (1960) thought was related to a major 1892–94 bird decline.The habits of both the dove and the mannikin would help to explain Henshaw’s (1902) observation that birds disappeared from entire forest tracts when only the edge or a few trails had been cut. Other species introduced at about the same time that have been implicated as possible malaria reservoirs include Common Myna, House Finch, and House Sparrow (Scott et al. 1986), but recently mynas and House Finches were surprisingly found to be free of malaria on O‘ahu (Cann and Douglas 1999; Shehata et al. 2001). The only reason given by van Riper et al. (1986) for doubting the presence of avian malaria before 1900 was the fact that P. H. Baldwin did not find it in native birds in Hawaii Volcanoes NP in 1938–39, and ‘one surely would have expected it to be common on Hawaii island during the late 1930’s’ if it had been the ‘causative factor in the demise of native birds at the turn of the century’ (van Riper et al. 1986: 340). However, Baldwin did find malaria in a Red-billed Leiothrix and a California Quail, introduced birds collected at 1350 m and 2000 m elevation, respectively. The fact that Baldwin failed to find malaria at lower elevations where van Riper et al. (1986) found high prevalence forty years later could simply reflect the fact that distributions of both parasites and vectors are patchy and episodic, as shown by several recent studies (C. T. Atkinson, pers. comm.VanderWerf 2001).Although van Riper et al. (1986) felt it unlikely that Baldwin could have overlooked the parasite, a recently developed and highly sensitive test for avian malaria (Feldman et al. 1995) has revealed that Plasmodium is far more widespread in both native and introduced birds

than previously thought, but often at very low levels not readily detected by traditional means (Shehata et al. 2001). Although a bit hackneyed, the saying ‘absence of evidence is not evidence of absence’ comes to mind here. Perhaps new molecular techniques will find Plasmodium in nineteenthcentury honeycreeper specimens. Another problem is that van Riper et al. (1986), like Warner before them, treat mosquito distribution as if static. If Warner was correct about the upper limit in the late 1950s, instead of simply mistaken as implied by van Riper et al.’s (1986) comments about the difficulty of detection of Culex quinquefasciatus at higher elevations, then mosquitoes obviously made a significant upward advance in two decades. Extrapolating backwards two decades, mosquitoes may not yet have been present in sufficient numbers in Baldwin’s study area to support an epizootic.We can never know what elevation would have been the initial upper limit for tropical Culex, or the rate at which they adapted to higher elevations in the nineteenth century. Another aspect of mosquito distribution is that it is strongly affected by climatic events year to year (C. T. Atkinson, pers. comm.; Atkinson et al. 1993b) and even prevailing wind patterns (Scott et al. 1986). Without avian malaria as a possible cause, van Riper and van Riper (1985) and van Riper et al. (1986) were left to speculate that avian pox alone might have been responsible for the earlier population crashes of birds. Although the presence of pox was obvious in the 1890s, its epidemiology fails to explain all the observations of sick birds made at the time, particularly the sudden deaths of birds driven into the lowlands by storms. Pox simply takes too long when it kills. It also does not account for the birds found listless and with other overt symptoms of malaria but without obvious lesions. Although slow to act, pox can be fatal by itself (Scott et al. 1986) and, importantly, birds with pox are more likely to have malaria than those without it (van Riper et al. 1982, 1986). Another anomaly in the ‘disease hypothesis’ involves the sudden disappearance of a community of finches in high elevation koa and mamane/naio forests of Kona shortly after their discovery (Atkinson 1977).The Greater and Lesser Koa-Finches, the

166 The Hawaiian Honeycreepers Kona Grosbeak, and the Kona population of Palila, all disappeared so quickly (Henshaw 1902) that only an epizootic could adequately account for it. Enigmatically, their known ranges were mostly in places that even today are above the mosquito zone. However, all of these species (or close relatives) have recently been found in low elevation fossil sites ( James and Olson 1991; S. L. Olson, pers. comm.), indicating that the Kona finches may have been remnant populations surviving in marginal habitat and as such more vulnerable to diseases than larger, more widespread populations, or that they seasonally visited lowland habitats. Even without mosquitoes, they could have contracted avian pox from such birds as Hawaiian Crow and ‘O‘u, which roamed widely into and out of mosquito territory (Perkins 1893). Perkins (1893), however, found pox-like lesions on Greater Koa-Finches and Palila that he shot at lower elevations (down to ca. 1000 m) ‘within the rain-belt’ but none on birds from higher, drier elevations. According to Dennis LaPointe (in Tummons 2001a), habitat fragmentation is an important and previously under-appreciated factor in mosquito distribution in Hawai‘i. Coffee farming in Kona fragmented forest habitat up to 1000 m before 1892 (Perkins 1893). That, coupled with strong sea-breezes that blow mosquitoes to higher elevations during the day, offer a possible explanation for the apparent occurrence of mosquito-borne disease at unusually high elevations in Kona, where today the refugia for endangered birds are in places sheltered from high winds (Scott et al. 1986). So now we have at least a plausible scenario for the ‘Kona finch anomaly’.

The whole story Based on a wealth of circumstantial evidence, much of it accumulated since Warner’s (1968) study, we can now construct a reasonable scenario to explain the pattern of the appearance and effects of epizootic diseases in Hawaiian birds. Many of the details are missing, others are speculative, and a few anomalies remain, but it is quite clear that a convergence of independent forces fostered a trimodal epizootic that, over a period of a little more than a century, caused the demise or endangerment of the

majority of Hawaiian honeycreeper species. The major points can be summarised as follows. 1. Hawaiian honeycreepers evolved in the absence of mosquitoes and therefore were totally naive to mosquito-borne infections. 2. A tropical strain of Culex quinquefasciatus was introduced to the Hawaiian Islands in 1826 and spread throughout the archipelago at elevations suited to its origin. 3. Avian pox and avian malaria are not host specific and are among relatively few diseases capable of infecting a broad spectrum of bird species. 4. Poxvirus avium, the avian pox pathogen that is spread by both mosquitoes and direct contact, was introduced some time after the late 1700s. 5. Plasmodium relictum, the cause of avian malaria, was brought to O‘ahu in birds, probably Nutmeg Mannikins and/or Spotted Doves, imported from the Orient around 1865. 6. Hawaiian honeycreepers are extremely vulnerable to avian malaria, which in many species can be 100% fatal. 7. As the vectors and diseases found each other, birds on O‘ahu, an island with relatively few areas high enough to be above the mosquito zone, began to succumb and by 1890 the island’s ‘Akepa, ‘akialoa, nukupu‘u, and ‘O‘u (plus non-drepanidine solitaire and ‘o‘o) were nearly or totally extirpated. Surviving species became restricted to high elevation forests.This was the first wave. 8. Nutmeg Mannikins and Spotted Doves spread to other islands in a stepwise fashion, reaching Hawai‘i by the 1880s and becoming common there by the 1890s. Their habits of entering even dense forests along trails and roads or in natural openings brought the deadly combination of pathogens, reservoir hosts, and vectors into contact with native forest birds, at least in the lowlands. 9. Native birds with no natural immunity to the diseases succumbed quickly to the combined effects of pox and malaria. Species confined to the lowlands or to the lowest forest belt at the time, or that regularly visited these areas (examples include Hawai‘i Mamo, Greater ‘Amakihi,

Diseases and parasites 167 Hawai‘i ‘Akialoa, and possibly ‘Ula-‘ai-hawane and the Kaua‘i and Maui nukupu‘us) went extinct. Sedentary species that inhabited a broader elevational range centred on the lowlands (such as Maui’s now-endangered species) became rare. Species that inhabited all elevations disappeared from the lower parts of their ranges. Elevations up to at least 400 m became pestilential death traps.These declines occurred over a period of about two decades, although effects in local areas could be seen in as little as 2 years. Events on Hawai‘i were fairly well documented, but those on Moloka‘i, Maui, and Kaua‘i must be surmised in retrospect because few observers were present at the time to document what happened. Some lowland species may have existed on those islands prior to the 1880s but died out before adequate ornithological explorations were done (Olson and James 1994a). These effects were delayed on Lana‘i by about two decades perhaps because the island had very few human inhabitants before 1923, but almost its entire native avifauna disappeared by 1940.This constituted the second wave. 10. As the tropical Culex slowly adapted to cooler elevations, the bird die-offs slowly advanced up the mountains, reaching 600 m by the 1950s, and 1500 m by the 1970s, sweeping away such mid-elevation species as ‘O‘u, Kaua‘i ‘Akialoa, Kakawahie, Black Mamo, and O‘ahu ‘Alauahio that had survived the turn-of-the-century epizootic, and restricting most of the other survivors to ever higher refugia, which in some cases are probably only marginal habitat.This is the third and ongoing wave.

Malaria and pox in honeycreepers today A few honeycreepers, or a few subpopulations of honeycreepers, are beginning to show evidence of resistance to both malaria and pox (van Riper 1991a; van Riper and van Riper 1985).Whether such resistance can increase in those species that have it and develop in others soon enough to prevent further

declines and extinctions remains to be seen ( Jarvi et al. 2001; Shehata et al. 2001).What is clear is that avian malaria and avian pox are still threats to surviving honeycreeper species (Tummons 2001b) and will become more acutely so as mosquitoes continue their slow upward advance (Tummons 2001a). Recent observations have shown that the prevalence of malaria and pox varies greatly from year to year, and is characterised by periodic epizootics in which large numbers of birds in the mosquito contact zone become ill. C. T. Atkinson pers. comm. observed such an epizootic in September–December 1992 at ca. 1200 m in Hawaii Volcanoes NP and nearby areas, apparently with an epicentre in the ‘Ola‘a Forest tract. It followed a dramatic increase in mosquitoes in the area, and infected Culex were found as high as 1700 m. Many ‘Apapane and a few Hawai‘i ‘Amakihi (the only honeycreepers still common in the area) were found dead or dying along roads and trails, the incidence of malaria in mist-netted birds increased to almost 100%, and sentinel birds in the same habitats became infected. Most of the dead birds had both acute malaria and secondary infections from necrotic pox lesions, a clear indication of the cumulative effect of the two diseases. Probably this same epizootic produced pox-like lesions in the non-drepanidine Hawai‘i ‘Elepaio at Hakalau Forest NWR on the windward slope of Mauna Kea (VanderWerf 2001). Infection rates in introduced birds in the Volcano/‘Ola‘a area never exceeded 10% even at the height of the epizootic. ‘Apapane and Hawai‘i ‘Amakihi exhibited mortality rates of ca. 50%, showing that some individuals could survive the diseases, but populations were slow to recover in the area. Surviving birds were immune to reinfection, but served as a reservoir for transmission of disease during future epizootics. Some resistance is clearly evolving and both Hawai‘i ‘Amakihi and ‘Apapane appear to be expanding their populations in low-elevation forests. In a study comparing subpopulations of Hawai‘i ‘Amakihi from dry forests on Mauna Kea and rainforests on Mauna Loa, van Riper (1991a) found that the rainforest populations, which lived in contact with the vectors, had developed greater tolerance of both malaria and pox than their conspecifics from

168 The Hawaiian Honeycreepers the mosquito-free zone. Other populations of Hawai‘i ‘Amakihi, which are sedentary, unlike ‘Apapane and ‘I‘iwi that return to higher elevations at night, are apparently also developing resistance.They are common below 600 m at Pu‘u ‘Anahulu in the South Kohala District and in the Manuka NAR in South Kona, the latter area particularly heavily infested with mosquitoes (pers. obs.). I have also observed a Hawai‘i ‘Amakihi at about 120 m in ‘Iao Valley,W. Maui, but that sighting was exceptional, and Pyle and Donaldson (2000b) cite reports from coastal localities in E. Maui. Pimm (1991: 342) reported an apparently resistant population of ‘several species in a small patch of native forest at 300 m on the island of Hawai‘i’ but did not give the exact location. The greatest amount of resistance to malaria, and the best documented, is found in the O‘ahu ‘Amakihi, which had been declining and contracting in range for decades (Ralph 1990b). Only a short time ago, the appearance of an O‘ahu ‘Amakihi in Manoa Valley was a noteworthy event (Howarth 1984). But the bird has become increasingly common in recent years at or below 300 m in the valleys behind Honolulu (Williams 1987; Pratt 2002b). Shehata et al. (2001) tested 40 ‘amakihi from Manoa Valley, a wet locality with high populations of mosquitoes, and found none with avian malaria. In a subset examined later by C. T. Atkinson (Cann and Douglas 1999), a single individual showed evidence of a past malarial infection.At the same time, other birds in the area, mostly introduced species, showed low but consistent levels of malaria. These findings show that although O‘ahu ‘Amakihi are still susceptible, they can somehow clear the parasite from their bodies. Just how they do it is the subject of ongoing research (Cann and Douglas 1999; Jarvi et al. 2001).These observations are exactly what we would expect if the above historical scenario is true. Perhaps simply by luck, ‘amakihi have a higher initial level of resistance to avian malaria than other honeycreepers or they would not be the great survivors they appear to be. Thus the initial exposure selects enough resistant individuals to maintain the species, but other species have too few survivors (or none at all) to avoid extinction. If malaria first appeared on O‘ahu, as I speculate above, then the O‘ahu ‘Amakihi

would have had longer than any other surviving honeycreeper species to evolve sophisticated means of resistance to Plasmodium. The loss of so many species from the island is indeed tragic, but the fact that at least one species of honeycreeper can now be easily seen only a few minutes from downtown Honolulu (Pratt 2002b) is heartening. Some evidence also exists to indicate that natural selection in Hawai‘i is favouring weaker or less virulent strains of both malaria and pox. Such extreme mortality of a host species, as has been demonstrated for Hawaiian honeycreepers, is clearly not in the best evolutionary interests of the parasite. If all the hosts die, so do the parasites and the cycle of transmission ends. Consequently, one might expect some equilibrium between virulence and susceptibility to evolve (Anderson and May 1982; Ewald 1994). Hawaiian strains of Plasmodium relictum are among the least virulent known (van Riper 1991b), despite their devastating effects in some species (strains of P. relictum in California are more virulent by several orders of magnitude). An unpublished report by R. Nakamura (discussed in Pimm 1992: 342–3) ‘found that substantial numbers of quite healthy native birds were infected with malaria and that an individual bird had lived for over five years with an infection’, so less virulent strains of Plasmodium are perhaps being selected. Avian pox occurs in two forms: the more common one that produces swellings and lesions on the extremities, and a more debilitating diphtheric form with less obvious external symptoms. In xeric habitats on Mauna Kea, where mosquitoes do not occur and pox is rare, the more severe form predominates in Hawai‘i ‘Amakihi, whereas the same species in rainforest within the mosquito zone of the same island exhibit mostly the less debilitating form (van Riper 1991a). Even the less virulent form apparently produces less debilitating lesions in infected birds today than it did historically ( Jarvi et al. 2001; van Riper et al. 2002). Obviously, those strains that could live without killing their hosts were favoured by natural selection, so less virulent ones have come to predominate in Hawaiian birds. This partly explains why introduced birds in Hawai‘i are rarely adversely affected by these diseases.

Diseases and parasites 169

Other diseases and parasites Known diseases and parasites of Hawaiian birds were catalogued by van Riper and van Riper 1985. Table 9.1 (and much of the following discussion) is compiled largely from their work. The first avian parasites in Hawai‘i undoubtedly arrived with colonising birds, and subsequently co-evolved with their hosts. Because selection favours less pathogenic strains of parasites, the endemic parasites have reached a benign equilibrium with their hosts.They also tend to be host-specific. Most introduced parasites likewise pose little threat to honeycreepers because they rarely have the ability to infect species other than their host. Nor are all parasites that infect

a large spectrum of hosts always detrimental. One of the most widespread avian parasites in Hawai‘i is the eyeworm Oxyspirura mansoni, which rarely causes serious harm. Although six species of Plasmodium have been reported in the Hawaiian Islands, Laird and van Riper (1981) showed that only P. relictum is present and that the other reports were based on misidentifications of unusual morphological types of that species. Several broad-spectrum diseases that could potentially reach Hawai‘i, including arboviruses, Newcastle disease, and avian influenza, are apparently not yet present (Quisenberry and Wallace 1959; Wallace et al. 1964; van Riper and van Riper 1985), but pose a continuing future threat.

Table 9.1 Known parasites and diseases of Hawaiian honeycreepers. Pathogen and classification Diseases, etc. Viruses Poxvirus avium Bacteria Citrobacter freundii Enterobacter cloacae Escherischia coli Klebsiella pneumoniae Pseudomonas aeruginosa Pseudomonas pseudoalcaligens Salmonella sp. Serratia marcescens Staphylococcus epidermis Streptococcus sp. Fungi Aspergillus fumigatus Candida albicans Endoparasites Protozoa Isospora loxopis Isospora sp.

Host species

References a

Probably all

1, 2

Drepanis coccinea Himatione sanguinea Himatione sanguinea Telespiza cantans (captive) Himatione sanguinea Telespiza cantans (captive) Himatione sanguinea Loxioides bailleui (captive) Telespiza cantans (captive) Telespiza cantans (captive) Drepanis coccinea Telespiza cantans (captive)

2 2 2 2 2 2 2 2 2 2 2 2

Hemignathus virens (captive) Telespiza cantans (captive) Himatione sanguinea

2 2 19

Hemignathus virens Magumma parva (captive)

3 4

170 The Hawaiian Honeycreepers Table 9.1 contd. Pathogen and classification

Host species

References a

Plasmodium relictum Trichomonas gallinae Nematoda Capillaria sp.

Probably all Himatione sanguinea

5, 18 2

Hemignathus virens Himatione sanguinea Drepanis coccinea Himatione sanguinea Hemignathus virens Himatione sanguinea Hemignathus virens Drepanis coccinea

2, 4 2, 6 2 6 4 6 6 6

Hemignathus stejnegeri

7

Hemignathus virens Himatione sanguinea Drepanis coccinea Hemignathus stejnegeri

2, 8, 9 2, 8, 9 2, 8, 9 7

Hemignathus virens Himatione sanguinea

2 2

Telespiza ultima Hemignathus virens Drepanis coccinea Himatione sanguinea Himatione sanguinea Himatione sanguinea Telespiza ultima Himatione sanguinea Drepanis coccinea Hemignathus virens Telespiza ultima Hemignathus virens Himatione sanguinea Telespiza ultima Drepanis coccinea Magumma parva Himatione sanguinea Telespiza ultima

2 12 12 12 12 12 2 12 12 12 11 12 12 2 12 13 12 11

Procyrnea longialatus Tetrameres sp. Viquiera hawaiiensis

Acanthocephala Apororhynchus hemignathi Cestoda Anonchotaenia brasilense

Drepanidotaenia hemignathi Trematoda Urotocus rosittensis Ectoparasites Acari Analges sp.

Androlaelaps sp. Anhemialges Boydaia agelaii Calcealges sp. Cheyletus eruditus Cheyletus malaccensis Eutrombicula conantae Harpyrhynchus sp. Ingrassiella sp. Megnina sp. Mouchetia sp. Neoschoengastia ewingi

Diseases and parasites 171 Table 9.1 contd. Pathogen and classification Ornithonyssus sylviarum Proctophyllodes sp.

Pterodectes sp. Ptilonyssus sp. Ptilonyssus (2 spp. nov.) Rhinonyssus sp. Schoengastia pobsa Toritrombicula nihoaensis Trouessartia sp. Mallophaga Colpocephalum hilensis Machaerilaemus hawaiiensis Myrsidea cyrtostigma Philopterus macgregori Hippoboscidae Ornithoica vicina a References: 1

Host species

References a

Hemignathus virens Telespiza ultima Hemignathus virens Drepanis coccinea Himatione sanguinea Hemignathus virens Himatione sanguinea Hemignathus virens Himatione sanguinea Hemignathus virens Telespiza cantans Telespiza ultima Hemignathus virens

12 2 12 12 12 12 12 12 12 4 10, 11 11 12

Drepanis coccinea Hemignathus virens Drepanis coccinea Himatione sanguinea Hemignathus virens

14 14 14, 17 14 14

Hemignathus virens Drepanis coccinea

15, 16 15, 16

Henshaw 1902; 2 van Riper and van Riper 1985; 3 Levine et al. 1980; 4 van Riper 1975; 5 van Riper et al. 1986; 6 Cid del Prado Vera et al. 1985; 7 Perkins 1903; 8 Baldwin 1948; 9 Voge and Davis 1953; 10 Brennan and Amerson 1971; 11 Goff 1984; 12 Goff 1980; 13 Haramoto in Berger 1981; 14 Zimmerman 1948; 15 Bequaert 1941; 16 Maa 1966; 17 Alicata 1969; 18 Navvab Gojrati 1970; 19 Miyahara in van Riper and van Riper 1985.

10 Status, conservation, and the future

When I first visited Hawai‘i in 1974, I was optimistic about the Hawaiian honeycreepers. After all, even though many were very rare, several had been rediscovered in the previous decade, and a new one had just been found on the windward slope of Haleakala. Surely, it seemed, more supposedly extinct species still clung to existence in some remote forest refuge where no one had looked for many years. I was naive enough to think that those that had lasted until ‘enlightened’ modern times would get the help they needed and survive. I never expected to study birds that would be extinct in my lifetime. But my youthful optimism has faded in the cold light of reality as one remote forest after another has been thoroughly searched with no new ‘rediscoveries’ and one species after another has declined and disappeared like candles winking out in the night. I never saw the Kakawahie (a 1975 expedition to Moloka‘i (Scott et al. 1977) in which I participated failed to find it only 12 years after the last sighting) or the O‘ahu ‘Alauahio (I participated in searches for it in 1977–78 that were successful (Shallenberger and Pratt 1978), but it was not I who won that particular birding lottery). I have been ‘a day late or a dollar short’ at three possible nukupu‘u sightings, two on Kaua‘i and one on Maui, and failed to find the Kaua‘i ‘Akialoa, seen only 6 years before by my colleague Phil Bruner. But I had wonderful never-to-be-forgotten field experiences tape-recording and photographing the ‘O‘u (as well as those other now lost nondrepanidine Alaka‘i birds, the Kama‘o and ‘O‘o‘a‘a (Conant et al. 1998)), the then newly discovered Po‘o-uli, and the ‘Akikiki, then relatively common, now becoming increasingly difficult to find (Pratt

2002b). Now I find myself talking to younger field ornithologists about ‘the good old days’, taking the role of Dave Woodside whose yarns fascinated me, a wide-eyed malihini those many years ago. What will remain of the Hawaiian honeycreepers a quarter century hence? The extinction dirge of Hawaiian honeycreepers, replayed in Chapters 2 and 9, has been widely publicised in the popular press (e.g. Berger 1975d, 1977; Gagne 1975; Yates 1984; Brower 1989; Wilcove 1989, 1999; Applebaum 1990; Bloch 1991; Collins 1993; Rogers and Jeffrey 1993). In this chapter we will examine where we are today, what is being done to slow the seemingly inexorable march to extinction, and what the future may hold. Table 10.1 lists the historically known honeycreepers that are either extinct or of conservation concern, and gives an assessment of current status. Sadly, this book may be the first to announce the probable extinction of ‘O‘u, O‘ahu ‘Alauahio, and the nukupu‘us, and to state flatly that Kakawahie and Kaua‘i ‘Akialoa, though still listed as endangered, are undoubtedly extinct. At some point, we have to admit defeat. Details of declines and current status for individual species are given in the species accounts (Part II).

Honeycreeper conservation after the Endangered Species Act The Endangered Species Act A major landmark in Hawaiian honeycreeper conservation was the US Endangered Species Act (ESA) of 1973 (Martin 1994). Over half of the

Status, conservation, and the future 173 original ‘default’ list of birds for the US were Hawaiian, and the majority of those were honeycreepers. The list was comprehensive at the time and only one species (Hawai‘i Creeper in 1975) has been added since (USFWS 1999a), although Jeff Foster (pers. comm.) says the ‘Akikiki has been proposed for listing. This landmark legislation

focused public attention nationwide on the concept of species loss, and brought the federal government into the conservation of Hawaiian species for the first time.1 The ensuing decade brought a flurry of interest on the part of state (DOFAW), federal (NPS, USFWS, USFS), and private (HAS, Sierra Club, TNCH) agencies and organisations,

Table 10.1 Conservation status of extinct and imperilled Hawaiian honeycreepers (historically known species only). Species or population

Federal a

State b

IUCN c

Last record d

Po’o-uli O‘ahu ‘Alauahio

E E

E E

CR CR

Kakawahie Laysan Finch Nihoa Finch Palila Greater Koa-Finch Lesser Koa-Finch Kona Grosbeak ‘O‘u

E E E E X X X E

E E E E X X X E

EX VU CR EN EX EX EX CR

Lana‘i Hookbill ‘Akikiki Hawai‘i Creeper ‘Akepa ‘Akeke‘e Greater ‘Amakihi Kaua‘i ‘Akialoa O‘ahu ‘Akialoa

NL NL E E NL X E X

NL NL E E NL X E X

EX CR EN EN EN EX EX EX

Maui-nui ‘Akialoa Lesser ‘Akialoa Kaua‘i Nukupu‘u

X X E

X X E

EX EX CR

O‘ahu Nukupu‘u Maui Nukupu‘u

X E

X E

EX CR

‘Akiapola‘au Maui Parrotbill Laysan Honeycreeper ‘Akohekohe Hawai‘i Mamo

E E X E X

E E X E X

EN VU NL VU EX

NA 1985 (1990) 1963 NA NA NA 1895 1892 1895 1989 (1996) 1918 NA NA NA NA 1901 1969 1892 (1939) 1894 1940 1899 (1995) 1860s 1890s (1996) NA NA 1923 NA 1898

Reference e

7 8 9

10 10 11 12 13 14

15 16 17 18 17 19 20 16 21 16 16

22 24

Extinct? f no probably yes no no no yes yes yes probably yes no no no no yes yes yes yes yes yes yes probably no no yes no yes

174 The Hawaiian Honeycreepers Table 10.1 Contd. Species or population

Federal a

State b

IUCN c

Last record d

Reference e

Extinct? f

Black Mamo ‘I‘iwi ‘Ula-‘ai-hawane

X NL X

X Eg X

EX NT EX

1907 NA 1892 (1937)

23

yes no yes

25 21

1999a. Code: E ⫽ Endangered; NL ⫽ not listed; X ⫽ presumed extinct. Carol J.Terry (pers. comm.), DLNR, DOFAW. Code as above. c BirdLife International (2000). Status Code (p. 21): EX ⫽ Extinct; CR ⫽ Critically endangered (very high extinction risk in the wild in immediate future); EN ⫽ Endangered (very high extinction risk in near future);VU ⫽ Vulnerable (high extinction risk in medium-term future); NT ⫽ Near Threatened (close to qualifying for Vulnerable). d Year only. Questionable subsequent reports in parentheses. NA ⫽ species frequently seen or closely monitored. e Author’s opinion based on information as of April 2004. f References: 7 Bremer 1986; 8 P. and H. Baker 2000; 9 Pekelo 1963; 10 Olson 1999; 11 Grant 1995; 12 Pyle 1989a; 13 Reynolds and Snetsinger 2001; 14 Perkins 1919; 15 Lindsey et al. 1998; 16 Conant et al. 1998; 17 Perkins 1903; 18 Northwood 1940; 19 Baldwin 1941; 20 Pratt and Pyle 2000; 21 Munro 1960; 22 Wetmore 1923; 23 Bryan 1908; 24 Henshaw 1902; 25 Rothschild 1893–1900. g O‘ahu; Moloka‘i; and Lana‘i only. a USFWS b Fide

sometimes in co-operation (Anon. 1991) and sometimes in contention (as in the case of conservation organisations suing the state over sheep in Palila habitat, Chapter 2).

Assessments and surveys Much of the early work simply involved documenting how little we actually knew (Ralph 1977; Scott et al. 1979, 1980) and developing data-gathering techniques (Reynolds et al. 1980; Ralph 1981a,b; Ralph and Scott 1981; Scott et al. 1981a).The huge HFBS effort to gather baseline data on endangered honeycreepers and other native birds (Scott et al. 1986), detailed in Chapter 2, resulted in a ‘classic in conservation biology’ (Pimm 1988) upon which future researchers could effectively build. As a result of these efforts, an impressive number of papers dealing with the problems and potential solutions were published (e.g. Kepler and Scott 1985; Scott et al. 1985, 1987a,b, 1988b, 1989; Stone and Scott 1985a; Mountainspring 1986; Stone and Loope 1987;Vitousek et al. 1987; Gagne 1988; van Riper and Quinn 1988; Freed and Cann 1989).We clearly know what the problems are, and we do not suffer from a shortage of recommendations as to how to proceed, but consensus has been elusive. Some

conservationists favour ecosystem-level approaches (Scott et al. 1988b; Conant 1998) while others emphasise the need for focused efforts to salvage critically endangered species within ecosystems (Freed and Cann 1989). I agree with Martin (1994) that this is a false dichotomy and, in fact, the two camps agree more than they disagree (Scott et al. 1989).Today’s efforts to save the remaining Hawaiian honeycreepers involve a mix of ecosystem-level and species-focused efforts (Banko et al. 2001). A limited 1994–96 reprise of the HFBS (Reynolds and Snetsinger 2001) found deterioration in virtually every parameter. For species such as the ‘O‘u, that might have been saved by heroic efforts a decade earlier, things have deteriorated so rapidly that any further efforts on their behalf would be futile. Nevertheless, some limited progress has been made as discussed in the following review.

Parks, refuges, and preserves Hawai‘i had fairly extensive areas in various parks and preserves before 1973. Hawaii NP, later divided into Hawaii Volcanoes NP on The Big Island and Haleakala NP on Maui, had been in existence since 1916. State parks included Koke‘e SP on Kaua‘i, where several species of honeycreepers were seen

Status, conservation, and the future 175 regularly (Berger 1972b), and on the same island the Alaka‘i Wilderness Preserve had been established by the state in 1961 following the bird discoveries of Richardson and Bowles (1964). Thus Kaua‘i was the only island whose best honeycreeper habitat was already protected in 1973. The Nature Conservancy had acquired Kipahulu Valley for Haleakala NP after a successful 1967 expedition (Banko 1968; Matthiessen 1970), but was stymied in further efforts to locate suitable places for acquisition by lack of information (Little 1984; Scott et al. 1987b). The state established a Natural Area Reserve System (NARS) in 1970, but the first areas set aside were of little interest to ornithologists and the entire system was neglected for years after it was begun (Dawson 2001). The 1976–83 HFBS (Scott et al. 1986) provided the needed data, and TNCH, who had been kept apprised of findings long before they were published, began an ambitious program of preserve acquisition. Their first success was Kamakou Preserve on Moloka‘i, which turned out not to harbour any endangered honeycreepers but to be valuable to other native organisms and a catalyst for cooperative efforts. Nearby Oloku‘i Plateau (Fig. 2.4b) became a natural area reserve (NAR) in 1986 and Kalaupapa National Historical Park, which includes a sizeable wilderness area adjacent to Kamakou Preserve, was established the same year. Later TNCH acquired Pelekunu Valley to tie all these areas together in a 7783 ha wilderness complex (Scott et al. 1987b). It did not do much for Moloka‘i’s honeycreepers, coming about two decades too late, but it set a valuable precedent for co-operative arrangements elsewhere. A similar scenario unfolded on Maui, where TNCH acquired Waikamoi Preserve adjacent to Haleakala NP in 1983 and the state established Hanawi NAR (Fig. 10.1), also adjacent to the national park, in 1986. This time, honeycreepers were big winners, with six species in Waikamoi and seven in Hanawi. These units now form the heart of the East Maui Watershed Partnership, a coalition of public and private agencies and individuals working together to preserve the forests of East Maui in the best interests of everyone, including Hawaiian honeycreepers in remote upland areas (Carol Gentz,TNCH, pers. comm.).

10.1 View looking west along windward slope of Haleakala, from the heart of Hanawi NAR, Maui. Ko‘olau Gap can be seen in the distance.

On Hawai‘i, results of the HFBS suggested the need for a refuge at mid-elevation on windward Mauna Kea, where the highest densities of endangered forest birds, mostly honeycreepers, were found (Scott et al. 1986).As a result of the work of landowners, politicians, biologists, and conservationists, Hakalau Forest NWR (Fig. 10.2) was established in 1986 (Scott et al. 1987a), the first in the nation solely for forest birds. Recently, a satellite refuge has been established in South Kona on a parcel formerly part of the McCandless Ranch (Pratt 2002b).Although its primary purpose was to provide habitat for the Hawaiian Crow, it also harbours small populations of ‘Akepa, Hawai‘i Creeper, and ‘Akiapola‘au. The remaining McCandless property adjacent to the refuge is also being managed for native species because their ecotourism business (Pratt & Jeffrey 1996b; Pratt 2002b) now earns more than their cattle operations (K. Unger, pers. comm.), or at least it did before the area was forsaken by the last Hawaiian Crows. Elsewhere on Hawai‘i, the ‘Ola‘a-Kilauea Management Area is an arrangement similar to that on East Maui in which Kamehameha Schools Bishop Estate, Hawai‘i Volcanoes NP, Hawai‘i DLNR, and Hawai‘i Dept. of Public Safety are co-operating with USFWS, USGS, and USFS in conservation management. This area is the keystone that connects the Hamakua forests with those of the Kilauea-Ka‘u area to form a continuous band of ‘essential habitat’ that may insure long term survival of the island’s seven remaining honeycreepers (Banko et al. 2001).

176 The Hawaiian Honeycreepers

10.2 View downslope from pasturelands above into upper forest edge of Pua ‘Akala section of Hakalau Forest NWR, Hawai‘i.

Management of limiting factors Disease By far the most serious limiting factor for honeycreepers is avian disease (Scott et al. 1986; Cann and Douglas 1999;Altizer et al. 2001; Jarvi et al. 2001; see also Chapter 9). Indeed, if this one factor were removed, all but a few species currently listed as endangered would be able to recover to healthy population levels and expand into formerly occupied lowland habitats in a relatively short time, despite all the other environmental perturbations ( Jarvi et al. 2001). (For that matter, were it not for disease, most of the species that have gone extinct since 1960 would probably still be with us.) Until recently, however, avian disease in Hawai‘i has been like the weather, frequently discussed but with no effort to manage or control it. Managers simply assumed that it was something the birds had to live (or die) with. Now, a series of studies planned by PIERC (C. T. Atkinson, pers. comm.) holds the promise of new ways to control malaria and pox in wild bird populations. One project, funded by the National Science Foundation with participants from PIERC, University of Hawai‘i, National Wildlife Health Center, Smithsonian Institution, and Princeton University, will use a holistic modelling approach to gather data on the epidemiology of avian diseases, demography of host populations, distribution and population dynamics of vectors, and

methods for controlling vector populations on the east slope of Mauna Loa. The study area includes large portions of Hawai‘i Volcanoes NP, Pu‘u Maka‘ala and Kahauale‘a NARs, and several state forest reserves as well as private land. Investigators will also examine possible co-evolution of parasite virulence, vector susceptibility, and host resistance.A second study, funded by NPS, will determine the prevalence of avian pox and malaria in both native and introduced birds in Kipahulu Valley of Haleakala NP on Maui and Kalaupapa NHP and adjacent Kamakou Preserve on Moloka‘i. An effort will be made to learn how weather patterns and geographic features influence distribution of mosquitoes, with the hope of determining effective control measures for remote areas. A third study, also funded by NPS, will focus specifically on source reduction to limit mosquito populations with the goal of creating an avian disease-free zone in Hawai‘i Volcanoes NP, in which three endangered honeycreepers (‘Akepa, Hawai‘i Creeper, and ‘Akiapola’au) that formerly occurred there can recover.

Feral ungulates Until methods are found to limit avian disease, preserve management will focus on limiting factors that can be controlled now. Considerable success has been achieved in the control or elimination of feral ungulates in a few relatively small areas (Stone and Loope 1987; Anderson and Stone 1993; Katahira et al. 1993; Banko et al. 2001), but ‘their removal should be the highest management priority in Hawaiian bird habitats’ (Banko et al. 2001: 372). Note that pigs have a synergistic effect on avian disease by providing breeding sites for mosquitoes. As discussed in Chapter 2, feral cattle are not the problem in Hawaiian forests that they once were, but have not been entirely eliminated. Hundreds of feral cattle have been removed from a fenced portion of Hakalau Forest NWR since 1988 (Maxfield 1998), but substantial numbers persist in South Kona forests (K. Unger, pers. comm.).The virtual elimination of goats from both national parks required the building of fences around the perimeters and intensive hunting (Stone and Loope 1987) but appears to have been effective (Fig. 10.3). Feral sheep and

Status, conservation, and the future 177

10.3 View toward Mauna Loa from Jaggar Museum on rim of Kilauea Caldera, showing clearly the boundaries of the fenced Mauna Loa Strip of Hawaii Volcanoes NP with recovering forests, in contrast to grazed lands on either side.

mouflon have been a problem mainly in the mamane–naio habitat of the Palila (Scowcroft and Giffin 1983), but now are being removed despite the protestations of hunters who, in the face of overwhelming evidence to the contrary, claim that ungulates benefit the Palila by reducing fire hazard from dry weeds and grasses (Tummons 1999b, d, 2001c; Dawson 2001). Catch-22, of course, is that the same ungulate that ‘controls’ weeds and grass also relishes mamane seedlings. Fig. 10.4 shows the result of uncontrolled grazing in mamane forest adjacent to the now fenced area around Pu‘u La‘au. Palila no longer are found in this area, but at least fire is not a concern, with little left to burn! Regeneration of the forest will require additional management, such as planting of mamane seedlings in some areas to overcome effects of past browsing and alien grass invasion (Hess et al. 1999). Such efforts require many years before benefits to avian habitat are realised (Scowcroft and Conrad 1988). Hunting interests have been particularly problematical in the case of feral pig control, opposing fencing and eradication efforts everywhere (Tummons et al. 1997; Joyce 1999a; Dawson 2001), even to the point of vandalising fences (Anon. 1997), despite the long history of such management ( Joyce 1999c). Though the damaging effects of pigs (Fig. 10.5) have been well documented (Ralph and Maxwell 1984; Stone and Loope 1987; Atkinson et al. 1995; van Riper and Scott 2001), hunters have expressed the

10.4 View ne. from Pu‘u La‘au on Mauna Kea, showing results of intensive grazing in mamane–naio forest.

belief that pigs are actually good for the forest and that evidence to the contrary is fabricated by researchers (Tummons et al. 1997; Joyce 1999a). (Exactly why researchers would want to fabricate evidence against a harmless or beneficial animal has apparently not been addressed.) One would think that hunters would be the conservationist’s ally in controlling feral pigs, but such is not the case. In portions of Hakalau Forest NWR that were open to public hunting, pig populations actually increased (Tummons et al. 1997). Hunted populations had an initial decrease, but then bounced back to prehunting levels because once pigs become scarce, hunters move to other more productive areas, leaving the remaining animals to regenerate rapidly ( Joyce 1999b). On the other hand, the effectiveness of fencing and systematic hunting and snaring for eliminating pigs in remote areas has been clearly demonstrated (Anderson and Stone 1993; Katahira et al. 1993; T. K. Pratt et al. 1997; Joyce 1999b). At least portions of Hawai‘i Volcanoes NP, Hakalau Forest NWR, Haleakala NP (upper Kipahulu Valley), and Hanawi NAR are now pig-free, as are TNCH’s Waikamoi Preserve and Kamakou Preserve (Banko et al. 2001), but removal of pigs from areas large enough to be ecologically significant remains a distant goal (van Riper and Scott 2001).

Mammalian predators Although rats may not have been the main cause of the late nineteenth-century bird declines, as some

178 The Hawaiian Honeycreepers suggested (Atkinson 1977; see Chapter 9), they are serious predators of Hawaiian honeycreepers and

also may impact them by competing for food and altering the plant community (Banko et al. 2001). The aerial broadcast of rodenticides, which has had great success in New Zealand, is being considered as a control measure, but gaining the necessary government permits for such actions is ‘a long and arduous process’ (Reilly 1998: 17). Rats are currently being reduced in several areas important to Hawaiian honeycreepers on Hawai‘i and Maui (Banko et al. 2001), and have been eradicated from Midway Atoll (R. J. Shallenberger, pers. comm.), a possible translocation site for Laysan Finch (see below). Eradication of rats from Kaho‘olawe now appears feasible, and will probably be necessary if that island is to become a translocation site for honeycreepers. As Banko et al. (2001) point out, control of rats does not engender public outcry, but feral cats are a different matter. Cats are probably predators of all surviving Hawaiian honeycreepers, but they have been particularly damaging to Palila (Snetsinger et al. 1994; T. K. Pratt et al. 1997). Control measures will require considerable diplomacy because Hawai‘i’s ‘cat lobby’ is probably as powerful as the pig hunters. Recently, the City and County of Honolulu appropriated US $30 000 to set up feeding stations for feral cats in city parks (Bickel 2001)! The only good news here is that the stations are to be used to round-up cats for neutering and release, small comfort to the birds they will consume after the operations, but perhaps a step toward control of feral populations. Mongooses, like rats, receive little public sympathy (no one is setting up feeding stations for them!). Despite their clearly documented damage to ground-nesting birds such as Nene, they are not known to impact any Hawaiian honeycreepers.

Population management Translocation 10.5 Damage to native forests by feral pigs: (a) forest floor on either side of exclosure fence comparing pigfree (left) and pig-damaged (right) areas; (b) downed trunk of hapu‘u tree fern with pithy core removed by pigs, creating ideal breeding sites for mosquitoes. Photos © Jack Jeffrey.

Translocation, physically moving birds into unoccupied but presumably suitable habitat, has had considerable success as a management tool in some parts of the world, most notably New Zealand (Armstrong and McLean 1995). There, however, native birds are not faced with the elevational

Status, conservation, and the future 179 limitations imposed by disease in Hawai‘i, and offshore islands are abundant and well vegetated. Hawai‘i simply does not have many potential translocation sites.The first experimental translocations of Hawaiian honeycreepers involved the introduction of Laysan Finches to Pearl and Hermes Reef in 1967 (Amerson et al. 1974), and Nihoa Finches to French Frigate Shoals in 1969 (Amerson 1971; Sincock and Kridler 1977). The former became established, but the Nihoa Finch experiment failed (USFWS 1984b). Despite the fact that it has reached carrying capacity (Morin and Conant 2002), the translocated Laysan Finch population is not large enough to be an effective backup, and other sites are needed in light of recent computer models of extinction risk of the original Laysan population (Tarr et al. 1998). Laysan Finches were established for several decades on Midway Atoll (Bailey 1956), and now that it is not only ratfree but a national wildlife refuge, Midway seems the logical next site. However, refuge personnel are reluctant to follow through until mosquitoes can be controlled at Midway (R. L. Shallenberger, pers. comm.), because Laysan Finches are extremely susceptible to mosquito-borne diseases (Warner 1968) that might be transmitted from migratory seabirds and shorebirds (but see previous chapter). Morin and Conant (2002) suggest that Nihoa Finches, which currently have no back-up population, be introduced to Midway, but in light of the birds’ natural history, in particular their apparent requirement of rock crevices as nest sites (see species account), this proposal is probably a bad idea. Nihoa Finches would likely attempt to nest in rocky breakwaters that are abundant at Midway (pers. obs.) with resultant losses to wave action, not to mention the potential for them to become a nuisance by nesting in buildings. Sincock and Kridler (1977) suggested Lisianski Island and Necker Island as possible translocation sites for Laysan and Nihoa Finches, respectively, but both are likely too small to support viable back-up populations (Morin and Conant 2002). Recent developments suggest that the main islands of Ni‘ihau and Kaho‘olawe would be better alternatives. Both are quite large and have limited fresh water sources (but no more so than Laysan or Nihoa), reducing the threat from

mosquitoes. Ni‘ihau has problems of access, although it is not exactly the ‘Forbidden Island’ of tourist industry hyperbole (Tabrah 1987). Nevertheless, the last time an ornithologist was allowed to work there was over a half-century ago (Fisher 1951), and even avian palaeontologists have been rebuffed (S. L. Olson, pers. comm.). Kaho‘olawe (see Chapter 2), on the other hand, might be ideal if vegetation were restored and rats eradicated (Morin et al. 1999). Importantly, it was likely within the original ranges of both Laysan and Nihoa Finches, which are known to have been sympatric on Moloka‘i, another island in the Maui-nui complex ( James and Olson 1991). Fears that the two might interbreed are probably unfounded, given their former sympatry and numerous potential reproductive isolating mechanisms (see species accounts). Kaho‘olawe would naturally provide the differing nest sites for each species, and its dry climate would likely keep the threat of avian malaria at bay or at least controlled. Of course, the biggest problem is the will to commit the necessary resources and actually do the job. Among main-island honeycreepers, the first experimental translocations involved Palila (Fancy et al. 1997).Although removal of feral ungulates had allowed vegetative recovery in places from which the birds had recently withdrawn, strong philopatry (Fancy et al. 1993b) was hindering their recolonisation of these areas. A total of 35 adult Palila from the high-density site at Pu‘u L a¯ ‘au on the western flank of Mauna Kea were translocated by helicopter to a low-density site at Kanakaleonui almost directly opposite on the eastern flank of the mountain, a distance of more than 20 km.A few were lost in the transport operation, so the initial release was 31 birds. Within a year, 16 had returned ‘home’ to Pu‘u La‘au, but four pairs nested at the release site during their first breeding season there, and two of those fledged young. Censuses during the ensuing 3 years showed that Palila density at Kanakaleonui had increased (but it was very low to begin with), so the experiment was deemed a success, and translocation was accepted as an effective tool for recovery of the Palila (Fancy et al. 1997). However, subsequent reports (Tummons et al. 1998b) indicate that this early optimism was premature. Homing of

180 The Hawaiian Honeycreepers translocated birds continues to be a problem that may be overcome by releasing captive-reared (see below) young birds that do not have the site tenacity of adults (P. C. Banko in Tummons et al. 1998b). Translocation of a single bird was recently attempted in the last-ditch effort to rescue the Po‘ouli from extinction (VanderWerf et al. 2003). The remaining male and 2 females (see species account) are scattered among noncontiguous home ranges and have little chance of meeting and forming a pair (Baker 2001). In an effort reminiscent of those that restored breeding populations of the equally imperilled Black Robin in New Zealand (Butler and Merton 1992) and the Mauritius Kestrel (Jones and Swinnerton 1997) from single breeding pairs, members of the Maui Forest Bird Recovery Project developed techniques for back-packing birds along forest trails to new sites, using Maui ‘Alauahio as surrogates (Tummons 2002a; VanderWerf et al. 2003). Then, one of the females was transported into the male’s home range, but, as in the case of the Palila, strong philopatry drew her back “home” the very next day, and she likely never encountered him. In light of the birds’ advancing age (all are at least 6 years old), the decision has now been taken to bring them all into captivity in a last resort effort to promote breeding (VanderWerf et al. 2003; see below). Conant (1988b) expressed misgivings about the use of translocation as a management tool because of potential unforeseen consequences, especially the possibility that rapid evolution, such as has occurred in translocated Laysan Finches (Conant 1988a) might produce a population distinct enough that it would no longer be the species being managed. In light of recent suggestions that conservationists adopt the phylogenetic species concept (Hazevoet 1996), this is not a trivial concern (Pratt and Pratt 2001). However, with the recent discovery that Laysan and Nihoa Finches once had much wider distributions in the archipelago ( James and Olson 1991), the proposed ‘translocations’ actually become reintroductions, a distinction clearly made by Conant (1988b) although the terms seem to have been used interchangeably by several other authors. So far, all ‘translocations’ of Hawaiian honeycreepers (and other Hawaiian birds) have actually been reintroductions.

Captive breeding The use of captive breeding and subsequent release into the wild as a means of restoring endangered species has been widely touted, but is plagued by ‘problems with (1) establishing self-sufficient captive populations; (2) poor success in reintroductions; (3) high costs; (4) domestication; (5) pre- emption of other recovery techniques; (6) disease outbreaks; and (7) maintaining administrative continuity’ (Snyder et al. 1996: 338). Captive breeding has a relatively short history in Hawai‘i with both a spectacular (though too early to evaluate completely) success with Puaiohi on Kaua‘i (Burnham 1998a; Pyle 1999d), and a spectacular apparent failure with the Hawaiian Crow (Burnham 1998b; Pyle 1999c; Pyle and Donaldson 2000a; pers. obs.), neither of which involved a Hawaiian honeycreeper. Research on rearing Hawaiian honeycreepers in captivity (Fig. 10.6) faltered for a time (Baker 1996), but exciting new efforts are now being made by the Hawaiian Endangered Bird Conservation Program at the Keauhou Bird Conservation Center (KBBC) in Volcano (Burnham 1998a,b, 1999; Kuehler et al. 2001) and the Maui Bird Conservation Center (MBCC) at Olinda. These facilities are operated by the Zoological Society of San Diego (Anon. 2000). KBCC began in 1995 with a surrogate research program to develop techniques using Hawai‘i ‘Amakihi (Kuehler et al. 1996). The methods developed have been used to rear Palila chicks for release to supplement the translocation program discussed above (Tummons et al. 1998b).The ‘quick-release’ method (Kuehler et al. 2001) developed may avoid some of the problems listed above, virtually all of which have been apparent in the Hawaiian Crow program (which KBCC inherited from earlier projects). In 2000, Palila, Hawai‘i Creeper, and Maui Parrotbill all bred for the first time in captivity at KBCC (Maxfield 2000), presumably a step toward maintenance of captive populations. Now, MBCC is faced with the daunting task of attempting to breed the Po‘ouli in captivity and reintroduce it to the wild (VanderWerf et al. 2003). Whether these projects will succeed in the long term remains to be seen. I agree with Conant (1998) that without real efforts to conserve habitat on a large scale, such programs may be

Status, conservation, and the future 181 translocation and captive rearing. It will be a long, labour-intensive, and expensive process, but at least provides a glimmer of hope for dealing with the most fundamental and pervasive threat honeycreepers face.

Birding and nature-oriented tourism

10.6 Hawaiian honeycreepers held in captivity for captive propagation: (a) Laysan Finch in Honolulu Zoo; (b) Hawai’i Creeper at Keauhou Bird Conservation Center.

little more than meaningless exercises.At least in the case of the Po‘o-uli, its habitat will remain available if reintroduction becomes possible.

Genetic engineering Based on hopeful signs of disease resistance in two ‘amakihi species, molecular biologists are now researching the genetic basis of both innate resistance of the birds and level of virulence in the pathogens. Shehata et al. (2001: 271) are attempting ‘to identify the genetic loci responsible for natural immunity, and to boost the numbers of individuals carrying these loci in natural populations.’ Jarvi et al. (2001: 254) hope ‘to provide population managers with new criteria for maintaining long-term population stability for threatened species through . . . maintaining genetic diversity . . . at loci important in immunological responsiveness to pathogens.’ The proposed methods are complex and will require the use of both

Another recent development is the increase in interest in Hawaiian honeycreepers on the part of recreational birders (Pratt 1993, 1995, 2002b; Pratt and Jeffrey 1996b), whose role in monitoring populations and distribution (see numerous citations by Pyle, Pyle and Ralph, Pyle and Donaldson, and Ralph and Pyle) is far too often under-appreciated (Banko et al. 2001; Pratt and Pratt 2001). Collar (1997) considers the birding community the ornithologists’ and conservationists’ most important ally. Far too often in Hawaiian parks and preserves, both public and private, birding has been discouraged (Pratt 2002b), primarily by overly restrictive or complicated rules of access and fees that discourage all but the fortunate few, to the ultimate detriment of the birds managers are ostensibly trying to protect (Pratt and Pratt 2001). The levels of public and private funding that will be necessary to support costly future research and restoration projects is such that managers simply cannot afford to alienate any large segment of the environmentalist community (Steiner 2001). Contrary to horror stories that seem to be rampant in Hawai‘i’s rumour mill, I find that recreational birding has little, if any, negative impact, especially in the Hawaiian context, and the good will engendered when birders have a realistic chance of seeing a flagship endangered species in a park or preserve has incalculable value in a democracy. But longentrenched myths are not easily discredited. Natureoriented tourism in general and birding in particular are becoming important enough in Hawai‘i’s economy to be noticed even by the popular press (e.g. Frierson 2000). Steiner (2001: 377) points out the need ‘to integrate future restoration efforts with tourism, the primary income generator for the Hawaiian Islands. One way to do this is through ecotourism and attraction of the birding community.’ Several ranchers on The Big Island already earn more from birding and other nature-oriented activities on

182 The Hawaiian Honeycreepers their land than from cattle (R. Pacheco, K. Unger, pers. comms.).When major landowners have a vested interest in preserving endangered honeycreepers so birders can see them, politicians and bureaucrats will listen. Ultimately, endangered species benefit because they come to be regarded as an asset rather than a burden and tax dollars spent on their behalf are seen as an investment rather than a liability.

Honeycreepers today and tomorrow Potential new threats Although Hawaiian honeycreepers are obviously overburdened, the story does not get any better. New threats are on the horizon that could exacerbate all the current problems with disease, habitat loss, and predation (Loope et al. 2001). The recent spread of West Nile virus, which affects both humans and birds, in North America has become a cause for concern in Hawaii (Tummons 2001d). Just as new human diseases seem to be appearing frequently in recent decades, as yet unknown bird diseases may be lurking in the shadows, waiting to devastate defenceless insular avifaunas. But we have enough known threats to contend with that we need not waste time contemplating mere possibilities.Avian malaria and pox in Hawai‘i are not static elements in the environment. As was discussed in Chapter 9, their mosquito vectors continue a seemingly inexorable advance upward in elevation as they adapt to colder temperatures. That relatively slow process could be circumvented overnight with the introduction of temperate zone strains of Culex quinquefasciatus. Even more insidious is the possible arrival of new vectors, especially several species of biting midges (Culicoides) known to transmit bird malaria, several of which occur in temperate climates (Loope et al. 2001) from which planes depart daily for Hawai‘i. What if a plant disease comparable to the chestnut blight that wiped out eastern North America’s dominant canopy tree (Cox 1999) were to strike ‘ohi‘a-lehua? The possible disaster is too horrible to contemplate, but Hawai‘i’s forests have already been infected with a highly virulent ‘disease’ of a different

sort.Alien plants are just as potentially dangerous as animals and microbes (Cuddihy and Stone 1990). Such invasives as kahili ginger, strawberry guava, blackberry, firetree, and banana poka have been discussed previously. Melastomes seem to be particularly invasive, and one in particular, Miconia calvescens, has the potential to totally displace native rainforests (Lövei 2001). In Tahiti, where Miconia is known as ‘purple plague’ and ‘green cancer’, native forests from seashore to mountaintop have been replaced by a Miconia monoculture in less than a quarter century (Kepler 1997). Kepler (1997: 8–9) believes this aggressive tree has potential impacts ‘far greater than all other noxious plants combined’. Since 1990, Miconia has made considerable headway in both Maui and The Big Island, despite a major control and public education campaign (Kepler 1997;Yuen 2001).Will ‘green cancer’ be the blight that deals the death blow to Hawai‘i’s forests, and with them the honeycreepers and virtually everything else uniquely Hawaiian? Or will something else be the coup de grâce? The brown tree snake Boiga irregularis, credited with devastating Guam’s avifauna (Savidge 1987), has become something of a media cause célèbre ( Jaffe 1994; Rodda et al. 1997; Burton 2000) and is a clear and present danger to Hawai‘i and other Pacific islands (Fritts 1987; McKeown 1996; Fritts and Rodda 1998; Rodda et al. 1998), requiring the expenditure of millions of dollars in an attempt to prevent further spread (Anon. 1998b). The native Hawaiian bird most immediately threatened by potential brown tree snake introduction is not a honeycreeper but the O‘ahu ‘Elepaio, a close relative of the Guam Flycatcher that was driven to extinction by the snake. It occurs at much lower elevations than most dreps and has undergone catastrophic declines in recent decades (VanderWerf et al. 1997). Hawaiian honeycreepers would undoubtedly succumb quickly to snake predation, but they may not be in as much danger from brown tree snakes as many fear. Perhaps the very same cool montane refuges that now afford some protection from lowland mosquitoes would also be effective against a tropical snake. Also, the recent discovery that acetominophen, a popular painkiller, is lethal to the snake (Burton 2000) may

Status, conservation, and the future 183 provide the long-sought key to eradication. Nevertheless, the O‘ahu ‘Amakihi, which seems to be dealing with avian disease better than most (Shehata et al. 2001), might see its recovery nipped in the bud if brown tree snakes become established on O‘ahu. As potentially damaging as the brown tree snake is, it is not the only serpentine threat, a fact that is often overlooked (Anon. 1998a; Kraus 1998; Loope et al. 2001). Of 63 snakes caught, turned in, or confiscated in Hawai‘i between 1994 and 1998, only one was a brown tree snake (Anon. 1998a). Most were pets that escaped or were liberated by their owners. Loope et al. (2001) list numerous other invasions of islands by snakes and numerous snake species that pose potential threats to Hawai‘i. Laws against such pets are often ineffectual or even exacerbate the problem as illustrated by the case of Jackson’s chameleon Chamaeleo jacksoni in Hawai‘i. This now well-established arboreal lizard (McKeown 1996) could not only impact the invertebrate food supply of insectivorous honeycreepers, but is potentially a direct predator of nestlings (Loope et al. 2001). It forms dense populations at low elevations in Hawai‘i (McKeown 1996), is now invading forested uplands, and could potentially reach tree line (Loope et al. 2001).As recounted by Loope et al. (2001: 294), the keeping of Jackson’s chameleons was illegal before 1994 (when the regulation was dropped because it had been ineffective) but sale and export were not, so people had ‘an economic incentive . . . to move the lizards around surreptitiously to begin new populations that could serve as a source of saleable animals. Consequently, the spread of the species to other islands and to new localities within islands was rapid, despite its illegality.’ Several species of tropical arboreal frogs are also spreading in Hawai‘i, with potentially disastrous results (Loope et al. 2001). Again, pet fanciers are partly to blame as are naive promoters such as McKeown (1996), who claimed that only one of these frogs was really damaging and that the others might be beneficial. Loope et al. (2001: 295) were quite correct to condemn such unscientific ideas as ‘statements of faith combined with a studious disregard for general ecological principles . . . deriving from a different agenda than the impartial descrip-

10.7 Heavily damaged native koa forest, previously with high, closed canopy, along Koke‘e Rd., Kaua‘i, October 1992, a few weeks after Hurricane Iniki.

tion of reality.’ Unfortunately, such attitudes are alive and well in Hawai‘i among aviculturists bent on ‘enriching’ Hawai‘i’s avifauna with everything from cockatoos to waxbills. Recently established parrots (Loope et al. 2001; Pratt 2002b) not only pose an agricultural threat, but could potentially carry new avian diseases. Threats from non-biological phenomena (droughts, hurricanes, lava flows, etc.) have a long history in Hawai‘i, but rapid ecological changes and population declines have made them far more damaging to Hawaiian honeycreepers than they used to be. The lava flow that was a dagger to the heart of the last significant population of ‘O‘u on The Big Island (see species account) would not have even been noticed a century earlier when the bird’s range and numbers were healthy. Hundred-year storms like Hurricanes Iwa and Iniki (Fig. 10.7) would have been physically devastating on Kaua‘i and would have reduced bird numbers temporarily, but would not likely have delivered a death blow to any species in the absence of avian malaria lurking in sheltered lowland valleys (Conant et al. 1998). The rules as well as the playing field are changing too fast for honeycreepers to stay in the game. One new physical threat is the possibility of global warming. With honeycreeper habitat reduced to tattered remnants, a slight climatic shift could destroy what little remains, either directly (Tummons 1999a), or by accelerating the further upward advance of mosquitoes (Tummons 2001a).

184 The Hawaiian Honeycreepers

Ongoing research Work on Hawaiian honeycreepers continues apace. Many of the names prominent in honeycreeper research in the 1970s and 1980s, including J. Michael Scott, C. J. Ralph, Cam and Angela Kay Kepler, Marie Morin, Stephen Mountainspring, Fred L. Ramsey, Paul W. Sykes, and Charles van Riper, III have moved on to mainland positions but continue to publish on Hawaiian subjects. Sadly, several important contributors of the period (Andrew J. Berger, C. Robert Eddinger,Wayne C. Gagné, John L. Sincock, Richard E.Warner, Alan Ziegler) are no longer with us. Others of that era still (or have returned to) work in Hawai‘i: Carter Atkinson, Paul C. Banko, Mark S. Collins, James D. Jacobi, Lloyd L. Loope, and Thane K. Pratt are all employed at the Pacific Islands Ecosystems Research Center (PIERC) of the USGS; Winston Banko has retired but continues to participate in USGS/BRD projects; Phillip L. Bruner heads his department at Brigham Young University Hawai‘i Campus;Tonnie L. C. Casey is a biologist with Kamehameha Schools Bishop Estate; Betsy Gagné is executive secretary for NARS; Jon G. Giffin continues to work for DOFAW on The Big Island; Jack Jeffrey has blossomed into a world-renowned bird photographer while continuing to work as a biologist at Hakalau Forest NWR (Tummons 2002b); Robert J. Shallenberger now works for The Nature Conservancy on the Big Island and David W. Woodside, even in his 80s, works for the Refuge Division of USGS. Storrs Olson and Helen James remain at the Smithsonian, analysing an exciting and productive new fossil site on Kaua‘i.At the University of Hawaii, three professors (Sheila Conant and Leonard Freed of the Zoology Department and Rebecca Cann of Department of Genetics and Molecular Biology of the School of Medicine) have graduate students involved in research on various aspects of honeycreeper biology. Freed has established a large new field station at Hakalau Forest NWR, fulfilling a long-held desire (Freed and Cann 1989) that will greatly facilitate field research there. At PIERC, a new generation of scientists has joined the old guard at work on the Kaua‘i and Maui Forest Bird projects, Palila restoration, and avian disease research. The Molecular Genetics Laboratory at the National Zoological Park

(Smithsonian Institution) in Washington, DC, under the direction of Rob Fleischer, continues to make progress in understanding honeycreeper genetics and evolution, and indeed produces publications at such a pace that review authors such as myself have difficulty keeping up. Obviously, in Hawaii at least, conservationists and scientists now seem to be working together for a common goal despite past difficulties (Pimm 2001;Tummons 2002e).

The future At some point, the Hawaiian honeycreepers and the ecosystems that support them may simply collapse from too many wounds that cannot heal. I hope we have at least a few more years or decades, and I can dream that we have several generations to enjoy them and learn what they can teach us. Four or five species have already gone extinct within my lifetime, and one hovers on the brink. As I write, everyone waits to see whether last-ditch efforts will save the Po‘o-uli or whether it will be the next Hawaiian honeycreeper lost. How many of the 17 other survivors will outlive me if I live another 25 years? I have seen the Alaka‘i go from a dawn greeted exuberantly by songs of ‘O‘o‘a‘a, Kama‘o, Puaiohi, Kaua‘i ‘Elepaio, ‘O‘u, ‘Akeke‘e, ‘Akikiki, Kaua‘i ‘Amakihi, ‘Anianiau, ‘I‘iwi, and ‘Apapane to mornings nearly silent except for alien voices of shama, Hwamei, and bush-warbler in just that length of time.With a little bit of luck and a few strategic research breakthroughs, most of the remaining honeycreepers can probably hang on for decades. But if any one of the potential disasters just discussed were to render today’s high elevation refugia inoperative, we can bid them all a wistful ‘aloha’. 1. Prior to the ESA, no Hawaiian honeycreeper enjoyed any federal legal protection; as non-migratory residents within the borders of a single state, they were not covered under the Migratory Bird Treaty Act. Now, because they are no longer considered a family of their own, they may enjoy protection through the bureaucratic sleight-of-hand that reinterpreted the law to cover all members of bird families that include some migratory species. At least the State of Hawaii protects them all unequivocally.

PART II Species accounts

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Po‘o-uli Melamprosops phaeosoma 187

Genus Melamprosops Casey and Jacobi Melamprosops Casey and Jacobi, 1974, Occas. Pap. Bernice P. Bishop Mus., vol. 24, no. 12, p. 217.Type, by original designation, Melamprosops phaeosoma Casey and Jacobi. Medium-sized, stocky, short-tailed birds with short, heavy, somewhat finchlike bills. Interorbital septum weak and sometimes fenestrated ( James and Olson 1991). Tongue non-tubular, with rounded

spoon-like tip (Bock 1978) and prominent lingual wings, unlike the truncate proximal end of most drepanidine tongues (Pratt 1992b). Food includes terrestrial molluscs and other invertebrates gleaned from trunks and branches of trees (Baldwin and Casey 1983;T. K. Pratt et al. 1997). Plumage colours atypical among honeycreepers, mostly brown above, buffy white below, with prominent black facial mask.

Po‘o-uli Melamprosops phaeosoma Casey and Jacobi PLATE 2 Melamprosops phaeosoma Casey and Jacobi, 1974, Occas. Pap. Bernice P. Bishop Mus., vol. 24, no. 12, p. 219. (Haleakala Volcano, Maui, Hawaii.) Other vernacular names: Poouli, Black-faced Honeycreeper (Berger 1975b)

Etymology Melamprosops ‘derived from Greek roots, meaning “black forehead” ’ and phaeosoma from same source, ‘meaning “brown body” ’ (Casey and Jacobi 1974). Actually, Greek prosops can also mean ‘a mask’. Hawaiian lexicographer Mary Kawena Pukui coined the vernacular name from Hawaiian words po‘o ⫽ head and uli ⫽ any dark colour (Pukui and Elbert 1971). Does not mean ‘black-faced’ contra Casey and Jacobi (1974), but rather ‘dark-headed’. According to Pukui and Elbert’s dictionary,‘blackfaced’ would be maka ‘ele‘ele. Apparently some breakdown in communication occurred at time of naming, but ‘Po‘o-uli’ now well entrenched and not inappropriate.

Systematics Whether this genus is drepanidine subject of considerable debate (see Ch. 4; Fackelmann 1992; Pratt 1992a). Now believed a very early offshoot from main line of Hawaiian honeycreeper descent (Fleischer et al. 2001; Pratt 2001b).

Description Small (14 cm) puffy round ball of a bird with elongated, somewhat finchlike bill.Tail so short that bird may look tail-less. Because both specimens (one in AMNH, the other in BPBM) believed immatures (Engilis et al. 1996) and few good photographs taken, exact colour descriptions and artists’ renderings conflicting. Colour photograph of holotype in flesh (Casey and Jacobi 1974) makes bird look green, rather than brown, dorsally because taken inside yellow tent (T. L. C. Casey, pers. comm.)! My earlier paintings (HAS 1975, 1993; Berger 1981; Shallenberger 1986; Pratt et al. 1987; Fackelmann 1992), based on two immature specimens, appear much darker below than as recently described (T. K. Pratt et al. 1997). However, one black and white photograph (Engilis et al. 1996) shows similar contrast. Adult plumage not represented among study skins but described from field observations and illustrated (Ching 1986; Engilis et al. 1996; T. K. Pratt et al. 1997) by artist/naturalist Patrick Ching (pers. comm.), who had extensive field experience with nesting pair. Recently published (Baker 1998, 2001; Scott 1998) colour photographs of mist-netted adult male reveal darker breast than depicted by Ching. Based on these sources, Definitive plumage shows sharply defined triangular black mask surrounding bill and extending to a point behind eye, broadly bordered above with pale grey

188 Po‘o-uli Melamprosops phaeosoma and below by a bold white ‘auricular patch’ that includes much of side of neck and extends onto the throat; grey above mask shades posteriorly through darker grey to olive brown on back; upperparts basically shades of brown; breast pale brownish grey shading to off-white on belly, sides tinged tawny or olive, darkening posteriorly to deep cinnamon on crissum and thighs. Female has slightly smaller and less intensely black mask and darker underparts (Engilis et al. 1996;T. K. Pratt et al. 1997). First Basic plumage like adult female with smaller mask (Engilis et al. 1996), lacks grey tones above mask reducing contrast, and is darker, more cinnamon brown above, buffier below. Juvenal plumage described by T. K. Pratt et al. (1997: 13) as differing from adult in being ‘paler gray brown, with less rufous and cinnamon’ above and ‘whitish’ below; black mask ‘smaller, restricted to area from dorsal ridge of bill, to distal corner of eye, to gape, and not extending onto chin.’ Bill black, iris reddish brown, legs and feet blackish brown. At least some individuals have pale tip to lower mandible that gradually darkens with age (Casey and Jacobi 1974, Engilis et al. 1996). For linear measurements of two specimens, see T. K. Pratt et al. (1997); of captured live bird, see Baker (1998).

Voice Calls are sharp chit notes, very similar to call note of Maui ‘Alauahio and Maui Parrotbill, possibly serving for mixed flock cohesion. Usually evenly spaced, but may be clustered in short bursts as chichi-chu-chit, which I once believed to be a ‘fragment’ of song as described by Engilis (1990 and pers. comm.), but which found to be alarm call (T. K. Pratt et al. 1997; Baker 2001). A ‘whistle call’ transliterated as chi-up, chew-ee, or chi-ii (T. K. Pratt et al. 1997) very similar to one call of Maui Parrotbill. Possibly the same call described by Mountainspring et al. (1990) as tschew. Song never recorded. Variously described as ‘a short buzzing series of chips, rushed and up-slurred toward the end: chitchit—chit-er, chit-er, chit-er, chit’ (Engilis 1990) or ‘chitchit chit-d chit-ter chit [pause] chit-ter chit-ter chit-ter’ (Kepler et al. 1996). Only male sings, in short bouts of up to eight songs. No whisper song or mimicry reported (T. K. Pratt et al. 1997). Typical and alarm

call recordings in Pratt (1996a), sonograms in Pratt (1992a) and T. K. Pratt et al. (1997).

Field identification Totally unlike any other native spp. on Maui. Bold black mask, emphasised by prominent pale cheek patches, presents striking pattern, especially viewed head-on (Engilis et al. 1996). Alien House Sparrow, also brown bird with black mask and bill and pale cheek patches (differently shaped, to be sure), causes confusion for casual or naive observers at Hosmer Grove in Haleakala NP (T. K. Pratt et al. 1997). Most frequent Po‘o-uli calls very similar to those of other Maui forest birds, so single bird in mixed flock easily overlooked.

Distribution, habitat, and population status Endemic to Maui. Originally widespread in several forested habitats, including dry forests, on E. Maui. Bones found commonly in fossil deposits from three caves on sw. slope of Haleakala (H. James in T. K. Pratt et al. 1997). Apparently disappeared from dry and mesic forests after arrival of humans, and thus not discovered by nineteenth-century naturalists who did not penetrate far into the rainforests. At discovery, population estimated at a few hundred birds in range of only 13 km2 between 1400 and

MAUI

Map 1

Po‘o-uli Melamprosops phaeosoma 189 2100 m elevation from west branch of Hanawi Stream east to headwaters of Heleleike‘oha Stream (T. K. Pratt et al. 1997; Baker 2001).A 1983 report of single bird near Ko‘olau Gap in Waikamoi Preserve (Mountainspring et al. 1990) never confirmed. Population declined and range contracted progressively. Population estimate for 1980 140 ± 280 (Scott et al. 1986). Disappeared from type locality (western end of range) by ca. 1989; only six believed to survive anywhere by 1997, those ominously including only one mated pair; no evidence of reproduction since 1996 (Baker 2001;VanderWerf et al. 2003). By 1998, only three birds located (Tummons et al. 1998a), one male and two females (Pyle 1998b; J. J. Groombridge, pers. comm.). Home ranges now so dispersed that birds’ chances of encountering each other essentially nil (Baker 2001); extinction seems imminent. As of this writing (March 2004), remaining three birds still survive with efforts underway for heroic goal-line stand against extinction (see Ch. 10). Attempt to translocate one female into the home range of the male failed (female returned to her home range the next day, probably without encountering the male). Decision has been made to take all 3 into captivity for possible breeding and release of offspring back into wild (VanderWerf et al. 2003). For popular accounts of plight of Po‘o-uli, see Carter (1995), Line (1997), Scott (1998), and Anon. (1999b).

Diet and foraging Foods include snails, insects, spiders, and rarely ‘olapa fruit. Forages on twigs (51%), branches (43%), and foliage (6%), but most prey captures (85%) on branches where insects predominate over snails (Casey 1978; Mountainspring et al. 1990;T. K. Pratt et al. 1997). One of few dreps to feed habitually in forest understorey (Fig. 11.1). Generally forages on plants at random, but shows preference for kanawao (Fig. 8.2c), which supports a high density of native snails, and kawau, and eschews pilo (Mountainspring et al. 1990).Tonnie Casey (in T. K. Pratt et al. 1997: 4) describes methodical foraging ‘hopping along substrate and turning from side to side or leaning over’ and capturing ‘prey by gleaning, reaching, or hanging’. Also excavates for prey by probing into substrate, flaking bark or lichens, or pulling away bark, epiphytes, or dead wood ‘using entire

11.1 Typical understorey foraging site for Po‘o-uli, Hanawi NAR, Maui.

body and legs for leverage’. Occasionally also taps substrate with bill, perhaps to detect prey.

Social and interspecific behaviour Often joins mixed-species foraging flocks (see Ch. 7) whose component spp. subdivide invertebrate-foraging niche (Ch. 8;T. K. Pratt et al. 1997). Mountainspring et al. (1990) believe Po‘o-uli and Maui ‘Alauahio may associate daily year-round.Territoriality very weakly expressed (T. K. Pratt et al. 1997).

Predation Presence of bones in subfossil owl pellets indicates prehistoric predation by stilt-owls (Olson and James 1991;T. K. Pratt et al. 1997), but no predation documented in recent times. Cryptic coloration and wing-flicking during foraging probably result of selection pressure from aerial predators (T. K. Pratt et al. 1997). Exhibits stronger response to human auditory lures (‘spishing’) and more obvious ‘approach and follow’ behaviour than most honeycreepers (Ch. 7; Pratt 1992a).

Breeding biology All information based on two nests built sequentially (the second following failure of the first) by a single pair in Hanawi NAR, Mar–June 1986 (Engilis et al. 1996; Kepler et al. 1996). In virtually all respects, nesting behaviour in step with that of other honeycreepers and cardueline finches (Baker 2001).

190 Genus Paroreomyza COURTSHIP AND MATING: Only mating display described involved male circling female in canopy of nest tree and nearby trees, singing vigorously 6–8 songs per 30 sec (Kepler et al. 1996). Pair-bond may be maintained by courtship feeding throughout nesting cycle (T. K. Pratt et al. 1997). NEST SITE AND TERRITORY: Canopy ‘ohi‘a trees 15 m tall. Nest 8 m above ground within terminal leaf cluster of secondary branches. NEST: Open cup built by both members of pair, with a base of bare pukiawe twigs and coarse mosses. Inner cup lined with fine fern rootlets. Average dimensions of two nests: outside diameter 158 mm; outer depth 100 mm; inner cup 78 ⫻ 60 mm diameter; depth 45 mm.

EGGS: Known only from old shell fragments, but apparently resemble those of other Hawaiian honeycreepers (see Ch. 8). Clutch size one or two eggs. INCUBATION AND HATCHING: Only female incubates, leaving nest only to defecate, forage, and receive food from male. Female solicits feeding by typical wing-fluttering display. Incubation period unknown. Eggs hatch in late morning, possibly at 2 d interval unlike the usual 1 d interval in most honeycreepers. Parents remove eggshells from nest. NESTLINGS, PARENTAL CARE, AND FLEDGING: Development little known. Male feeds both chicks and female, but female does bulk of chick feeding (presumably passing food regurgitated to her by male). Nest sanitation by parents very efficient throughout nest cycle. Only chick observed fledged at 21 d.

Genus Paroreomyza Perkins Paroreomyza [subgenus] Perkins, 1901, Ibis, p. 583. Type, by original designation, Oreomyza [⫽ Himatione] maculata Cabanis. Small, warbler-like honeycreepers with short, straight, warbler-like bills, with dark maxilla and pale mandible, never plumbeous or black as in Hemignathus. Adults sexually dichromatic, males much brighter than females, immatures (and adult females in one sp.) may have pale wing-bars. Differ from typical Hawaiian honeycreepers in wide variety of aspects: lack drepanidine odour (Pratt 1992a,b); possess thin non-tubular tongue, resembling those of wood warblers (Parulidae), with notched, frayed tip and prominent lingual wings at base (Richards and Bock 1973; Pratt 1992b); routinely use spider silk in nest construction and have nest helpers (H. and P. Baker 2000; P. and H. Baker 2000); actively mob predators (Pratt 1992b); defend large ‘home range’ territory (H. and P. Baker 2000); and have fenestrated interorbital septa (Richards and Bock 1973; Pratt 1992b). Foraging movements

resemble those of Black-and-White Warbler, not those of nuthatches (Pratt 1979a, 1992b). Call a loud, explosive chick or chip, song (only one sp. known) a lively jumble incorporating call note, very different from any other drepanidine song. Three spp., one each on O‘ahu, Maui/Lana‘i, and Moloka‘i, the latter two sympatric on Maui-nui. Unknown from Kaua‘i or Hawai‘i even in fossil record. Perkins (1903) first established Paroreomyza as subgenus for ‘alauahios and Kakawahie, but it became the genus name when the original Oreomyza was found to be preoccupied. Amadon (1950) considered Paroreomyza a subgenus of Loxops. Raikow (1977b) returned it to full generic status. Pratt (1979a), Berger (1981), Olson and James (1982b), and AOU (1983), restricted it to the current spp. Pratt (1992b) and Tarr and Fleischer (1995) questioned whether this genus was drepanidine, but recent studies (Fleischer et al. 2001; Pratt 2001b; James 2004) place it among the Hawaiian honeycreepers.

O‘ahu ‘Alauahio Paroreomyza maculata 191

O‘ahu ‘Alauahio Paroreomyza maculata (Cabanis) PLATE 5 Himatione maculata Cabanis, 1850, Mus. Heineanum 1: 100 (footnote). (Oahu.) Himatione maculata Wilson and Evans (1890–99) Oreomyza maculata Rothschild (1893–1900), Henshaw (1902), Perkins (1903) Paroreomyza maculata maculata Bryan and Greenway (1944), Raikow 1977b, Olson and James (1982b) Loxops maculata maculata Amadon (1950), Carlquist (1970), Berger (1972a) Other vernacular names: Alauwahio, Oahu Creeper, Creeper

Etymology Genus name combination of Greek para, beside, subsidiary, and Oreomyza (see under Oreomystis bairdi). Species epithet is Latin for ‘spotted’, erroneous allusion to supposedly spotted breast of first specimens (spots probably artefacts of skin preparation that allowed dark feather bases to show).

Systematics Pratt (1992b) reviewed tangled taxonomic history of ‘creeper complex’ to which this sp. belongs. Bryan and Greenway (1944) considered all current members of genus conspecific, but separated them as a sp. from what are now the two Oreomystis. Amadon (1950) lumped all into polytypic sp. called the Creeper, which Pratt (1979a) re-split into two genera and five spp. Olson and James (1982b) distributed these five spp. among three genera, considering the two ‘alauahios conspecific, but later ( James and Olson 1991) split them as have all authorities since.

Description A small (11 cm) yellow and green bird, males brighter than females and juveniles, with short, straight bill. Definitive male basically yellow-green above, darkest in centre of back, with yellow underparts, forehead, and supercilium, blackish lores, and olive green postocular stripe. Definitive female

duller throughout than male, with underparts white washed with pale yellow, flanks washed olive-grey, and two bold off-white or yellowish-white wingbars formed by pale tips of wing coverts, and without bold pattern of dark lores and postocular stripe of male (lores have only a tiny fleck of dark feathering immediately in front of eye and appear pale. Juvenal plumage (both sexes) much duller, greyish olive-brown above, off-white below, lacking adult face pattern (face mostly pale), but with two bold buff wing-bars. First Basic plumage similar but tinged yellow below. Second Basic plumage intermediate between previous and adult coloration, with streaks of yellow below (less so in female), and lores of male same colour as postocular stripe. Both sexes have brown eyes, dark brown legs and feet, and dull brown maxilla, paler brown below with mandible nearly white at base (Rothschild 1893– 1900). For standard measurements, see P. and H. Baker (2000).

Voice Call apparently very similar to that of congeners, a loud chirk or chip (Bryan 1905b; Shallenberger and Pratt 1978). Song never described.

Field identification One of several ‘little green birds’ on O‘ahu that cause difficulty for birders (Pratt et al. 1987). Very similar in overall coloration to O‘ahu ‘Amakihi, but male with much bolder superciliary stripe and dark postocular stripe along upper edge of auriculars, female with pale lores. Straight bill is diagnostic when only ‘amakihi compared, but not so easy to see. Both Japanese White-eye and Japanese BushWarbler are straight-billed ‘little green birds’ that might be mistaken for ‘alauahio, but neither has wing-bars.

Distribution, habitat, and population status Endemic to O‘ahu. Probably originally widespread and common, but by 1890s uncommon and

192 Maui ‘Alauahio Paroreomyza montana restricted to Wai‘anae and Ko‘olau Mountains above 500 m (Perkins 1903). Found in mesic to wet forests with dense undergrowth; unlike Kakawahie (Bryan 1908), did not eschew introduced trees such as kuku‘i (Bryan 1905b). By 1935, Munro (1960) could find none in extensive searches. Shallenberger and Pratt (1978) reviewed 41 published reports and concluded that most were misidentifications of O‘ahu ‘Amakihi, with only three nearly certain and another six probable, mostly from Poamoho and ‘Aiea trails in the Ko‘olau Mts. G. Swedberg (in Shallenberger and Pratt 1978) and W. Donaghho saw a large flock of ‘alauahio 9 Sep 1968, and proved their identification by collecting one selected randomly! Three identified in surveys in 1977–78 in N. Halawa Valley, Moanalua Valley, and a valley south of Manana Trail (Shallenberger and Pratt 1978; Shallenberger and Vaughan 1978). Only subsequent report with convincing details from 1985 Christmas Bird Count on Poamoho Trail (Bremer 1986). Surveys conducted by DOFAW in 1991 failed to find any, despite various reports and rumours (P. and H. Baker 2000; Ellis et al. 1992a) and E.VanderWerf (pers. comm.), a highly competent observer, has failed to find it in extensive recent searches. Sadly, probably recently extinct.

Diet and foraging Insectivorous. Foraging movements apparently similar to those of Maui ‘Alauahio but less well documented. Took beetles that live in tree cavities or under bark in decaying wood (Perkins 1903). Reported higher in the canopy than congeners, often in koa (Perkins 1903; Bryan 1905b; Shallenberger and Pratt 1978; Bremer 1986).

Social and interspecific behaviour Most observations have been of pairs or small groups, with groups up to six probably representing families.The large flock of 30–50 birds reported by Swedberg and Donaghho (above) almost certainly was a postbreeding mixed flock that included other spp. (P. and H. Baker 2000).

Breeding biology NEST SITE: Of two nests discovered at sites in Wai‘anae Mountains, one was 2.5 m from ground in naupaka kuahiwi and one 7 m up in kuku‘i tree (Bryan 1905b). NEST: Open cup of fine twigs, moss, and lichens bound with spider silk and egg cases; inner cup lined with fine rootlets, twigs, and leaf litter (P. and H. Baker 2000). EGGS: Two eggs collected by A. Seale in 1901 and described by Bryan (1905b) were typical of Hawaiian honeycreepers, cream with pale brown and reddishbrown spots concentrated around the larger end. Measured 20.4 ⫻ 14.8 mm and 19.9 ⫻ 15.3 mm (metric conversions by P. E. Baker from Bryan’s figures, P. and H. Baker 2000). Clutch size unknown.

Life cycle and demography Apparently breeds earlier than congeners, Jan–June with peak in Apr, perhaps because of warmer climate at the lower elevations where nests have been found (P. and H. Baker 2000).

Maui ‘Alauahio Paroreomyza montana (Wilson) PLATES 5, 8 Himatione montana S. B. Wilson, 1890, Proc. Zool. Soc. London (1889), p. 446. (Lanai.) Himatione montana Wilson and Evans (1890–99) Oreomyza (Paroreomyza) montana Perkins (1903)

Paroreomyza maculata Bryan and Greenway (1944), Munro (1944), Raikow 1977b, Olson and James 1982b Loxops maculata Amadon (1950), Carlquist (1970), Berger (1972a)

Maui ‘Alauahio Paroreomyza montana 193 Other vernacular names:‘Alauahio,‘Alauwahio, Maui Creeper, Maui and Lanai Creeper (Berger 1981), Creeper

Etymology See O‘ahu ‘Alauahio. Specific epithet means ‘montane’, an apparent reference to the birds’ occurrence in mountain forests. Maui subsp. named in honour of Sir Alfred Newton, an early supporter of research on Hawaiian birds.

Systematics See O‘ahu ‘Alauahio. Two subspp. recognised, P. m. montana on Lana‘i and P. m. newtoni on Maui. Pratt and Pratt (2001) suggest these may be fragments of a former morphocline on Maui-nui because the relatively few W. Maui specimens are somewhat intermediate. Otherwise, differentiation of subspp. must have occurred entirely since break-up of Maui-nui at end of last glaciation.

Description A small (11 cm) Hawaiian honeycreeper with shortest straight bill in the genus. Exhibits delayed plumage maturation, with Definitive plumage attained at Second Prebasic moult. Adults somewhat sexually dichromatic, immatures distinctive. Definitive male (P. m. newtoni) bright golden yellow on head and underparts, often fading posteriorly to cream on undertail coverts; upperparts, including wing coverts and edges of primaries and rectrices, olive green blending smoothly into yellow of head on nape; loral spot black, which, combined with feather shadows, may appear as a dark eye line, but never the ‘bandit mask’ of members of Hemignathus. Definitive female much duller yellow, with dorsal colour extending forward through crown to define a diffuse yellow superciliary stripe; pale tips to greater secondary coverts of some individuals may form single wing-bar. Juvenal and First Basic plumages dull greyish-green, palest below with no yellow, with two cream wing-bars. Second Basic similar, but yellower in face with reduced wingbars. Bill yellowish-pink with dusky culmen, legs pinkish-brown or flesh-coloured in all ages and sexes. Nominate race on Lana‘i slightly smaller, with adults slightly brighter green on back and

juveniles with broader wing-bars (H. and P. Baker 2000).

Voice Most frequent call note a loud cheek or chip, possibly a flock contact note.Alarm call a loud, plaintive, one- or two-syllable note resembling call of House Finch (H. and P. Baker 2000). Juvenile begging call monotonously repeated four-syllable chi-chi-chi-chee (H. and P. Baker 2000). Whisper song, apparently having some territorial function (H. and P. Baker 2000), resembles those of other honeycreepers (Ch. 7), but includes chip calls. Primary song (Fig. 11.2) variable, one form a lively and distinctive jumble of notes incorporating chip call at regularly spaced intervals: chip, whichy-wheesee-whurdy-whew, chip, whichy-wheesee-whurdy-whew, chip, etc. Also a jumbled warble very similar to song of House Finch but lacking burry notes at end, often uttered as part of flight display. H. and P. Baker (2000: 6) transliterate one song as ch-k-chewy-k-wee k-chewykwee k-chewy-k-wee-k, another as k-weedy k-weedy k-weedy k-weet the latter resembling song of parulid Common Yellowthroat. Singing highly seasonal, most intense between Mar and July, with pronounced dawn chorus (H. and P. Baker 2000) unlike many other honeycreepers that tend to spread singing throughout day (pers. obs.). For sonograms, see Pratt (1992b) and H. and P. Baker (2000), for recordings see Pratt (1996a).

Field identification Duller plumages surprisingly difficult to distinguish from Hawai‘i ‘Amakihi, but adults much yellower. Note much less black in lores and straight bill. Often overlooked where common (i.e. Hosmer Grove, Haleakala NP) because of similarity of overall colour to ‘amakihi (Pratt 2002b). Introduced Japanese Bush-Warbler resembles juvenile ‘alauahio

kHz 8 4 1

2

11.2 Song of Maui ‘Alauahio, LNS 05150.

3

sec

194 Maui ‘Alauahio Paroreomyza montana but is much duller grey-brown above with more sharply defined superciliary stripe. Song could be confused with that of House Finch, but not with any other honeycreeper. Sharp call notes not only diagnostic, but best cue to presence of birds.

Distribution, habitat, and population status Endemic to Maui-nui complex, and prehistorically widespread ( James and Olson 1991; H. and P. Baker 2000). However, James (in H. and P. Baker 2000) emended earlier reports of this sp. from Moloka‘i ( James and Olson 1991) to state that Paroreomyza bones found were too fragmentary for positive ID but within size range of P. montana. Unless third congener present on Maui-nui, these remains seem almost certain to represent Maui ‘Alauahio. Historically found throughout forests of Lana‘i and Maui, but now restricted to three disjunct populations above 900 m on E. Maui: abundant in rainforest belt on n. and e. slopes of Haleakala, common in forest plantation at Polipoli Springs (w. slope), and scarce in relict dry forest on sw. slope at Kahikinui (H. and P. Baker 2000). Sometimes occurs above treeline in subalpine scrub (Pratt 2002b). Extirpated from W. Maui (Scott et al. 1986). Lana‘i subsp. extinct, last seen 1937 (Munro 1960).

_ MAUI and LANA'I

Diet and foraging From H. and P. Baker (2000) plus additional references as noted. Eats mainly arthropods including moths, beetles, spiders, leafhoppers, lacewings, and ichneumonid wasps captured mostly by gleaning bark and leaves, but occasionally by ‘flycatching’ (pers. obs.). Sometimes follows Maui Parrotbill to clean up ‘scraps’. Also takes some nectar (Carothers 1982). Foraging substrates in Waikamoi Preserve, in descending order of importance, were ‘ohi‘a, koa, pilo, alani, ‘ohelo, kolea, pukiawe, and ‘olapa. Reported taking nectar from ‘ohi‘a, ‘ohelo, mamane, and ha‘iwale (Berger 1981; Carothers 1982).

Social and interspecific behaviour From H. and P. Baker (2000). Family groups comprising several generations remain together in home range throughout year, foraging together often in close proximity, but chasing members of other such groups when encountered. Other spp. (Maui Parrotbill, Hawai‘i ‘Amakihi, Po‘o-uli, and Maui Nukupu‘u) may join family groups to form mixed-species flocks.

Predation Bones in pellets ( James and Olson 1991) evidence of predation by stilt-owls and possibly Wood Harrier in prehuman times. Actively mob Short-eared Owls and feral cats (see Ch. 7 for details), but actual predation by them undocumented. Rats take eggs and nestlings, and may be major predators in forest edges (H. and P. Baker 2000).

Breeding biology All information from H. and P. Baker (2000) unless otherwise noted.

Map 2

COURTSHIP AND MATING: Monogamous, pairing for life although cuckoldry occurs occasionally. Displays usually accompany song. In song-flight, singing male ascends, flapping continuously or with pauses, to ⬎20 m above canopy, then descends to different perch at end of song or sings during descent. Singing perched male paces back and forth along branch with tail cocked up. Male also courts

Maui ‘Alauahio Paroreomyza montana 195 female with ‘gifts’ of pieces of vegetation. Soliciting female points bill and tail up, flutters wings, and gives begging calls. Members of pair may mutually preen after copulation. Non-breeding second-year birds usually serve as nest helpers to their parents. NEST SITE AND TERRITORY: Site chosen by female, usually in leafy terminal cluster but also on horizontal branches or cone clusters of introduced conifers. In native forest, most commonly in ‘ohi‘a, but also alani, ‘ohelo, kolea, pukiawe, and ‘olapa. Does not disdain alien trees, even conifers such as sugi and pines, but never uses eucalyptus. Male defends nest itself from close intruders. NEST: Female builds open cup nest in three phases: (1) base and thin outer wall; (2) inner wall and base of cup; and (3) lining of inner cup; each phase using different materials. Only rarely attached to supporting branches, but rather situated among them and not often supported from below; sometimes funnel-shaped and taller than wide (Pratt 1992b).The Bakers describe seven distinct varieties based on two sizes (small and tight vs. large and loose) and three groups of materials (based mainly on what is available in different habitats), plus a ‘pulu-ball’ type composed largely of pulu, a soft, fibrous, silky or woolly substance produced by Cibotium tree ferns (Kepler 1983). See H. and P. Baker (2000) for complex measurements of nests. ‘Ohi‘a forest nests built primarily of mosses and lichens, and occasionally pulu, lined with aerial fern rootlets and strips of cambium. Nests in subalpine scrub have little moss and more lichens often mixed with grass. Nests in forests of alien trees made of moss, lichen, and grass in varying proportions and may include such exotic materials as pine needles, horse hair, and manmade fibres (nylon). For detailed list, see original source. All nests (except for one held together by sticky pine sap!) bound with spider silk, usually taken from egg cases. Claim that spider silk unique to Paroreomyza nests among honeycreepers (H. and P. Baker 2000) overlooks report of spider silk in one nest of Hawai‘i Creeper (Sakai and Ralph 1980). Nest kept clean of faecal material throughout breeding cycle, but never reused.

EGGS: White with lavender, brown, and grey blotches concentrated at larger end. Average size 18.6 ⫻ 13.85 mm. Clutch apparently always two, laid immediately after nest completion, one per day. INCUBATION AND HATCHING: Female incubates, fed by mate and helpers, usually off nest but sometimes on it. Approaching mate or helper gives chip calls, female answers with begging calls. Birds approach nest indirectly, leave by plunging directly over nest rim out of canopy. Incubation period 16–18 d. Both chicks hatch on morning of same day. Female removes eggshells. NESTLINGS: Helpless at hatching, eyes closed, nearly naked with sparse grey down, orange-pink skin, purple gape with yellow rictal flanges. Feathers of various tracts begin to unsheathe at 6–10 d. PARENTAL CARE AND FLEDGING: Brooding by female decreases progressively through 11 d after hatching. Both adults and helpers feed chicks, usually by regurgitation, food sometimes passed first to female. Chicks fledge at 17–19 d, but may flee nest earlier to escape predators, with some ‘jumplings’ known to survive. Fledglings weak and short-tailed at first, fully mobile within 2 weeks.

Life cycle and demography From H. and P. Baker (2000). Breeding season Mar–Aug. Immatures remain with parents and continue to beg (usually unsuccessfully as they age) for up to 20 mo., do not breed until third year. Percentage of pairs with helpers varies from 29% in relatively disturbed habitats (Hosmer Grove and upper Waikamoi Preserve) to 95.6% in more pristine localities (Hanawi NAR). The differences reflect striking differences in nesting success (20–30% and 96%, respectively). Nest helpers (one or two depending on previous year’s outcome) always offspring of female if not both parents, except in cases of death of female and ‘adoption’ of helpers by ‘stepmother’. Advantages of co-operative breeding not readily apparent from survival statistics. Life span unknown, but at least 8 years.

196 Kakawahie Paroreomyza flammea

Kakawahie Paroreomyza flammea (Wilson) PLATES 5, 8 Loxops flammea Wilson, Proc. Zool. Soc. London (1889), p. 445. (Kalae, Molokai.) Oreomyza flammea Henshaw (1902), Perkins (1903) Paroreomyza maculata flammea Bryan and Greenway (1944) Other vernacular names: Creeper, Moloka‘i Creeper

Etymology See O‘ahu ‘Alauahio. Epithet flammea refers to flame-red colour of male. Hawaiian name means ‘wood chopping’ (Pukui and Elbert 1971), a reference either to sharp call notes that sound like distant wood chopping or to birds’ pecking at trees.

Systematics See O‘ahu ‘Alauahio.

Description Small (13 cm) birds with short, straight bills larger than those of ‘alauahios. Definitive male entirely flame scarlet, darker on lores, back, and scapulars. Definitive female rusty brown above, buffy white below variably washed or mottled with bright orange on forehead and throat. Immatures like female but less orange below. Moult patterns poorly understood. Juvenal plumage dark brown above, buffy below with indistinct facial markings, tinge of orange on chin, and two tawny wing-bars. First and Second Basic plumages progressively more like adult. Bill colour varies, but always darker on maxilla, grey to brown, with mandible ‘creamy yellow to deep rose-colour’ (Rothschild 1893–1900: 121–2). Iris brown, feet brown with variable pinkish tinge (P. and H. Baker 2000). For standard measurements, see P. and H. Baker (2000).

Field identification Only all red (male) or rusty brown (female) bird on Moloka‘i. Male could only be mistaken for ‘Apapane, which often cocks tail to display white undertail feathers, or ‘I‘iwi if bill not seen, but behaviourally distinct from both, rarely foraging in canopy with nectar-feeders.

Distribution, habitat, and population status Extinct. Widespread and common in forests of e. Moloka‘i from near sea level to mountaintops in nineteenth century (Rothschild 1893–1900; Perkins 1903). Frequented heavily forested gulches (Schauinsland 1906, translated by Yoshinaga 1997) and followed them down into otherwise deforested areas (Perkins 1903). Bryan (1908) found it still common in dense ‘ohi‘a forest, but by 1930s could not be found in searches specifically for it (Banko 1984a). Rediscovered by Pekelo (1963) who saw at least two pairs and a single male 1961–63. Expedition in 1975 in which I participated failed to find it (Scott et al. 1977), as did the HFBS (Scott et al. 1986) and subsequent searches through 1995. Pristine

MOLOKA'I

Voice Call apparently very similar to that of congeners, a loud chirk or chip (Bryan 1908; Pekelo 1963). Song never described.

Map 3

Genus Telespiza 197 Oloku‘i Plateau not surveyed since 1988 (Reynolds and Snetsinger 2001).

Diet and foraging Apparently entirely insectivorous (Bryan 1908), gleaning leaves and picking at bark or moss (Bryan 1908; Pekelo 1963) on trunks and limbs of trees, often quite near ground (Perkins 1903). Bryan (1908) reported feeding on a large moth held in feet and torn apart.

Breeding biology From P. and H. Baker (2000) except as noted. Open cup nest in terminal leaf clusters of ‘ohi‘a trees, mostly fairly vertical, 3–5 m above ground usually supported by interweaving with vertical branches rather than from below (Pratt 1992b). Outer shell neatly woven from mosses or lichens, with various other plant fibres, twigs, and spider silk. Inner cup lined with fine cambium strips, grasses, and fibres from ‘ie‘ie. Outside diameter 6.0–9.1 cm, inner cup 3.5–5.9 cm. Eggs unknown.

Social and interspecific behaviour From Bryan (1908) and P. and H. Baker (2000). No large flocks reported, but most observations were of small, presumably family, groups. Several such groups might come together uttering loud call notes to mob an intruder. No actual predation reported but may have been preyed upon prehistorically by Moloka‘i Stilt-Owl.

Courtship and mating Pairs form at approximately age 15–20 mo., remain together throughout year (P. and H. Baker 2000).

Life cycle and demography From Bryan (1908) and P. and H. Baker (2000). Although collections have disproportionate number of male specimens, sex ratio probably 1:1. Pairs form before birds reach full adulthood, but first breeding not until second year.Young remain with parents for long period as in Maui ‘Alauahio, so probably produce one brood/year. Females noted coming into breeding condition in Feb. Nests found under construction or completed Apr–June, recently fledged birds in June and Aug.

Genus Telespiza Wilson Telespyza [⫽ error for Telespiza] S. B.Wilson, 1890, Ibis, p. 341. Type, by monotypy, Telespyza [sic] cantans Wilson. Small- to medium-sized Hawaiian finches with heavy bills slightly hooked at tip. Tongue thick, fleshy, non-tubular, with ventral surface cornified and curving upward at sides (Raikow 1977b) and lacking lingual wings (Pratt 1992a). Nasal opercula lacking (Raikow 1977b). Sexually dichromatic, males brightly coloured in yellow and grey, females and immatures more or less streaked with brown. Highly vocal with complex canary-like songs and finchlike calls. Osteological characters reflect bill

shape. Among other Hawaiian finch genera, closest to Loxioides in morphology, but with longer, narrower bill ( James and Olson 1991). Four spp., two known historically from NWHI and as subfossils from main islands, and two from remains only in main islands.Additional spp. may be forthcoming as new fossil material is gathered. At least three spp. formerly sympatric. All qualitatively similar osteologically, differing in size in stepwise hierarchy. (Information on osteology and prehistoric spp. from James and Olson (1991).) Long-used spelling Telespyza shown to be erroneous by Olson and James (1986).

198 Laysan Finch Telespiza cantans

Laysan Finch Telespiza cantans Wilson PLATES 1, 8 Telespyza [sic] cantans S. B.Wilson, 1890, Ibis, p. 341, pl. 9. (Midway Island, North Pacific, error ⫽ Laysan Island.) Telespyza flavissima Rothschild (1892) Telespyza cantans cantans Delacour (1928), Bryan and Greenway (1944) Telespyza cantans Banks and Laybourne (1977), Pratt (1979a), Berger (1981), AOU (1983) Loxioides cantans Greenway (1968) Psittirostra cantans cantans Amadon (1950), Berger (1972a), Raikow (1977b) Psittirostra (Telespyza) cantans Olson and James (1982b) Psittacirostra cantans cantans Carlquist (1970) Other vernacular names: Laysan Canary (Munro 1960), Laysan Finch Bill (Throp 1970)

Etymology Telespiza from Greek tele, far (off ) ⫹ spiza ⫽ finch, thus ‘distant finch’ in reference to its occurrence in the remote Northwest Hawaiian Islands. Epithet cantans is from Latin and refers to the bird’s singing abilities.

Systematics All Hawaiian finch genera were combined by Amadon (1950) into Psittirostra; Greenway (1968) restricted that genus to the ‘O‘u and placed the others in Loxioides; Banks and Laybourne (1977) and Pratt (1979a) advocated the present treatment. Telespiza may ultimately be lumped with Loxioides ( James and Olson 1991; James 1998). Delacour (1928), Bryan and Greenway (1944), Amadon (1950), and subsequent authors considered the two Telespiza finches subspp., but Banks and Laybourne (1977) found sufficient differences in size, bill morphology, coloration, plumage sequences, and sexual dimorphism to warrant sp. status, now confirmed by discovery of former sympatry on the main islands ( James and Olson 1991). Further potential isolating mechanisms include differences in displays and nesting (see below).

Description A large (19 cm) sexually dichromatic finch. Adult male: entire head and breast bright yellow, shading into dull off-white below with flanks tinged grey, crown tinged olive; ‘collar’ on upper back grey, remainder of upperparts yellowish-grey; wing and tail feathers darker, edged with yellow.Adult female similar but duller throughout, with stronger olive tinge to crown and auriculars, diffuse brown streaks on flanks, and dark brown central spots on scapulars and lower back. Females come to resemble males progressively with each succeeding moult (Morin and Conant 2002). Both sexes have complex series of plumages before adult stage, beginning with a yellow Juvenal plumage heavily streaked on head, back, breast, and sides with dark brown (males more streaked than females). First Basic plumage retains flight feathers from Juvenal, but with faded edges (less yellow); streaking somewhat reduced on head and throat. Second Basic plumage further reduces streaking and approaches adult appearance Legs and feet black, iris dark brown, orbital ring black. Bill brownish-grey or ‘horn colour’ in all ages and sexes (pers. obs.) but paler in younger birds. For more detailed colour description and precise measurements, see Morin and Conant (2002).

Voice Recordings published (Pratt 1996a). Calls melodious, some resembling those of Palila (Berger 1981; Morin and Conant 2002). Alarm call usually a harsh cheruup (Berger 1981) but also when being handled a cry resembling that of Bristle-thighed Curlew (Morin and Conant 2002). Song loud, complex, varied, canary-like in complexity. Wetmore (in Olson 1996) compared it to songs of House Wren and thrashers. Possible vocal mimicry suggested by Freeth (in Munro 1960).

Distribution, habitat, and population status Historically restricted to Laysan Island, except for populations introduced and established on Midway

Laysan Finch Telespiza cantans 199 Atoll ca. 1905–44 (Alsatt 1945; Fisher and Baldwin 1945b; Bailey 1956), and Pearl and Hermes Atoll 1967 to present (Sincock and Kridler 1977; Morin and Conant 2002). Prehistoric bones found on O‘ahu and Moloka‘i ( James and Olson 1991), indicating a much wider distribution in prehuman times. Although considered endangered because of highly restricted range (Table 10.1), probably not under any imminent threat, but computer analyses indicate unacceptably high extinction risk (Morin and Conant 2002).

Diet and foraging Omnivorous diet discussed in Ch. 8. Following details from Morin and Conant (2002) unless otherwise attributed. In over 2000 feeding observations, 10 most frequent food items were: seeds of alena (8%), sandbur (6.8%), makaloa (8%), kawelu (8.7%), pigweed (6.2%), and ‘akulikuli (5%); flowers of ‘akulikuli (4%) and puaokama (8.3%); fruits of puaokama (2.5%); and invertebrates (10.8%). All parts of above plants, including roots and leaves, eaten occasionally. Ely and Clapp (1973) reported feeding on flowers of non-native tobacco and coconut palms. Also trace amounts of carrion (⬍1%) and eggs (⬍2%) as well as numerous other plants including several alien weeds, button sedge, kipukai, pohuehue, hinahina kahakai, beach naupaka, kupala, and nohu. Introduced population feeds more heavily on nohu than parent population (Conant 1988b). Forages in all plant associations on Laysan, but prefers areas of bunchgrass (kawelu), beach naupaka shrubs, and beach vines alena, pohuehue, and puaokama. Sometimes digs in sand with the bill to expose roots, seeds, and invertebrates. May enter seabird burrows, purpose unknown. Deliberately cracks small eggs, but depends on fortuitous breakage for larger ones. Learns to follow humans to take eggs of birds flushed from nests (Ely and Clapp 1973; Sincock and Kridler 1977; Conant et al. 1981).

Social and interspecific behaviour Extremely tame and tolerant of other spp., including humans, to the point of making themselves nuisances to researchers (Morin and Conant 2002).

Predation Typical predators not normally present on Laysan, but a Bristle-thighed Curlew once seen with freshly killed Laysan Finch (Woodside in Morin and Conant 2002), and a Peregrine Falcon, a rare winter visitor to the Hawaiian Islands (Pratt et al. 1987), preyed heavily on Laysan Finches in January 1961, while a Short-eared Owl present at same time apparently did not (Boswell and Keitt in Morin and Conant 2002). Introduced population on Midway Atoll wiped out by rats that gained access during Second World War (Fisher and Baldwin 1946a,b).

Breeding biology COURTSHIP AND MATING: From Morin (1992a), and Seto (1990) as summarised by Morin and Conant (2002). Courtship displays by both sexes include rapid raising and lowering of wings and tail along with distinctive vocalisation (apparently not recorded). Females may solicit males or even initiate pair formation with such displays. Sexes defend each other against intruders, probably an example of mate-centred territoriality characteristic of cardueline finches and Hawaiian honeycreepers (Ch. 4, 8). Males ward off other males with a ‘face-off ’ display in which the birds crouch facing each other, raise their dorsal feathers, and move toward each other with beaks open. Pairs mostly monogamous within a year, except when mates are lost, but polygyny might occur occasionally (Tarr et al. 2000). NEST SITE AND TERRITORY: Nest usually built near ground (Crossin 1966) in kawelu clumps (Fig. 11.3) or beach vines, but formerly often in ‘aweoweo (Fisher 1903, 1906), lost in rabbitplague but possibly becoming re-established (Morin and Conant 1990). Both sexes defend the nest site only; no other territoriality except mate defence (above). NEST: Open cup built by female only, mostly from various parts of kawelu (Crossin 1966; Morin and Conant 2002). Outside diameter ca. 16 cm, inner cup ca. 7 cm.Very rarely a pair will reuse previous nest (Morin 1992a).

200 Laysan Finch Telespiza cantans nest rim. Young fledge when 22–6 d old. Dependence period ca. 3 weeks, but fledglings beg until 40 d old or older.

Life cycle and demography

11.3 The bunchgrass kawelu, which provides both nest sites and food for Laysan Finches; Laysan Albatrosses rest nearby.

EGGS: From Morin (1992a) and Crossin (1966). Average size ca. 22 ⫻ 16 mm. Light cream or white with brown or maroon spots concentrated at larger end, dull or slightly glossy. Clutches up to 5 eggs recorded, mean 3.19 and mode 3. INCUBATION AND HATCHING : From Morin (1992a). One egg laid per day, incubation starting with first egg, producing sequential hatching. Only female incubates; usually sits tight on nest, but leaves to be fed by mate. Incubation period usually 16 d. NESTLINGS: From Morin (1991, 1992a). Chicks hatch with pale grey down feathers growing from all feather tracts except crural and caudal tracts. Bill and rictal flanges yellow, mouth lining lavender with red spots on the palate. Pin feathers show on d 4, and feathers unsheathe after d 10. PARENTAL CARE AND FLEDGING: From Morin (1992a) and Crossin (1966). Fed by both parents, but mostly by male. Faecal sacs removed until chicks ca. 16 d old, then allowed to accumulate on

Pair formation begins in Jan (Ely and Clapp 1973), nest-building by Mar with peak usually in May (Morin and Conant 2002). Weather perturbations may delay breeding in some years by at least 1 mo.; may renest if first destroyed; some raise two broods/year; can breed in first year, but rarely does so on Laysan (Morin 1992a). Survivorship not determined, but apparently long lived; one captive individual lived 12 years (Santos in Morin and Conant 2002) and wild birds known to reach at least age 10 (Mostello in Morin and Conant 2002). Population has undergone huge fluctuations in the past, and continues to fluctuate to a lesser extent year to year. Population estimated at 2700 in 1911 (Dill and Bryan 1912), 4000 in 1915 (Munter 1915), but dropped to ca. 100 at the height of the rabbit plague (see Ch. 2; Ely and Clapp 1973). Rebounded to over 5000 in 1951 (Brock 1951). Since 1968 (for survey methods see Conant et al. 1981), population has fluctuated between a high of 20 802 in 1976 and a low of 5201 in 1988, with some year-to-year declines and subsequent rebounds of as much as 50% (Morin and Conant 2002), not unusual for small closed ecosystems (Dennis et al. 1991). Fluctuations result primarily from weather conditions, as when a storm caused 90% mortality of eggs and chicks in 1986 (Morin 1992a). Usual carrying capacity of Laysan estimated at 10 000 (USFWS 1984a). Introduced population undergoes similar fluctuations (Sincock and Kridler 1977; USFWS 1984a; Morin and Conant 2002), with population between 108 and 876 over 20 years (373 in 1998, carrying capacity ca. 500).

Nihoa Finch Telespiza ultima 201

Nihoa Finch Telespiza ultima Bryan PLATES 1, 8 Telespiza ultima Bryan, 1917, Auk 34: 70, 71. (Nihoa Island, Hawaiian Group.) Telespyza ultima Pratt (1979a), Berger (1981), AOU (1983) Telespyza cantans ultima Delacour (1928), Bryan and Greenway (1944) Loxioides cantans ultima Greenway (1968) Psittirostra cantans ultima Amadon (1950), Carlquist (1970), Berger (1972a), Raikow (1977b) Psittirostra (Telespyza) ultima Olson and James (1982b) Psittacirostra cantans ultima Carlquist (1970)

Etymology See Laysan Finch. Epithet ultima given because Bryan (1917) believed this would be the last Hawaiian passerine discovered.

Systematics See Laysan Finch.

Voice Recordings published (Pratt 1996a). Following descriptions from Berger (1981).Two different melodious calls, a two-syllable up-slurred one and a threesyllable up-slurred then down-slurred call. Alarm note a harsh chip. Song loud, melodious, and complex with a variety of whistled notes.

Distribution, habitat, and population status Historically restricted to Nihoa. Population introduced to French Frigate Shoals in 1967 failed by 1984 (Sincock and Kridler 1977; USFWS 1984b). Prehistoric bones from Moloka‘i ( James and Olson 1991) indicate much wider distribution in prehuman times. Considered endangered because of highly restricted range (Table 10.1) and probably not under immediate threat, but population shows slight downward trend since 1968. Computer analyses reveal unacceptably high extinction risk (Morin and Conant 2002).

Description Like Laysan Finch but smaller (17 cm), with proportionally smaller bill that lacks strong overhanging hook at tip. Moult sequence shorter, with only Juvenal, First Basic, and Definitive plumages. Definitive male differs from Laysan Finch in less golden tone to yellow of head and breast, slightly bluer tone to dorsal grey, yellow not extending onto grey-tinged flanks. Definitive female distinctive, much more heavily streaked with dark brown than Laysan Finch, especially on breast, and emphasised by brighter edgings to back feathers; flanks greyish or cinnamon. First Basic plumage less heavily streaked than adult female, but with bold malar streak. Juvenal plumage even less heavily streaked, especially below, but darker on crown and back. Bill browner than in Laysan Finch. For precise measurements see Morin and Conant (2002) or Banks and Laybourne (1977).

Diet and foraging Omnivorous, diet differing from that of Laysan Finch only to the extent that habitats differ (Nihoa lacks some plants found on Laysan, and vice versa). On Nihoa reported to eat seeds, flower heads, fruit, and leaves of ‘aweoweo,‘ihi,‘ilima, kakonakona, and ‘ohai (Richardson 1954; Clapp et al. 1977; Conant 1985; Morin and Conant 2002). For the latter, said to feed on seed pods in a manner reminiscent of Palila eating mamane seeds (Morin and Conant 2002). Birds introduced to French Frigate Shoals ate seeds of nohu and introduced ironwood trees (Amerson 1971). Foraging behaviour similar to that of Laysan Finch (Morin and Conant 2002). Like Laysan Finches, learn to follow humans to take eggs of birds flushed from nests (Clapp et al. 1977; Conant et al. 1981), but able to crack only smaller eggs.

202 Makawehi Finch Telespiza persecutrix

Breeding biology COURTSHIP AND MATING: Courtship display involves male standing on a rock, partially extending, raising, and quivering (not fluttering) wings in a ‘shoulder-hunch’ posture while singing.Also has a song-flight display (Morin and Conant 2002). NEST SITE AND TERRITORY: Nests in rock crevices or niches in rock piles (Richardson 1954; Sincock and Kridler 1977; Morin and Conant 2002). Absence of suitable nest sites may have doomed introduced population on French Frigate Shoals, where birds nested in artificial piles of concrete blocks (Amerson 1971; Sincock and Kridler 1977). Male defends area within 3–4 m of nest site (Conant 1983); no other territoriality reported. NEST: Both sexes gather nest material, but whether both build unknown (Morin and Conant 2002). Nest ‘sprawling’ and ‘loose’ (Amerson 1971), made of grass, twigs, and feathers of large seabirds such as boobies and frigatebirds (Richardson 1954; Sincock and Kridler 1977). EGGS: Two eggs from captive birds measured 21.1 ⫻ 15.7 mm and 21.45 ⫻ 16.0 mm (Berger

1981). Coloration as in Laysan Finch. Clutches from 2–5 reported, most often 2 (Morin and Conant 2002). INCUBATION AND HATCHING: Based on captive birds (Berger 1981). Incubation delayed until clutch complete. Only female incubates, fed on or off nest by male. Incubation period of one egg 15 d 9 h, hatching in early morning. PARENTAL CARE AND FLEDGING: Both sexes feed young. One chick fledged at 24 d age (Berger 1981).

Life cycle and demography Timing of breeding probably similar to that of Laysan Finch, based on captive birds (Berger 1981) and near-fledgling observed in Mar (Richardson 1954). Population estimated at over 1000 in 1915 (Munter 1915), and 500–1000 in 1940 (Vanderbilt and Meyer de Schauensee 1941). Censuses (for survey methods see Conant et al. 1981) since 1968 reveal wide year-to-year variation as in Laysan Finch, between 1318 and 6686 birds (Morin and Conant 2002).

Makawehi Finch Telespiza persecutrix James and Olson Telespiza persecutrix James and Olson, 1991, Ornithol. Monogr. 46: 30. (Barber’s Point, Oahu, Hawaiian Islands.) All information from James and Olson (1991) except discussions of vernacular names and speculations about coloration.

Etymology See Laysan Finch. Species epithet from Latin persecutrix, stated to mean ‘one who follows pertinaciously’ ( James and Olson 1991: 30), but alternately (OED) ‘a female persecutor’, is a ‘sequel’ to the name ultima for the Nihoa Finch (q.v.). Makawehi refers to the lithified dune locality on Kaua‘i where sp. first found, and follows widespread practice of naming

birds for their site of discovery (as in Cape May Warbler). So far, only Telespiza from that locality.

Systematics See Laysan Finch. Although only slightly smaller than Nihoa Finch, T. persecutrix cannot be regarded as a subsp. because that sp. occurred on islands both east and west of Kaua‘i and O‘ahu, and may have had a continuous distribution producing sympatry.

Description Similar osteologically to Nihoa Finch but bill (and presumably bird) somewhat smaller. Because it

Genus Loxioides 203 occurred in relatively dry lowland habitats, probably looked much like a small Nihoa Finch. Among ground-finches of the Gal a¯ pagos, which have a size hierarchy reminiscent of that in Telespiza, plumage varies little and spp. differentiate on the basis of bill size.

Distribution, habitat, and population status Extinct following arrival of humans but before discovery by Europeans. Known from subfossil deposits at lowland sites on Kaua‘i and O‘ahu. Sympatric with Laysan Finch on O‘ahu.

Maui-Nui Finch Telespiza ypsilon James and Olson Telespiza ypsilon James and Olson, 1991, Ornithol. Monogr. 46: 35. (Ilio Point, Molokai, Hawaiian Islands.) All information from James and Olson (1991) except discussions of vernacular names.

Etymology See Laysan Finch. Specific epithet refers to the twentieth letter of the Greek alphabet, chosen because the holotype was found at ‘Site 20’. Vernacular name continues the tradition of locality names for spp. in this genus.A possible even smaller additional sp. known from fragmentary remains from Lua Lepo, an upland (808 m) locality on E. Maui. If the smallest Telespiza, it could be called the ‘Least Finch’, although ‘Lua Lepo Finch’ sounds better.

Description Smallest of the four described Telespiza. Bill smaller with relatively larger narial openings than in the

others. May well have resembled Nihoa Finch in plumage colour (see Makawehi Finch), but the undescribed sp. from more humid uplands could have been darker, perhaps with orange or red replacing yellow.

Distribution, habitat, and population status Extinct following arrival of humans but before discovery by Europeans. Probably occurred throughout Maui-nui complex, but specimens only from Moloka‘i and Maui. Abundance of bones indicates bird was common in arid lowland habitats, but also occurred in upland dry forest.

Social and interspecific behaviour An important prey item for the Moloka‘i StiltOwl, comprising 53% of the drepanidine bones from owl pellets at ‘Ilio Point.

Genus Loxioides Oustalet Loxioides Oustalet, 1877, Bull. Sci. Soc. Philom. Paris (7)1: 99. Type, by monotypy, Loxioides bailleui Oustalet. Large Hawaiian finches with short, heavy bills adapted for feeding on fruits of mamane. Culmen arched and gonys convex. Nasal opercula lacking, tongue similar to that of Rhodacanthis (Gadow 1899). Shares several cranial features with Orthiospiza ( James and Olson 1991). Sexual dimorphism slight but males brighter than females. Songs complex and

canary-like, calls whistled. Known from upland dry forest on Hawai‘i and recently found in lowland fossil deposits on Kaua‘i (S. L. Olson, pers. comm.). Currently monotypic, but Olson (1999) suggests both Telespiza and Rhodacanthis might eventually be merged with Loxioides. James (1998) shows Loxioides embedded within Telespiza, which would be paraphyletic without inclusion of the former, but shows Rhodacanthis as more closely related to Chloridops.

204 Palila Loxioides bailleui

Palila Loxioides bailleui Oustalet PLATES 1, 8 Loxioides bailleui Oustalet, 1877, Bull. Sci. Soc. Philom. Paris (7)1: 100. (Hawaii.) Psittirostra bailleui Amadon (1950), Berger (1972a) Psittacirostra bailleui Carlquist (1970) Psittirostra (Loxioides) bailleui Olson and James (1982b)

Etymology Genus name means ‘like Loxia’, alluding to the genus of the cardueline crossbills. Species named by Oustalet (1877) for Théodore Ballieu, the French Consul at Honolulu who collected the first specimens, but name misspelled (Olson and James 1994)! Hawaiian name apparently onomatopoetic (see Voice, below).

Systematics Placed by Amadon (1950) in an enlarged genus Psittirostra, that included all drepanidine finches. See Laysan Finch for further discussion of generic limits in drepanidine finches. James (1998) found that osteology indicated close relationship between Loxioides and Telespiza.

Description A large (19 cm) Hawaiian finch with an arched, blunt bill resembling those of bullfinches (Pyrrhula). Plumage data from Jeffrey et al. (1993) and pers. obs. Sexually dichromatic, but both sexes in Definitive plumage have grey upperparts (rump paler) and white underparts, the flanks sometimes tinged grey; wing and tail feathers dark brown with golden yellow edgings (very broad on coverts), producing a greenish-yellow appearance; and yellow head and breast with dark lores.Yellow of male more golden, that of female more lemon yellow. Male’s yellow head sharply demarcated from back at nape (although some grey feathers may appear above the nape line), but grey extends variably through nape to crown of female. Lores black in male, dark grey in female. Crown of First Basic females may

be streaked with grey (yellow feathers with grey centres). Definitive plumage (both sexes) reached in second year. Juvenal plumage resembles adult female but has double wing-bars formed by pale green tips to greater and middle coverts, retained in First Basic plumage. Legs and feet black, iris dark brown. Bill dull yellow in Juvenal, darkening to entirely black in First Basic plumage. For standard measurements, see Banko et al. (2002).

Voice Typical call is an onomatopoetic, sweetly warbled pa-li-la, sometimes compared to calls of such carduelines as Pine Grosbeak. Primary song (Fig. 7.1) relatively quiet but long, complex, and canary-like, incorporating whistles, warbles, chirps, and trills. For recordings of both, see Pratt (1996a). Also a whisper song that includes mimicry and a short, slurred contact note (Banko et al. 2002).

Field identification Palila is one of the largest passerines in its habitat. Flying away, it can be identified by its pale rump (darker in similar-sized Sky Lark) when other colours are not seen. Habitat now shared with yellow-variant House Finches and Yellow-fronted Canaries, with Saffron Finches not far away and expanding (Pratt 2002b). Palila larger than all, with distinctive contrasting grey back. Coloration surprisingly concealing among mamane trees with their yellow flowers and silvery-green foliage. Local Hawai‘i ‘Amakihi utter a call very similar to Palila’s (possible mimicry), but which usually slurs up rather than down at the end. Some vocalisations of Yellow-fronted Canary also resemble those of Palila.

Distribution, habitat, and population status Historically known only from upland dry forests of Hawai‘i between 1500 m and treeline (ca. 3000 m), but subfossil remains found at lowland dry forest sites on O‘ahu (Olson and James 1982b)

Palila Loxioides bailleui 205 and Kaua‘i (Burney et al. 2001). Formerly widespread in ‘saddles’ between the island’s three main peaks including e. slope of Hualalai and nw. slope of Mauna Loa; now restricted to narrow discontinuous band at 2000 to 2750 m from sw. to n. slopes of Mauna Kea (Scott et al. 1986;T. K. Pratt et al. 1997; Gray et al. 1999). Distribution coincides closely with that of mamane and mamane–naio forest (Scott et al. 1984). Highest population concentrated at w. end of range around Pu‘u La‘au, where greatest elevational range achieved (Scott et al. 1986; Jacobi et al. 1996). Population, best documented of all drepanidines, exhibits wide annual fluctuations (Scott et al. 1984; Jacobi et al. 1996; Banko et al. 1998; Gray et al. 1999) that make detection of longterm trends difficult. Mean population 1980–95 3390 ⫾ 333, but ranged from 1584 ⫾ 324 in 1985 to 5685 ⫾ 535 in 1981 ( Jacobi et al. 1996). Declines result primarily from reduced availability of mamane seed pods during drought conditions associated with El Niño events (Lindsey et al. 1997;

HAWAI'I

Map 4

T. K. Pratt et al. 1997; Gray et al. 1999). Overall range unchanged since 1975, but populations outside main centre declining drastically ( Jacobi et al. 1996). Genetic studies (Fleischer et al. 1994) reveal that present scattered populations are fragments of recently panmictic population rather than result of recent movements, as expected for this strongly philopatric (Fancy et al. 1993b, 1997) sp. Although not presently declining, Palila faces high risk of extinction because single viable population subject to annual fluctuations that could reduce numbers below easily recoverable limits, and it occupies a fire-threatened habitat with high potential for catastrophe (Gray et al. 1999).

Diet and foraging Feeds mostly on mamane seeds extracted from green pods (Perkins 1903). Pods plucked, then carried to an interior perch of the tree to be held by the feet against a branch and ripped open along a longitudinal seam (Figs. 6.2, 11.4). Also eats naio fruit and seeds, alien poha fruit, and mamane flowers and (very rarely) leaves (Munro 1960; van Riper 1980a; Berger 1981; Banko et al. 2002). Strongly attracted to flowering sandalwood (pers. obs.), but what is eaten there unknown. Also eats insects (van Riper 1980a).Young fed caterpillars, and adults sometimes gorge themselves on a pyralid with bold warning colours (Perkins 1903).Young also receive mamane pods and flowers (Pletschet and Kelly 1990).

11.4 Palila in typical feeding posture with mamane pod held in feet. Photo © Jack Jeffrey.

206 Palila Loxioides bailleui

Social and interspecific behaviour SOCIAL BEHAVIOUR: Generally found in small parties of ca. five birds. May be family groups, but often several males move about together.

Predation Plumage colour cryptic, more pronounced in female, reflecting predation by Short-eared Owl and Hawaiian Hawk ( Jeffrey et al. 1993). Tolerant of human approach, often seeming oblivious to potential threats from below. Predation on eggs, nestlings, and incubating females by black rats and feral cats reported (van Riper 1980a; Pletschet and Kelly 1990) but significance controversial. Amarasekare (1993) considered it minor; difference in predation rates found by Pletschet and Kelly (1990) and T. K. Pratt et al. (1997) possibly the result of different definitions of predation. Predation much more severe on nestlings than on eggs (Pletschet and Kelly 1990).

Breeding biology All information from van Riper (1980a) and Pletschet and Kelly (1990) except as noted. COURTSHIP AND MATING: Male performs low flutter-and-glide advertising flights and sexual chases early in season. Courtship feeding occurs throughout breeding cycle, but decreases progressively and replaced by mutual billing with no food exchange by end of season. Female solicits feeding by wing fluttering. NEST SITE AND TERRITORY: Nests built in site selected by female in terminal forks or on horizontal branches of mamane (or rarely other) trees. Male defends only the female and immediate vicinity of nest, but forages in much broader area (Fancy et al. 1993b). Uses song and direct chasing to exclude conspecifics from vicinity of mate. Female may also defend nest site. NEST: Both sexes build, but most work done by female. Construction time varies between 7 and 20 d. Base made of mamane twigs broken off nest tree. Bowl of grasses but may opportunistically contain sheep wool (van Riper 1977). Lining of Usnea

lichens or fine grass and rootlets. Diameter ca. 15 cm, height ca. 8 cm; bowl 7.5 cm wide, 4 cm deep. EGGS: Typical honeycreeper eggs with white ground and reddish-brown spots concentrated at larger end. Average ca. 25 mm long and ca. 17 mm wide. Interval between nest completion and laying usually 1–2 d but can be as long as 20 d. Eggs laid in morning, one per day until clutch (1–3 eggs, usually 2) complete. INCUBATION AND HATCHING: Only female usually incubates and broods, but male may occasionally brood for short periods. Incubation period ca. 16 d. Berger’s (1981) report of longer periods based on confusion with nestling periods. Hatching rate ca. 50% overall but ca. 89% in nests with some eggs that hatch (Pletschet and Kelly 1990). Hatching failure primary cause of nest mortality. Eggshells removed immediately after hatching. Daytime brooding decreases with age of nestlings but female always on nest at night and during heavy rain through d 18. NESTLINGS: Helpless at hatching. Skin pinkishorange with sparse black down in dorsal feather tracts. Bill dull yellow with increasingly white tip with age ( Jeffrey et al. 1993), turning orange and then black. Eyes open at 7 d, grasping reflex appears at 9 d, ability to stand at 7 d. Young gape readily through 12 d, thereafter cower in nest when disturbed. Preening develops gradually following 9 d, becomes vigorous after 15 d.Young may jump from nest after 15 d (Banko et al. 2002) and begin flapping wings at 21 d. Nestling periods vary between 21 and 29 d, long periods for passerines, apparently the result of slow growth rates of chicks. PARENTAL CARE AND FLEDGING: Both parents feed young, but 63% by females. Nest helpers rarely also feed if one parent lost. Feeding by regurgitation. Adults remove faecal sacs until young ca. 4 d old, then allow them to accumulate on nest rim; nest becomes very untidy by fledging. Parents may remove young from nest if banded before 10 d age, apparently a nest sanitation response triggered by bands on legs of nestlings (Banko et al. 2002). Pletschet and Kelly (1990) report fledging rate of 51%, much lower than the 80% reported by van Riper (1980a), who trapped potential predators around nest trees.

Greater Koa-Finch Rhodacanthis palmeri 207

Life cycle and demography From Lindsey et al. (1995) except as noted. Wide annual population fluctuations result in similar variations in age ratios. Proportion of year-old birds relatively constant, but hatch-year birds range from as low as 3% to almost a quarter of population; survivorship of hatch-year birds much lower than for

older ones. Sex ratio 1:1 for nestlings and secondyear birds, but males outnumber females among hatch-year birds and adults.Adult male bias possibly result of differential predation on incubating females (Pratt et al. 1997). Survivorship comparable to that of other honeycreepers. Breeding season Apr–Sep with peak in mid-June (Pletschet and Kelly 1990).

Genus Rhodacanthis Rothschild Rhodacanthis Rothschild, 1892, Ann. and Mag. Nat. Hist. (6)10: 110. Type, by subsequent designation (Bryan and Greenway, 1944), Rhodacanthis palmeri Rothschild. Large, heavy-billed Hawaiian finches adapted for feeding on green pods of koa. Tongue nontubular, compact, scooped out dorsally, the cornified ventral surface curving upward laterally (Gadow 1899). Sexually dimorphic, males brightly coloured, females cryptic. Vocalisations loud whistles, distinctive in the drepanidine context. Only two historically known spp., from the Kona region of Hawai‘i, but bones of a Kaua‘i sp. as yet

undescribed have been recently found (Olson 1999), and remains of Rhodacanthis close to, but possibly not conspecific with, the historically known spp. known from O‘ahu and Maui ( James and Olson 1991; Olson 1999). Rothschild (1893– 1900) was the first to suggest Rhodacanthis, Telespiza, and Loxioides similar enough to be lumped (Loxioides has priority). Olson (1999) suggests that if intermediates found in fossil deposits, combination still appropriate. James’s (1998) topology reveals closer relationship with Chloridops, a grouping that may also include one sp. of Xestospiza (H. James, pers. comm.)

Greater Koa-Finch Rhodacanthis palmeri Rothschild PLATES 1, 8 Rhodacanthis Palmeri Rothschild, 1892, Ann. and Mag. Nat. Hist. (6)10: 111. (Kona, Hawaii, Sandwich Islands.) Psittirostra palmeri Amadon (1950), Berger (1972a) Loxioides kona Greenway (1968) Psittiacirostra palmeri Carlquist (1970) Psittirostra (Rhodacanthis) palmeri Olson and James (1982b) Other vernacular names: Hopue, Orange Koa Finch (Henshaw 1902; Bryan and Greenway 1944), Greater Koa Finch, Koa Finch (Pratt 1979a)

Etymology Genus name is from classical roots rhodos ⫽ red or ‘a rose’ and acanthis ⫽ ‘a bird fond of thistles’ or

‘linnet’ ( Jaeger 1955) that together mean, roughly, ‘red finch’. Species named in honour of Henry Palmer (see Ch. 3). English name reflects dependence on koa. Validity of Hawaiian name hopue strongly questioned by Perkins (1903), hence not widely used.

Systematics See Laysan Finch for discussion of generic limits in drepanidine finches. James and Olson (1991) suggested that both Rhodacanthis and Telespiza could be lumped with Loxioides, but James’s (1998) phylogeny places Rhodacanthis as close relative and possible congener of Chloridops. On other characters Pratt (2001b) clustered Rhodacanthis, Loxioides, and Chloridops as a sister group to Telespiza.

208 Greater Koa-Finch Rhodacanthis palmeri

Description A huge (23 cm), big-billed finch. Adult male with striking flame-scarlet head and breast with satinygold highlights. According to Perkins (1903: 437) ‘the largest and most beautiful of the [drepanidine finches]. Unfortunately the golden sheen of the orange-reddish crown partially loses its lustre [in specimens] . . . In the winter months this plumage is at its best, and at this season the bird in life must be counted one of the most beautiful of Hawaiian species.’ Orange colours become duller posteriorly on underparts to yellowish-buff undertail coverts, upperparts darker olive-brown with orange highlights, brightest on rump and uppertail coverts. Females cryptically coloured in dull yellowish-green, paler below and fading posteriorly to off-white undertail coverts. Bill and feet (both sexes) bluishgrey (Rothschild 1893–1900). Juvenal plumage similar to adult female but duller above, yellower below, feathers of lower breast and belly tipped with dusky producing a mottled or streaked appearance. Bill dark brownish-grey, browner below, with yellow rictal flanges. Transitional (First Basic?) male much duller than adult but distinct from female. Upperparts lack orange tones of adult, head mostly same colour as back, forecrown and short streak above eye yellowish-orange, throat and upper breast orange fading posteriorly to cream-coloured undertail coverts. For measurements, see Olson (1999).

female was reported to have a distinctive ‘rather deep single note when alarmed’ (Perkins 1893: 104).

Field identification Male unmistakable, a huge gleaming red-orange grosbeak. Females paler than Kona Grosbeak with different bill shape, slimmer profile. Female ‘O‘u has smaller, hooked, pink bill. Female Lesser Koa-Finch very similar in all respects except size and slightly darker underparts; not reliably separable in field (Pratt 1979a).

Distribution, habitat, and population status Extinct. Likely originally inhabited all mesic koa forests on Hawai‘i. Historically found in koa woodlands on various substrates in several localities as summarised by Olson (1999). Heart of historical

Voice Call and song not well differentiated by observers and reports somewhat conflicting. Palmer (in Rothschild 1893–1900) described a low, melancholy, descending whistle of two or three notes. Perkins (1903: 438) characterised the song as ‘entirely different from that of any other native bird’ comprising 4–6 prolonged clear whistles, variable within and among individuals. On the other hand, Munro ( journal quoted in Olson 1999: 10) stated that the initial notes of the ‘O‘u’s song were ‘very like that of the [Greater Koa-Finch], but not so strong, (the note . . . seems to flood the whole surrounding bush & is difficult to locate).’ Other authors also mention the ventriloquial nature of the song. Both sexes responded to human imitations (Rothschild 1893–1900; Perkins 1903). The

11.5 ‘A‘ali‘i, showing papery fruits that were an alternative food source of Greater Koa-Finch.

Lesser Koa-Finch Rhodacanthis flaviceps 209 range was koa belt between 900 and 1200 m north and south of Pu‘u Lehua (see Kona Grosbeak for locality details) where most specimens obtained. Palmer apparently collected two specimens on s. leeward slope of Mauna Kea, and Perkins observed but did not collect Greater Koa-Finches on the s. flank of Mauna Loa ca. 10 km above present-day headquarters of Hawaii Volcanoes NP. Only 65 specimens in world collections (Banko 1979 as updated by Olson 1999).

Diet and foraging Food almost entirely green pods of koa, sometimes entire pod (in pieces), occasionally only seeds. Munro’s journal (quoted in Olson 1999) described bird hanging tail downward from pod while ripping it open to expose seeds. Also occasionally ate ‘a‘ali‘i

(Fig. 11.5) seeds (Munro 1960). Seasonally fed on caterpillars, sometimes gathered from shorter trees such as naio (Perkins 1903).

Breeding biology Courtship and mating virtually unknown, but pairbond apparently strong; one female called to dead mate 5 min after collection (Perkins 1893). Olson (1999) deduced possible breeding cycle of eggs in spring months and young birds present July–Oct. Perkins (1903) saw male pick up nest material from the ground and carry it to top of koa tree. Later, several large nests became exposed in koa crowns after caterpillars had stripped leaves.Young fed pieces of koa pods by both parents (Perkins 1903). Fledglings apparently dependent ⬎1 mo. (Olson 1999).

Lesser Koa-Finch Rhodacanthis flaviceps Rothschild PLATE 1 Rhodacanthis flaviceps Rothschild, 1892, Ann. and Mag. Nat. Hist. (6)10: 111. (Kona, Hawaii, Sandwich group.) Psittirostra flaviceps Amadon (1950), Berger (1972a) Loxioides flaviceps Greenway (1968) Psittiacirostra flaviceps Carlquist (1970) Psittirostra (Rhodacanthis) flaviceps Olson and James (1982b) Rhodacanthis palmeri (Pratt 1979a) Other vernacular names:Yellow-headed Koa Finch (Henshaw 1902; Bryan and Greenway 1944)

among historically known specimens ( James and Olson 1991) support existence of two koa-finch spp.

Description A fairly large (19 cm), big-billed finch. Definitive male head and underparts bright golden yellow shading posteriorly to yellowish-olive; undertail coverts paler olive-yellow; upperparts olive green. Definitive female nearly identical to female Greater Koa-Finch but darker on lower abdomen. Bill and feet as in Greater Koa-Finch. Juvenal plumage like Greater Koa-Finch but with less mottling on belly (Olson 1999). For measurements see Olson (1999).

Etymology See Greater Koa-Finch. Species name Latin for ‘yellow-headed’.

Systematics See Greater Koa-Finch. Original collectors of koafinches were surprised when two spp. named (Perkins 1903; Munro 1960); Pratt (1979a) concluded that smaller, yellower birds could be accommodated within the variational range of one sp.. However, subfossil remains of two spp. (Olson and James 1982b), as well as qualitative osteological differences

Field identification ‘Young male Greater Koa-Finches are yellower than adults but never as clearly yellow on the whole head as [Lesser Koa-Finch]’ (Pratt et al. 1987: 298). Palila occurred in same habitat and localities, but easily identifiable by paler grey back, dark lores and bill. ‘O‘u has yellow head sharply demarcated from dark breast. I have heard reports from novice birders of what they thought were Lesser Koa-Finches (one can dream, after all), but in every case observer could not eliminate possibility of yellow variant

210 Kona Grosbeak Chloridops kona House Finch or Saffron Finch, both now abundant in Kona region (Pratt 2002b). See Greater KoaFinch for comments on female ID.

Distribution, habitat, and population status Extinct. Known historically only from type locality near Pu‘u Lehua (see Kona Grosbeak for locality details) where all eight known specimens collected in 1892 (Olson 1999). All taken from same trees in which Greater Koa-Finches were feeding. Fossil remains reported from Barber’s Point, O‘ahu (Olson and James 1982b; Scott et al. 1986) and from

Maui are no longer certain to be R. flaviceps ( James and Olson 1991; Olson 1999), but may still indicate that this sp. had a wider distribution in lowland dry forest in prehuman times.

Diet and foraging Apparently similar to that of Greater Koa-Finch, but Munro (in Perkins 1903) thought it might also feed in ‘a‘ali‘i.

Interspecific behaviour Associated with Greater Koa-Finch in koa trees, but whether they interacted unknown.

Genus Chloridops Wilson Chloridops S. B.Wilson, 1888, Proc. Zool. Soc. London, p. 218. Type, by monotypy, Chloridops kona Wilson. Large Hawaiian finches with massive arched bills (almost as large as remainder of head) adapted for crushing hard dry fruits or seeds.Tomial crest (bony support under the tomium) exhibits ventral bulge about a third of the distance from the tip that sets this genus apart from other drepanidine finches ( James and Olson 1991) whose tomial crests (but not necessarily their rhamphothecae) are evenly decurved. Rhamphotheca of single historically known sp. is also sinuated (Olson 1999). Thick walls of underlying bones ‘encase a dense network of uncommonly

sturdy bony trabeculae that presumably functioned to disperse the strong forces generated when cracking hard seeds with the bill’ ( James and Olson 1991: 36). Tongue thick and fleshy like that of a parrot (Munro in Olson 1999). Nasal opercula lacking (Gadow 1899). No sexual dimorphism in size (Olson 1999) or coloration. Pratt (2001b) suggested possible relationship to Loxioides, but James and Olson (1991) and Olson (1999) regard it as distinctive, and allied Loxioides with Telespiza and Rhodacanthis. James (1998) groups them as Loxioides/Telespiza and Chloridops/Rhodacanthis, but the two are not sister groups. Recent analysis (H. James, pers. comm.) adds Xestospiza conica to latter grouping.

Kona Grosbeak Chloridops kona Wilson PLATES 1, 8 Chloridops kona S. B.Wilson, 1888, Proc. Zool. Soc. London, p. 218. (Kona, Hawaii.) Psittirostra kona Amadon (1950), Berger (1972a) Psittacirostra kona Carlquist (1970) Loxioides kona Greenway (1968) Psittirostra (Chloridops) kona Olson and James (1982b) Other vernacular names: Kona Finch, Grosbeak Finch, Palila (Wilson and Evans 1890–99; Henshaw 1902; apparently in error), Chloridops (Munro 1944, 1960)

Etymology Latin Chloridops could mean either ‘green face’ or better, ‘green-looking’. The epithet refers to the Kona region where the bird was discovered.

Systematics Elsewhere (Pratt 2001b) I hypothesised that Chloridops and Loxioides were sister taxa on the basis of qualitative similarities of bill shape, and suggested their possible merger. I incorrectly ( James 2001)

Kona Grosbeak Chloridops kona 211 cited James and Olson (1991) as suggesting same merger, but they instead suggest Loxioides closer to Telespiza and Rhodacanthis.

Description A medium-sized (15 cm), chunky, short-tailed, dull olive green finch with disproportionately large head and huge bill. No sexual dichromatism, but Juvenal plumage shades to greenishwhite on belly and undertail coverts, with slight orange tinge to breast. Bill brownish-grey, the lower mandible paler and greyer (based on summarised descriptions in Olson 1999). Frohawk (in Wilson and Buff 1989) and Pratt et al. (1987) show the bill as ‘dull flesh-colour’ as the holotype was described by Wilson (1888), but Munro’s unpublished notes (Olson 1999) reveal this plumage to be Juvenal and not simply a variation. Legs and feet Sepia (119). For measurements see Olson (1999).

Voice Call note as described by Palmer a ‘low prolonged “cheep,” not at all loud’ (Rothschild 1893–1900: 210). Munro (1960: 131 and in journal quoted in Olson 1999) describes a ‘light sweet song . . . long with a variety of notes’. That so few songs were heard by early naturalists may simply indicate that their observations took place outside the breeding season (see below).

Distribution, habitat, and population status Extinct; last seen 1894. Historically known only from a 10-km2 area (Perkins 1903) centred around the cinder cone Pu‘u Lehua in the N. Kona Dist. near the S. Kona border, between ca. 1400 and 1500 m elevation, plus a single sighting by Palmer 16 km to the south above Honaunau (Munro 1960; Olson 1999). Disappeared from this area within a decade of discovery. Only 56 specimens in world collections (Banko 1979 as updated by Olson 1999). Specialised for naio, a successional tree on very rough lava flows and in forest openings (Perkins 1903; Munro 1960), often associated with mamane. Olson (1999) suggests fruit size may have been a limiting factor in distribution because naio within the range has smaller fruits with more seeds per fruit than elsewhere (Wagner et al. 1990).

Diet and foraging Ate dried naio fruits almost exclusively, although young undoubtedly fed invertebrates.This fruit has an extremely hard endocarp enclosing 4–10 cells containing one seed each (Wagner et al. 1990). According to Munro’s journal (in Olson 1999), naio fruits cut into pieces in bill and then further processed between edges of mandibles using ‘tooth’ in lower mandible. Journal also reports a few stomachs containing pieces of green naio fruits and bits of green leaves. May have opportunistically also taken softer fruits because pollen from ‘ie‘ie found in feathers of specimens (Cox and Elmqvist 2000).

Field identification

Courtship and breeding

A sluggish and inconspicuous bird easily overlooked except for the cracking sounds of its feeding (Perkins 1903). In coloration very similar to female koa-finches and ‘O‘u, but bill much bigger and more bulbous, the whole head disproportionately large. Calls and songs as inconspicuous as other features. Former range now invaded by several alien finches (House Finch, Saffron Finch, Northern Cardinal,Yellow-fronted Canary), none of which would cause confusion for an experienced birder, but which might confound the beginner bent on rediscovering an extinct bird.

Usually observed in pairs or small family groups. Probably had female-centred territoriality characteristic of carduelines and drepanidines as evidenced by Munro’s description (in Olson 1999: 9) of a male that ‘kept coming round chirping and singing’ after his mate was shot. I once witnessed similar behaviour by a male ‘Akepa whose mate had been mist-netted. Breeding probably took place fall–spring as in many other honeycreepers, because specimens taken July–Oct not in breeding condition (Olson 1999) and birds encountered in small groups that Perkins (1903) regarded as adults leading young.

212 King Kong Grosbeak Chloridops regiskongi

Wahi Grosbeak Chloridops wahi James and Olson Chloridops wahi James and Olson, 1991, Ornithol. Monogr. No. 46: 36. (Barbers Point, Oahu, Hawaiian Islands.) All information from James and Olson (1991) except discussion of vernacular names and as indicated.

Appearance

Etymology

Extinct following arrival of humans but before discovery of Hawai‘i by Europeans (within last 1500 years). Would have been contemporaneous with historical spp. Ranged from Kaua‘i to Maui (Olson 1999). Bones come from Barber’s Point and ‘Ulupau Head on O‘ahu, Pu‘u Naio Cave on Maui, and Makawehi on Kaua‘i, all arid localities that probably supported dry forest (remnants still present near Maui site).

See Kona Grosbeak. Specific epithet from ‘Hawaiian wahi, to cleave or break in pieces, in reference to the function of the bill in cracking hard seeds’ ( James and Olson 1991: 37). A likely Hawaiian name for members of this genus would have been manu wahi.

Systematics Closely related to Kona Grosbeak and to an as-yet undescribed congener from Maui. Kaua‘i population not named by James and Olson (1991) shown by additional specimens to belong to this sp. (Olson 1999).These three share some features that set them apart from King Kong Grosbeak.

Considerably smaller billed than Kona Grosbeak, but otherwise similar.

Distribution, habitat, and population status

Diet and foraging Smaller and weaker bill implies feeding on seeds less difficult to crack than the hard dry fruits of naio on which Kona Grosbeak fed (Olson 1999).

King Kong Grosbeak Chloridops regiskongi James and Olson Chloridops regiskongi James and Olson, 1991, Ornithol. Monogr. No. 46: 40. (Barbers Point, Oahu, Hawaiian Islands.) Other vernacular names: King Kong finch ( James and Olson 1991) All information from James and Olson (1991).

Systematics Several qualitative differences show that the other spp. of Chloridops are more closely related among themselves than any is to C. regiskongi.

Appearance Etymology See Kona Grosbeak. The vernacular name, upon which the epithet regiskongi based, originated with a reporter’s misquotation of Olson as calling the as-yet-unnamed fossil ‘a giant, gargantuan, a King Kong finch’ (Benson 1977). However, the name was popular and has come to be widely used.

Has the largest bill of any known Hawaiian finch and differing in shape from those of congeners in that ‘maxilla is much deeper and is sharply pointed, the sides of the maxilla rising steeply to form a blunt crest that extends along the dorsal midline from the tip through the dorsal nasal bar’ ( James and Olson 1991: 41).

‘O‘u Psittirostra psittacea 213

Distribution, habitat, and population status Extinct following arrival of humans but before discovery of Hawai‘i by Europeans (within last 1500

years). Certainly contemporaneous with surviving spp. Remains come from Barber’s Point and ‘Ulupau Head, arid localities that probably supported dry forest.

Genus Psittirostra Temminck Psittirostra Temminck, 1820, Man. Ornithol. (ed. 2) 1: 70.Type, by monotypy, Loxia psittacea Gmelin. Large, stocky Hawaiian honeycreepers with unique bill morphology, vaguely parrot-like or finchlike, with strongly hooked maxilla overhanging mandible. Has no analogue among other passerines

but some tanagers possess somewhat similar bills without the overhang. Moderate nasal operculum, tongue as in Telespiza (Raikow 1977b). Sexually dimorphic, males brighter than females. Calls loud whistles, song highly complex and loud. Single monotypic sp. known from all forested islands.

‘O‘u Psittirostra psittacea (Gmelin) PLATES 2, 8

Psittacirostra psittacea Wilson and Evans (1890–99), Perkins (1903), Carlquist (1970) Psittirostra olivacea Rothschild (1893–1900), Bryan (1901a), Henshaw (1902)

Bangs (1911) P. p. oppidana from Moloka‘i, Bryan and Greenway (1944) argued both based on specimens within normal range of variation; sp. considered monotypic ever since. Lack of inter-island differentiation attributable to habit of flying high over canopy for long distances, presumably sometimes between islands, or may simply indicate early stage of taxon cycle (see Ch. 4; Tarr and Fleischer 1995).

Etymology

Description

Scientific name translates roughly from the Latin as ‘parrot-like parrotbill’.

A medium-sized (17 cm), finchlike bird (Fig. 11.6). Definitive male bright olive green, greyer on breast and belly, shading to olive-yellow on sides and flanks and off-white on undertail coverts; head sharply demarcated bright golden yellow. Definitive female similar but with head concolour with body and undertail coverts with olive-grey centres producing scaly pattern. Bill and feet orange-pink in both sexes, female duller. Juvenal plumage similar to female, but wing coverts tipped off-white, forming wing-bars. Moult sequence unknown, but individuals may have wing-bars more than one year (Snetsinger et al. 1998). For measurements, see James et al. (1989).

Loxia psittacea Gmelin, 1789, Syst. Nat. 1(2): 844. Based on the ‘Parrot-billed Grosbeak’ Latham, Gen. Synop. Birds 2(1): 108, pl. 42. (Sandwich Islands ⫽ Hawaii.)

Systematics Genus now and originally regarded as monotypic, but expanded by Amadon (1950) to include all Hawaiian finches. Greenway (1968) again made it monotypic and put other finches in Loxioides. Olson and James (1982b) recognised five subgenera within Amadon’s Psittirostra and later Olson and James (1991) raised all to generic level to agree with Banks and Laybourne (1977), Pratt (1979a), Berger (1981), and AOU (1983). Although Rothschild (1905) named P. p. deppei from O‘ahu and

214 ‘O‘u Psittirostra psittacea

¯ ‘u, 11.6 One of the few photographs ever taken of the ‘O Alaka‘i Plateau, July 1975.

native and introduced can sound somewhat like ‘O‘u. Many notes of White-rumped Shama, now well established in the ‘O‘u’s former range on Kaua‘i (pers. obs.), could be mistaken for it and does not always break into more complex songs after slow single whistles. ‘O‘u never varies its beacon-like call once a calling bout begins; an identical whistle may be uttered dozens of times in succession (pers. obs.). On Hawai‘i, cardueline Yellow-fronted Canaries now widespread in potential ‘O‘u habitat (Paton 1981; pers. obs.) and utter very similar, though much quieter, calls and songs. Wishful thinking could even make the cardueline House Finch’s song seem like that of an ‘O‘u.

Voice

Distribution, habitat, and population status

Call notes loud, far-carrying, slow, up-slurred or occasionally down-slurred, rather human-like whistles given from high perch at an even cadence for long periods (Conant et al. 1998; pers. obs.), possibly as ‘sound beacon’ for pairs or family groups.The down-slurred whistle has a particularly melancholy quality. Songs (Fig. 11.7) long and complex, including whistles, warbles, and trills, usually introduced with 2–3 loud whistles tweee-tweeetweee. Henshaw (1901: 123) regarded ‘O‘u as ‘the most beautiful songster of the Hawaiian forest’, saying ‘that it is the generally received opinion among the settlers that the forest is full of escaped cagebirds; yet in purity, sweetness and power the song of the [‘O‘u] far surpasses the Canary’s best efforts.’ However, ‘O‘u not lavish with song, singing only during presumed breeding season. For recordings see Pratt (1996a), for additional sonograms see Snetsinger et al. (1998).

Field identification Because of critical status, accurate field ID of ‘O‘u more important than ever. Colour combination of both male and female, especially with the pink bill, really unmistakable among dreps, and ‘O‘u larger and plumper than any potentially confusing introduced birds (although female Northern Cardinals have been called ‘O‘u!). Because most recent reports have all been audio only, this aspect of ID is especially important. Unfortunately, many spp. both

Originally widespread at all elevations except barren alpine zones on all islands and in all kinds of forest.Wandered widely to exploit seasonally available resources, perhaps even between islands. Possible occurrence of ‘home base’ controversial. Snetsinger et al. (1998) summarised information and considered the ‘ ‘ie‘ie belt’, an artificial mesic to wet belt at 800–1500 m elevation (extended to the coast in many places before extensive clearing of lowlands by Hawaiians), to be ‘true home’. ‘O‘u believed main pollinator of ‘ie‘ie before replacements introduced (Cox and Elmqvist 2000). Bryan (1908) possibly correct that ‘O‘u preferred very wet ‘ohi‘a forest because last remnant populations in that habitat (Alaka‘i Plateau on Kaua‘i and Ola‘a region on Hawai‘i). Probably predominantly low elevation sp., thus highly vulnerable to mosquitoborne diseases (see Ch. 9). ‘O‘u among most abundant Hawaiian birds historically. Declines first noticed on O‘ahu where rare by the 1890s and never reliably reported in twentieth century. Underwent rapid population crashes on several islands. Still numerous at about 1300 m (Banko 1986) on Maui in 1901 (Henshaw 1902) and Molokai in 1907 (Bryan 1908), but not reported from either island again (Banko 1986). On Lana‘i increased as forests recovered in early 1920s, but extirpated there by 1931 (Munro 1944). Still present in low numbers above 800 m around Hawaii Volcanoes NP through 1970s (van Riper

‘O‘u Psittirostra psittacea 215 8

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¯ ‘u, recorded by Gauthey et al. (1968). Numbers refer to LNS catalogue. 11.7 Vocalisations of the recently extinct ‘O

1978a; Katahira 1979; Conant 1980, 1981b). HFBS found remnant population thinly spread between park (Ola‘a Tract) and Saddle Rd., with tiny zone of concentration in kipukas between Kulani Tract and Saddle Rd. between 1300 and 1500 m elevation (Scott et al. 1986). This tiny remaining pocket hit head-on by major lava flow from Mauna Loa in 1984, after which only scattered reports and rumours of ‘O‘u on island (Pratt 1995; Snetsinger et al. 1998; Reynolds and Snetsinger 2001).The tiny ‘current range’ mapped by Snetsinger et al. (1998)

overly optimistic. Last confirmed sighting on Hawai‘i in 1987 in Ola‘a Tract (arrow on map), with only one unconfirmed audio report (1996) since (Reynolds and Snetsinger 2001). Sadly parallel story unfolded on Kaua‘i where population in the low hundreds persisted into the 1970s, then dwindled to a handful by early 1980s (Scott et al. 1986; Conant et al. 1998; Reynolds and Snetsinger 2001). Hurricane Iwa in 1982 (Pyle 1983b; Engilis and Pratt 1989) decimated the remaining population and even more powerful

216 ‘O‘u Psittirostra psittacea

KAUA'I

NI'IHAU

O'AHU MOLOKA'I MAUI _ LANA'I KAHOOLAWE

HAWAI'I

Map 5

Hurricane Iniki in 1992 devastated the high elevation forests (Walker 1993; Pratt 1993) and probably drove native birds into the pestilential lowlands where they could not survive (Conant et al. 1998). Last confirmed sighting of ‘O‘u on Kaua‘i in 1989 (Engilis and Pratt 1989; Pyle 1989a). None found after Iniki (Telfer 1993) and audio reports in the mid-1990s have remained unconfirmed. Reynolds and Snetsinger (2001) consider it extirpated on Kaua‘i. Likely went extinct in last decade of twentieth century.

be one of main pollinators of Freycinetia under pristine conditions (Cox and Elmqvist 2000). When ‘ie‘ie out of season, ‘O‘u wander widely in small groups (Perkins 1903) seeking out other native plants (Figs 8.2, 8.3) that produce soft fruits such as haha, ‘olapa, mamaki, kawa‘u, ‘ohi‘a-ha, and probably ‘ohe (Snetsinger et al. 1998; Mull and Mull 1971). However, ‘O‘u opportunistic, not eschewing such introduced fruits as mountain apple, guava, banana, kuku‘i, wauke, peach, and Citrus. Small fruits eaten whole, larger ones devoured piecemeal.

Diet and foraging A highly eclectic feeder, primarily a frugivore but also eating caterpillars (Wilson and Evans 1890–99), often preferentially during irruptions (Perkins 1903), and nectar (Perkins 1903; Conant et al. 1998). Favoured food plant ‘ie‘ie (Figs 2.9, 8.1), including fruit, flowers, and leaf bracts around inflorescences (Perkins 1903). Believed to

Breeding biology Little known. Nest never discovered, but birds with enlarged ovaries noted in spring (Rothschild 1893–1900). Perkins (1903) observed many juveniles, some barely able to fly, in Kona in June. Begging young seen as late as August (Snetsinger et al. 1998).

Lana‘i Hookbill Dysmorodrepanis munroi 217

Genus Dysmorodrepanis Perkins Dysmorodrepanis Perkins, 1919, Ann. Mag. Nat. Hist., ser. 9, 3, p. 250.Type, by monotypy, Dysmorodrepanis munroi Perkins. Medium-sized Hawaiian finches with peculiar short, heavy bill. Upper mandible strongly hooked, lower mandible strongly up-curved and striking the upper so that a large diastema forms between the

mandibles of the closed bill. In the diastema, the edges of the bill roll inwards forming ‘broad, blunt surfaces that lie roughly parallel to each other at opposite sides of the diastema’ ( James et al. 1989). Tongue non-tubular, ‘not acute at the tip and is apparently emarginate there, with the edges microscopically serrulate’ (Perkins 1919: 251).

Lana‘i Hookbill Dysmorodrepanis munroi Perkins PLATES 2, 8 Dysmorodrepanis munroi Perkins, 1919, Ann. Mag. Nat. Hist., ser. 9, 3, p. 251. (Kaiholena Valley, Island of Lanai.) Psittirostra psittacea Amadon (1950), Greenway (1968), Carlquist (1970) Other vernacular names: Dysmorodrepanis (Munro 1944)

Etymology Genus name from Greek, dusmoros, ⫽ ill-fated, ⫹ drepanis for honeycreeper (Snetsinger et al. 1998). Species named in honour of George C. Munro who collected the holotype.

Systematics Greenway (1937) and Amadon (1950) regarded this bird as an aberrant specimen of ‘O‘u, despite clear evidence from Munro’s comments in original description (Perkins 1919) that sp. valid.To be honest, I also examined type specimen and accepted earlier judgement uncritically. James et al. (1989) extracted the skull from the specimen to convincingly demonstrate sp. validity, possibly related to Psittirostra but with many autapomorphies.

Appearance A chunky bird slightly smaller than ‘O‘u, greyisholive above, very pale below, with bizarre hooked upper mandible and upturned lower mandible that

leave a noticeable gap (diastema) when closed. Head (except nape and crown) and underparts offwhite, tinged yellow, most strongly on forehead and diffuse superciliary. Inner webs and broad tips of tertials white. Sex of single specimen (Fig. 3.2) thought to be possibly female ( James et al. 1989) but Munro (1960, in Perkins 1919) saw other individuals on Lana‘i with same coloration, one of which behaved as male (see below). Large amount of off-white, unusual among Hawaiian honeycreepers, led Amadon (1950) to regard specimen as partially albinistic. Brown bill colour unlike that of any age ‘O‘u. For measurements, see Snetsinger et al. (1998).

Voice Munro (in Perkins 1919: 252) specifically states vocalisations of the hookbill were different from those of ‘O‘u: ‘I heard two or three birds calling to one another, the cry being less sweet and not so loud as that of the [‘O‘u], and I watched one on the bare branch of a tree-top a short distance away. It called regularly at intervals and kept moving its head, stretching its neck and turning on its perch without changing its place on the branch.’ This behaviour nearly identical to that of an ‘O‘u I observed on Kaua‘i in 1975 (Conant et al. 1998). Munro described voice of another individual 2 years later as having some notes like ‘O‘u whistles, but others distinctive, ‘especially a low squeak or whistle’.

218 Genus Oreomystis

Field identification White tips and inner webs of the tertials and inner secondaries form striking white patch on folded wing, reminiscent of smaller white patch on wings of ‘I‘iwi. Munro adamant that hookbill smaller than an ‘O‘u. The bird described calling from a tree-top above was ‘more active [than ‘O‘u] , but less so than [an ‘amakihi]’ (Perkins 1919: 252). For his subsequent sighting Munro ‘near enough to note the light colouring round the eye but not the form of the beak’ and said the bird ‘was too small for [an ‘O‘u], not so thick-set, and with a very short tail’. Considering number of supposed sightings of nukupu‘u in the twentieth century that did not include seeing the unique bill (Pratt and Pyle 2000), above statement not surprising. Times different then, most birdwatching done down gun barrel without optical aids. Frankly, unfortunate that Munro could not obtain another specimen; sp. doomed in any case and might not have languished in obscurity for so long if a second one had been collected.

Distribution, habitat, and population status Extinct; last seen 12 Aug 1918 (Perkins 1919). Inhabited rather small native forest of Lana‘i. Holotype

collected in densely wooded upper Kaiholena Valley above what is now Ko‘ele; subsequent sightings higher up same valley and at Waiakeakua, dry-forest locality some distance to the se., indicating that hookbill may have preferred drier habitats than ‘O‘u. All localities between 600–800 m elevation (Snetsinger et al. 1998). Munro (1960) believed it formerly inhabited large akoko forest replaced by pineapple fields just as last birds were seen. So far, no trace of sp. among subfossils from other islands of Maui-nui, although bones of ‘O‘u abundant (H. James in Snetsinger et al. 1998).

Diet and foraging Single specimen had eaten fruits of opuhe, and Munro (1960) thought the similar-sized fruits of akoko would have also been taken. However, I agree with James et al. that simple frugivory could not provide selection pressure for such a bizarre bill.They speculated that the hookbill may have fed on terrestrial snails, with the diastema in the bill serving as a platform to hold the prey while tongue and bill extracted body of animal from shell. However, only other snail-eating honeycreeper, Po‘o-uli, usually crushes snails and eats them shell and all (T. K. Pratt et al. 1997).

Genus Oreomystis Stejneger Oreomystis (not Pokorny, February 1887, Insecta) Stejneger,April 1887, Proc. U. S. Natl. Mus. 10: 99.Type, by original designation, Oreomyza bairdi Stejneger. Oreomystis Stejneger, 1903, Proc. Biol. Soc.Washington 16: 11. New name for Oreomyza Stejneger, preoccupied. Small, short-tailed Hawaiian honeycreepers with short, straight bills, the gonys of which is concave in profile (Fig. 6.9).Tongue non-tubular, similar to tongues of many insectivorous passerines

(Gardner 1925), but lacking lingual wings (Richards and Bock 1973; Gadow 1899). Primarily insectivorous, foraging nuthatch-like on trunks and branches. No plumage dimorphism, but immatures different from adults, colours muted. Songs simple descending trills, juvenile calls short, in series, unlike those of most other honeycreepers (Pratt 2001b). Two spp., one on Kaua‘i, one on Hawai‘i. Oreomystis replaced Oreomyza (Stejneger 1888), later found to be preoccupied (Stejneger 1903).

‘Akikiki Oreomystis bairdi 219

‘Akikiki Oreomystis bairdi (Stejneger) PLATES 5, 8 Oreomyza bairdi Stejneger, 1887, Proc. U. S. Natl. Mus. 10: 99. (Kauai.) Paroreomyza bairdi bairdi Bryan and Greenway (1944) Loxops maculata bairdi Amadon (1950) Viridonia maculata bairdi Greenway (1968) Paroreomyza maculata bairdi Raikow (1977b) Other vernacular names: Kaua‘i Creeper, Creeper, ‘Akikihi, ‘Akikeke

Etymology Original genus Oreomyza from Greek roots oreo ⫽ mountain and myza ⫽ sucker (a mistaken reference to nectarivory) modified by Stejneger (1903) to Oreomystis so that new name would not look very different from the preoccupied one (Foster et al. 2000). Epithet posthumously honours Spencer Fullerton Baird, zoologist and second secretary of the Smithsonian Institution (Mearns and Mearns 1992).

Systematics See Ch. 5 and O‘ahu ‘Alauahio.Taxonomic history tracked by above synonymy. Current classification not controversial, but whether Oreomystis is monotypic or includes the Hawai‘i Creeper (as here) remains contentious (Pratt 2001b).

feet pale pink, culmen sometimes with dusky grey tinge. Iris dark brown. For measurements, see Foster et al. (2000).

Voice Call notes variable, the most distinctive an up-slurred sweet. Also a sharp whit (Foster et al. 2000) similar to calls of ‘Anianiau and ‘Akeke‘e that may be flock cohesion notes, and a see-u-weet indistinguishable from typical call of ‘Anianiau (Foster et al. 2000). Juvenile calls when following parents syncopated clusters of raspy chit notes very similar to juvenile calls of Hawai‘i Creeper (Pratt 2001b): chi-di-dit, chit-it, chit-chi-dit, etc.Whisper song indistinguishable from those of other Hawaiian honeycreepers. Primary song, rarely uttered, a descending trill that trails off at the end (Fig. 11.8). For additional sonograms see Foster et al. (2000), and (Pratt 2001b); for recordings, Pratt (1996a).

kHz

Oreomystis bairdi 05055

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Description A small (13 cm) grey and white Hawaiian honeycreeper with conical straight bill with concave gonys profile (Fig. 6.9) and relatively short tail. Sexes alike, but juveniles differ from adults. Definitive plumage olive-grey above, sharply demarcated from white underparts tinged grey posteriorly including flanks. May have very slight yellowishcream wash on breast. Juvenal and First Basic plumage similar but with broad white ring encircling eye, sometimes meeting over bill, and pale wing-bars not present in all individuals, possibly the result of wear (Foster et al. 2000). Bill, legs, and

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11.8 Comparison of songs of two species of Oreomystis; numbers refer to LNS catalogue.

220 ‘Akikiki Oreomystis bairdi

Field identification Not likely to be visually confused with any other Kaua‘i forest bird except Kaua‘i ‘Elepaio, which is also mainly grey but has white rump, wing-bars, and tail tip, and very different shape and postures. Vocalisations problematic. Similar song sometimes sung by Kaua‘i ‘Amakihi, and calls mirrored in repertoires of ‘Akeke‘e and ‘Anianiau.

Distribution, habitat, and population status Endemic to Kaua‘i, and probably found islandwide in prehuman times as indicated by subfossil remains from Makawehi Dunes at sea level (Olson and James 1982b). Collectors in the 1890s found it common in forests above 300 m, and it remained ‘abundant’ in the Alaka‘i region above 1140 m into the 1960s (Richardson and Bowles 1964). Gauthey et al. (1968) reported a decline in numbers, and 1968–73 surveys by John Sincock found it only in the Koke‘e and Alaka‘i regions and on isolated La‘au Ridge (Sincock et al. 1984; Scott et al. 1986). By early 1980s had virtually disappeared from temperate mixed forest and koa forest of Koke‘e SP (Conant et al. 1998). Now found mostly in more

KAUA'I

remote part of Alaka‘i Plateau, from the Alaka‘i Swamp Trail eastward, in greatly reduced numbers (Pratt 2002b), though ‘ohi‘a rainforest may not be optimal habitat. Major limiting factor probably mosquito-borne disease (Foster et al. 2000; see Ch. 9). Decline continues apace, sp. recently proposed for classification as endangered sp. ( J. Foster, pers. comm.).

Diet and foraging Diet almost entirely arthropods, primarily caterpillars and spiders, but including myriapods, slugs, and beetles (Perkins 1903). Very rarely takes nectar (Conant et al. 1998; Foster et al. 2000). Forages mainly on larger trunks and branches of ‘ohi‘a and koa trees, often on dead branches. Forages occasionally in forest understorey on kawa‘u, kalia, ‘olapa, haha‘aiakamanu, and ferns (Bryan and Seale 1901; Perkins 1903; Foster et al. 2000). Virtually all observers have compared movements to those of nuthatches (Sittidae), but does not brace with tail. Creeps along and around substrates, often hanging upside down.Tarsus shorter and hallux better developed than in other honeycreepers (Stejneger 1887). Crouches low with legs and feet usually hidden, as does Hawai‘i Creeper but unlike Paroreomyza.

Social and interspecific behaviour Often encountered in small family groups or postbreeding assemblages of up to 12 individuals (Wilson and Evans 1890–99; Munro 1960; Richardson and Bowles 1964), although groups usually smaller today (Foster et al. 2000). Joins mixed-species flocks, but not ‘nuclear species’ as suggested by observations of Munro (1960) because flocks often form without ‘Akikiki (pers. obs.). Historically, such flocks included ‘Anianiau, Kaua‘i ‘Amakihi, ‘Akeke‘e, Kaua‘i ‘Akialoa, Kaua‘i Nukupu‘u, and Kaua‘i ‘Elepaio (Perkins 1903).

Predation

Map 6

Actual predation never observed, but undoubtedly prey of Kaua‘i Stilt-Owl as evidenced by remains in pellet deposits (Olson and James 1982b). Flocking behaviour probably evolved in response to aerial predators.

Hawai‘i Creeper Oreomystis mana 221

COURTSHIP AND MATING: Probably monogamous, but data few.

EGGS: From Eddinger (1972b). Only one known, 18.3 ⫻ 23.3 mm, white speckled brown with spots concentrated at larger end. Clutch size unknown but one nest held two chicks.

NEST SITE AND TERRITORY: Only five nests ever reported, all 8–10 m high, four in ‘ohi‘a and one in ‘ohi‘a-ha (Eddinger 1972b; Foster et al. 2000). Territoriality unknown.

INCUBATION AND HATCHING: Only female incubates, sitting tightly on nest despite disturbance (Berger 1981). Incubation period unknown.

NEST: Both sexes build, but proportions unknown (Eddinger 1972b) and male may not participate (Foster et al. 2000). Open cup made mostly of mosses (especially outer shell), cup lined with plant fibres including strips of ‘ohi‘a and ‘olapa bark and rootlets (Eddinger 1972b; Foster et al. 2000).

NESTLINGS: From Eddinger (1972b). Chicks already present when only nest studied discovered, so ages at particular events unknown. Remnants of natal down white, contour feathers patterned as in First Basic plumage. Gape bright pink with yellow rictal flanges.

Breeding biology

Hawai‘i Creeper Oreomystis mana ( Wilson) PLATES 5, 8 Himatione mana S. B. Wilson, 1891, Ann. and Mag. Nat. Hist. (6)7: 460. (Hawaii.) Oreomyza mana Perkins (1903) Paroreomyza bairdi mana Bryan and Greenway (1944), Munro (1944) Loxops maculata mana Amadon (1950) Viridonia maculata mana Greenway (1968) Paroreomyza maculata mana Raikow (1977b) Loxops mana Olson and James (1982b), James and Olson (1991) Other vernacular names: Olive-green Creeper, Creeper

Etymology See ‘Akikiki for genus. Species apparently named for Mana, a locality on the Island of Hawai‘i.

Systematics For Oreomystis, see O‘ahu ‘Alauahio and Ch. 5. Relationships of this sp. the most controversial among authorities today (Pratt 2001b). For the classification used here, based largely on external phenotypic characters, see Ch. 5 and Pratt (1992b, 2001b). Olson and James (1995), presumably on

osteological grounds, allied O. mana with ‘amakihis, but James’s (1998) osteologically based phylogeny (Fig. 4.7) places it as sister group to virtually all the thin-billed honeycreepers. Genetic studies inconsistent: Feldman (1997) considers it the sister group of the ‘red and black’ honeycreepers (Fig. 4.6c); Fleischer et al. (1998, 2001) hypothesise a relationship to ‘akepas (Figs 4.6d, e). All the alternative hypotheses require that the large number of similarities in plumage dimorphism and maturation, bill shape, tongue morphology, foraging behaviour, and juvenile vocalisations between this sp. and O. bairdi be the result of convergence.

Description Small honeycreeper with relatively short tail and short, conical, very slightly decurved bill. No sexual plumage differences, but immatures distinct. Definitive adult greyish-green above, paler and tinged olive-yellow below, with white throat and pale yellowish-olive cheeks and diffuse superciliary (none of these sharply demarcated); dark grey triangular mask extends from base of maxilla to point behind eye. First Basic plumage similar but paler below with little yellow tinge, prominent welldefined yellowish-white superciliary, and mask

222 Hawai‘i Creeper Oreomystis mana much narrower and paler. Juvenal plumage similar but paler still, approaching white, below, greyer above and with somewhat broader superciliary. Bill very pale horn colour, often appearing nearly white, legs and feet darker horn colour. Iris dark brown. Standard measurements given by Lepson and Woodworth (2002).

HAWAI'I

Voice Call note an up-slurred quiet sweet. Begging calls of juveniles wheezy whit notes in syncopated bursts: whit-whit, whi-whi-whit, whit-whit, etc. Song a rattling descending trill,‘drier’ and faster than those of other spp. (Fig. 8).Whisper song rarely heard, but described by VanderWerf (1998: 542) as containing ‘fragments of typical descending trills mixed with a variety of soft calls and mimicry of other spp., including ‘Elepaio . . . and ‘I‘iwi’. For additional sonograms see Scott et al. (1979); Pratt (1992b) and Lepson and Woodworth (2002); for recordings, Pratt (1996a).

Field identification One of the more difficult ID problems among honeycreepers (Peterson 1961; Scott et al. 1979).All ages easily confused with female and immature Hawai‘i ‘Amakihi, but note white throat, pale bill, and broader mask, giving creeper a wide-eyed look as compared to squinty-looking ‘amakihi. Creeper’s green tones of a less yellow hue than those of ‘amakihi, a good mark once appreciated. Immature creeper very similar to juvenile ‘Akepa, but note latter’s lack of dark mask and very different behaviour and voice. Note that shadows among fluffed feathers can produce effect of dark lores even when no dark feathers are present; masks of some creepers mottled with paler feathers.

Distribution, habitat, and population status Originally occupied a wide variety of forest habitats on Island of Hawai‘i (endemic) from mamane–naio forest above 1000 m in Kona down to below 600 m in very wet rainforests above Hilo, but distribution was uneven at lower elevations (Henshaw 1902; Perkins 1903). Common in Hawaii Volcanoes NP above 1100 m into 1940s (Baldwin 1953), but disappeared very suddenly from vicinity of Kilauea Caldera in late 1930s

Map 7

(Dunmire 1961; Scott et al. 1986) and from rest of park in 1950s (Banko and Banko 1980). By 1980, population of ca. 12 400 in four fragments, the largest in N. and S. Hilo districts between 900 and 2100 m and in Ka‘u FR between 700 and 1900 m, with low-density relicts in Kona above 1500 and on nw. slope of Hualalai; highest densities at higher elevations (Scott et al. 1986). Not known to have declined since 1980. Today found mostly in koa/‘ohi‘a forest, but still occasionally, at least, visits mamane forest on Mauna Kea (Snetsinger 1996).

Diet and foraging Diet mainly arthropods (insects, spiders, myriapods, etc.), worms, and snails (Perkins 1903; Sakai and Johanos 1983) picked from cracks and crevices in the bark of larger limbs and trunks of trees. Larger vermiform prey killed by beating against branch and torn into pieces before being consumed (Sakai and Johanos 1983). Favours koa (Perkins 1903; Pratt 1979a) but also forages in other trees; tends to forage over smaller branches than its congener on Kaua‘i

Hawai‘i Creeper Oreomystis mana 223 (Pratt 1992b), choosing larger branches of subcanopy (Scott et al. 1979). Movements resemble those of nuthatches (Sittidae) as described for ‘Akikiki.

Social and interspecific behaviour Family groups forage together for ⬎3 weeks after breeding season but eventually break up and join large mixed-species flocks with ‘Akepa, Hawai‘i ‘Amakihi, ‘Akiapola‘au, Hawai‘i ‘Elepaio, and ‘I‘iwi (pers. obs.; Pratt et al. 1977; VanderWerf 1998). Can be most numerous species in such flocks (Lepson and Woodworth 2002). Predation suspected but undocumented (Woodworth et al. 2001). Gives alarm call and takes cover when Hawaiian Hawk nearby (Lepson and Woodworth 2002). Nests may be raided for materials by ‘Apapane, ‘I‘iwi, and Hawai‘i ‘Amakihi (VanderWerf 1998;Woodworth et al. 2001).

Breeding biology COURTSHIP AND MATING: Mating display similar to that of other dreps, with male hopping or flying back and forth in front of female, but usually accompanied by whisper song (VanderWerf 1998). Courtship feeding, solicited by female crouching, quivering wings, and uttering begging notes, observed during nest construction (VanderWerf 1998). Monogamous, pairs remaining together for more than 1 year (Ralph and Fancy 1994b). NEST SITE AND TERRITORY: In tall canopy trees, mostly ‘ohi‘a but also koa in proportion to its availability (Sakai and Johanos 1983; VanderWerf 1998; Woodworth et al. 2001). Usually an open cup supported from below (as on horizontal branch or in crotch), but sometimes in cavities, terminal leaf clusters, or space between bark and trunk (VanderWerf 1998;Woodworth et al. 2001). Family groups highly philopatric (Lepson and Woodworth 2002), but individual wanderers also reported (Snetsinger 1996). Home ranges not defended and broadly overlap those of conspecifics (Ralph and Fancy 1994b; VanderWerf 1998 Lepson and Woodworth 2002). Males, and sometimes females, defend area within 20 m of nest, a fairly large territory for a Hawaiian honeycreeper, but much smaller than ⬎100 m distance between nests (VanderWerf 1998). NEST: Construction requires 9–19 d (Sakai and Johanos 1983;VanderWerf 1998; Woodworth et al.

2001). Female builds with minimal help from male (Sakai and Johanos 1983; VanderWerf 1998). Materials appear to vary depending on locality: body of nest mainly ‘ohi‘a bark strips in Hakalau Forest NWR (VanderWerf 1998;Woodworth et al. 2001), bryophytes, ‘olapa twigs, and fern parts (pulu, trunk fibres, and rhizomes), and sometimes spider silk in Keauhou/Kilauea region above Hawaii Volcanoes NP (Sakai and Ralph 1980; Sakai and Johanos 1983). Exterior of both camouflaged with lichens and bryophytes. EGGS: Bluish-white with brown splotches concentrated at larger end, average (n ⫽ 2) size 19.65 ⫻ 14.35 mm (Sakai and Johanos 1983).Average (n ⫽ 11) clutch size 2.1 (Woodworth et al. 2001). INCUBATION AND HATCHING: Female only incubates, fed by male, usually off nest, but also foraging on her own, giving begging display at male’s approach while still on nest (Sakai and Johanos 1983) and sometimes fed there (VanderWerf 1998). Incubation period 13–14 d in wild birds (Sakai and Johanos 1983; VanderWerf 1998), enigmatically longer (16–17 d) in artificial incubation (Woodworth et al. 2001). NESTLINGS: From Sakai and Johanos (1983). Naked (except for grey down) and helpless at first, eyes closed, gape orange-pink, bill yellow. Eyes open and pin feathers present 5–8 d. Fully feathered by 11 d. PARENTAL CARE AND FLEDGING: Female only broods; majority of feeding (by regurgitation) by male, either of female or directly to chicks (Sakai and Johanos 1983; VanderWerf 1998). Nestling period 18–20 d (Sakai and Johanos 1983;VanderWerf 1998; Woodworth et al. 2001). Parents remove faecal sacs.

Life cycle and demography Ralph and Fancy (1994a) report protracted breeding season (Nov–Sep) with peak in May, Woodworth et al. (2001) report season Jan–June, with peak Feb– May. Breeding and moulting overlap in few individuals (Ralph and Fancy 1994a; Woodworth et al. 2001). Nest success rate 30%, average of 1.7 chicks fledging per successful nest. Adult annual survival very high, 70% (Ralph and Fancy 1994a) to 88% (Woodworth et al. 2001). Limiting factors mainly low reproductive potential and high rates of nest failure, possibly due to predation (Woodworth et al. 2001).

224 Pololei Shovelbill Vangulifer neophasis

Genus Vangulifer James and Olson Vangulifer James and Olson, 1991, Ornithol. Monogr. 46: 62.Type, by original designation, Vangulifer mirandus James and Olson. From James and Olson (1991) except as noted. Hawaiian honeycreepers having long and delicate but broad and deep bills with unique blunt or rounded tip. Retroarticular processes short. Has no

living analogue among passerines, so how bill was used difficult to speculate. Bill too weak for seedeating, too broad for bark probing, too blunt for nectarivory, and unsuited to forceful gaping. Possibly used to capture insects on the wing. Two spp., both as yet found only on Maui. James’s (1998) phylogenetic topology suggests genus may be paraphyletic.

Kiwi Shovelbill Vangulifer mirandus James and Olson Vangulifer mirandus James and Olson, 1991, Ornithol. Monogr. 46: 63. (Maui, Hawaiian Islands.) All information from James and Olson (1991).

Etymology Latin name of genus translates ‘little shovel bearer’; specific epithet means wonderful or strange. Vernacular name from Hawaiian word for ‘curved’ or ‘bent’, undoubtedly connected etymologically to name of New Zealand Kiwis (Apterigidae), but pronounced with ‘v’ sound.

Description A medium-sized honeycreeper with an odd, slightly decurved, blunt-tipped bill.

Distribution, habitat, and population status Extinct. Known only from subfossil mandibles and maxillae from Pu‘u Naio Cave and Lua Lepo, Maui.

Diet and foraging Has some features in common with ‘flycatchers’ of various kinds, but curved bill not one of them.

Pololei Shovelbill Vangulifer neophasis James and Olson Vangulifer neophasis James and Olson, 1991, Ornithol. Monogr. 46: 65. (Maui, Hawaiian Islands.) All information from James and Olson (1991).

Etymology See Kiwi Shovelbill. Specific epithet from Greek meaning ‘new-looking’.Vernacular name is Hawaiian word for ‘straight’.

Systematics Differences between this sp. and V. mirandus possibly sufficient to warrant separate genera ( James and

Olson 1991). V. neophasis may be more closely related to Oreomystis and/or Paroreomyza than to V. mirandus ( James 1998).

Description Fairly large honeycreeper with straight, blunttipped bill.

Distribution, habitat, and population status Extinct. Known only from subfossil mandibles and maxillae from Pu‘u Naio Cave, Pu‘u Makua Cave, and Lua Lepo, Maui.

Curve-billed Gaper Aidemedia zanclops 225

Genus Aidemedia James and Olson Aidemedia James and Olson, 1991, Ornithol. Monogr. 46: 66.Type, by original designation, Aidemedia lutetiae James and Olson. Hawaiian honeycreepers with long, straight to somewhat decurved bills, at least superficially similar

to that of Greater ‘Amakihi (Olson and James 1991) with which James (1998) groups this genus in single clade, independent of that containing other ‘amakihis but allied with ‘akepas. Long retroarticular processes indicate forceful gaping motions in feeding.

Straight-billed Gaper Aidemedia chascax James and Olson Aidemedia chascax James and Olson, 1991, Ornithol. Monogr. 46: 67. (Barbers Point, Oahu, Hawaiian Islands.) All information from James and Olson (1991).

Description Honeycreepers with long, straight bills that may have resembled those of European Starling or American meadowlarks.

Etymology Genus named in honour of Joan Aidem, who pioneered in the collecting of subfossil bird bones on Moloka‘i. Spelling does not follow usual conventions in order to form a palindrome. Specific epithet is Greek for ‘gaper’.

Distribution, habitat, and population status Extinct. Known only from subfossil remains from Barber’s Point, O‘ahu.

Curve-billed Gaper Aidemedia zanclops James and Olson Aidemedia zanclops James and Olson, 1991, Ornithol. Monogr. 46: 69. (Barbers Point, Oahu, Hawaiian Islands.) All information from James and Olson (1991).

Distribution, habitat, and population status Extinct. Known only from subfossil remains from Barber’s Point, O‘ahu.

Etymology See Straight-billed Gaper. Specific epithet based on Greek words for ‘sickle’ and ‘face’ in reference to the down-curved bill.

Description Honeycreepers with long, shallowly curved bills (not as strongly decurved as in Hemignathus or Drepanis).

Diet and foraging Because the bills of this and preceding sp. are more similar, except for curvature, to each other than to that of following sp., possibility exists that they were male and female of same sp., with different or even co-operative foraging techniques, as seen in the New Zealand Huia Heteralocha acutirostris (Callaeidae).

226 ‘Akepa Loxops coccineus

Maui-nui Gaper Aidemedia lutetiae James and Olson Aidemedia lutetiae James and Olson, 1991, Ornithol. Monogr. 46: 71. (Moomomi Dunes, Molokai, Hawaiian Islands.) All information (except comments on name) from James and Olson (1991).

Description Honeycreepers with long, straight bills convergent in morphology with those of meadowlarks.

Etymology

Distribution, habitat, and population status

See Straight-billed Gaper. Specific epithet based on Latin name for Paris in an attempt at humour in the form of an extremely convoluted pun.

Extinct. Known only from subfossil remains from Moloka‘i (Mo‘omomi) and Maui (Pu‘u Naio and Pu‘u Makua caves and Lua Lepo).

Genus Loxops Cabanis Loxops Cabanis, 1847, Arch. Naturgesch. 13: 330. Type, by original designation, Fringilla coccinea Gmelin. Chrysomitridops Wilson, 1889, Proc. Zool. Soc. London, p. 445.Type, by monotypy, C. caeruleirostris Wilson. Small Hawaiian honeycreepers with short, conical, finchlike bills, the tips of which are crossed as an adaptation for opening imbricated leaf buds of ‘ohi‘a and seed pods of koa. Possesses drepanidine tubular

tongue and occasionally takes nectar, but primarily insectivorous.Tail long and rather deeply notched at tip. Sexually dichromatic, but degree varies among spp. High-pitched whistled calls, song of varied trills.Two spp., one polytypic with three subspp., but Pratt and Pratt (2001) suggest that all forms may prove to be full spp.Taxonomic history of this genus is discussed in Ch. 5. Loxops subsumes the name Chrysomitridops proposed by Wilson (1889) for the ‘Akeke‘e (Perkins 1903).

‘Akepa Loxops coccineus (Gmelin) PLATES 6, 8 Fringilla coccinea Gmelin, 1789 Syst. Nat. 1(2), p. 192. Based on the ‘Scarlet Finch’ Latham, Gen. Synop. Birds, 2(1), p. 270. (in insulis Sandwich ⫽ Hawaii.) Other vernacular names: ‘Akakane, ‘Akepeuie

1997; Lepson and Pratt 1997). Hawaiian name often translated as ‘lively’ or ‘nimble’, but more likely based on kepa, ‘to cut obliquely’ or ‘turn to one side’ (Pukui and Elbert 1971) an allusion to bill shape (Lepson and Freed 1997).

Etymology Loxops from Greek loxos ⫽ crosswise and ops ⫽ face, usually translated as ‘twisted face’ in reference to the crossed bill tips (Berger 1981); or possibly based on opsis ⫽ having appearance of, a comparison with crossbills of the genus Loxia (Lepson and Freed

Systematics Three ‘megasubspecies’ phylogenetic spp. and possibly biological spp. (Pratt and Pratt 2001): L. c. coccineus on Hawai‘i; L. c. ochraceus on Maui; and L. c. wolstenholmei on O‘ahu. Status moot at present

‘Akepa Loxops coccineus 227 (Pratt 1989b); resolution dependent on DNA studies because O‘ahu and Maui representatives extinct or nearly so (see below). For use of name wolstenholmei instead of longstanding rufus, see Olson (1986).

Description Very small (10 cm) honeycreeper with long notched tail and short conical bill whose mandible is curved to one side (ca. 1:1 ratio of left- to right-billed birds). Sexual dichromatism striking. Males (on Hawai‘i at least) exhibit delayed plumage maturation, not reaching Definitive plumage until third year (Lepson and Freed 1997).Three subspp. differ in colour. HAWAI‘I: From Lepson and Freed (1997) and pers. obs. Definitive male bright red-orange or vermilion over whole body, wings and tail dark brown, edged red-orange. Definitive female grey-green or pale olive, much paler below but without striking contrast, with broad yellow-orange breast band (a few with more orange below), wing and tail feathers dark brown edged grey-green. Juvenal sexes alike, olive-grey above, off-white to cream below, with paler face and superciliary line, wing feathers edged olive-yellow, some with olive-yellow shoulders, sides of breast, and flanks. First Basic plumage similar (sexes alike), but supercilium less pronounced and olive-yellow colour lacking. In Second Basic plumage, males begin to differ from females, becoming blotchy with more than half of body feathering some shade of orange or brownish-orange (variable), underparts cream to orange. Second Basic female lacks supercilium and may have trace of orange on breast. Juvenal bill brownish-grey, darkest along culmen. Bill of post-Juvenal variable (not age related), most often straw yellow but sometimes grey. MAUI: From Lepson and Freed (1997), Pratt et al. (1987), and Pratt (1989b). Definitive males dichromatic, ca. 45% orange, not as brilliant as in Hawai‘i males, 45% a peculiar mustard yellow unlike the yellow of any other Hawaiian honeycreeper, 10% intermediate. Females similar to but duller than Hawai‘i females and lack orange in breast. Juveniles average

slightly darker than same-age Hawai‘i birds. First and Second Basic plumages poorly known. Bill grey. O‘AHU: Definitive males brick red (Pratt 1989b), dull rufous, or ochraceous-orange (Lepson and Freed 1989). Definitive females like Maui birds but duller and slightly darker below. Juvenal as for Maui, other plumages poorly known. Bill grey.

Voice All subspp. have similar vocalisations, although those on Maui and O‘ahu poorly known (Perkins 1903; Engilis 1990). Calls may have 1–3 notes, the most characteristic a high-pitched quick teedle-eedee and a simpler cheedlee; a single sweet very similar to call of Hawai‘i Creeper. Alarm call (in response to aerial predators) a rapid series of seet notes (Lepson and Freed 1997).Whisper songs similar to those of other honeycreepers uttered by both sexes, usually while perched but sometimes while foraging (pers. obs.; Lepson and Freed 1997). Primary song (Fig. 11.9) a variable slow, listless trill that may shift cadence or pitch in mid-strophe (Pratt et al. 1987), usually descending, the effect enhanced by decreased volume toward end (Lepson and Freed 1997). For additional sonograms, see Scott et al. (1979), Pratt (1989b), and Lepson and Freed (1997); for recordings, Pratt (1996a).

Field identification Males of all subspp. very distinctive except for yellow Maui ones, which are yellower above with

kHz

Loxops coccineus 8 06046 4 Loxops caeruleirostris 8 05038 4 1

2

3

sec

11.9 Comparison of songs of two species of Loxops; numbers refer to LNS catalogue.

228 ‘Akepa Loxops coccineus distinctive tawny tinge as compared to Maui ‘Alauahio. Females could be confused with various other ‘little green birds’ (Scott et al. 1979; Pratt et al. 1987), especially juvenile Hawai‘i Creeper, most easily distinguished by behaviour (Lepson and Freed 1997). ‘Akepa never has dark lores like Hemignathus spp. ‘Alauahios both have longer, warbler-like bills. Female and juvenile Yellow-faced Grassquit easily mistaken for female O‘ahu ‘Akepa but browner, with darker bill, and pale yellow throat often bordered by mottled dark malar streak, tail not notched. Grassquit feeds near ground, ‘Akepa in canopy.

Distribution, habitat, and population status HAWAI‘I: Historically found throughout island in suitable forest habitat, but apparently more common at higher elevations (Lepson and Freed 1997). Formerly in lowland koa/‘ohi‘a forest (Henshaw 1902) and temperate mixed forest (Perkins 1903; Richards and Baldwin 1953). Locally common at high elevations into 1950s (Richards and Baldwin 1953). Current distribution patchy above 1100 m (Scott et al. 1986) on e.

and s. slopes of Mauna Kea and Mauna Loa correlated with old-growth ‘ohi‘a and koa forest that provides suitable nest sites (Freed 2001; Hart 2001). MAUI: Probably originally widespread in forests to sea level, but by time of first collection in 1879 (Finsch 1880) restricted to E. Maui but locally common (Perkins 1903). Very rare in twentieth century, most reports (summarised by Conant 1981b; Scott et al. 1986) from ‘ohi‘a rainforest at high elevation on e. and n. slopes of Haleakala. Last sighting of two birds in 1988 (Engilis 1990), only unconfirmed audio detections since (Reynolds and Snetsinger 2001). May be extinct.World collections hold 62 specimens (Banko 1979; Lepson and Freed 1997). O‘AHU: Extinct. Only 20 specimens known in collections (Banko 1979; Lepson and Freed 1997). Common in early nineteenth century but rare by 1890s when last specimens taken (Perkins 1903). Only one ‘convincing’ report after turn of twentieth century (Lepson and Freed 1997), that of Donaghho (1963) in 1930s.

O'AHU MOLOKA'I MAUI _ LANA'I KAHOOLAWE

Map 8

HAWAI'I

‘Akepa Loxops coccineus 229

Diet and foraging Eats primarily spiders and insects, especially psyllid larvae and eggs, lepidopteran larvae, leafhoppers, lacewings, and bugs (Nabidae; Lepson and Freed 1997), but also sometimes takes nectar (Perkins 1903). Forages by methodically probing in terminal leaf clusters, apparently opening leaf buds in the manner described for ‘Akeke‘e (q.v.), and also pries apart koa leaves and pods bound together by spider or caterpillar silk (Lepson and Freed 1997).

Social and interspecific behaviour From Lepson and Freed (1997).After breeding season ( June–Oct) family groups form nucleus of mixed-species foraging flocks (Ch. 7) that may include >40 ‘Akepa. Remainder of year solitary or in pairs (except during displays; see below).

Predation Fledglings taken by Hawaiian Hawk (Lepson and Freed 1997), but apparently not adults.

Breeding biology All information from Lepson and Freed (1995, 1997) in reference to Hawai‘i form except as noted. COURTSHIP AND MATING: Unique (among honeycreepers) prebreeding group displays by males resemble lek behaviour, though pair-bonds monogamous and long-lasting. Named group displays include: Song Bouts; Grappling Fights (males attack each other with bills and feet while calling vigorously, sometimes fall to ground); Arboreal Display (males sing and make short hops in canopy in sight of female); and Aerial Display (groups of males sing and engage in high (up to 100 m) ‘twisting dogfights’. Perkins (1903) reported possible Aerial Display on Maui. Courtship Display typical of Hawaiian honeycreepers, male hopping back and forth on tree branch in front of female. Both sexes explore for potential sites as much as 6 mo. prior to nesting, but form pair-bond before choosing. NEST SITE AND TERRITORY: Always in cavities in ancient koa and ‘ohi‘a trees on Hawai‘i, but only observation of Maui nest (Perkins 1903) was in

terminal ‘ohi‘a branch. Cavities mostly openings in wood, but also bark crevices and underneath bark (Freed et al. 1987b). Dependent on natural cavities, but will nest in suitably camouflaged artificial cavities ( J. K. Lepson, pers. comm.; Pyle and Donaldson 2001). No site-dependent territoriality; male practices ‘roving mate defense’. NEST: Cup-shaped nest built entirely by female from rootlets, twigs, Usnea lichens, bark strips, moss, grasses, and even non-native pine needles (for detailed list see Sincock and Scott 1980; Collins 1984). May reuse nest cavity from previous year. Removal of faecal sacs fastidious until just before fledging, when sacs may accumulate on nest rim. EGGS: Dull white with brown and black blotches concentrated at larger end, 18 ⫻13.4 mm (averaged from data in Lepson and Freed 1997; Sincock and Scott 1984; Collins 1984). Clutch 1–3, median 2. INCUBATION AND HATCHING: Only female incubates, but is fed by male. Female responds to male calls while on nest with distinctive double-note calls. Incubation period not well known, 14–16 d. NESTLINGS: Helpless at hatching, with eyes closed, naked except for sparse grey down on dorsal tracts. Skin orange, bill dark orange, gape reddish-pink with yellow rictal flanges. Maturation little known because of inaccessibility of nests. Fully feathered by 12 d. PARENTAL CARE AND FLEDGING: Female broods up to 8 d after hatching. Both parents feed young by regurgitation. Fledging at 16–20 d. Premature departure of young from nest inferred from indirect evidence. Earliest fledging 20 Apr, latest 30 June. Parents may feed chicks up to 10 weeks after fledging.

Life cycle and demography From Lepson and Freed (1997). Breeding season Mar–Oct, eggs laid Mar–June. Both sexes usually breed in third year, but may attempt it earlier. Annual reproductive output low (usually one/pair), but success rate of nests 79%. Females breed up to 7 years. Adult annual survival rate can be ⬎80% in prime habitat. Potential life span at least 10 years.

230 ‘Akeke‘e Loxops caeruleirostris

‘Akeke‘e Loxops caeruleirostris (Wilson) PLATES 6, 8 Chrysomitridops caeruleirostris Wilson, 1890, Proc. Zool. Soc. London, p. 373. (Kauai.)

Other vernacular names:‘O‘u-holowai, Kaua‘i ‘Akepa

Kaua‘i. Songs variable high-pitched, vigorous trills with abrupt shift ca. midway through (Fig. 9). Singing most frequent Mar–June (breeding season). Whisper song very similar to those of Kaua‘i ‘Amakihi, ‘Anianiau, and ‘Akikiki. For additional sonograms see Pratt (1989b), Lepson and Pratt (1997); for recordings, Pratt (1996a).

Etymology

Field identification

See ‘Akepa for genus. Epithet Latin for ‘pale blue bill’. Hawaiian name incorporates ke‘e ⫽ crooked or bent, alluding to crossed bill tips.

Black face mask and contrasting cap and rump separate ‘Akeke‘e from other ‘little green birds’ on Kaua‘i. Yellow rump especially useful for birds in flight, notched tail for distant perched birds. Methodical probing in leaf clusters a useful behavioural cue. Songs distinctive, but characteristic call very similar to possible flock cohesion notes uttered by several other Kaua‘i honeycreepers.

Loxops coccineus caeruleirostris Amadon (1950), Greenway (1968), Berger (1981), AOU (1983)

Systematics Originally placed in own genus (Wilson 1889), which Bryan (1901a) regarded as a subgenus of Loxops, and Henshaw (1902) and Perkins (1903) synonymised with it. Specific distinctness maintained by all authors until Amadon (1950). Re-split on basis of numerous isolating mechanisms (Pratt 1989b; AOU 1991).

Distribution, habitat, and population status Endemic to Kaua‘i, where widespread in upland forests in nineteenth century (Bryan and Seale

Description Small (10 cm) Hawaiian honeycreeper with conical, cross-tipped bill thicker than that of ‘Akepa. Finchlike, with noticeably notched tail. Olive green above, yellow below, with contrasting yellow ‘cap’ (forecrown and forehead) and rump, and black or dark grey triangular mask extending from base of bill to point just behind eye. Bill very pale blue, legs and feet dark brown or black, iris dark brown appearing black in field. Sexual dichromatism slight, females less golden yellow than males with reduced face mask not encircling base of mandible. First Basic plumage Definitive. Juvenal plumage much greyer than adults with buffy-yellow underparts and no distinct mask. For detailed colour notes and standard measurements, see Lepson and Pratt (1997).

KAUA'I

Voice Call ‘a loud, up-slurred sweet! or peek!’ with piercing or ringing quality (Lepson and Pratt 1997).Also has calls similar to those of other flocking spp. on

Map 9

Genus Magumma 231 1901; Perkins 1903; Munro 1960). Remains common to uncommon in much reduced area above 1000 m in Koke‘e and ‘Alaka‘i regions (Lepson and Pratt 1997; Conant et al. 1998). Probably more common than reports by inexperienced birders indicate (Lepson and Pratt 1997; Pratt 2002b). Whether isolated population present in 1970s in Makaleha Mts. (Scott et al. 1986) persists unknown. Presently occupies 10–12% of historical range (Scott et al. 1986).

Diet and foraging From Pratt (1989b) and Lepson and Pratt (1997). An ‘ohi‘a specialist, foraging on outer branches and terminal leaf clusters of canopy. Rarely, forages in other plants such as kolea, ‘ohelo, and ‘ohi‘a-ha. Feeds mostly on arthropods, especially spiders, but including psyllids and caterpillars among others. May forage in flowers but not known to take nectar. Extracts prey from between scales of leaf buds in manner similar to that of crossbills feeding on seeds of conifers: inserts bill between scales and gapes, laterally abducting lower mandible to part scales; extends brushy-tipped tongue to entangle prey. Does not twist entire head contra Richards and Bock (1973). See also Ch. 6.

Social and interspecific behaviour Usually in pairs or family groups, formerly conspecific flocks of up to 15 individuals (Bryan and Seale 1901; Perkins 1903). Joins loose mixed-species flocks with ‘Akikiki, Kaua‘i ‘Amakihi, ‘Anianiau (Lepson and Pratt 1997) and formerly other now

extinct spp. (Henshaw 1902; Perkins 1903; Munro 1960).

Breeding biology COURTSHIP AND MATING: Courtship display similar to that of other hemignathine honeycreepers, with male hopping back and forth on branch in front of female while uttering monosyllabic calls (Eddinger 1972a). Copulation may occur without full display. NEST SITE AND TERRITORY: In terminal crowns of ‘ohi‘a trees 9–12 m above ground (Eddinger 1972a; Pratt 1989b). Defends only small area around nest. NEST: Not well known, data from only five nests. Open cup woven around or wedged between small branches (Lepson and Pratt 1997). Constructed by both sexes equally (Eddinger 1972a) of mosses and Usnea lichens. Lined with grass or soft strips of bark (Eddinger 1972a; Berger 1981). One nest measured 8.2 ⫻11.3 cm outside (11.3 cm high), with inner cup 4.4 ⫻ 6.3 cm, depth 4.4 cm (Eddinger 1972a). All nests observed in spring, mostly in April (Lepson and Pratt 1997). EGGS: Only two reported (Eddinger 1972a), from single clutch, measured 16.6 ⫻ 13.3 mm and 16.6 ⫻ 13.1 mm.White with brown splotches, concentrated at larger end. Apparently laid on successive days. Nest failed after second egg laid. No nests have been monitored through incubation period to hatching or fledging.

Genus Magumma Mathews Rothschildia Wilson and Evans, 1890–99, p. xxi. Magumma Mathews, 1925, Bull. British Ornithologists’ Club 45: 93. New name for Rothschildia Wilson and Evans, nomen nudum. Type, by monotypy, Himatione parva Stejneger. The smallest Hawaiian honeycreeper, a warblerlike leaf-gleaner and nectarivore with short, slightly

decurved bill dark above, pinkish-grey below. Plumage yellow to yellow-green, without dark lores and with little dorso-ventral contrast. Sexual dimorphism and age variation slight. Sings trills with complex internal elements; call notes simple. Monotypic, endemic to Kaua‘i.

232 ‘Anianiau Magumma parva

‘Anianiau Magumma parva Mathews PLATES 4, 8 Himatione parva Stejneger, 1887, Proc. U. S. Natl. Mus. 10: 94. (Kauai.) Oreomyza parva Rothschild (1893–1900) Chlorodrepanis parva Henshaw (1902), Perkins (1903), Bryan and Greenway (1944), Munro (1944) Loxops parva Amadon (1950), Berger (1972a) Viridonia parva Greenway (1968) Hemignathus parvus Pratt (1979a), Berger (1981), AOU (1983, 1998) Loxops parvus Olson and James (1982b), James and Olson (1991) Other vernacular names: Lesser ‘Amakihi, Anauani‘i, Alawi (young birds)

Etymology Meaning of genus name not stated by its author (Mathews 1925), and it appears not to have classical connections; possibly a patronym. Specific epithet from Latin for ‘small’. Meaning of Hawaiian name obscure; Perkins (1903) thought it a reference to the nearly straight bill, but modern sources do not support that idea (Pukui and Elbert 1971; Lepson 1997).

Systematics First placed in Himatione (Wilson and Evans 1890–99), but in same publication Perkins said to favour separate monotypic genus Rothschildia, a name never validly published (and preoccupied). Mathews (1925) offered Magumma as replacement. Although long associated with the ‘amakihis in various classifications, recent genetic and biochemical studies ( Johnson et al. 1989; Tarr and Fleischer 1995; Fleischer et al. 1998, 2001) as well as more critical plumage comparisons (Conant et al. 1998) suggest Perkins on right track. For now, monotypic genus warranted (see Ch. 5 for details).

darker on back and rump; tail and flight feathers dark brownish grey with narrow edgings of body colour; no dark feathering in lores. No distinctive First Basic plumage. Juvenal plumage dull olive green, tinged yellow on throat and breast. Bill mostly dull brownish-pink, palest below, shading to horn colour along culmen. Legs and feet pale horn colour. Iris dark brown, appearing black in field. For standard measurements, see Lepson (1997).

Voice Call notes varied, the most distinctive a two-syllable tew-weet (Pratt et al. 1987) or three-syllable see-u-weet (T. Snetsinger in Lepson 1997) with terminal note up-slurred. Other calls, possibly flock contact notes (see Ch. 7), resemble those of several other Kaua‘i honeycreepers and not always distinguishable (Lepson 1997). Song is a sweet, lively, high-pitched trill with the individual elements usually doubled or tripled (Fig. 11.10): weesee-weesee-weesee etc. or weesityweesity-weesity-etc); rarely monosyllabic and slower. Drops very slightly and steadily in pitch, but does not have a sudden pitch shift (contra Lepson 1997).Whisper song similar to those of other hemignathine honeycreepers. For recordings, see Pratt (1996a); for additional sonograms see Lepson (1997).

Field identification Kaua‘i’s other surviving ‘little green birds’ (Kaua‘i ‘Amakihi, ‘Akeke‘e, Japanese White-eye) all show greater dorso-ventral contrast than ‘Anianiau, and have distinctive bill shapes. Kaua‘i Nukupu‘u (q.v.) similarly brightly coloured, but much larger, with black lores and bill. ‘Anianiau calls problematic; Lepson (1997) considers them not as ‘sweet’ and more forceful than those of ‘Akeke‘e, but I would reverse the comparison.

Description

Distribution, habitat, and population status

The smallest (10 cm) Hawaiian honeycreeper. Bill thin, short, very slightly decurved. Nearly uniformly bright yellow with slight mustard tinge (male) or greenish-yellow (female), very slightly

Endemic to Kaua‘i, where probably the most widespread of surviving honeycreepers despite inhabiting only 15% of original range (Scott et al. 1986; Lepson 1997). Inhabits forests above 600 m, occasionally

‘Anianiau Magumma parva 233 kHz 05120

KAUA'I

8 4

05051 8 4

05057 8 4

05057 Map 10

8 4 1

2

3

sec

11.10 Four representative primary songs of ‘Anianiau; numbers from LNS catalogue.

lower in sheltered valleys (USFWS 1983; Scott et al. 1986). Formerly common in forests at all elevations, but rare in lowlands by early 1900s (Bryan and Seale 1901). Still common above 1100 m in Koke‘e and ‘Alaka‘i regions in temperate mixed forest and native rainforest dominated by ‘ohi‘a-lehua, koa, and Cheirodendron spp. (Conant et al. 1998; Pratt 2002b). Subfossil bones found in coastal sites (Olson and James 1982b; James and Olson 1991). Population currently stable (Lepson 1997).

Diet and foraging From Lepson (1997) and pers. obs. Forages in flowers and leaves of outer canopy and subcanopy shrubs, vines, and ferns for nectar and invertebrates (about equal proportions in diet). Takes nectar from short-flowered ‘ohi‘a, ‘ohelo, ‘alani, kanawao, koa, native mints (Stenogyne and Phyllostegia spp.), and such alien plants as blackberry and Methley plum. Steals nectar (see Ch. 8) from haha‘aiakamanu, ‘ohe naupaka, and banana poka. Gleans

invertebrates, most frequently from leaves and branches of ‘ohi‘a-ha and ‘olapa. Occasionally eats (or extracts juices from) soft fruits such as Cheirodendron spp. and kanawao.

Social and interspecific behaviour Joins loose foraging flocks with Kaua‘i ‘Amakihi, ‘Akeke‘e, ‘Akikiki, and formerly other spp. (Wilson and Evans 1890–99; Perkins 1903). Bryan and Seale (1901) report conspecific flocks of up to 12 individuals. Also joins feeding aggregations of nectarivores at prime feeding sites.

Predation Formerly preyed upon by Kaua‘i Stilt-Owl and possibly Wood Harrier ( James and Olson 1991), both now extinct. Possibly preyed upon by Short-eared Owl and introduced Barn Owl,as well as rats and feral cats, but direct observations lacking (Lepson 1997).

Courtship and mating From Eddinger (1970). Monogamous. Courtship display involves male extending wings parallel to the ground, hopping up and down in front of female, and uttering primary and whisper songs.

234 Genus Hemignathus Female chases male from tree repeatedly following display, then follows him. Copulation (one observation) accompanied by whisper song, with primary song given afterwards. Courtship feeding occurs before and during nest construction.

Breeding biology All information from Eddinger (1970) except as noted. NEST SITE AND TERRITORY: Nest built in dense terminal leaf cluster of ‘ohi‘a tree, 3.3–9.5 m high. Male defends only immediate vicinity of nest, with territory as small as 9 m diameter or including only nest tree. Male chases conspecifics, but tolerates other spp. unless nest approached closely. Female may also defend nest. NEST: Open cup, ca. 9 cm outside, 5 cm inside, 7.5 cm high, woven around vertical branches. Both sexes build; female does more of construction but is fed by male. Materials include mosses and liverworts (for spp., see Lepson 1997), Usnea lichens, leaves, twigs, and bark of ‘ohi‘a, and pukiawe, fibres of grasses and sedges, and rootlets. May steal nest material from other honeycreeper nests. EGGS: Average size 18 ⫻ 22 mm. White, with variable amounts of tan and red-brown spots con-

centrated at larger end. Clutch size 2–4 eggs, median 3. INCUBATION AND HATCHING: Only female incubates. Female solicits feeding by male away from nest by fluttering wings and giving fledgling-like calls. Incubation period ca. 14 d. Eggs hatch in morning, usually entire clutch by noon. NESTLINGS: Helpless at first with eyes closed. Skin pink, exposed except for dark grey down on dorsal tracts. Gape pink with ‘cream-yellow’ rictal flanges. Eyes open at 3 d, contour feathering present by 10 d, flight feathers begin to unsheathe by 13 d. PARENTAL CARE AND FLEDGING: Female broods continuously early in nestling period (less frequently thereafter) as well as at night and during heavy rain. Both sexes feed nestlings by regurgitation, female more frequently (ca. 60%). Faecal sacs removed throughout nestling period. Young fledge at ca. 18 d, may jump from nest if disturbed after 11 d .

Life cycle and demography Eddinger (1970) found that 72.5% of eggs laid hatched and 60% produced fledglings. One brood per season. Life span of wild birds unknown, but captives live up to 9.5 years (Berger 1981).

Genus Hemignathus Lichtenstein Hemignathus Lichtenstein, 1839, Abh. Phys. Kl. Akad. Wiss. Berlin (1838), p. 449. Type, by subsequent designation (G. R. Gray, 1841), Hemignathus lucidus Lichtenstein. Small to very large Hawaiian honeycreepers with more or less down-curved sharp-pointed bills. Length varies widely, but culmen always arched and mandible, with one exception, always concave in profile. All spp. have what Pratt (2001b: 82) termed ‘ ‘amakihi coloration’ characterised by: (1) adult dorsal plumage olive green; (2) adult underparts yellow to olive green, paler than dorsum; (3) lores narrowly dark grey or black; (4) bill

dark brownish-grey to black, usually with pale (often bluish) base to mandible; (5) adult females like males but not as bright; and (6) immatures ( Juvenal and First Basic plumages) greenish-grey above, off-white below with at least faint wingbars (see also Ch. 6). Coloration so uniform across spp., sex, and age lines that bill must be seen to confirm field ID in most cases (Conant et al. 1998; Pratt and Pyle 2000).Tongue of drepanidine tubular type. Diet ranges from generalist to nearly totally insectivorous. Primary songs short trills or warbles, rather stereotyped, whisper songs more complex. Four subgenera recognised, sometimes

Kaua‘i ‘Amakihi Hemignathus (Chlorodrepanis) kauaiensis 235 elevated to generic rank, based primarily on bill morphology: Chlorodrepanis for the ‘amakihi group with short, curved bills; Viridonia for the straightbilled Greater ‘Amakihi; Akialoa for the very long-

billed ‘akialoas; and Hemignathus for the heterobills, whose lower mandible is half the length of the upper. See Ch. 5 for further discussion of taxonomic history.

Subgenus Chlorodrepanis Perkins Chlorodrepanis Perkins, 1899, in Wilson and Evans 1890–99, p. xxi. Type, by subsequent designation (Richmond, 1902, Proc. U. S. Natl. Mus. 24: 673), Himatione stejnegeri Wilson.

Bill short, decurved, mandibles of about equal length. Omnivorous. Songs simple trills.Three spp., one each on Kaua‘i and O‘ahu and one on the remaining large islands, with subspp. on Maui-nui and Hawai‘i.

Kaua‘i ‘Amakihi Hemignathus (Chlorodrepanis) kauaiensis Pratt PLATES 3, 8 Himatione stejnegeri Wilson, 1890, Proc. Zool. Soc. London, 1889, p. 446. Hemignathus kauaiensis Pratt, 1989, ‘Elepaio 49: 14, New name for Himatione stejnegeri Wilson, preoccupied. Chlorodrepanis stejnegeri Perkins (1903) Chlorodrepanis virens stejnegeri Bryan and Greenway (1944), Munro (1944) Loxops virens stejnegeri Amadon (1950), Berger (1972a), Olson and James (1982b) Viridonia virens stejnegeri (Greenway 1968) Viridonia stejnegeri Sibley and Monroe (1990) Hemignathus stejnegeri Pratt (1979a), Pratt et al. (1987) Hemignathus virens stejnegeri Berger (1981), AOU (1983) Other vernacular names:‘A‘alawi, Kihikihi (Munro 1960), ‘Alawi, Kihi, ‘Amakihi ‘awa‘awa (Pukui and Elbert 1971)

Etymology For genus, see Kaua‘i Nukupu‘u. Chlorodrepanis from Greek khloros ⫽ green and drepanon ⫽ sickle, presumably meaning ‘green sickle[bill]’. Epithet

means ‘of Kaua‘i’. ‘Amakihi possibly from Hawaiian ‘ama ⫽ talkative or bright and kihi ⫽ corner or sharp tip, as in ends of crescent moon or corner of the eye, a allusion to bill shape (Pukui and Elbert 1971).

Systematics See Ch. 4 and 5 for systematics of the genus. All ‘amakihis in this subgenus first considered conspecific by Bryan and Greenway (1944), although many writers, including Peterson (1961) and Bock (1970), remarked about the distinctiveness of the Kaua‘i form. On the basis of numerous biological isolating mechanisms, summarised by Conant et al. (1998) and Pratt and Pratt (2001), Pratt (1979a) and Pratt et al. (1987) returned it to sp. rank, as did Sibley and Monroe (1990), and James and Olson (1991), but not Berger (1981) or AOU (1983).After Johnson et al. (1989) and Tarr and Fleischer (1993) found genetic differences equivalent to sp. level in other honeycreepers, AOU (1995) concurred. See Ch. 5 for further discussion.

Description Small (11 cm) Hawaiian honeycreeper with relatively short tail and medium-length, (17–21 mm) sickle-shaped bill. Bill longer and deeper than those of other ‘amakihi spp. with no overlap in length

236 Kaua‘i ‘Amakihi Hemignathus (Chlorodrepanis) kauaiensis measurements, narrow overlap in depth (Conant et al. 1998). Sexually dimorphic in size of bill as well as coloration. Typical ‘ ‘amakihi coloration’ as described for genus. Definitive adults yellower than Hawai‘i ‘Amakihi, greener and less contrasting than O‘ahu ‘Amakihi. Undertail coverts dingy grey or greenish-grey. Moult sequence undescribed, but apparently lacks distinctive greyish immature stage with prominent wing-bars. Bill colour browner than in other ‘amakihis, darkest at tip and palest at base of lower mandible, looking fairly pale in the field. For measurements, see Lindsey et al. (1998).

(a) kHz 05118 8 4 (b) 8 4 (c) 05109 8

Voice Calls varied, the most common a loud tseet (pers obs.; Munro 1960; Eddinger 1970), often indistinguishable from calls of other Kaua‘i honeycreepers (Conant et al. 1998) and thus long overlooked. Also sometimes utters cat-like mewing note virtually identical to calls of O‘ahu and Hawai‘i ‘Amakihi. Whisper song long, complex, including possible mimicry (see Ch. 7). Primary song a loud, lively, somewhat variable trill that typically seems to drop in pitch, with distinctive introductory note different from the rest of the trill. Individual elements simpler and duration shorter than trills of other ‘amakihis (Fig. 11.11; Conant et al. 1998). For additional sonograms, see Lindsey et al. (1998), for recordings see Pratt (1996a).

Field identification Easily distinguished by bill shape from most of Kaua‘i’s ‘little green birds’ except Kaua‘i Nukupu‘u. Pratt and Pyle (2000) demonstrated that most recent reports of the latter were actually this sp. on basis of plumage colour. Nukupu‘u not only much yellower, but has bright white undertail and belly and intensely black bill. Confusion results from observers’ lack of appreciation for size of Kaua‘i ‘Amakihi bill as compared to others more often illustrated and observed. Song can be virtually identical to that of ‘Akikiki.

05035

4 (d) 05028 8 4 1 (e)

2

05265 8 4

11.11 Examples of primary songs of Kaua‘i ‘Amakihi (a–d) compared with a typical song (e) of Hawai‘i ‘Amakihi. Note shorter duration and distinctive introductory note of the former, the ‘fading in’ quality of the latter.

by 1970s restricted to areas above 600 m in Koke‘e/Alaka‘i region and Makaleha Mts. (Scott et al. 1986). Range may have contracted upward since then (pers. obs.) but no data available; still seen at lower elevations than other Kaua‘i native birds (Pratt 2002b). Common where it occurs, although least numerous of frequently seen honeycreepers in depths of Alaka‘i, apparently because it prefers forests with large component of koa (Conant et al 1998).

Distribution, habitat, and population status

Diet and foraging

Endemic to Kaua‘i and originally distributed throughout island’s forests (Munro 1960). Richardson and Bowles (1964) found it down to 450 m but

Forages mainly in understorey and on trunks and limbs, picking at bark for insects (pers. obs.;Wilson and Evans 1890–99) in manner similar to that of

Kaua‘i ‘Amakihi Hemignathus (Chlorodrepanis) kauaiensis 237

KAUAI

NEST SITE AND TERRITORY: In non-blooming ‘ohia, av. 5.6 m from ground, in terminal crown, horizontal branch clusters, or cluster of branches near trunk. Like other dreps (but unlike Hawai‘i ‘Amakihi), defends small territory (radius 5.5 m in one example) around nest. NEST: Self-supporting open cup built by both sexes, female doing most construction. Body (n ⫽ 2) of twigs and aerial roots of ‘ohi‘a, stems of ‘uluhe fern, mosses, bark, leaves, and lichens. Cup lined with shredded grass and bark. Mean dimensions: outside diameter 11.4 cm, outside vertical thickness 6.3 cm, cup diameter 6.3 cm, depth of cup 3.8 cm.

Map 11

‘Akikiki (Conant et al. 1998).Also takes nectar from ‘ohi‘a, kanawao, ‘ohe naupaka, and Kaua‘i koli‘i (pers. obs.) as well as alien blackberry, banana poka, and Methley plum (Berger 1981). Usually inserts bill into curved corollas of ‘ohe naupaka (Conant et al. 1998) and koli‘i (pers. obs.), serving as pollinator, but also steals nectar (see Ch. 8) especially from banana poka (pers. obs.; Conant et al. 1998). Perkins (1903) reports feeding on ‘akia fruit.

Social and interspecific behaviour Component of loose mixed-species foraging flocks.

EGGS: White with brown splotches all over, concentrated at larger end. Mean dimensions: 24.1 ⫻ 18.8 mm (n ⫽ 16). Clutch 1–4, mean 3 (n ⫽ 20). Laid 2–3 d following nest completion, one per day until clutch complete. INCUBATION AND HATCHING: Incubation by female only for 14 d. All eggs hatch morning of same day. NESTLINGS: Helpless, with eyes closed at hatching, skin pinkish-yellow with grey down on dorsal tracts, gape bright pink with cream-yellow rictal flanges. Eyes open at 4 d, feathers begin to unsheathe by 7 d. PARENTAL CARE AND FLEDGING: Female broods at night and during storms. Both parents feed (female 73%) and remove faecal sacs. Fledging at 17–20 d, mean 18.8 (n ⫽ 20).

Breeding biology All information from Eddinger (1970).

Life cycle and demography

COURTSHIP AND MATING: Displays similar to those of other hemignathine dreps. In Flitting Display male sings and rapidly flits up and down in branches below female. Courtship chasing frequent early in breeding cycle (Mar–Apr). Courtship feeding occurs before and during nest construction and especially during incubation.

Breeding season Mar–Jul (Eddinger 1970), with pair formation in Feb (Lindsey et al. 1998). Mean hatching success 88.8% (n ⫽ 63), fledging success 91.1% (n ⫽ 56), reproductive success 81% (Eddinger 1970). Life span up to 9 years in captivity (Berger 1980). Population estimated 15 000–20 000 in late 1980s (Ellis et al. 1992a), higher than estimate from 1970s by Scott et al. (1986).

238 O‘ahu ‘Amakihi Hemignathus (Chlorodrepanis) flavus

O‘ahu ‘Amakihi Hemignathus (Chlorodrepanis) flavus (Bloxam) PLATE 3 Nectarinia flava Bloxam, 1827, in Byron, Voyage of H. M. S. Blonde, App. 3, p. 249. (Oahu.) Himatione chloris Cabanis, 1850, Mus. Heineanum, I, p. 99. (Oahu.) Chlorodrepanis chloris Bryan (1901a) Chlorodrepanis virens chloris Perkins (1903), Munro (1944) Loxops virens chloris Amadon (1950), Berger (1972a), James and Olson (1991) Viridonia virens chloris Greenway (1968), Sibley and Monroe (1990) Hemignathus virens chloris Pratt (1979a), Berger (1981), AOU (1983) Other vernacular names: Common ‘Amakihi

Etymology See Kaua‘i Nukupu‘u and Kaua‘i ‘Amakihi for genus and subgenus. Latin epithet flavus ⫽ yellow; Greek root khloros ⫽ green.

Systematics See Kaua‘i Nukupu‘u and Kaua‘i ‘Amakihi. Originally described as distinct sp., but no one since Bryan and Greenway (1944) had suggested that this form was anything other than well-marked subspp. when mtDNA studies by Tarr and Fleischer (1993) indicated not only that O‘ahu bird equally distinct, but sister taxon to Kaua‘i ‘Amakihi, on which basis AOU (1995) gave it sp. rank. However, later studies by Fleischer et al. (1998) placed it as sister taxon of Hawai‘i ‘Amakihi, as might be predicted on morphological grounds, but still sufficiently distinct genetically to be considered full sp. In retrospect, distinctive plumage of H. flavus should not have been denigrated as Amadon (1950) and others did. Pratt and Pratt (2001) consider plumage colour one of better phenotypic clues to relationships among Hawaiian honeycreepers.

Description Small (11 cm) Hawaiian honeycreeper with short decurved bill. Sexual dimorphism more pronounced than in other ‘amakihis. See description of ‘‘amakihi coloration’ under genus. Definitive male yellower throughout than others, with much stronger, more sharply defined dorso-ventral contrast; underparts golden yellow from chin to undertail; cheeks yellow, but superciliary reduced to tiny fleck of yellow above narrow black lores. Female not as bright above, straw yellow below tinged olive on throat and abdomen, with two prominent cream-coloured wing-bars. Juvenal plumage greyish-green above, yellowishwhite below, with off-white wing-bars. Moult sequence not well studied; birds intermediate between above plumages possibly in First Basic plumage (Lindsey et al. 1998). Bill not as bluish as that of Hawai‘i ‘Amakihi, more like that of Kaua‘i bird in colour. For measurements, see Lindsey et al. (1998).

Voice Call a low-pitched, buzzy, mewing cheee virtually identical to homologous call of Hawai‘i ‘Amakihi. Rothschild’s (1893–1900) report of call as sweet, and Bryan‘s (1905b) as tswe-et may refer to same call. Primary song a loud trill much like those of Hawai‘i ‘Amakihi but somewhat slower with notes more distinct (E. A. VanderWerf in Lindsey et al. 1998). Whisper song not reported. For sonogram, see Lindsey et al. (1998), for recording see Pratt (1996a; presented under Common ‘Amakihi).

Field identification See O‘ahu ‘Alauahio. Curved bill diagnostic for ‘amakihi, but also note lack of pale supercilium.

Distribution, habitat, and population status Endemic to O‘ahu, where originally in forests at all elevations. Retreated upwards with clearing of lowland forests, later restricted to higher elevations by

O‘ahu ‘Amakihi Hemignathus (Chlorodrepanis) flavus 239

O'AHU

and drinking at sap flows (pers. obs.; Lindsey et al. 1998). Prefers outer foliage of canopy trees, but also forages in understorey shrubs (Bryan 1905a; VanderWerf and Rohrer 1996; pers. obs.). Readily feeds in non-native trees such as kuku‘i (Perkins 1903), golden shower tree and horticultural Hibiscus (Howarth 1984), various eucalyptus, Formosan koa, paperbark, schefflera, and wiliwili haole (pers. obs.; Shallenberger and Vaughan 1978; Pratt 2002b).

Social and interspecific behaviour

Map 12

Move about in small, apparently well-organised flocks of 4–6 (Bryan 1905a). May gather in large aggregations of up to 30 at rich nectar sources (Shallenberger and Vaughan 1978; Lindsey et al. 1998). Chases Japanese White-eye away from flowers (E. A.VanderWerf in Lindsey et al. 1998).

Breeding biology mosquito-borne diseases (Lindsey et al. 1998). Range now divided into two widely separated segments in Wai‘anae and Ko‘olau Mts. ‘Uncommon and sparsely distributed’ above 500 m in Wai‘anae Range, but found down to lower forest edge in Ko‘olaus (Lindsey et al. 1998). Surprisingly, more common and at lower elevations at se. end of range above heavily populated areas (Shallenberger and Vaughan 1978; VanderWerf 1993; Lindsey et al. 1998). Declined slowly 1958–85 (Williams 1987), but then began apparent recovery (Conry 1991; Lindsey et al. 1998) and now common in Honolulu suburbs to 70 m in Moanalua and 50 m in Makiki and Manoa valleys, and as low as 30 m in windward Waiahole and Waianu valleys (VanderWerf 1997; Lindsey et al. 1998; Pratt 2002). Repopulation of lower elevations apparently result of developing resistance to avian malaria (Cann et al. 1996). Now occurs frequently in exotic as well as native forests and suburban plantings (Howarth 1984; Pratt 2002b).

Based on only three active nests. Courtship and mating unknown. NEST SITE AND TERRITORY: Aggressive encounters between males suggest some form of territoriality (Lindsey et al. 1998). Nest located in dense terminal clumps mostly 7–10 m high, although Bryan (1905a) reported one (not confirmed) at the base of a tree fern ca. 30 cm from the ground. Trees used include native ‘ohi‘a (Bryan 1905a; Russell and Ralph 1981) and koa (Eddinger 1984), and alien lychee (VanderWerf 1997).

Diet and foraging

NEST: Built by female but male may bring material (Russell and Ralph 1981). Nests (summarised by Lindsey et al. 1998) similar to those of better known Hawai‘i ‘Amakihi, with outer shell of grasses, including native pili, and/or twigs with inner cup lined with fine fibrous material such as tree fern rootlets and pulu (see Maui ‘Alauahio account). Bryan’s (1905a) low nest made entirely of mosses and pulu. Outside diameter ca. 7.5–15 cm, inside 4.5–6, cup depth 2.7–3.1.

A feeding generalist that eats nectar, fruit, and arthropods (Perkins 1903). Foraging techniques include probing flowers, leaf-gleaning, bark-picking,

INCUBATION AND HATCHING: Only female incubates, fed off nest by male (Eddinger.

240 Hawai‘i ‘Amakihi Hemignathus (Chlorodrepanis) virens

Hawai‘i ‘Amakihi Hemignathus (Chlorodrepanis) virens (Gmelin) PLATES 3, 8 Certhia virens Gmelin, 1788, Syst. Nat. 1(1): 479. Based on the ‘Olive-green Creeper’ Latham, Gen. Synop. Birds 1(2), p. 740. (in insulis Sandwich ⫽ Hawaii, restricted to Kona district, Island of Hawaii by Medway, 1981, Pacific Science 35: 105–175.) Himatione virens Wilson and Evans (1890–99) Chlorodrepanis virens Bryan (1901a), Perkins (1903), Bryan and Greenway (1944), Munro (1944) Loxops virens Amadon (1950), Berger (1972a), Olson and James (1982b), James and Olson (1991) Viridonia virens Greenway (1968), Sibley and Monroe (1990) Other vernacular names: Common ‘Amakihi, ‘Amakihi, Maui ‘Amakihi, Lana‘i ‘Amakihi, Moloka‘i ‘Amakihi

Etymology See Kaua‘i Nukupu‘u and Kaua‘i ‘Amakihi. Modifier ‘Common’, which I suggested for this sp. plus the O‘ahu ‘Amakihi (Pratt 1979a), used by the 1983 AOU Check-list, first to include Hawaiian birds, as counterpart to Greater ‘Amakihi without splitting Kaua‘i form. Nevertheless appropriate because spp. numerically common as well as common to more than one island. It remained appropriate as the Kaua‘i and O‘ahu spp. were split, but the AOU (1998) in its wisdom chose to restrict use of ‘Common ‘Amakihi’ to the 1983 construct (H. virens ⫹ flavus ⫹ kauaiensis) and applied the single-island modifier even though workers on Maui and its satellite islands find it an extremely confusing designation. Thus we are burdened with such ridiculous circumlocutions as ‘Hawai‘i ‘Amakihi from Maui’ or ‘Hawai‘i Hawai‘i ‘Amakihi’ and discussion is rendered very difficult.

Systematics See Kaua‘i ‘Amakihi and O‘ahu ‘Amakihi for spp. limits in this complex. H. virens has two subspp., H. v. virens on Hawai‘i and H. v. wilsoni on

Maui-nui islands. The Moloka‘i and Lana‘i populations were also named (kalaana and chloridoides, respectively), but Amadon (1950) correctly synonymised them under wilsoni. In my own study, I found only one statistically significant difference (slightly longer bill in Maui birds based on only eight specimens) among them, not enough for taxonomic separation (Pratt 1979a). However, males from Hawai‘i (described below) have a number of statistically significant, if overlapping, difference that set them apart from Maui-nui ‘amakihi. C. van Riper (pers. comm.) and others have suggested that populations of H. v. virens from dry habitats, such as at Pu‘u La‘au, are brighter and yellower than those from wet forests. My comparisons of a large sample of freshly collected and older specimens from all over the island (Pratt 1979a) revealed no intra-island variation such as seen in Hawai‘i ‘Elepaio (Pratt 1981). ‘Amakihi at Pu‘u La‘au do, indeed, look very yellow in this open and sunny habitat, but in the same light are no brighter than rainforest birds.

Description Small (11 cm) Hawaiian honeycreeper with short curved bill.The prototype for ‘ ‘amakihi coloration’ detailed in genus introduction.The two subspp. differ on average only, Maui-nui males being slightly paler below with narrower dark lores, females greyer and less yellow, especially in throat, but only extreme individuals could be correctly assigned to island on coloration alone, and no differences are apparent in the field. Sexually dimorphic in bill length (female shorter). Juvenal plumage greenishgrey above, pale grey below, variably streaked with pale yellow; wing-bars pale grey to white. First Basic plumage similar, but wing-bars reduced by wear and irregular replacement with new feathers; may be nearly absent (Lindsey et al. 1998). Some Juvenal or First Basic individuals very grey, with almost no yellow pigmentation (hence no green). For measurements, see Lindsey et al. (1998).

Hawai‘i ‘Amakihi Hemignathus (Chlorodrepanis) virens 241

Voice Calls almost infinitely varied, but most common a cat-like mew similar to calls of Blue-gray Gnatcatcher of North America. Others include squeaks, short chips and chirps, an inquisitive, up-slurred quee-e, and on Mauna Kea a multisyllabic call that may be imitation of Palila, distinguished, if at all, by rising rather than descending pitch at end (Pratt 2002b). Foraging birds outside breeding season utter highly varied repertoire of calls, function unknown. Whisper song long and complex, typical for hemignathine honeycreepers. Primary song variable but always a loud ringing trill, the elements of which, though more complex (Fig. 7.2, Fig. 11.11) in sonograms, almost always sound like a single note.Tends to fade in as if volume were being turned up gradually rather than start abruptly as in Kaua‘i ‘Amakihi.

Field identification Because of wide variation in plumage colour, Hawai‘i ‘Amakihi easily transformed into rarer sp. by wishful thinking, but bill shape diagnostic. Throat never clear white as in Hawai‘i Creeper. Vocalisations can be annoying for those searching for Palila on Mauna Kea (Pratt 2002b).

Distribution, habitat, and population status Originally found in virtually all forest and scrub types on Hawai‘i and Maui-nui, and remains most widespread (but not necessarily most abundant) honeycreeper on Maui and Hawai‘i. On Hawai‘i, favours drier habitats and reaches greatest density in mamane–naio forest on Mauna Kea, and in some dry forest still occurs as low as 100 m (Scott et al. 1986). Common in lowland dry forest remnants in N. Kona Dist. (Pratt 2002b). On Maui, greater densities occur in mesic to wet forests, probably because of damage to dry forests by feral ungulates, but is one of only two honeycreepers still present in dry forest remnants on s. slope of Haleakala (Scott et al. 1986). Common in subalpine scrub on n. slope of Haleakala, and also present in conifer plantations (Fig. 11.12) such as Hosmer Grove and Polipoli Springs (Pratt 2002b). Recently observed near sea

MOLOKA'I MAUI _ LANA'I KAHOOLAWE

HAWAI'I

Map 13

level along windward Hana Hwy. (Pyle and Donaldson 2000b). Still present in moderate numbers above 1100 m on W. Maui, occasionally wandering into ‘Iao Valley at ca. 300 m (pers. obs.) On Moloka‘i, Bryan (1908) considered it second most abundant bird, but scarce by 1949 (Richardson 1949) and rare by 1975 (Scott et al. 1977) in upland forest. HFBS found it locally common in backs of lowland valleys in mesic to dry native forest (Scott et al. 1986). On Lana‘i, common before 1920 (Munro 1960), rare by 1960s, and last seen 1976 (Hirai 1978); presumed extirpated. Recent sightings in lowlands of both Maui and Hawai‘i indicate possible development of resistance to avian malaria as documented for O‘ahu ‘Amakihi.

Diet and foraging Broad generalist feeder, with ca. one-third of feeding in flowers for nectar, remainder gleaning or pecking bark or leaves for insects (Ralph 1990a). Also takes small proportion of fruit (Baldwin 1953) and tree sap (Lindsey et al. 1998; J. K. Lepson, pers. comm.). For arthropods, prefers small, soft-bodied flightless kinds such as caterpillars (Baldwin 1953). Favoured nectar sources ‘ohi‘a and mamane, but feeds in virtually any native or introduced flower (for detailed list, see Lindsey et al. 1998). Seems particularly fond of wiliwili and silk oak in N. Kona

242 Hawai‘i ‘Amakihi Hemignathus (Chlorodrepanis) virens rising arc and fluttering at apex; Low Courtship Chase faster, in straight line nearer the ground; Advertising Display, high or low concentric circles with hovering or fluttering; and Flitting Display, in which male flies toward female, hops around her, or flies back and forth below her with or without singing whisper song.

11.12 Eclectic habitat and foraging tastes are demonstrated by this Hawai‘i ‘Amakihi in exotic slash pine, Hosmer Grove, Maui. Maui ‘Alauahio may also use pine and other alien plantations for foraging and nesting.

dry forests (pers. obs.). Steals nectar from flowers with incompatible configurations, such as garden nasturtium (Henshaw 1902). Park-tame birds in Haleakala NP even beg for handouts of human ‘junk food’ (Pratt 1999b, 2002b). See more detailed discussion of omnivory in Ch. 8.

Social and interspecific behaviour During non-breeding season, forages in small conspecific groups or joins mixed-species flocks (pers. obs.; Lindsey et al. 1998). Home ranges large and overlapping, but include nesting territory (van Riper 1987).

Predation Remains found in stomachs of feral cats and mongooses (van Riper 1978b; Snetsinger et al. 1994; Lindsey et al. 1998), and predation of nestlings by roof rats documented by van Riper (1978b). Documented avian predators include native Shorteared Owl and introduced Barn Owl (Snetsinger et al. 1994).

Breeding biology COURTSHIP AND MATING: From van Riper (1987). Monogamous, pairs remaining together multiple seasons. Named displays include: High Courtship Chase with male pursuing female in slow, gently

NEST SITE AND TERRITORY: Both parents search for nest site, female testing sites with grass blade (van Riper 1987). Placed in terminal or lateral forks or clusters in tree spp. dominant in the area: mamane (88%) and naio (12%) in mamane–naio forest on Mauna Kea (van Riper 1987); ‘ohi‘a in mesic to wet forest (Kern and van Riper 1984; Lindsey et al. 1998), and introduced conifers in plantations on Maui (Lindsey et al. 1998). Unlike most Hawaiian honeycreepers, defends large mutually exclusive territories (Baldwin 1953; van Riper 1987). NEST: Female builds open cup nest; opportunistic in choice of materials (Kern and van Riper 1984). In mamane–naio forest, outer shell of mamane leaves and twigs and grass stems, lining of Usnea lichen, rootlets, plant fibres, and even pig hair and sheep wool (van Riper 1977; Kern and van Riper 1984). In ‘ohi‘a forest, frame of twigs,‘ohi‘a rootlets, and hapu‘u tree fern leaves and rootlets, lining of plant fibres, rootlets, animal hair, and some Usnea (Kern and van Riper 1984). In Maui conifers, frame of grass, pine needles, moss, and twigs, lining of pine needles and grass (Lindsey et al. 1998). Mean dimensions (from van Riper 1987): length 10.9 cm, width 9.0 cm, cup diameter 5.2 cm, cup depth 3.5 cm, overall height 6.9 cm. EGGS: From van Riper (1987). Creamy white variably splotched with purple and brown, colour concentrated at larger end. Mean dimensions (n ⫽ 90): 19.0 ⫻ 13.9 mm. Clutch 1–4, mean 2.5 (n ⫽ 20). Laid one egg per day until clutch complete. INCUBATION AND HATCHING: From van Riper (1987). Only female incubates. Incubation period mean 14 d. First egg usually hatches at night, followed by remainder before noon next day.

Greater ‘Amakihi Hemignathus (Viridonia) sagittirostris 243 NESTLINGS: From van Riper (1978b) and Lindsey et al. (1998). Helpless, with eyes closed at hatching, skin pink, down grey, bill dull yellow, gape rose red. Eyes open at 5–7 d. Feathers begin to unsheathe at 8–11 d. Young will jump from nest prematurely if disturbed by 14 d. PARENTAL CARE AND FLEDGING: From van Riper (1978b) and Lindsey et al. (1998). Only female broods. Both parents feed by regurgitation, but female does most. Faecal sacs removed throughout cycle in most (but not all) nests on Hawai‘i, but Maui birds allow them to accumulate on nest rim after 14 d. In most nests on Hawai‘i, chicks defecate

over nest rim beginning at 10 d. Mean nestling period 16.8 d.

Life cycle and demography From van Riper (1987) and Lindsey et al. (1998). Breeding season highly attenuated, eggs or young recorded Nov–Aug, peak Mar–June. Second broods frequent on Mauna Kea, infrequent on Maui. May breed in first year. In Mauna Kea population, mean hatching success 53.5%, fledging success 64.9%, overall reproductive success 34.7%. Potential life span at least 12 years. From van Riper’s data, Freed (1988) calculated average survival rate of 85%.

Subgenus Viridonia Rothschild Viridonia Rothschild, 1892, Annals and Mag. Nat. Hist. (6)10: 112. Type by monotypy, Viridonia sagittirostris Rothschild. Bill longer than in previous subgenus, nearly straight, used for probing and gaping to extract insect prey. Retroarticular processes pronounced (Richards and Bock 1973; James and Olson 1991). Sexual dimorphism, age-related plumage differences, and dorso-ventral contrast slight compared

to other Hemignathus. Songs simple trills, resembling those of ‘amakihis (Perkins 1903). Sometimes combined with Chlorodrepanis, which then becomes junior synonym. Treatment herein tentative; may not belong within Hemignathus at all. James (1998) indicates that Viridonia could well absorb the newly named Aidemedia and become a full genus allied with ‘akepas. Monotypic, endemic to island of Hawai‘i.

Greater ‘Amakihi Hemignathus (Viridonia) sagittirostris (Rothschild) PLATES 4, 8 Viridonia sagittirostris Rothschild, 1892, Annals and Mag. Nat. Hist. (6)10: 112. (Mauna Kea, Hawai[i], Sandwich group.) Loxops sagittirostris Amadon (1950), Carlquist (1970), Berger (1972a) Other vernacular names: Green Solitaire

Etymology See Kaua‘i Nukupu‘u for genus. Viridonia from Latin viridis ⫽ green. Epithet from the Latin for

‘arrow-bill’. In fact, if this subgenus is shown not to be related to the ‘amakihis, Arrowbill would be a suitable and appropriate English name for this sp.

Systematics Long regarded as allied with ‘amakihis, but closer look (Pratt 2001b) suggests resemblances superficial and differences substantial. Possibly related to the subfossil genus Aidemedia ( James and Olson 1991; James 1998).

244 Subgenus Akialoa

Description Medium-sized (17 cm) Hawaiian honeycreeper with long, sharp-pointed, nearly straight bill somewhat like that of an American oriole (Henshaw 1902). No age or sex differences in plumage known. Plain olive green throughout, slightly darker on dorsum, yellower on throat and breast; no prominent markings other than narrow black lores. Bill and legs black, base of mandible pale blue. For measurements, see Lindsey et al. (1998).

Voice Call described by Henry Palmer (in Rothschild 1893–1900) as repeated chirrup, high and clear. Henshaw (1902) described a low plaintive note, which Perkins (1903) said resembled the whistle of the Hawai‘i Mamo. Song a trill similar to those of ‘amakihis with several distinctive additional notes (Perkins 1903).

Distribution, habitat, and population status Extinct. Endemic to Hawai‘i; known only from wet ‘ohi‘a forest a few km on either side of Wailuku R. near Kaumana, above Hilo, at 152–1220 m elevation (Rothschild 1893–1900; Henshaw 1902; Perkins 1903). Last collected apparently by A. M.Woolcott in 1901 near Kaiwiki Stream, north of Wailuku R. and not reported since (Berger 1981). Low-elevation distribution unique among historically known honeycreepers (Scott et al. 1986).

Diet and foraging From Perkins (1903). Mainly insectivorous but occasionally took nectar. Foraged by probing into bark crevices of ‘ohi‘a trees and leaf axils of ‘ie‘ie and gleaning from ferns for arthropods, especially crickets of the genus Paratrigonidium, and including carabid beetles, spiders, and caterpillars.

Social and interspecific behaviour Field identification Easily overlooked because of similarity of voice and appearance to Hawai‘i ‘Amakihi, but straight bill and calls distinctive.

Palmer (in Rothschild 1893–1900) and Perkins (1903) reported birds in pairs.Apparently had home ranges if not territories, and responded to imitations of calls as if defending territory (Perkins 1903).

Subgenus Akialoa Olson and James Akialoa Olson and James, 1995, Proc. Biol. Soc. Wash. 108: 384. Type, by original designation, Certhia obscura Gmelin. Bill unusually long with mandible nearly as long as maxilla. Maxilla has blunt probing tip, mandible sharp point. Possesses ‘deep medial groove on the ventral maxilla, . . . shorter retroarticular processes of the mandible, . . . abruptly constricted nasals and long,

attenuated premaxillae’ (Olson and James 1988: 13), shared with the heterobills (subgenus Hemignathus). Plantaris muscle absent. Number of spp. problematical (see Ch. 5) and complicated by incomplete prehistoric record. Several islands had more than one sp. Two spp. as yet undescribed from Hawai‘i ( James 2004).Taxonomy here recognises all named forms as spp., following Olson and James (1995).

Kaua‘i ‘Akialoa Hemignathus (Akialoa) stejnegeri 245

Kaua‘i ‘Akialoa Hemignathus (Akialoa) stejnegeri Wilson PLATE 3 Hemignathus stejnegeri Wilson, 1889. Ann. Mag. Nat. Hist. 6(4): 400. (Kauai.) Hemignathus procerus Cabanis (1890), Wilson and Evans (1890–99), Henshaw (1902), Amadon (1950), Carlquist (1970), Berger (1972a, 1981), AOU (1983) Hemignathus obscurus procerus Bryan and Greenway (1944), Munro (1944), Amadon (1950), Pratt (1979a), Olson and James (1982b), Pratt et al. (1987) Hemignathus obscurus ellisianus Sibley and Monroe (1990) Hemignathus ellisianus stejnegeri AOU (1998) Akialoa stejnegeri Olson and James (1995) Other vernacular names: Akialoa, Greater Akialoa (AOU 1998)

Etymology See Kaua‘i ‘Amakihi for genus. Subgenus name adopts Hawaiian name for the birds, which combines ‘akihi or ‘akia, apparently a generic term for green birds with curved bills, perhaps related to kihi (see Kaua‘i ‘Amakihi), with loa ⫽ long, a reference to the bill. Epithet honours Leonhard Stejneger, an ornithologist at Smithsonian Institution who described several Hawaiian birds from specimens sent from Kaua‘i by fellow Norwegian Valdemar Knudsen (see Ch. 3).

Description A very large (19 cm) honeycreeper, darker above, paler below, with extremely long down-curved bill. Definitive male yellow-green above, with bright yellow supercilium and underparts, flanks tinged olive, black lores, and auriculars yellow-green at upper edge shading to yellow below, and crown feathers with dark centres producing scaly or spotted effect. Definitive female patterned similarly but with less crown spotting and duller coloration. Immature plumage (may be Juvenal or First Basic) duller still, olive-grey above, off-white below with

tinge of yellow posteriorly, with two prominent offwhite wing-bars. Bill and feet dark plumbeous grey (Lepson and Johnston 2000), pale grey at base of lower mandible. Iris dark hazel (Wilson and Evans 1890–99; Bryan and Seale 1901). For standard measurements, see Lepson and Johnston (2000).

Voice Calls include a chirp similar to but louder than call of cardueline House Finch (Munro 1960) or canary (Rothschild 1893–1900),‘a different chirp, evidently a breeding season call’, and a loud keewit as described by Donaghho (Munro 1960: 116). Huber (1966) described a down-slurred whistle given by a bird perched in a tree with an ‘O‘u and stated that it was identical to that bird’s whistle but with opposite inflection. Conant et al. (1998) questioned this because ‘O‘u gives both up-slurred and downslurred whistles, but in light of Palmer’s description of hollow whistled call given by Lana’i ‘Akialoa (Rothschild 1893–1900), Huber may have been correct.A ‘light sweet song’ given by both sexes (Munro 1960: 116) probably a whisper song. Full song apparently unknown.

Field identification The large size and extremely long bill separate this bird from all others on Kaua‘i. Much brighter yellow-green colour than Kaua‘i ‘Amakihi, sex for sex. Wing-bars of immature unique among Kaua‘i honeycreepers.

Distribution, habitat, and population status Extinct. Originally widespread and common in wet forests of Kaua‘i, from 200 m to 1300 m (Rothschild 1893–1900). Mostly in ‘ohi‘a forest, but Munro (1960) reported them also in koa. In 1891, many noted with pox lesions and other signs of disease (Munro 1960), coinciding with start of decline. Rare by 1920s (Donaghho 1941) and unreported until ‘rediscovered’ in heart of Alaka‘i

246 O‘ahu ‘Akialoa Hemignathus (Akialoa) ellisianus Plateau in 1960 (Richardson and Bowles 1964). Subsequently seen by Huber (1966). Last sighting by P. L. Bruner (pers. comm.; Conant et al. 1998) in 1969. Approximately 120 specimens in world collections (Banko 1979; Lepson and Johnston 2000).

Diet and foraging Actively hopped on short legs along lichen and moss-covered trunks and branches of trees, probing for invertebrates with the long bill deep into cracks and crevices, or into leaf bases of ‘ie‘ie and halapepe (Munro 1960). Sometimes inserted bill up to its base, so that shot bird might hang from bill (Perkins 1903). P. L. Bruner (pers. comm.) described bird hanging head-down to reach underside of branch with bill. Bryan and Seale

(1901) compared the birds’ position on tree trunk with that of North American flickers. Also took nectar from haha‘aiakamanu or other flowering lobelioids. Munro (1960: 116) states ‘the long [nectar-adapted] tongue is useful in extracting honey from the deep tubes of the lobelia flowers and in working out grubs and insects from their hiding places.’

Social and interspecific behaviour Joined mixed-species foraging flocks with Kaua‘i ‘Amakihi and ‘Akikiki (Rothschild 1983–1900).

Breeding biology Little information, but Munro (1960) reported peak of breeding season as Mar–Apr.

O‘ahu ‘Akialoa Hemignathus (Akialoa) ellisianus (Gray) PLATE 3 Drepanis (Hemignathus) ellisiana G. R. Gray, 1860, Cat. Birds Trop. Islands Pacific, p. 9. (Oahu.) Hemignathus lichtensteinii Wilson and Evans (1890–99), Perkins (1903), James and Olson (1991) Hemignathus obscurus ellisianus Bryan and Greenway (1944), Munro (1944), Amadon (1950), Pratt (1979a), Olson and James (1982b), Pratt et al. (1987) Akialoa ellisiana Olson and James (1995) Other vernacular names: ‘Akialoa, Kipi (Bryan 1901a), Greater Akialoa (AOU 1998)

Etymology See Kaua‘i Nukupu‘u and Kaua‘i ‘Akialoa for genus and subgenus. Epithet ellisianus honours surgeon’s mate William Ellis of the Cook expedition, who first illustrated several Hawaiian birds.

Description A very large (19 cm) yellow and green honeycreeper with extremely long down-curved bill. For this and the following sp. (q.v.), depictions on

Plate 3 based on Rothschild (1893–1900). I cannot explain how he could depict two plumages when, apparently, only one specimen was at hand, except perhaps as an extrapolation based on incoming feathers of the single bird. Lepson and Johnston’s (2000) description of the presumably adult male in the collection of the Academy of Natural Science of Philadelphia does not match anything portrayed by Rothschild (1893–1900) or Wilson and Evans (1890–99), both of which must have been based on the holotype now in Berlin’s Museum f ür Naturkunde der Humboldt-Universität, believed to be an immature male (Lepson and Johnston 2000). Soft part colours not recorded in life. For standard measurements, see Lepson and Johnston (2000).

Distribution, habitat, and population status Known from two specimens taken in 1830s when uncommon. Only one of several specimens collected by Deppe (Lichtenstein 1839) remain (Lepson and Johnston 2000). J. K. Townsend collected another in Nu‘uanu Valley. Rare by 1840s, several

Maui-nui ‘Akialoa Hemignathus (Akialoa) lanaiensis 247 expeditions having failed to find any. Perkins (1903) saw a pair above Nu‘uanu Valley in 1892 and shot one, but was unable to retrieve specimen. Three reports in 1930s (Donaghho 1963) have

been treated with scepticism, but one by J. d’A. Northwood at Palehua in 1939 was fairly detailed and believable by modern standards (Banko 1984b).

Maui-nui ‘Akialoa Hemignathus (Akialoa) lanaiensis Rothschild PLATE 3 Hemignathus lanaiensis Rothschild, 1893, Bull. Brit. Orn. Club 1: 24. (Lanai.)

Other vernacular names: Lana‘i ‘Akialoa, Greater Akialoa (AOU 1998)

specimens, nor can I determine which illustration applies to which specimen. Rothschild thought the three specimens represented a male, a young male, and a female. Lepson and Johnston (2000) consider both males immature. Perkins (1903) observed a very bright yellow Maui-nui ‘Akialoa that he presumed to be a Definitive male, but no specimens of that plumage exist. Because of the uncertainties, I decline to offer a detailed description here but refer the reader to the colour plate. For standard measurements, see Lepson and Johnston (2000).

Etymology

Voice

See Kaua‘i Nukupu‘u for genus, Kaua‘i ‘Akialoa for subgenus. Specific epithet means ‘of Lana‘i’. English name proposed by Olson and James (1995) to more accurately reflect former range on several component islands of Maui-nui.

From Palmer in Rothschild (1893–1900: 90). Resembles voice of Lesser ‘Akialoa but much clearer; also ‘a whistle, but yet with something hollow in it, very difficult to express in words’ apparently resembling a whistle attributed to Kaua‘i ‘Akialoa by Huber (1966).

Hemignathus obscurus lanaiensis Bryan and Greenway (1944), Amadon (1950), Carlquist (1970), Pratt (1979a), Olson and James (1982b), Pratt et al. (1987) Hemignathus ellisianus lanaiensis AOU (1998) Akialoa lanaiensis Olson and James (1995)

Description A very large (19 cm) yellow and green honeycreeper with extremely long down-curved bill. Illustration on colour plate based secondarily on colour lithographs in a copy of Rothschild’s magnum opus in the McIlhenny Collection of the Louisiana State University Library.They agree with his original description (Rothschild 1893b), quoted in Wilson and Evans (1890–99), but not with recent description by Lepson and Johnston (2000). I cannot account for the discrepancies, which do not appear to be the result of aging of the century-old

Distribution, habitat, and population status Extinct. Three specimens collected Nov 1892 (Rothschild 1893–1900), one seen in 1894 (Perkins), and another probable sighting by Munro (1960) between 1911 and 1930.

Diet and foraging Little information, but Rothschild (1893–1900: 90) states ‘very quick in their movements from flower to flower’ so fed at least partly on nectar.

248 Lesser ‘Akialoa Hemignathus (Akialoa) obscurus

Lesser ‘Akialoa Hemignathus (Akialoa) obscurus (Gmelin) PLATES 3, 8 Certhia obscura Gmelin, 1788, Syst. Nat. 1(2): 470. Based on the ‘Hook-billed green Creeper’ Latham, Gen. Synop. Birds 1(2): 703, pl. 33, fig. 1. (in insulis Sandwich ⫽ Hawaii.)

and Evans 1890–99: 69) or ‘dark plumbeous grey’ ( J. K. Lepson, pers. comm.) with pale base to lower mandible. For standard measurements, see Lepson and Johnston (2000).

Akialoa obscura Olson and James (1995)

Voice

Other vernacular names: Hawai‘i ‘Akialoa,‘akihi-aloa, ‘akihi-loa, Akialoa, Hawaiian Akialoa (AOU 1983)

Call distinctive, apparently somewhat like that of ‘amakihis but louder and deeper (Perkins 1903). Perkins (1903: 424) described song as ‘a short vigorous trill, recalling that of both [‘Akiapola‘au] and [Hawai‘i ‘Amakihi] but distinct from either.’

Etymology See Kaua‘i Nukupu‘u for genus, Kaua‘i ‘Akialoa for subgenus. Epithet obscurus (‘dark’) a reference to relative plumage colour. I retain English name Lesser ‘Akialoa (AOU 1998), based on diminutive size relative to other ‘akialoas, even though I disagree with AOU (1998) taxonomy because name will remain appropriate once two recently discovered much larger spp. of ‘akialoa from Hawai‘i (Olson and James 1995; James 2004) are described. The modifiers ‘Hawai‘i’ and ‘Hawaiian’ will then be misleading.

Field identification

Description

Extinct. Endemic to Hawai‘i, where common and widespread into the 1890s (Wilson and Evans 1890–99; Rothschild 1893–1900; Perkins 1903), originally occurring in all wet forests down to sea level (first specimens taken at Kealakekua by Cook’s expedition). Wilson (1890b) found it primarily at low elevations (350–800 m) but Perkins (1903) found it up to 2000 m. Henshaw (1902) found it common in mesic koa/‘ohi‘a forest above Hilo but rare in very wet ‘ohi‘a/tree fern forests of ‘Ola‘a. Timing of decline and extinction not well documented because so few observers present in the early twentieth century. Last three specimens taken by C. E. Blacow and L. E. Miller at ‘Umikoa on the ne. slope of Mauna Kea Sep 1903 ( J. K. Lepson pers. comm. based on specimen labels). Apparently declined precipitously but may have persisted for several decades. Most recent reports are of one heard

A medium-sized (17 cm), all olive green bird with a very long, thin, decurved bill. Bill more sharply curved than in other ‘akialoas. Definitive male olive green above, rump and underparts slightly paler and yellower, undertail coverts olive-yellow, lores dark grey, supercilium bright olive-yellow and more defined than in Hawai‘i ‘Amakihi. Definitive female has similar pattern with more sharply defined supercilium and duller colours throughout: upperparts grey-green, underparts and supercilium dull yellow, fading posteriorly. Immature plumage (may be Juvenal or First Basic) similar to adult female, but all bright yellow tones replaced by dull white, brightest on supercilium and undertail coverts, with dull yellow wash on breast, olive-grey wash on sides; upperparts greyish-olive with two offwhite wing-bars.‘Bill and feet dark brown’ (Wilson

Looks like an overgrown Hawai‘i ‘Amakihi with a very long bill; coloration almost identical except the breast is much darker and pale superciliary more sharply defined. ‘Akiapola‘au has biggerheaded profile and is much yellower (male) or paler olive (female), has hammering feeding method. Song apparently closer to ‘amakihi than to ‘aki, but distinctive.

Distribution, habitat, and population status

Hoopoe-billed ‘Akialoa Hemignathus (Akialoa) upupirostris 249 (based on similarity to call of Kaua‘i ‘Akialoa) ca. 1937 near Keanakolu (Donaghho 1963) and one seen in 1940 in the same general area by Blacow (Baldwin 1941). Last specimens and reports all from area just north of present-day Hakalau Forest NWR. Specimens total 113 in the world’s museums (Banko 1979, as updated by J. K. Lepson, pers. comm.).

Diet and foraging For detailed description of foraging and diet by Perkins (1903), see Ch. 8. Mixed diet of invertebrates and nectar varied seasonally (Munro 1960). Foraged by hopping along trunks and branches of canopy trees and tree ferns (Wilson and Evans 1890–99; Rothschild 1893–1900; Perkins 1903; Munro 1960), frequently choosing decaying wood (Wilson 1890b). Probed with long bill into organic matter caught in leaf bases of ‘ie‘ie (Henshaw 1902; Munro 1960) or picked and probed in bark crevices and epiphytes.Took nectar

from ‘ohi‘a and tubular lobelioid flowers (Perkins 1903).

Breeding biology Only information comes from Perkins (1903: 424–5) who described a nest: built towards the extremity of one of the largest spreading branches of a Koa, placed above a fork and well concealed. It contained only one young one, and that already able to fly, while a second one was seen sitting on the branch outside the nest, with the old birds. The nest itself . . . appeared to be quite similar in form to that of [Hawai‘i ‘Amakihi], but was better concealed amongst the lichens covering the branch, and these appeared to have been partly used in its construction. This nest was found at the end of June, and in the same district (Kona) many young birds were noticed at the same time being fed by the parents.

Hoopoe-billed ‘Akialoa Hemignathus (Akialoa) upupirostris ( James and Olson) Hemignathus upupirostris James and Olson, Ornithol. Monogr. No. 46: 60. (Kauai.) Akialoa upupirostris Olson and James (1995)

Etymology See Kaua‘i Nukupu‘u and Kaua‘i ‘Akialoa.The epithet combines the Latin name of the Hoopoe upupa with rostrum, ⫽ beak.

Description From James and Olson (1991). A slightly larger ‘akialoa than those historically known from O‘ahu and Kaua‘i. It lacks lingual trough on the dorsal

surface of mandible characteristic of all other drepanidines, possibly indicating shorter tongue. Condition convergent with those in some other birds with long sickle-shaped bills such as scythebills (Campylorhamphus, Dendrocolaptidae), woodhoopoes (Phoeniculidae), and hoopoes (Upupidae).

Distribution and population status Extinct. Known only from prehistoric bones found at sites on Kaua‘i and O‘ahu ( James and Olson 1991). Bones of a smaller undescribed sp. with similar bill morphology found on Maui and of very large undescribed sp. with intermediate condition on Hawai‘i (H. James in Lepson and Johnston 2000).

250 Kaua‘i Nukupu‘u Hemignathus (Hemignathus) hanapepe

Subgenus Hemignathus Lichtenstein Hemignathus Lichtenstein, 1839, Abh. Phys. Kl. Akad. Wiss. Berlin (1838), p. 449. Type, by subsequent designation (G. R. Gray, 1841), Hemignathus lucidus Lichtenstein. Heterobills, with mandible about half the length of the maxilla. Mandible curved (three spp.) or straight. Bills used independently in foraging. Jaw muscles modified, intermediate between those of congeners and Pseudonestor (Zusi 1989). Plumage brightest and yellowest of all Hemignathus. Insectivorous. Songs short warbles.

Comprises three historical spp. of nukupu‘u (Pratt and Pratt 2001), one sp. each on Kaua‘i, O‘ahu, and Lana‘i; the recently described Giant Nukupu‘u known from prehistoric remains on Hawai‘i (Olson and James, 2003); and ‘Akiapola‘au, endemic to Hawai‘i. Note that a study skin of a nukupu‘u reported to have been taken on Hawai‘i (Olson and James 1994b) turned out to be an O‘ahu Nukupu‘u on the basis of DNA comparisons (Fleischer in T. K. Pratt et al. 2001).

Kaua‘i Nukupu‘u Hemignathus (Hemignathus) hanapepe Wilson PLATES 4, 8 Hemignathus hanapepe Wilson, 1889, Annals and Mag. Nat. Hist. (6)4: 401. (Kauai.) Heterorhynchus hanapepe Rothschild (1893–1900), Bryan (1901a), Henshaw (1902), Perkins (1903) Heterorhynchus lucidus hanapepe Olson and James (1982b) Hemignathus lucidus hanapepe Bryan and Greenway (1944), Munro (1944), Amadon (1950), Berger (1972a, 1981),AOU (1983, 1998), Pratt et al. (1987), Sibley and Monroe (1990), James and Olson (1991) Hemignathus (Heterorhynchus) lucidus hanapepe Pratt (1979a)

Etymology Genus name from Greek meaning ‘half-jaw’ in reference to the lower mandible of nukupu‘us (to which genus was at first restricted), which is half length of upper. Species named for Hanapepe Dist., where Wilson (1889) found the bird. Nukupu‘u from Hawaiian nuku ⫽ nose (or beak) ⫹ pu‘u ⫽ hill, often applied to round-topped cinder cones, roughly ‘hillbill’ or ‘hump-bill’, from the shape of the beak.

Systematics Generic and spp. limits discussed in more detail in Ch. 5. All nukupu‘us and ‘Akiapola‘au lumped as

one sp. by Bryan and Greenway (1944), but Amadon (1950) and all subsequent authors considered the latter separate on basis of distinctive bill morphology. Pratt and Pratt (2001) recognised three phylogenetic spp. of nukupu‘u, and considered biological spp. status likely, position recently reinforced by large genetic differences between them, comparable to differences among other drepanidine spp. (R. C. Fleischer, pers. comm.). Noteworthy that rather striking colour differences not segments of morphocline.This book first modern publication to present nukupu‘us as three spp.

Description Medium-sized (15 cm) honeycreepers with ‘hetero’ bill, the mandible ca. half length of maxilla, both strongly down-curved. Sexually dimorphic as adults, with immatures of both sexes resembling adult female. Adult plumage probably attained in Second Prebasic moult. Definitive males have bright golden yellow head and breast, yellowish-green dorsum, white belly, and black lores. Definitive female and immatures mainly dull grey above (extending onto crown and forehead to define broad pale superciliary), with wing feathers edged pale yellow; underparts and superciliary white, some individuals showing yellow tinge in anterior portion of

Kaua‘i Nukupu‘u Hemignathus (Hemignathus) hanapepe 251 superciliary, chin, and throat; lores dark grey or black merging with grey postocular patch in upper auriculars. Some immatures show slightly paler wing-bars. Moult sequence not well known. Bill, legs, and feet intense black, iris brown. For measurements, see T. K. Pratt et al. (2001).

Voice Contact call a whistled keewit uttered by both sexes (Perkins 1895, 1903; Munro 1960). Song ‘a short but rather sweet warble, consisting of some half a dozen notes’ (Palmer in Rothschild 1893–1900: 102), which Perkins (1895) considered indistinguishable from song of Maui Nukupu‘u.

Field identification Easily confused with smaller Kaua‘i ‘Amakihi, which has much heavier and longer bill than other ‘amakihis, and often picks at bark. Bill of nukupu‘u actually thinner at base and absolutely black, the colour merging imperceptibly with black lores (Kaua‘i ‘Amakihi lores not intensely black). Underparts of ‘amakihi much darker in both sexes, undertail coverts never white. Head of nukupu‘u either all brilliant yellow (male) or nearly without yellow (female and immature), never the dull yellow-green of female Kaua‘i ‘Amakihi. In bright light some male ‘amakihi can look very yellow, but note colour of bill, lores, and undertail. Most convincing detail for Kaua‘i Nukupu‘u would be white undertail of male and white underparts of female, neither of which mentioned in any recent reports (see below).

Distribution, habitat, and population status Very likely extinct. Known historically from mesic to wet forests of koa and ‘ohi‘a from 600 to over 1200 m, between Waialae Str. and Hanapepe R. (Wilson and Evans 1890–99; Rothschild 1893– 1900; Perkins 1903). Subfossil remains from coastal Maha‘ulepu Cave indicate broader habitat and elevational range in prehuman times (Burney et al. 2001). Nineteenth-century naturalists considered it the rarest Kaua‘i honeycreeper (Perkins 1903). Last collected in 1890s and not observed again until a

‘fleeting glimpse’ of two birds in 1960 (Richardson and Bowles 1964: 30) and report by King (1961) in same area. John Sincock (in Scott et al. 1986) reported only two, without details, in 17 years of field work in the Alaka‘i region. S. L. Conant had a 30 sec glimpse in 1975, but others working in same area could not locate bird (Conant et al. 1998). Virtually all modern sightings by competent observers lack convincing details, and those with details usually reveal themselves to be misidentifications by observers unfamiliar with plumage of Kaua‘i Nukupu‘u (Pratt and Pyle 2000). For example, one supposed sighting in 1995 (reported in Conant et al. 1998 and repeated by Reynolds and Snetsinger 2001) had mixture of nukupu‘u vocal and bill details with plumage colours of Kaua‘i ‘Amakihi; others failed to eliminate that sp. as contender (based on notes from T. Casey and J. Jeffrey, in litt.). No twentieth-century report entirely convincing (Pratt and Pyle 2000).

Diet and foraging Insectivorous, taking prey from bark of trees in manner similar to ‘Akiapola‘au, but with movements less vigorous (Perkins 1903). Munro (unpublished journal entry reported by T. K. Pratt et al. 2001) observed that lower mandible not used to pry up bark or dig into wood contra Amadon (1947). Had ‘habit of keeping along the upper surface of a branch and examining the sides within its reach’ (Wilson and Evans 1890–99: 82) in contrast to ‘Akiapola‘au, which often hangs upside down from underside of branches. Foraged in ‘ohi‘a and koa, but also understorey trees such as kalia and ‘olapa, especially when dead and decaying (Perkins 1903). Only report of nectarivory from Keawe, Wilson’s Hawaiian assistant, who stated that the bird ‘feeds on bananas and oranges’ (Wilson and Evans 1890–99), which Perkins (1903) took to mean taking nectar from flowers rather than feeding on the fruit.

Social and interspecific behaviour Frequently joined mixed-species flocks (see Ch. 7), among which it kept to lower forest strata, leaving canopy to other spp. (Perkins 1903).

252 O‘ahu Nukupu‘u Hemignathus (Hemignathus) lucidus

O‘ahu Nukupu‘u Hemignathus (Hemignathus) lucidus Lichtenstein PLATE 4 Hemignathus lucidus Lichtenstein, 1839, Abh. Phys. Kl. Akad.Wiss. Berlin (1838), p. 451, pl. 5. (Oahu.) Heterorhynchus lucidus Rothschild (1893–1900), Bryan (1901a), Henshaw (1902), Perkins (1903), Olson and James (1982b) Hemignathus (Heterorhynchus) lucidus Pratt (1979a)

Etymology See Kaua‘i Nukupu‘u. Species name is Latin for shining or bright.

Systematics See Kaua‘i Nukupu‘u.

Description Medium-sized (ca. 14.5 cm), heterobilled honeycreeper (see Kaua‘i Nukupu‘u). Bill proportionally longest of three nukupu‘us, with slightly shallower curvature. Plumage sequence little known. Adults sexually dimorphic, immatures resemble adult female.Adult male bright golden yellow from cheeks to underparts, fading posteriorly to yellowish-white undertail coverts, fairly well-defined superciliary of same colour; upperparts including nape and crown uniform olive green, wing and tail feathers darker with olive green edgings; postocular stripe olive green, lores black. Female and immature very similar to Kaua‘i Nukupu‘u but darker, browner grey above, wing coverts edged yellow; dingier underneath, with yellowish-grey breast and flanks, yellowish-white undertail coverts, and golden yellow tinge to superciliary, cheeks, and throat. Some immatures have slightly paler wing-bars. Bill brownish-black with paler base to lower mandible, legs and feet black, iris brown. For measurements, see T. K. Pratt et al. (2001).

Field identification Male separable from O‘ahu ‘Amakihi on basis of bold yellow superciliary, from O‘ahu ‘Alauahio by curved, much darker bill. Female distinguished from both by lack of prominent wing-bars.

Distribution, habitat, and population status Extinct. Known only from small number of specimens taken in 1830s in and above Nu‘uanu Valley but reported abundant as late as 1860s (Perkins in Munro 1960). Late nineteenth-century naturalists believed it extinct (Wilson and Evans 1890–99; Rothschild 1893–1900), and Perkins (1903: 426) wrote: No doubt at the time [1837] the valley was densely forested for the greater part of its length, since Bloxam records that in 1825 he visited its head by a narrow path through a dense and shady forest. Lying as it does immediately behind the city of Honolulu this fine valley is now practically denuded of its forest and largely filled with unsightly guava scrub; and the cliffs on either side, which to Bloxam appeared bare in comparison with the valley itself, now alone retain their native vegetation. Today, Nu‘uanu Valley is the route of the freeway known as Pali Hwy. Lichtenstein’s (1839; translated in Rothschild 1893–1900) report that Deppe found the bird commonly feeding at flowers in banana plantations has not been taken very seriously (T. K. Pratt et al. 2001), but occasional nectarivory among congeners, and especially the native report of feeding on banana flowers by the Kaua‘i bird (Wilson and Evans 1890–99, Perkins 1903), lend credence.

Maui Nukupu‘u Hemignathus (Hemignathus) affinis 253

Maui Nukupu‘u Hemignathus (Hemignathus) affinis Rothschild PLATE 4 Hemignathus affinis Rothschild, 1893, Ibis, p. 112. (Maui.) Heterorhynchus affinis Rothschild (1893–1900), Bryan (1901a), Henshaw (1902), Perkins (1903) Heterorhynchus lucidus affinis Olson and James (1982b) Hemignathus lucidus affinis Bryan and Greenway (1944), Munro (1944), Amadon (1950), Berger (1972, 1981),AOU (1983, 1998), Pratt et al. (1987), Sibley and Monroe (1990), James and Olson (1991)

Rothschild 1893–1900: 104) as resembling song of House Finch but ‘lower in tone’. Note that Maui ‘Alauahio also has song like that of House Finch.

Field identification See Maui Parrotbill. Same size as Maui Parrotbill and female, in particular, easily confused with it because of bold yellow superciliary. Superciliary of nukupu‘u not as sharply defined as that of parrotbill. Male unmistakable if bill well seen, but otherwise resembles smaller Maui ‘Alauahio rather closely.

Other vernacular names: Nukupu‘u, Maui ‘Akiapola‘au

Distribution, habitat, and population status

Etymology

Endemic to Maui. Historically found in a restricted zone 1220–2100 m elevation above Makawao (Rothschild 1893–1900; Perkins 1903). Not reported for seven decades, then rediscovered in 1967 in Kipahulu Valley (Fig. 3.3; Banko 1968). HFBS found only one individual and inferred habitat to be wet ‘ohi‘a and mesic koa/‘ohi‘a forest with dense native understorey (Scott et al. 1986). Now possibly extinct, repeated intensive surveys having failed to locate any population (Reynolds and Snetsinger 2001), despite reports of juvenile in 1983 (Mountainspring in Scott et al. 1986) and of a pair showing courtship behaviour in 1989 (Fleischer in Reynolds and Snetsinger 2001). Last report with any published details that of Engilis (1990). All reports in last decade of adult males from Hanawi NAR. No recent report includes any physical evidence (one recording of supposed call note lost (R. C. Fleischer, pers. comm.)) and none could be confirmed by follow-up visits to locality, in contrast to ‘Akiapola‘au, Po‘o-uli, and other honeycreepers that are easily relocated within home range once found (although Reynolds and Snetsinger (2001) report ‘resightings’, again without details). Pratt and Pyle (2000) sceptical of all twentieth-century records. Sightings would be more believable if they involved the more distinctive female.Whether Maui Nukupu‘u still exists or is Loch Ness Monster of Hawaiian honeycreepers impossible to say.

See Kaua‘i Nukupu‘u. Latin affinis means related or adjacent.

Systematics See Kaua‘i Nukupu‘u.

Description Smaller (ca. 14 cm) than other nukupu‘us (above) but with same bill morphology and apparent plumage sequence. Adult males greyest of group dorsally, the colour of back extending through nape to hindcrown, with wing feathers edged pale dull olive; remainder of plumage bright rich yellow, becoming pale yellow on undertail coverts, lores black. Female and immature resemble adult male O‘ahu Nukupu‘u, with similar bold yellow superciliary and slightly more olive colouring in auriculars. Bill and feet black, iris brown. For measurements, see T. K. Pratt et al. (2001).

Voice Contact call keewit apparently identical to that of Kaua‘i Nukupu‘u and similar to that of ‘Akiapola‘au (Perkins 1903).Two songs, one described by Perkins (1895) as very similar to warbled song of ‘Akiapola‘au and identical to that of Kaua‘i Nukupu‘u, the other described by Palmer (in

254 ‘Akiapola‘au Hemignathus (Hemignathus) munroi

Social and interspecific behaviour When more numerous, frequently joined mixedspecies flocks (see Ch. 7). Has a close association

with Maui Parrotbill, the two foraging together but not competing because of different feeding apparatus (Perkins 1903).

‘Akiapola‘au Hemignathus (Hemignathus) munroi Pratt PLATES 4, 8 Heterorhynchus wilsoni (not Himatione wilsoni Rothschild, April 1893) Rothschild, November 1893, Avifauna Laysan, p. 75. (Hawaii.) Hemignathus munroi Pratt, 1979a, Dissertation Abstracts 40: 1581. New name for Heterorhynchus wilsoni, preoccupied. Heterorhynchus wilsoni Rothschild (1893–1900), Bryan (1901a), Henshaw (1902), Perkins (1903) Hemignathus olivaceus Wilson and Evans (1890–99) Hemignathus lucidus wilsoni Bryan and Greenway (1944), Munro (1944) Hemignathus wilsoni Amadon (1950), Carlquist (1970), Berger (1972a), Sibley and Monroe (1990), James and Olson (1991) Other vernacular names: Hawai‘i Nukupu‘u, Akialoa (Wilson and Evans 1890–99),‘aki’ (birders’ nickname, Pratt 2002b)

Etymology See Kaua‘i Nukupu‘u for genus. Species first named in honour of Scott Wilson, Rothschild’s first collector (see Ch. 3), renamed to honour George C. Munro, prominent in Hawaiian ornithology for seven decades (Pratt 1979a,b) but who had, at the time, no sp. named for him (Dysmorodrepanis munroi was then regarded as a freak). Hawaiian name from akihi ⫽ small green bird ⫹ po‘o ⫽ head ⫹ la‘au ⫽ war club,‘club-headed green bird’ an allusion to its hammering motions when foraging.

Systematics See Kaua‘i Nukupu‘u for genus. Except for Bryan and Greenway (1944), all authors have considered this form a separate sp. from the nukupu‘us.Amadon (1950) considered the two groups a superspecies, but Pratt (1979a), on basis of striking morphological

differences, questioned that designation. Recent discovery of Giant Nukupu‘u as subfossil from a cave on Hualalai (James and Olson 2003) precludes designation of any superspecies in subgenus Hemignathus. Mayr (1963) used ‘‘aki’ as example of how new morphotypes can arise. See Maui Parrotbill.

Description Medium-sized (15 cm) honeycreeper with heterobill, the lower mandible straight, leaving small gap (diastema) when bill closed. Tail short relative to congeners. Sexual dimorphism striking in bill size (male larger), moderate in plumage. Plumage sequence from T. K. Pratt et al. (1994), colours from pers. obs. Males exhibit delayed plumage maturation with Definitive plumage reached in Third Prebasic moult. Definitive male yellowisholive above, bright golden yellow on face (may be tinged orange) and underparts, with black lores; olive of dorsum extends slightly paler over top of head to base of bill, similar colour forming indistinct postocular stripe, but facial markings (especially yellow supercilium) not sharply defined. Definitive female greyer olive dorsally, extending onto cheeks and sides of neck with hint of pale superciliary; throat and breast yellow, shading to off-white on belly and undertail, lores dark grey. Bill and legs of adults black. Juvenal plumage yellowish-grey or olive above, with yellowish-tawny rump, wing feathers with broad pale edgings tending to off-white at tips, those of coverts forming narrow wing-bars; paler olive-grey below with flanks tinged tawny, feathers with darker greyish tips producing mottled effect; facial markings indistinct, bill tan or dull yellow at base. Immature plumages confusing, with sexes identifiable only by bill size. First Basic plumage similar to Juvenal, but without mottling on breast, sometimes with

‘Akiapola‘au Hemignathus (Hemignathus) munroi 255 reduced wing-bars due to wear or irregular replacement. Second Basic plumage usually lacks wing-bars, resembles adult female but may have less yellow in throat. For measurements, see T. K. Pratt et al. (2001).

HAWAI'I

Voice From Pratt et al. (1987),T. K. Pratt et al. (2001) and pers. obs. Calls include a variable whistle of 2– 4 notes, transliterated by T. K. Pratt as pitiur, churree, ker-ree, chureeur, chederee, or cheerweewee, used as contact note between members of pair. Other calls during foraging include a swit or chip and an upslurred sweet or squeet. Older immatures still begging from parents during learning period utter a distinctive sharp, evenly paced ‘beacon’ call variously described as chirp, tsew, or tseoop. Primary song (Fig. 7.3) a variable warble, tu-tu-WHEE-wheer-tudu-WHEE-yoo, that sometimes ends with a trill. Whisper song includes quiet versions of primary song (making it more recognisable to sp. than other drepanidine whisper songs), short random notes, and apparently mimicry of other spp. in home range ( fide T. K. Pratt, J. K. Lepson). For recordings, see Pratt (1996a).

Field identification Likely to be confused only with Hawai‘i ‘Amakihi; considerably smaller and not as bright yellow (males only). Overhanging maxilla of ‘aki can be hard to discern, but base of bill always looks comparatively heavy, and head looks disproportionately large, the latter impression often enhanced by fluffing of feathers. Hammering behaviour diagnostic (see below).

Distribution, habitat, and population status From Scott et al. (1986) and T. K. Pratt et al. (2001). Endemic to Island of Hawai‘i, with eclectic tastes in habitat, from dry mamane–naio forest, to mesic koa/‘ohi‘a forest and ‘ohi‘a rainforest. Originally island-wide, but now fragmented into ca. eight relict populations at 1340–2700 m elevation, some of which are now represented by only a few old males. Largest populations in mesic

?

?

Map 14

to wet forest on windward slopes of Mauna Kea and Mauna Loa from Hakalau Forest NWR to kipukas of Upper Waiakea FR and KulaniKeauhou area ne. of Hawaii Volcanoes NP, and in e. end of Ka‘u FR. Only two small populations remain in Kona in koa/‘ohi‘a forest, and only individuals remain in high elevation mamane–naio forest of e. and w. slopes of Mauna Kea.

Diet and foraging Diet almost entirely arthropods, but occasional taking of nectar noted by all observers. Prey categories include caterpillars, spiders, grubs and other larvae, and adult beetles (Perkins 1903; Ralph and Fancy 1996;T. K. Pratt et al. 2001). Use of peculiar bill discussed in detail in Ch. 6. Also uses feet to hold mamane pods against perch so that larvae can be extracted, or to restrain prey (T. K. Pratt et al. 2001). Larger prey items may be masticated, but smaller items swallowed whole. Has clear preference for koa even when ‘ohi‘a more common, and also for-

256 ‘Akiapola‘au Hemignathus (Hemignathus) munroi ages on naio and kolea, but prefers mamane over naio when present (Ralph and Fancy 1996). In all kinds of trees, prefers lichen-covered or dead branches males and females performing intraspecific niche partitioning with males preferring larger trunks and branches, females twigs and small branches (Ralph and Fancy 1996). Has unique (among honeycreepers) trait of drilling sap wells in ‘ohi‘a trees (Fig. 8.6), from which sap drips into bird‘s open mouth below well, guided by licking tongue (T. K. Pratt et al. 2001).

Social and interspecific behaviour From T. K. Pratt et al. (2001) except as noted. Aggressive encounters (mostly territorial disputes) usually involve countersinging and display only, but may become more violent with chasing and foot-grappling and involve both sexes in same-sex battles. Lion Head display directed toward either sex has head feathers ruffed out for long periods. Helicopter display involves a circular, hovering song-flight directed toward another male. Maintain family groups year-round, but may join postbreeding flocks with ‘Akepa, Hawai‘i Creeper, and other spp. (see Ch. 7). Unusually pugnacious toward other spp., displacing or chasing Lesser ‘Akialoa (Munro 1960), ‘Apapane, ‘Hawai‘i ‘Amakihi, Hawai‘i Creeper, ‘Akepa, ‘I‘iwi, and Japanese White-eye, but displaced by Red-billed Leiothrix.

Predation Probably prey to same avian and mammalian predators as other honeycreepers, but documentation limited. Short-eared Owl observed pursuing ‘Akiapola‘au (T. K. Pratt et al. 2001) and remains found in one Barn Owl pellet at Hakalau Forest NWR (Mostello 1996). Dive into underbrush (adults) or cease calling (immatures) when aerial predators pass overhead or when Hawai‘i ‘Elepaio give alarm calls (T. K. Pratt et al. 2001).

Breeding biology COURTSHIP AND MATING: Monogamous, pairs remaining together multiple years in stable popu-

lations (T. K. Pratt et al. 2001), but in declining populations pair-bonds may change annually (Ralph and Fancy 1996). Courtship rarely observed and little known; no uniquely associated displays reported. NEST SITE AND TERRITORY: Pairs maintain yearround home range, defended in part or totally. In some habitats, somewhat colonial with territories adjacent and clustered (T. K. Pratt et al. 2001). Nests built predominantly in terminal leaf clusters of ‘ohi‘a trees, 7–22 m above ground (Banko and Williams 1993;T. K. Pratt et al. 2001), rarely in cavities (Ralph and Fancy 1996). NEST: Gathering of materials and construction entirely by female, but male may play minor role in shaping nest. Thin strips of ‘ohi‘a bark peeled off by grasping with bill and pulling (Sakai and Ralph 1980; T. K. Pratt et al. 2001). Body of nest primarily of ‘ohi‘a twigs and hapu‘u tree fern rachises, rhizomes, and pulu (Fig. 8.22), with a few leaves in upper part; uniquely, ‘ohi‘a bark strips 2 cm wide project 5 cm above rim irregularly around nest cup, facing outward (Banko and Williams 1993). Average outside dimensions 110 ⫻ 140 mm, thickness 150 mm; inner cup diameter 65–79 mm, depth 45 mm (T. K. Pratt et al. 2001). EGGS: Only one egg described, 22.7 ⫻ 17.0 mm, pale cream splotched with brownish red, most heavily about larger end (Banko and Williams 1993).T. K. Pratt et al. (2001) consider clutch size as one, based on typical number of fledglings, but rarely broods of two are seen (Fancy and Ralph 1996) and I once observed three begging juveniles being fed by a pair at Kulani. INCUBATION AND HATCHING: Female alone incubates, is fed by male off nest only until young ca. 3 d old (limited data summarised by T. K. Pratt et al. 2001). Nestlings never described. PARENTAL CARE AND FLEDGING: From Ralph and Fancy (1996) except as noted. Female alone broods. First 3 d female feeds young and removes faecal sacs,

Maui Parrotbill Pseudonestor xanthophrys 257 male feeds female off nest; 3–6 d male feeds female on nest, ⬎9 d feeds young directly although female still primary feeder. One chick fledged at 21 d. Period of post-fledging dependence unusually long (4–13 mo.), perhaps because complex feeding apparatus difficult to master ( J. K. Lepson, pers. comm.).

Life cycle and demography Because of long dependency period, probably only one brood per year. Reproductive rate unknown, survivorship in one study 71% (Ralph and Fancy 1996). One wild male lived at least 13 years (T. K. Pratt et al. 2001).

Genus Pseudonestor Rothschild Pseudonestor Rothschild, 1893, Bull. British Ornithologists’ Club 1: 35. Type, by monotypy, Pseudonestor xanthophrys Rothschild. Medium-sized Hawaiian honeycreepers with massive, laterally compressed bill, parrot-like in profile. Jaw muscles modified for manipulation of large bill (Zusi 1989).Tongue non-tubular, slightly

curled longitudinally, with lateral and terminal laciniae (Rothschild 1893–1900). Insectivorous. Resembles heterobills in anatomy (Zusi 1989), behaviour, and ecology, and now believed allied with them and not with drepanidine finches as in most earlier classifications. Monotypic, endemic to Maui.

Maui Parrotbill Pseudonestor xanthophrys Rothschild PLATES 2, 8 Pseudonestor xanthophrys Rothschild, 1893, Bull. British Ornithologists’ Club 1: 36. (Island of Maui, Sandwich Islands.) Other vernacular names: Pseudonestor, Parrotbilled Koa Finch

Etymology Nestor is a genus of large parrots found in New Zealand, hence Pseudonestor translates roughly as ‘false parrot’. Specific epithet is based on Greek roots that mean ‘yellow eyebrow’.

Systematics Long regarded as related to drepanidine finches (Amadon 1950; Berger 1981; AOU 1983; Simon et al. 1997), especially Psittirostra (Raikow 1977b). Perkins (1903) and Pratt (1979a) suggested that relationships instead lay with heterobills (Hemignathus sensu stricto) and that similarities with psittirostrine honeycreepers superficial. Zusi (1989) found synapomorphic conditions of jaw muscles that support such a position, Fleischer et al.’s (1998)

DNA research placed Pseudonestor and ‘Akiapola‘au in same clade (but enigmatically allied parrotbill most closely with ‘Anianiau), and recent studies have found numerous homologies in plumage colour and sequence (Berlin et al. 2001) and breeding ecology (Simon et al. 2000) between parrotbill and ‘Akiapola‘au. Pratt (2001b) suggested possible future merger of Pseudonestor with Hemignathus. See also Ch. 5.

Description Medium-sized (14 cm) Hawaiian honeycreeper with short tail and massive parrot-like bill. Sexual dimorphism of bill size striking, coloration less so (Berlin et al. 2001a). Body coloration and maturational sequence essentially same as in Hemignathus, but dorso-ventral contrast much more pronounced. Facial features unique to genus. Definitive male olive green above, golden yellow below, with broad golden yellow eyebrow or superciliary stripe isolated from rest of underparts by dark eyestripe connecting nape with base of bill through

258 Maui Parrotbill Pseudonestor xanthophrys upper auriculars and lores, posteriorly colour of upperparts, black in lores. Upper mandible dark brown to black above, changing abruptly at bottom of eye-stripe to yellowish-ivory; lower mandible entirely yellowish-ivory. Female identified by duller plumage and smaller bill. Juvenal and First Basic plumages replace yellow with dingy white, and most (but not all) juveniles have offwhite wing-bars, and some have faint grey mottling on breast (Berlin et al. 2001a). Some evidence of distinctive Second Basic plumage (immatures intermediate in yellow coloration), but sequence not well known. For measurements, see Simon et al. (1997).

Voice Varied calls given by adults include sharp chick very similar to typical call of Maui ‘Alauahio but usually louder and more slowly paced (pers. obs.; Simon et al. 1997), a thin squeee similar to one call of Hawai‘i ‘Amakihi (Perkins 1903; Simon et al. 1997), and one- or two-note up-slurred whistles (Simon et al. 1997) probably uttered by adult males only (pers. obs.). First-year birds still following parents utter loud, regularly spaced tseeoop very similar to homologous call of ‘Akiapola‘au (pers. obs.) sometimes characterised as ‘sound beacon’ (Pratt 2001b; T. K. Pratt et al. 2001). Whisper song (Simon et al. 1997: 4) described as similar to primary song ‘but quieter, followed by series of short, soft, warbling whistles’ and chip, squeee, and whit notes may be true subsong (see Ch. 7) as in ‘Akiapola‘au. Primary song a descending cascade of short warbles tweetwee-twee-twee-twee or chewEE-chewEE-chewEE-chewEE-chewEE with a distinctive plaintive quality reminiscent of songs of Canyon Wren (Troglodytidae) or Ivory-billed Woodcreeper (Dendrocolaptidae). Individual elements resemble portions of song of ‘Akiapola‘au.

bill, leading to misidentification as female Maui Nukupu‘u, which also has yellow superciliary though less sharply defined.

Distribution, habitat, and population status Endemic to Maui-nui and probably widespread in variety of forest types in prehuman times. Subfossil remains found at ‘Ilio Point, Moloka‘i, and at Kahikinui and Pu‘u Naio, Maui, all lowland dry forest sites (Olson and James 1982b; James and Olson 1991). When first discovered in 1890s, restricted to E. Maui above 1000 m (Henshaw 1902; Perkins 1903). Lost to science for nearly half a century, rediscovered in 1950 (Richards and Baldwin 1953). HFBS found it restricted to ca. 50 km2 of rainforest on windward slopes of Haleakala from 1200 m to tree line at 2350 m (Scott et al. 1986), from Waikamoi Preserve clockwise to Kipahulu and Manawainui valleys, a distribution that remains unchanged today (Simon et al. 1997; Reynolds and Snetsinger 2001). Rare to locally uncommon, but regularly observed in Waikamoi Preserve (Pratt 2002b). Favours forests with substantial component of koa (Perkins 1903), so present range may be marginal (Simon et al. 2000).

MAUI

Field identification Parrotbill’s pale superciliary easily seen even in poor light when outline of bird may be obscure (pers. obs.), but dark eye-stripe appears to continue uninterrupted from back of auriculars to tip of bill and can produce the optical illusion of thin black

Map 15

Maui Parrotbill Pseudonestor xanthophrys 259

Diet and foraging Feeds almost entirely on larvae of woodboring beetles (ⱕ77% of prey captures, Mountainspring 1987), caterpillars, and other invertebrates (Rothschild 1893–1900; Perkins 1903; Simon et al. 1997). Simon et al. (1997) give detailed taxonomic summary of prey. Prey extracted from live and dead wood, dry twigs, and ripe fruits. For details of foraging manoeuvres, see Ch. 6. Rarely, takes nectar from ‘ohi‘a, ‘akala, and ‘ohelo (Simon et al. 1997). Whether parrotbill actually eats small fruits or simply opens them in search of insect larvae (see Ch. 6, 8) problematical.

Social and interspecific behaviour Juveniles continue to follow and beg from parents 5–8 mo. as bill-strength foraging efficiency increase (Simon et al. 2000). Pairs highly philopatric in mutually exclusive home ranges (ca. 9 ha) throughout year; may or may not show strong aggression at borders (Simon et al. 1997, 2000, 2001b). In rare aggressive encounters, adult males pursue each other along a branch, turn and face each other with heads lowered and rumps raised, and sing alternately; alternate singing continues after birds part (Baker and Baker 1997). Males also pursue each other and occasionally attack with bill (Mountainspring 1987; Simon et al. 1997). Not aggressive toward other spp. except in defence of nest materials that may be stolen by ‘Akohekohe and ‘Apapane (Simon et al. 1997). Often forages in company with Maui Nukupu‘u (Perkins 1903) and Maui ‘Alauahio, the latter feeding opportunistically at parrotbill excavations (Baker and Baker 1997).

Breeding biology All information from Simon et al. (1997, 2000) except as noted. COURTSHIP AND MATING: Monogamous, pairs remaining together multiple seasons. Courtship rituals include sexual chasing, with male sometimes carrying twig in bill, wing-fluttering by both sexes, and courtship feeding with begging display by

female, the latter occurring intermittently throughout year to maintain pair-bond. NEST SITE AND TERRITORY: Breeding territory includes much of home range. Nest in outer canopy of large ‘ohi‘a trees ca. 0.5 m below foliage in branch forks ca. 12 m above ground. Perkins (1903) reported one possible nest in koa. NEST: Built by female, with male contributing material early in process, requiring 7–18 d for completion. Made mostly of Usnea lichen and/or mosses (Lockwood et al. 1994) interspersed with pukiawe twigs. Mean dimensions: outside diameter 9.6–11.4 cm, overall height 7.5 cm, inner cup diameter 4.4–5.6 cm, cup depth 3.3 cm. EGGS: Off-white to tan mottled with lavenderbrown mostly around larger end, lightly flecked with black throughout. Size (mean, n ⫽ 6) 21.7 ⫻ 15.4 mm. Clutch usually one (87% of observed nests or pairs with fledglings, n ⫽ 40), rarely two. Laying begins 2–4 d after completion of nest. INCUBATION AND HATCHING: Female alone incubates, fed by male on or off nest. Incubation period estimated 16 d. Hatching never observed in wild. NESTLINGS: Helpless with eyes closed at hatching, nearly naked with thin grey down or dorsum, skin reddish-pink. PARENTAL CARE AND FLEDGING: Female alone broods, but male primary feeder. Parents remove faecal sacs through 9 d, afterwards chicks defecate over rim with some faecal accumulation on nest. Chicks fledge at 18–22 d. Broods with two fledglings may be divided between parents (Baker and Baker 1997).

Life cycle and demography Long attenuated breeding season, with first nests in Nov.Adults with young present all year. May renest if first attempt fails (Simon et al. 2000), but long fledgling dependency precludes second nesting by successful pairs.

260 ‘Apapane Himatione sanguinea

Genus Himatione Cabanis Himatione Cabanis, 1850, Museum Heineanum, vol. 1: 99. Type, by monotypy, Himatione sanguinea (Gmelin). Small Hawaiian honeycreepers with mediumlength, thin, pointed, slightly down-curved bills, nectar-adapted, drepanidine tubular tongue, and full nasal opercula. No sexual dichromatism, but

juveniles distinctive. Adult body plumage mainly red, Juvenal brown.Vocalisations highly varied and complex, with tonal qualities like those of meliphagids. Two spp., one geographically undifferentiated among main Hawaiian Islands, another isolated on Laysan (NWHI).

‘Apapane Himatione sanguinea (Gmelin) PLATES 6, 8 Certhia sanguinea Gmelin, 1788, Syst. Nat. 1(1): 479. Based on the ‘Crimson Creeper’ Latham, Gen. Synop. Birds 1(2): 739. (in insulis Sandwich ⫽ Hawaii.) Other vernacular names: Akapane, Akakani

Etymology Genus name from Greek himation, a crimson war cape worn by the Spartans, specific epithet from Latin word for blood, both references to the bird’s colour.

Systematics Genus Himatione used for several other small Hawaiian honeycreepers by early workers (see synonymies), but restricted to this and following spp. since ca. 1900.

Description A medium-sized (13 cm) honeycreeper with short down-curved, but not sickle-shaped, bill, tail often held cocked up, and outer primaries with truncate tips that produce noticeable note in flight. Sexes distinguishable only by measurements (Fancy et al. 1993a). Definitive plumage reached at Second Prebasic moult. Adults bright crimson or blood red over most of head and body, including shoulders and wing coverts, brightest on head; belly and posterior flanks grey, undertail coverts fluffy and white; wings and tail black, tertials narrowly edged crimson on outer web, primaries narrowly edged

white on broad portion of outer web. Juvenal plumage with basal colour of upperparts and breast tawny or ochraceous (close to the artists’ raw sienna), most feathers with broad dusky tips producing spotted pattern heaviest on breast; upper back greyer, close to artists’ raw umber, feathers of forehead and auriculars with darker centres producing streaks; outer scapulars, face, and throat variably tinged orange-red; posterior underparts white, including flanks; wings and tail dull black, lesser and marginal coverts broadly edged buff, greater coverts, secondaries, and tertials edged buff shading to white at tips, primaries edged as in adult. First Basic plumage intermediate, with body colouring of adult, often with some Juvenal feathers scattered through, but retaining very faded flight feathers of Juvenal. Bill and feet black at all ages, but juvenile has pinkish base to lower mandible. For measurements, see Fancy and Ralph (1997).

Voice and other sounds Most frequent sound mechanically produced by primaries in flight, a low, rapid, dove-like cooing trill that inexperienced observers often believe to be a vocalisation. Call, often uttered in flight, an inconspicuous down-slurred teerp or tewp. Songs comprise nearly infinite array of complex patterns that include whistles, trills, warbles, burrs, squeaks, clicks, and bell-like sounds. May be canary-like in

‘Apapane Himatione sanguinea 261 pattern, but not in tone quality, which tends toward the metallic or reedy. One song sounds like a person dialling an old-fashioned rotary telephone zhuup! tic-tic-tic-tic-tic, etc.; another is a lively shurTEE, ta-da-TEE, quoit-quoit-quoit; but others may be only single sustained notes. Ward (1964) described some of the variations, which may be individual as well as geographic. At given time at given locality, all ‘Apapane seem to sing same song for a while, then switch to another. Novice birders in Hawai‘i complain that every new song heard seems to be sung by an ‘Apapane. For recorded samples, see Pratt (1996a) and Kellogg et al. (undated); for sonograms see Ward (1964) and Fancy and Ralph (1997).

Field identification White undertail coverts contrasting strongly with darker body plumage, especially useful mark in overhead flight, distinguish ‘Apapane from all other Hawaiian forest birds (Pratt et al. 1987). Despite confusing vocal array, does not mimic other spp.

Distribution, habitat, and population status The most abundant Hawaiian honeycreeper today, reaching densities of 2000/km2 in Ka‘u Dist., Hawai‘i, among highest densities known for any non-colonial bird (Scott et al. 1986). Originally distributed throughout ‘ohi‘a forests of all main islands, but now nearly extirpated from Moloka‘i and Lana‘i (Scott et al. 1986; Pratt 2002b) and much reduced from former numbers on O‘ahu (Fancy and Ralph 1997). Still occurs, perhaps as daytime visitor only, near sea level at a few places on Hawai‘i, but most of population found above 1250 m there and on Maui and Kaua‘i. O‘ahu population occurs 300 m to summit (946 m) in Ko‘olau Mts., above 600 m in Waianae range (Fancy and Ralph 1997). Nomadic in search of nectar sources, especially outside breeding season (Perkins 1903; Baldwin 1953; MacMillen and Carpenter 1980; Ralph and Fancy 1995). Makes daily flights of several km between low-elevation foraging sites and high-elevation roosts (Macmillen and Carpenter

KAUA'I

NI'IHAU O'AHU

MOLOKA'I MAUI _ LANA'I KAHOOLAWE

Map 16

HAWAI'I

262 ‘Apapane Himatione sanguinea 1980; Berger 1981). Visits mamane–naio forest on Mauna Kea when trees in bloom, but returns to ‘ohi‘a forest to roost (Ralph and Fancy 1995).

Diet and foraging Mainly nectarivorous, but also takes spiders and insects from foliage. A major pollinator of ‘ohi‘a (Carpenter and MacMillen 1975), source of most nectar taken. Also reported feeding in native koa, naio, mamane, kolea, alani, kanawao, koki‘o ke‘oke‘o, and olapa as well as alien tagasaste, schefflera, fuchsia (Berger 1981), and even formerly in coconut (summarised by Ralph and Fancy 1997). For details of arthropods taken, see Baldwin (1953) and Fancy and Ralph (1997).

Social and interspecific behaviour Highly gregarious even during breeding season (except in immediate vicinity of nest). Holds low position in dominance hierarchy of nectarivores (see Ch. 8). Flocking allows access to food resources from which small groups or individuals would be excluded (Carothers 1986a). May join postbreeding mixed-species flocks (Fancy and Ralph 1997).

Predation Actual cases poorly documented, but believed to suffer predation by introduced rats, feral cats, mongooses, and Barn Owls as well as native Hawaiian Hawk and Short-eared Owl (Fancy and Ralph 1997), although enigmatically rare in pellets of extinct native owls (Olson and James 1982b).

Breeding biology COURTSHIP AND MATING: Courtship display involves singing male hopping branch to branch with tail held cocked to display white undertail (Berger 1981). Singing and sexual chasing occur during pair formation Jan–Mar. Pairs monogamous. Female solicits copulation and courtship feeding by lowering and fluttering wings (Eddinger 1970). NEST SITE AND TERRITORY: Both sexes defend small area around nest only. Nest sites highly variable, most often terminal clump in canopy of ‘ohi‘a, but also in upper branches of kawa‘u, koa,

and hapu‘u ferns (Sakai 1983), and in tree cavities and lava tubes (van Riper 1973a; Sakai 1983). NEST: Open cup built by both sexes predominantly of moss but including lichens, ‘ohi‘a leaves and twigs, bark, and rootlets, cup lined with grass and sedge fibres. May use material from abandoned nests (Eddinger 1970; Sakai 1983) or build atop previous year’s nest (van Riper 1973a). Dimensions average 9.5 cm outside diameter, 5.1 cm cup diameter, 10.2 cm height, 3.8 cm cup depth (Fancy and Ralph 1997, based on data from Eddinger 1970). EGGS: From Eddinger (1970) and Berger (1981). White with spots of various shades of brown concentrated at larger end. Laying begins 1–6 d after nest complete, one egg per day in morning. Clutch 1–4, median 3. INCUBATION AND HATCHING: From Eddinger (1970). Only female incubates, fed off nest by male. Incubation period averages 13 d. Eggs all hatch morning of same day. NESTLINGS: From Eddinger (1970).Young helpless at first, with eyes closed, naked except for sparse down dorsally, gape bright pink with cream rictal flanges. Eyes open at 4 d. Feathers begin unsheathing at 8 d, fully feathered by 14 d. PARENTAL CARE AND FLEDGING: From Eddinger (1970). Female only broods, almost continuously at first, later mainly at night and during heavy rain. Both parents feed chicks, but ⬎70% of feedings by female. Nest kept clean of faecal sacs throughout cycle.Young fledge at 15–17 d, mean 16 d.

Life cycle and demography Breeding cycle in first half of year, with peak numbers of juveniles in July (Fancy and Ralph 1997). Breed at 1 year (Baldwin 1953). Success rates 79.8% for hatching, 70.2% for fledging, 88.0% for nestling survival (Eddinger 1970). Ralph and Fancy (1995) found annual survival rate (i.e. probability of survival) 0.72 for adults, 0.13 for immatures (figures include permanent emigration as well as mortality). Longevity in wild unknown; one captive lived at least 11 years (Fancy and Ralph 1997).

Laysan Honeycreeper Himatione freethii 263

Laysan Honeycreeper Himatione freethii Rothschild PLATE 6 Himatione fraithii (sic) Rothschild, 1892, Annals and Mag. Nat. Hist. (6)10: 109. (Laysan.) Himatione fraithii Henshaw (1902) Himatione sanguinea fraithii Bryan and Greenway (1944) Himatione freethi Rothschild (1893–1900), Wilson and Evans (1890–99), Bryan (1901a), Perkins (1903), Munro (1944), Olson and James (1982b) Himatione sanguinea freethii Amadon (1950), Greenway (1968), Carlquist (1970), Pratt (1979a), Berger (1972a, 1981), AOU (1983, 1998) Other vernacular names: Laysan Honeyeater, Laysan ‘Apapane, Laysan Akakani

Etymology For genus, see ‘Apapane. Species named for Captain Freeth who managed the guano operation on Laysan when Palmer and Munro visited in 1892 (Bailey 1956). Rothschild (1892) initially misspelled the name as fraithii but later in same publication corrected it.

Systematics First described as separate sp. but lumped with ‘Apapane by Amadon (1950) and most subsequent authors. Olson and James (1982b) and Olson and Ziegler (1995) split the two on basis of osteological differences, and Pratt and Pratt (2001) discuss numerous potential isolating mechanisms that argue for separate sp. status (see Ch. 5).

Description Similar to ‘Apapane but with shorter, stouter bill and primaries without oblique truncation.Adults mostly orange-vermilion, tinged scarlet on the head, with dusty brown belly and undertail coverts, dark brown wings and tail edged as in ‘Apapane. Juvenal plumage dull brown above, light ashy-brown below, with feathers tipped rich buffy-brown (or darker tones) producing mottled pattern; undertail coverts pale buff, chin and throat orange-buff (Rothschild

1893–1900). Olson and Ziegler (1995) consider the plumage differences the result of fading in intense atoll light, but white undertail feathers cannot ‘fade’ to brown because they already lack pigment.

Voice Palmer (in Rothschild 1893–1900) described song as low and sweet, with several different notes. Apparently uttered only during the breeding season.

Distribution, habitat, and population status Endemic to Laysan and extinct. Found throughout the small island, but concentrated in grassy and bushy vegetation adjacent to the island’s interior lagoon (Fisher 1903, 1906). Destruction of island’s vegetation by introduced European rabbits led to demise of this sp. (see Ch. 2 and Pratt 1994a for details).

Diet and foraging From Fisher (1903, 1906). Took nectar from flowers of ‘akulikuli and ihi and also fed on insects such as caterpillars and miller moths. Often foraged by walking on the ground.

Breeding biology NEST SITE AND TERRITORY: In grass (apparently kawelu based on Fisher’s photograph in Bailey 1956) clump. No information on territoriality. NEST: Open cup of grass blades and rootlets, lined with rootlets and albatross down; bowl 6.9 cm diameter, 4.2 cm deep (Schauinsland 1899; Fisher 1903). EGGS: White with greyish spots concentrated at larger end (Fisher 1903). Size 19.7 ⫻ 14.2 mm averaged from data in Schauinsland (1899) and Fisher (1903). Clutch 3–5, median probably 3 (Fisher 1903; Ely and Clapp 1973).

264 ‘Akohekohe Palmeria dolei

Genus Palmeria Rothschild Palmeria Rothschild, 1893, Ibis, p. 113. Type, by monotypy, Palmeria mirabilis Rothschild ⫽ Himatione dolei Wilson. Large Hawaiian honeycreeper similar to Himatione in bill shape and body proportions but with highly autapomorphic plumage and unusual derived con-

dition of flexor digitorum longus (Raikow 1977b). Plumage mainly black, with red-orange spots and nape patch, white-tipped primaries and rectrices. Stiff, recurved, yellowish-white feathers of forehead form bulbous crest that probably serves as pollen brush. Nectarivorous. Monotypic, endemic to Maui-nui.

‘Akohekohe Palmeria dolei (Wilson) PLATES 7, 8 Himatione dolei S. B. Wilson, 1891, Proc. Zool. Soc. London, p. 166. (Maui.) Other vernacular names: Crested Honeycreeper, Crested Honeyeater

Etymology Genus named for Henry Palmer (see Ch. 3). Species epithet honours kama‘aina politician and jurist Sanford B. Dole1 who, as amateur ornithologist, published first organised list of Hawaiian birds (Dole 1869). Hawaiian name often considered simply onomatopoeic (Berlin and VanGelder 1999), but that does not explain why early naturalists created an English name for this sp. alone when valid Hawaiian one available. Hawaiians had sense of humour and loved puns; name includes kohe ⫽ vagina (Pukui and Elbert 1971), allusion to resemblance of bird’s crest to pubic hair, apparently too much for Victorian sensibilities.

Systematics Although closely related to and originally considered congeneric with ‘Apapane, Rothschild’s (1893) genus has stood test of time undoubtedly because of highly autapomorphic plumage. Amadon (1986) stated that if Vestiaria and Drepanis merged, Palmeria and Himatione would have to follow (but he never attributed much importance to plumage differences among honeycreepers). Recent studies ( James 1998; Pratt 2001b) fail to support unequivocally merger of Palmeria with Himatione, unlike case of Vestiaria and

Drepanis. Point currently moot. Future merger on genetic grounds would not be surprising.

Description Large (18 cm) honeycreeper with sharp, slightly decurved bill above which a globular crest of stiff, recurved feathers. Adult sexes alike, separable only by measurements or presence of cloacal protuberance or brood patch (Simon et al. 1998); immatures distinctive. Adult only drepanidine describable as ornate. Base colour throughout (except for grey undertail coverts) black, body feathers with silvergrey shaft streaks, tipped silver on face and upper breast, orange posteriorly both above and below, feathers of head and underparts lanceolate; wing and tail feathers margined and tipped white, forming white band at tail tip and sometimes single wingbar; marginal coverts (‘shoulders’) mottled white, orange, and black; broad comma-shaped buffyorange patch surrounds eye and extends posteriorly to sometimes connect with irregularly margined flame orange nape patch; thighs buffy-orange; crest feathers mostly white but variably tipped dark slate grey or tinged golden yellow. Crest appears ragged and unkempt when wet, but fluffs up when dry to resemble cotton ball stuck to forehead. First Basic plumage mostly dark brownish slate, paler and buffy on belly and undertail coverts, breast feathers with off-white or buff shaft streaks; feathers of forehead grey and bristly, hinting at future crest; some head feathers have hint of orange tips; wing and tail feathers slaty brown, tips of tertials and outer webs of

‘Akohekohe Palmeria dolei 265 primaries edged white. Juvenal plumage much browner, with head and breast feathers tipped oliveyellow, forehead, auriculars, and chin silvery, undertail coverts greyish-buff; wing and tail feathers dark grey edged tawny or olive-yellow (Berlin and VanGelder 1999). Bill yellow in Juvenal plumage, darkening progressively from tip back to become intense black in adults. Legs dark brown ( juveniles) to black (adults). Iris brown. For measurements, see Berlin and VanGelder (1999).

Voice and other sounds Although differences possibly only semantic, I disagree with Berlin and VanGelder’s (1999) categorisation of all vocalisations as calls with no primary song, despite association of some with territoriality. Repertoire intermediate between that of ‘Apapane, with obviously distinguishable songs, and ‘I‘iwi with calls and songs undifferentiated. Most frequent call a loud, human-like, up-slurred whistle (like the first note of the ‘wolf whistle’) easily imitated and nearly identical to similar call of ‘I‘iwi. More distinctive is four-note peter-peter, reminiscent of similar call of Tufted Titmouse but with hollow quality. Other vocalisations, most of which probably songs because seasonal, comprise highly varied array with common qualitative theme: notes harsh or screechy, low-pitched, dissonant (with interspersed clicks and ‘smacking sounds’ like human tsk-tsk tongueclicking, and bell-like notes, entire repertoire much quieter than songs of ‘Apapane and ‘I‘iwi, and with ventriloquial quality that makes them sound further away than they are.Variants characterised as evenly paced AH, ko-hay ko-hay (namesake), kerr-kerr ko-ko or gluck-gluck (Berlin and VanGelder 1999), ah-churg-churg-churg or greee-tawk-tawk (Pratt et al. 1987). Some songs have double-note elements that sound like oh-oh, possibly the source of recent audio reports of Bishop’s ‘O‘o. Although lacking specialised primary tips of ‘Apapane and ‘I‘iwi, ‘Akohekohe produces similar but quieter wing note.

and may have pale patches about the head, but never yellow plumes from auriculars. Fan-shaped rather than graduated tail diagnostic.

Distribution, habitat, and population status Originally probably inhabited all of Maui-nui in suitable habitat, but known historically only from ‘ohi‘a rainforests of e. Moloka‘i and E. Maui. Locally abundant in 1890s (Perkins 1903) but very rare on Moloka‘i by 1907 (Bryan 1908) and believed extirpated by 1944 (Richardson 1949), though Pekelo (1967) thought it still present in 1960s; now certainly gone from Moloka‘i (Scott et al. 1977, 1986). Maui range originally included Kula and Makawao regions (Wilson and Evans 1890–99), now given over to ranch lands. Remains locally common on n. and e. slopes of Haleakala at 1100–2300 m from Waikamoi Preserve clockwise around to Manawainui Valley and restricted to very wet, epiphyte-laden forests with dense understorey (Berlin and VanGelder 1999). Apparently wander widely after breeding in search of nectar resources (Conant 1981a;VanGelder and Smith 2001).

Diet and foraging Apparently an ‘ohi‘a nectar specialist, but seasonally feeds ⬍40% on kolea,‘alani, kanawao,‘ohelo, pukiawe, and ‘akala (Berlin and VanGelder 1999; Berlin et al. 2001;VanGelder and Smith 2001). Also takes MAUI and MOLOKA'I

Field identification Unlikely to cause confusion in field except for hopeful observers searching for Bishop’s ‘O‘o. Juvenile ‘Akohekohe can look almost uniformly dark,

Map 17

266 ‘Akohekohe Palmeria dolei invertebrates such as caterpillars, flies, and spiders mainly from leaves and branches of ‘ohi‘a (Carothers 1986a,b; Berlin and VanGelder 1999; Berlin et al. 2001b), and very rarely may take ‘olapa fruit during periods of low bloom (Carpenter 1976). Immatures take higher percentage of animal food than adults, perhaps because of protein demands of growth (Carothers 2001). Feeding movements reported as 63% probing in flowers or leaves, 23% gleaning leaves and bark for invertebrates, and 12% biting (Berlin et al. 2001b).Trapline feeding discussed in Ch. 8.

Social and interspecific behaviour Solitary most of the time, paired only during breeding season. Highly aggressive and dominant today in hierarchy of nectarivores (see Ch. 8), but formerly dominated by Bishop’s ‘O‘o and Black Mamo (Perkins 1903). Territorial around favoured trees (Carothers 1986b), maintaining mutually exclusive territories with ‘I‘iwi (Carothers 1986a). May also defend all-purpose home-range territory, unlike most other honeycreepers (Simon et al. 2001b).

Predation Probably preyed upon by prehistoric stilt-owls (Olson and James 1991). Short-eared Owl suspected as occasional predator today (VanGelder 1996). Other documented predators include Barn Owl and (presumably roof ) rats (Berlin and VanGelder 1999).

Breeding biology

ily same as feeding territory. Territory diameter 160–240 m. Sites dense stands of ‘ohi‘a with heavy canopy cover and understorey with large amount of kolea. Most nests within 80 m of nearest neighbour, producing clustered distribution. Nest in upper 20% of canopy, ca. 14 m above ground, with dense foliage above. NEST: From Berlin and VanGelder (1999). Built mostly by female. Base of twigs from ‘ohi‘a,‘olapa, and pukiawe, topped with moss and lichen clumps. Materials may be reused from old nests. Inner cup lined with fern rootlets and ‘ohi‘a stamens. Outside measurements (mean; n ⫽ 29): 15 ⫻ 12.4 cm, thickness 8.1 cm, inner cup 7.3 ⫻ 5.4 cm, cup depth 3.3 cm. EGGS: From Berlin and VanGelder (1999) and Simon et al. (2001a). Drab grey spotted, streaked, and splotched with shades of brown, mostly around larger end. Size (n ⫽ 6) 24.2 ⫻ 15.3 mm. Clutch size one (30%) or two (70%). INCUBATION AND HATCHING: From Berlin and VanGelder (1999), VanGelder and Smith (2001), and Simon et al. (2001a). Incubation period 14–19 d. Female alone incubates. Male feeds female on or near nest. NESTLINGS: From Berlin and VanGelder (1999). Young helpless with eyes closed at hatching, mostly naked with pale grey down dorsally. Eyes open at 4 d. Chicks defecate over nest rim after 6 d. Pinfeathers present by 12 d, other maturational events poorly known.

COURTSHIP AND MATING: From Berlin and VanGelder (1999). Have spectacular aerial displays prior to breeding season in which 2–6 individuals fly ⬍50 m above canopy making wide circles, arcs, dives, and turns or hovering briefly, singing all the while. Males chase females into air for displays, may present gift of nesting material. Courtship feeding occurs throughout breeding cycle.

PARENTAL CARE AND FLEDGING: From Berlin and VanGelder (1999) and Simon et al. (2001a). Female only broods, but both parents feed nestlings by regurgitation. Nest kept clean of faecal sacs by both. Nestling period 21 d (mean, n ⫽ 35). Fledglings independent 10–14 d after fledging.

NEST SITE AND TERRITORY: From Berlin and VanGelder (1999) and VanGelder and Smith (2001). Territory maintained around nest site not necessar-

From Berlin and VanGelder (1999), Berlin et al. (2001), and Simon et al. (2001b). Protracted breeding season Nov–June coincides with peak ‘ohi‘a

Life cycle and demography

Hawai‘i Mamo Drepanis pacifica 267 bloom but also with coldest and wettest weather. Renesting frequent compared to other honeycreepers. Reproductive success highly variable (36–80%)

year-to-year, depending on rainfall. Annual probability of adult survival as high as 0.95. Population apparently stable.

Genus Drepanis Temminck Drepanis Temminck, 1820, Manuel d’ornithologie, ed. 2, vol. 1, p. 86. Type, by subsequent designation of Gray (1840), Certhia pacifica Gmelin. Drepanorhamphus Rothschild, 1900,The avifauna of Laysan, pt. 3, p. 163. Type, by monotypy, Drepanis funerea Newton. Vestiaria Jarocki, 1821, Zoologia 2: 75. Type, by monotypy, Certhia vestiaria Latham ⫽ Certhia coccinea Forster. Medium to large Hawaiian honeycreepers with down-curved, falcate or sickle-shaped bills supported by bone throughout most of their length (Baldwin 1953). Nostrils fully operculate.Tongue of drepanidine tubular configuration (Raikow 1977b). Primarily nectarivorous, all spp. feeding on flowers of arborescent Lobeliaceae when available, as well as

other flowers (Perkins 1903; Spieth 1966; Fancy and Ralph 1998; Pratt 2002a). Bills apparently coevolved with ornithophilous flowers of several small trees and shrubs (see Ch. 4, 8). Adults coloured in bold patterns of red, yellow, black, and white. Sexes similar, but at least one sp. has distinctive Juvenal plumage (plumages of others poorly known).Vocalisations loud and far-carrying, possibly convergent with songs of Hawaiian meliphagids (see Ch. 4, 7). Three spp.: Hawai‘i Mamo of the Island of Hawai‘i; Black Mamo of Maui-nui ( James and Olson 1991); and ‘I‘iwi, known from all forested islands. Includes monotypic genera Drepanorhamphus Rothschild, 1900 for Black Mamo and Vestiaria Jarocki, 1821 for ‘I‘iwi. See Ch. 5 for discussion of merger of Vestiaria and Drepanis.

Hawai‘i Mamo Drepanis pacifica (Gmelin) PLATES 7, 8 Certhia pacifica Gmelin, 1788, Syst. Nat. 1(1): 470. ‘Great Hook-billed Creeper’ Latham, Gen. Synop. Birds 1(2): 703. (in insula amicis maris australis ⫽ Friendly (Tonga) Islands, error for Hawai‘i). Drepanidis pacifica Carlquist (1970) Other vernacular names: Mamo, mamo, hoha, hoho

Etymology Drepanis is Latin for sickle, in reference to bill shape. Specific epithet pacifica given when exact locality of origin unknown. Hawaiians used ‘o‘ o as

generic term for all mostly black passerines, not restricting it to meliphagid genus Moho.

Systematics Monotypic. See discussion of genus, above.

Description One of largest (23 cm) Hawaiian honeycreepers with huge sickle-shaped bill. No sexual dimorphism or age differences known. Entirely glossy black except lower back, rump, uppertail coverts, upper flanks, undertail coverts, and thighs brilliant golden yellow; marginal coverts form yellow leading edge above and below wing, remaining underwing coverts yellowish-white; greater upper primary

268 Black Mamo Drepanis funerea coverts white, alular quills black, centres white; primaries, secondaries, and tertials brownish-black shading to grey on distal portions of outer webs.All but central four rectrices brownish-black with white spots near tips, progressively larger toward outside edge. Bill and legs dark brownish-black. For measurements, see Pratt (2002a).

Voice Much less vocal than congeners (Henshaw 1902). Perkins (1903: 399) describes ‘single, rather long and plaintive note, as imitated by natives who have been familiar with it.’ Responded to imitations of the note, which may therefore have been a song rather than a call.

Distribution, habitat, and population status Extinct. Endemic to Island of Hawai‘i where probably widespread in wet forests before alteration by humans. Probably mostly a low elevation sp., although Brigham (1892, 1899) reported a sighting high on Hualalai volcano in 1893. Most reports from below 1200 m (Pratt 2002a). Last known specimen captured alive in ‘Ola‘a by native collector working for Henry Palmer in 1892 (Munro 1960). Last sighting by Henshaw (1902) in 1898 at Kaumana above Hilo, now converted to suburban, mostly non-native vegetation.

Diet and foraging Field identification Not likely to be confused with any other bird on Hawai‘i. Hawai‘i ‘O‘o larger, longer tailed, with much less yellow posteriorly and much shorter bill. Flight said to resemble that of a cuckoo, smooth but not rapid (Henshaw 1902).

Mostly indirect evidence of specialisation on nectar of haha, but probably took any nectar available (Pratt 2002a). Henshaw (1902: 52) saw pair that ‘were very active, and evidently were in pursuit of insects which they were hunting in the very tops of the tall ohias.’

Black Mamo Drepanis funerea Newton PLATE 7 Drepanis funerea Newton 1893. Proc. Zool. Soc. London, p. 690. (Moloka‘i) Drepanorhamphus funereus Rothschild (1893–1900), Perkins (1903),W. A. Bryan (1901a) Drepanidis funerea (Carlquist 1970) Other vernacular names: Mamo, hoa, ‘O‘o-nukuumu, ‘O‘o-nuku-mu

Etymology See Hawai‘i Mamo. Specific epithet funerea suggested by R. C. L. Perkins to the describer (Newton 1893: 690) as an allusion to the bird’s ‘sombre plumage and the sad fate that too probably awaits the species’.

Systematics Monotypic. See discussion of genus, above. Newton (in Rothschild 1893–1900) proposed the monotypic genus Drepanorhamphus for this sp., but relationship to Hawai‘i Mamo obvious and all authors since Bryan and Greenway (1944) consider them congeneric.

Description Large (20 cm) Hawaiian honeycreeper with huge decurved bill (more strongly curved than that of Hawai‘i Mamo). No plumage variation known. Entirely dull sooty black, except for narrow dull white or pale grey outer webs of primaries that produce a pale streak on folded wing. Feathers have soft and velvety surface, unlike glossy surface of

‘I‘iwi Drepanis coccinea 269 Hawai‘i Mamo feathers. Bill and legs black except for dull yellow nasal operculum. For measurements, see Pratt (2002a).

Voice Apparent call note clear querulous whistle ho-a (Bryan 1908). Possible song ‘a rollicking whistle of five or six clear notes’ (Bryan 1908: 153–4) or ‘a loud cry of extraordinary clearness, repeated at short intervals’ differing in degree rather than kind ‘from the more gentle call note’ (Perkins 1903: 402).

Kamakou region, but by 1907 found only further east in upper reaches of Moanui land division (ahupua‘a) where Bryan (1908) collected last specimens. Rumoured to have survived longer on Oloku‘i (Banko 1981). Probably restricted to wet forests.

Diet and foraging Specialised on ornithophilous flowers of oha (Fig. 11.13) and fed secondarily on lehua blossoms (Perkins 1903; Bryan 1908). No insects found in any collected stomachs (Bryan 1908).

Field characters Unlikely to be confused with any other bird on Moloka’i or Maui except Bishop’s ‘O‘o, which has shorter bill, much longer tail, and patches of yellow in plumage.

Distribution, habitat, and population status Extinct. Known historically only from Moloka‘i, but prehistoric remains found on Maui ( James and Olson 1991); probably inhabited all of Maui-nui. R. C. L. Perkins (1903) collected first specimens 1893 in

11.13 Flowers of ‘o¯ha¯ with superimposed head of specimen of Black Mamo, showing close fit of bill and corolla. Photo and montage © Jaan K. Lepson.

‘I‘iwi Drepanis coccinea (Forster) PLATES 7, 8 Certhia coccinea Forster, Goettingisches Magazin Wissenschaften, etc., 1, 1780, p. 347. Drepanis coccinea, G. R. Gray, Gen. B. i. p. 96, (1847); Dole (1869); Sclater (1871); Pratt (1979a) Vestiaria coccinea Wilson and Evans (1890–99), Rothschild (1893–1900), Perkins (1903), Bryan and Greenway (1944), Amadon (1950); Berger (1972a, 1981), Pratt et al. (1987), Sibley and Monroe (1990), James and Olson (1991), AOU (1983, 1998) Other vernacular names: ‘I‘iwi polena and ‘I‘iwi popolo for immature plumage.

Etymology See Hawaii Mamo. Specific epithet a reference to the birds scarlet coloration. Hawaiian name possibly onomatopoeic.

Systematics Monotypic. See discussion of genus, above. Bangs (1911) described the subspecies V. c. suavis from Moloka‘i, long disregarded.

Description A medium-sized (15 cm) honeycreeper with long, thick, sickle-shaped bill. Primary tips broadly

270 ‘I‘iwi Drepanis coccinea truncate. Adults can be sexed on basis of measurements but not plumage (Fancy et al. 1993a). Definitive head and body feathers bright scarlet-vermilion, brilliance enhanced in life by transmitted light; bases of feathers grey, sometimes showing as small grey patch in front of folded wing at sides of breast. Birds may appear lighter on head (Rothschild 1893–1900), result of faded feathers from previous year contrasted with newly moulted ones. Moult begins on belly, ends on the head (Fancy et al. 1993a). Undertail coverts tinged buffy-grey; thighs and narrow eye-ring pale orange, contrasting with surrounding plumage; wings and tail glossy black, greater secondary coverts narrowly edged and tipped scarlet (quickly lost to feather wear); tertials with variable amounts of white or grey, innermost usually all white, next white or grey on inner web only; two innermost greater secondary coverts white edged scarlet, tipped black; wing linings dull white, tinged scarlet. Iris ‘dark hazel’ (Wilson and Evans 1890–99) to medium brown. Bill and legs scarlet-orange, slightly paler than body colour. Juvenal contour feathers between greenish-yellow and mustard yellow, variably tipped black, most prominently on back and top and sides of head; may have scarlet tinges to cheeks, throat, and scapulars. Wings and tail dark grey, wing coverts broadly edged mustard yellow. Outer webs of primaries narrowly edged mustard yellow to dull white. Tertials may have grey inner webs, innermost can be all pale grey. As black tips wear, pattern becomes more diffusely spotted or scalloped and underlying colour fades to tawny yellow. First Basic plumage retains most primaries and secondaries from Juvenal plumage, but body feathers like Definitive plumage with some pale Juvenal feathers intermixed (Fancy et al. 1993a). Bill dull brown at first, gradually lightening through shades of yellow and orange until adult colour reached. Bill colour change not necessarily co-ordinated with moult (pers. obs.). Legs and feet orange-pink, with dusky tinge on toes, gradually lightening as bill. For standard measurements, see Fancy et al. (1993a).

Voice and other sounds ‘An almost infinitely varied repertoire of creaks, whistles, gurgles, and reedy notes’, some resembling

notes played on an old harmonica (Pratt et al. 1987: 309). Other calls reminiscent of sound of a rusty hinge. Also utters human-like, up-slurred whistle easily confused with similar note of ‘Akohekohe (pers. obs.). May imitate other birds, especially Hawai‘i ‘Elepaio, in places (see Ch. 7). Song and call notes essentially undifferentiated. Truncate primaries produce trilling coo in flight indistinguishable from that of ‘Apapane. For recordings see Pratt (1996a); for sonogram see Fancy and Ralph (1998).

Field identification Irregularities of moult can produce birds that appear like adults with yellow heads or juveniles with scarlet vests (pers. obs.). Might possibly be confused with short-billed ‘Akepa (much smaller) or ‘Apapane, but latter is more crimson or blood red, with prominent white under tail, the latter feature especially useful if birds fly overhead.

Distribution, habitat, and population status Map 18. Originally widely distributed in forests of all islands, but now largely restricted to upland ‘ohi‘a and koa forest above the ‘mosquito zone’ (see Ch. 9). Most abundant on Hawai‘i where densities may reach 2000/km2 in good habitat (Scott et al. 1986; Ralph and Fancy 1995). E. Maui populations lower and less dense (Scott et al. 1986); Engilis (1990) estimated 318 ⫾ 135/km2 in Hanawi NAR. Population on Kaua‘i, estimated at ca. 5000 in 1981 (Fancy and Ralph 1998), greatly reduced by Hurricane Iniki in 1992 (Telfer 1993; Pratt 1995), but appears to be recovering (D. Kuhn, pers. comm.).Tiny relict populations exist on O‘ahu (VanderWerf and Rohrer 1996), W. Maui, and Moloka‘i (Scott et al. 1986; Reynolds and Snetsinger 2001). Extirpated on Lana‘i by 1929 (Munro 1960; Hirai 1978).Vagrants reported on Ni‘ihau (F. Sinclair in Wilson and Evans 1890–99) following ‘gales’ on nearby Kaua‘i. Has lowest survivorship of any extant spp. for which data available (see Ch. 8, Table 8.1), perhaps because of effects of avian pox and malaria to which ‘I‘iwi extremely susceptible (Fancy and Ralph 1998;Tummons 2001b).

‘I‘iwi Drepanis coccinea 271

KAUA'I

NI'IHAU

O'AHU MOLOKA'I MAUI _ LANA'I KAHOOLAWE

HAWAI'I

Map 18

Diet and foraging Primarily nectarivorous but also takes invertebrates (summarised by Fancy and Ralph 1998). Main nectar sources ‘ohi‘a and mamane, but also feeds on native koa, naio, kolea, ‘akala, alani, kanawao, kokio ke‘oke‘o, ohelo, and lobelioids Clermontia and Cyanea (Carothers 1982; VanderWerf and Rohwer 1996; Fancy and Ralph 1998), as well as introduced banana poka (Conant et al. 1998; Fancy and Ralph 1998; Fig. 8.12) and tagasaste (Waring et al. 1993). Spieth (1966: 470–1) describes movements of ‘I‘iwi feeding on flowers of C. arborescens: The Iiwi flies to the flower and quickly perches on the flower-bearing twig just basad of the flower pedicel. The bird then quickly swings into an upside-down position under its perch and simultaneously twists its body so that its head is under the open corolla, with the bill pointing upwards. It then easily and with precision slips its slender decurved beak

into the corolla.The nicety of fit between the bird’s head and bill and the fleshy corolla of the flower is indeed striking. However, most recent observations of ‘I‘iwi foraging on falcate corollas have involved nectar-stealing (see Ch. 8; Engilis 1990; Conant et al. 1998). At least half of foraging time spent in ‘ohi‘a and other open flowers (Fancy and Ralph 1998; Fig. 8.9) may represent adaptive shift resulting in evolution of shorter bills (Smith et al. 1995). More efficient feeder on short corollas than other nectarivores with shorter bills (Carothers 1982). Defends feeding territories only during times of intermediate flower abundance (Carpenter and MacMillen 1976a).

Social and interspecific behaviour Feeding territories defended against other nectarivores as well as conspecifics (Carothers 1986a), but aggression reduced by dense canopy foliage

272 ‘I‘iwi Drepanis coccinea (Carothers 1986b). Originally intermediate in feeding hierarchy of Hawaiian nectarivores, now much altered by extinctions. Formerly dominated by meliphagids and mamos, ‘I‘iwi now dominant on all islands except Maui, where ‘Akohekohe takes top position (W. A. Bryan 1908; Pimm and Pimm 1982; Carothers 1986a). May face competition from introduced Japanese White-eye (Mountainspring and Scott 1985). Small numbers may join postbreeding mixed-species foraging flocks on Hawai‘i (see Ch. 7).

Predation Presumably subject to the same native and introduced predators as other Hawaiian birds (Ch. 10), but specific documentation rare. Predation by feral cats and Short-eared Owls (Snetsinger et al. 1994), as well as Hawaiian Hawks ( J. K. Lepson, pers. comm.) reported.

Breeding biology All information from Eddinger’s (1970) study of Kaua‘i birds except as noted. COURTSHIP AND MATING: Monogamous when breeding, but pairs apparently separate after young fledge (Fancy and Ralph 1998). Sexual chasing begins well before nest construction; pair bond maintained by frequent courtship feeding, solicited by female lowering and quivering wings; similar display with a crouch initiates copulation. Courtship display in top of tree involves male singing and slowly swaying back and forth while fluttering wings apparently in the manner of a soliciting female, fluttering of wings while hopping from branch to branch, and short song flights (Berger 1981). NEST SITE AND TERRITORY: In terminal branches or crown of ‘ohi‘a tree, average height 7.2 m.Territory larger than that of most other honeycreepers, defended against conspecifics only. NEST: Both sexes participate in nest building, but female does most. Nests vary in size and composition, but generally have an outer body of ‘ohi‘a

twigs and mosses and inner cup lined with lichens and fibres from leaves or bark. Average (n ⫽ 7) outside diameter 9.5 cm, depth 7.44 cm, cup diameter 5.4 cm, cup depth 3.5 cm. EGGS: First eggs laid in early morning 1–5 d after nest completion, then one each day until clutch complete (1–3 eggs, usually 2). Eggs average 20.7 mm long, 15.5 mm wide, colour white variably spotted, dotted, and blotched with dark chocolate brown mostly around larger end. INCUBATION AND HATCHING: Only female incubates, silently, beginning when clutch complete. Incubation period 14 d. Male feeds female by calling her away with single loud note; does not visit nest. Eggs hatch before noon. NESTLINGS: Altricial young with eyes closed at hatching, bright orange-pink skin tinged yellow, naked except for mingled light (80%) and dark grey down mostly on dorsum, gape rose pink tinged orange on roof of mouth, rictal flanges bright yellow. Eyes open at 5 d. Bill short and straight at first, gradually lengthening and developing curvature. First contour feathers unsheathe on sides of breast at 7 d, wing feathers unsheathed by 15 d. After 12–14 d, young will jump out of nest at slightest disturbance. PARENTAL CARE AND FLEDGING: Only female broods, at night and during heavy rain. Female feeds nestlings (71% of feedings) more than male, but male feeds female. Chicks solicit feeding by fluttering wings.Young fledge at 21–2 d.Young continue to follow one or more parent and beg for up to 4 mo. (Fancy and Ralph 1998).

Life cycle and demography Longevity of wild birds unknown. Breed during first year and annually afterwards. Breeding season very attenuated but probably shorter in any given locality or any particular year, apparently earlier (Feb–June peak) on Hawai‘i, coinciding with seasonal availability of ‘ohi‘a nectar (Baldwin 1953; Ralph and Fancy 1994a). Males with enlarged testes present all year; eggs, young, and

‘Ula-‘ai-hawane Ciridops anna 273 females with brood patches reported Dec–July (Fancy and Ralph 1998). Hatching success (n ⫽ 30) 83.3% (Eddinger 1970). Low adult annual

survivorship of 55% (Ralph and Fancy 1995) possibly biased by high rate of dispersal and low level of philopatry (Fancy and Ralph 1998).

Genus Ciridops Newton Ciridops Newton, 1892, Nature, vol. 45, p. 469. Type, by monotypy, Fringilla anna Dole. Small Hawaiian honeycreepers with short finchlike or tanager-like bills, but skeletal support weak as in other nectarivorous honeycreepers and unlike that of drepanidine or cardueline finches; several features of mandible, including upturned retroarticular process, distinguish Ciridops from closely related genera ( James and Olson 1991). Tongue of tubular drepanidine type (Bock 1972). Hindlimbs

much stouter that those of other honeycreepers ( James and Olson 1991). For historically known spp., sexes alike in plumage, adults with bold patterns of red, black, white, and grey, immatures very drab olive green and buff. Some feathers stiffened along midrib, lanceolate. Distributed throughout main Hawaiian Islands, one sp. on Hawai‘i and Maui-nui, a second on Kaua‘i, and a possible third (undescribed) on O‘ahu ( James and Olson 1991; Pratt 2002a).

‘Ula-‘ai-hawane Ciridops anna (Dole) PLATES 2, 8 Fringilla anna Dole, in Thrum, Hawaiian Almanac, 1879, p. 49. (Hawaii). Other vernacular name:Waaihawane (Bryan and Greenway 1944)

Etymology Latin Ciridops most often translated as ‘shining face’, but ciris also was a bird into which the mythological Scylla was transformed, and ops can mean appearance or aspect ( Jaeger 1955), so the name probably was intended to mean ‘looking like Scylla’s ciris’. Species named for Sanford B. Dole’s wife, Anna. Hawaiian name translates literally ‘red eat palm-nuts’ but a more mellifluous and likely translation would be ‘red palmcreeper’ because we do not know exactly what it ate in the palms and it is a honeycreeper.

Systematics Belongs to the ‘red and black’ group of mainly nectarivorous honeycreepers. Bones of a Ciridops found

at Barber’s Point, O‘ahu, are close to, but shorter than, those of C. anna, but whether they belong to a different sp. has not been determined.

Description Small (11 cm) honeycreeper with short, slightly elongated, finchlike bill. Bony support of bill weak as compared to true Hawaiian finches ( James and Olson 1991). Three distinct plumages represented among five known specimens, only four of which usable for colour comparisons (Olson 1992). Sexes apparently similar. Two similar specimens (Plate 2, 1a) have entire head, throat, and centre of back pale grey darkening in face to black lores; feathers of forehead, throat, and auriculars stiffened distally and paler along the shaft producing scaly effect as in throat feathers of ‘Akohekohe and ‘I‘iwi; broad black breast band extends dorsally to inner scapulars; lateral scapulars, rest of upperparts and underparts, and secondary coverts bright crimson; shoulder, upper primary coverts, primaries, secondaries, inner webs of tertials, and tail black; outer webs of tertials white,

274 ‘Ula-‘ai-hawane Ciridops anna narrowly edged proximally with crimson, forming prominent white patch on folded wing (Pratt 2002a). Another specimen (Plate 2, 1c) may represent First or Second Basic plumage, although no other ‘red and black’ honeycreeper has distinctive subadult plumage (but note differing plumages and sequences of two spp. of Loxops). Cannot be transitional because of feather colours present in neither the above-described plumage nor immature (below). Black of the breast extends forward to the chin, but the black in face less extensive; crimson of lateral scapulars, greater upper secondary coverts, belly and crissum and white in tertials replaced with tawny.This plumage better known because depicted by F. W. Frohawk for Wilson and Evans (1890–99), reprinted in colour in Wilson and Buff (1989), and obviously copied in Munro (1960). Bill and foot colour ‘pinkish brown’ (Wilson and Evans 1890–99). Presumed immature (Plate 2, 1b), either Juvenal or First Basic, mostly yellowish-olive, shading to grey on breast and belly; flanks and head, except nape, tawny; facial feathers stiffened distally as in adult and with buff shaft streaks broadening distally; wings and tail dull black broadly edged yellowish-olive. Only standard measurements available of single specimen in Wilson and Evans (1890–99).

11.14 Flowers and fruits of loulu-lelo, both possible food sources of ‘Ula-‘ai-ha¯ wane.

Field identification Adult combination of red, black, and grey unique among Hawaiian birds, was basis of Munro‘s (1960) ID of bird glimpsed on Kohala Mountain in 1937. Note white wing patch shared with ‘I‘iwi. Immatures much greater field challenge as ‘little green birds’; can easily be confused by the inexperienced with several introduced finches and Red-billed Leiothrix.

all islands of Maui-nui complex. Exact distribution on Hawai‘i unknown, but anecdotal and circumstantial evidence indicates a wide distribution in native wet forests with patchy groves of native loulu palms (Pratt 2002a). Last specimen taken by Hawaiian collectors working for Palmer on Kohala Mountain in 1892. Munro (1960) may have seen one there in 1937.

Distribution, habitat, and population status

Diet and foraging

Extinct. Known historically from Hawai‘i and possibly Moloka‘i (Olson 1992), where prehistoric remains have been found that do not differ significantly from those taken from Hawai‘i specimens ( James and Olson 1991) contrary to preliminary reports (Olson and James 1982b). Likely inhabited

Never observed in life by ornithologists. Name and local folklore imply feeding in some way on palm ‘nuts’ called hawane (Pukui and Elbert 1971). Pritchardia fruits are fairly large (up to 6 cm), globose and fleshy with a hard seed within (Wagner et al. 1990). Because of weak bill (see above), seed probably not main food. Nectar-adapted tongue

Mauka Grosbeak Orthiospiza howarthi 275 possibly used to extract juices from fruits as do some other honeycreepers, to take nectar from Pritchardia flowers, or both (Fig. 11.14; Pratt 2002a).

One preserved stomach held only foliage insects (S. L. Olson in Scott et al. 1986), but nectar would have been difficult to detect.

Kaua‘i Palmcreeper Ciridops tenax James and Olson Ciridops tenax James and Olson, 1991, Ornithological Monographs No. 46, p. 74. (Kaua‘i.)

possibly a ‘ “missing link” between two distinct morphotypes’ ( James and Olson 1991: 77).

Etymology

Description

See ‘Ula-‘ai-hawane (above). Epithet tenax Latin for ‘holding firmly; so named for the muscular leg and large, presumably grasping foot possessed by members of this genus’ ( James and Olson 1991: 74).

Possibly somewhat smaller than ‘Ula-‘ai-hawane ( James and Olson 1991).

Systematics

Extinct. Known only from bones found in lithified sand dunes at Makawehi, Kaua‘i ( James and Olson 1991).

Because of this sp.’s intermediacy of characters between C. anna and other ‘red and black’ dreps,

Distribution and population status

Genera insertae sedis Genus Orthiospiza James and Olson Orthiospiza James and Olson, 1991, Ornithol. Monogr. no. 46: 46. (Maui, Hawaiian Islands.) From James and Olson (1991).Very large Hawaiian finches with arched heavy bills. Narial openings unusually large, bony support of roof of mouth flat between weak ventral ridges, lacking a medial

trough. Interorbital septum and details of wall of cranial cavity of the cardueline/drepanidine type (Zusi 1978). Numerous autapomorphies make the relationships of this distinctive genus unclear. Helen James (pers. comm.) still regards it as drepanidine, but could not determine its position within the subfamily.

Mauka Grosbeak Orthiospiza howarthi James and Olson Orthiospiza howarthi James and Olson, 1991, Orthinol. Monogr. 46: 47. (Lua Manu, Kipahulu Valley, Maui, Hawaiian Islands.)

Etymology Genus name from Greek orthios, high, lofty, + spiza (see Laysan Finch). Species named in honour of

Dr Francis G. Howarth of the BPBM ‘for his studies of the entomology of Hawaiian caves and for his attentiveness to potential fossil deposits in lava tubes’ ( James and Olson 1991: 49). Vernacular name, a well known Hawaiian word for ‘toward the mountains’, chosen in spirit of generic name.‘Grosbeak’ a taxonomically noncommittal name for finches with large bills.

276 Cone-billed Finch Xestospiza conica

Systematics Although two synapomorphies indicate this sp. drepanidine ( James and Olson 1991), it falls outside the group in an overall osteological analysis (James 1998, 2004). Possibly an independent colonization by a cardueline ancestor, but retained as a honeycreeper pending further data.

Description From James and Olson (1991). A huge finch, as large as Greater Koa-Finch, with high-arched bulbous or inflated bill. Nostrils huge compared to those of other honeycreepers.

Distribution, habitat, and population status From James and Olson (1991). Extinct following arrival of humans but before discovery by Europeans. Known from subfossil deposits in three high elevation caves (lava tubes) in the rainforests of Kipahulu Valley, Maui (Fig. 3.3). Not represented in cave deposits on leeward slope of Haleakala, so possibly restricted to montane wet forests. Two of three specimens are juveniles that entered the caves unaided, perhaps reflecting some species-specific foraging or breeding behaviour.

Genus Xestospiza James and Olson Xestospiza James and Olson, 1991, Ornithol. Monogr. No. 46, p. 52. Type, by original designation, Xestospiza fastigialis James and Olson. From James and Olson (1991). Similar in many respects to Melamprosops but characters apparently symplesiomorphies. Differs from that genus mostly in

quantitative rather than qualitative comparisons. Bill cone-shaped with flat or nearly flat dorsal profile. A recent phylogenetic analysis that includes newly discovered subfossils from Maha‘aulepu, Kaua‘i, suggests that this genus may not be monophyletic (H. James, pers. comm.), hence uncertain placement herein.

Cone-billed Finch Xestospiza conica James and Olson Xestospiza conica James and Olson, 1991, Ornithol. Monogr. No. 46, p. 53. (Kauai.)

Etymology From James and Olson (1991: 52–3). Xestospiza from Greek, xestos ⫽ planed or scraped plus spiza ⫽ a finch, from the dorsal profile of the maxilla, which is flattened as if planed; epithet conica ‘Latinized from Greek, konikos, conelike; from the shape of the maxilla’. English name from Olson and James (1982b: 40). Ziegler (2002) refers to members of this genus as conebills, but that name should not be used because of confusion with tanagers of the genus Conirostrum, also called conebills.

Systematics H. James’s (pers. comm.) most recent phylogenetic analysis fails to support the monophyly of

Xestospiza and places this sp. as sister to Chloridops regiskongi, and in turn closely related to Rhodacanthis rather than to X. fastigialis.

Description Known only from the holotype and paratype, a pair of maxillae. Slightly larger than X. fastigialis and differing in osteological details. Cone-shaped bill with dorsal profile slightly curved, not as flattened as that of X. fastigialis ( James and Olson 1991).

Distribution and status Extinct. Known only from prehistoric bones found in lithified dunes at Makawehi, Kaua‘i ( James and Olson 1991).

Ridge-billed Finch Xestospiza fastigialis 277

Ridge-billed Finch Xestospiza fastigialis James and Olson Xestospiza fastigialis James and Olson, 1991, Ornithol. Monogr. No. 46, p. 55. (Maui.)

Etymology See Cone-billed Finch. Epithet fastigialis based on Latin for ‘a slope to a point’, or ‘a gable’ in reference to the bill ridges ( James and Olson 1991). English name from Olson and James (1982b: 40).

Systematics James and Olson (1991) discuss the superficial similarities between this sp. and cowbirds (Molothrus), and show that the cranial fenestra and interorbital septum are of the cardueline/drepanidine type (Zusi 1978) and unlike those of icterids. See Conebilled Finch.

Description From James and Olson (1991: 57). A finch about same size as Nihoa Finch, with cone-shaped pointed bill resembling that of a cowbird in profile. Dorsal profile line nearly flat, with a bulge at the apex above the nostrils marked by ‘two converging rugose

ridges’. Between the ridges the surface is flat rather than convex as in most other drepanidines. ‘The mandible lacks retroarticular processes’ and is generally similar to that of other drepanidine finches.

Distribution and status Extinct. Found in subfossil deposits at Barber’s Point and ‘Ulupau Head, Oahu; Mo‘omomi Dunes and ‘Ilio Point, Moloka‘i; and Pu‘u Naio Cave, Lua Lepo, and Crystal Cave, Maui; the widest distribution of any of the recently described finches ( James and Olson 1991). From the abundance of remains, apparently once common. 1. Son of missionaries, in 1884 elected to Hawaiian legislature.Active in 1887 movement to force democratic reforms upon Hawaiian monarchy, served as justice of Supreme Court (1887–93). President of Republic of Hawaii (1893–1900) following overthrow of Queen Liliuokalani. Governor of Territory of Hawaii (1900–03) following US annexation, federal district judge (1903–15). From Allen (1988) and Mearns and Mearns (1992).

Appendix 1: Honeycreepers in Hawaiian material culture Sheila Conant The feathers of birds were the most valued possessions of the ancient Hawaiians.

David Malo (1951: 76)

and Rose et al. (1993) provided detailed analyses of the cultural significance, construction, and composition of particular groups of objects.

Introduction

Identification of feathers

The feathers of birds have been used by many cultures to create objects of ornamentation. Often such objects served as indicators of wealth and social status, as was certainly the case in Hawaiian culture. Although the feathers of many species of birds were used in Hawaiian artefacts, objects made with feathers of honeycreepers appear to have been made for, and used primarily by, the chiefly class or ‘ali‘i (Malo 1951; Buck 1957; Kaeppler 1985; Rose et al. 1993). Among the objects collected on the voyages of Cook and Vancouver are various objects featuring honeycreeper feathers: capes, cloaks, helmets, religious images (e.g., of human heads), ka¯ hili (feathered standards), lei (necklaces), and feather aprons.These first collections of feather artefacts tell us much of what we know about how the feathers of birds, including the honeycreepers, were used in the ceremonial artefacts of war, religion, and other forms of social intercourse in early Hawai‘i. William T. Brigham (1899, 1903), of the Bernice P. Bishop Museum (BPBM), published two treatises on Hawaiian feather work at the turn of the century, followed by a supplement (Brigham 1918). These works constituted the most extensive treatment of feather artefacts and are still important today. However, Malo (1951), Buck (1957), and Force and Force (1968) also provided general treatments of feather work in their papers on material culture, while Kaeppler (1970, 1985), Rose (1978, 1979),

Although earlier works on Hawaiian feather artefacts (see above) usually identified the origin of the feathers in various objects, these identifications were not always correct, and so, for my own descriptions of feather work, I have relied strictly on my own identifications. During the last 25 years, I have examined dozens of Hawaiian feather artefacts, and had the opportunity to study the structure of these objects, to estimate the numbers of feathers used in individual objects, and to identify the birds from which feathers were taken. Feather identification, particularly of honeycreeper feathers, has turned out to be relatively simple because so few species were used and the colours of the feathers are quite distinctive. Among the honeycreepers, only a few species have red feathers, and those of the ‘I‘iwi are a very distinct vermilion, while those of the ‘Apapane are a deep scarlet. Of all the objects I have examined, none have contained red feathers that I have judged to belong to any other honeycreeper species, and the vast majority of red feathers were from ‘I‘iwi. The yellow feathers in Hawaiian artefacts can be distinguished by both colour and structure. The yellow feathers of the Hawai‘i Mamo, which are uppertail and undertail coverts (ko‘omamo), were an orangish-yellow, and the yellow feathers of the meliphagid ‘o¯‘o¯s a lemon yellow. The yellow axillary feathers (‘e‘e) of Moho nobilis had a short rachis and long vane, giving them

Appendix 1: Honeycreepers in Hawaiian material culture 279 a fluffy pompom-like appearance, while the rachis of mamo feathers extended to the distal end of the feather.The undertail coverts (pue) of the ‘o¯‘o¯s also yield some yellow feathers. Although both Hawai‘i ¯ ‘o¯ and Hawai‘i Mamo feathers were used, the ‘O ¯ ‘o¯ are much yellow axillary feathers of Hawai‘i ‘O more common in feather work than are ko‘omamo (yellow mamo feathers). Fading, wear, and ageing affect these attributes to some extent, but the feather structure usually remains sufficiently intact to allow identification.

Purpose of feather collecting Feathers were a form of taxes paid to the ‘ali‘i or chiefs in Hawaiian society, particularly during the makahiki, an annual festival featuring sports and religious ceremonies during which war was kapu or forbidden. The more powerful the chief, the more feathers he could command.The dominance of red feathers in religious objects, such as images of gods, suggests that red is associated with sacredness.Yellow feathers, the rarest and most difficult to obtain, apparently signified political power (Kaeppler 1985).

Pre-contact artefacts Captain James Cook made his first landing at Waimea Bay, Kaua‘i on 18 January 1778. Although his ships remained only 2 days, Cook and HMS Resolution’s surgeon (William Anderson) and artist ( John Webber) hiked onshore, and Anderson, who noted the Hawaiians’ feather capes and cloaks, recorded his observations of ‘I‘iwi brought for sale by the Hawaiians (Cook and King 1784, v. 2: 207–8): We were at a loss to guess from whence they could get such a quantity of these beautiful feathers: but were soon informed, as to one sort; for they afterward brought great numbers of skins of small red birds for sale, which were often tied up in bunches of twenty or more, or had a small wooden skewer run through their nostrils. At the first, those that were bought, consisted only of the skin from

behind the wings forward; but we, afterward, got many with the hind part, including the tail and feet. The red bird of our island was judged by Mr. Anderson to be about the size of a sparrow; of a beautiful scarlet colour, with a black tail and wings; and an arched bill, twice the length of the head, which, with the feet, was also of a reddish colour. In 1978, Adrienne Kaeppler, after years of anthropological detective work, assembled a collection of material culture objects, or ‘artificial curiosities’ as they were called, collected on the three Pacific voyages of Cook.This collection, which was exhibited at the BPBM, included objects from cultures throughout the Pacific Basin, ranging from the Pacific Coast of North America through Hawai‘i to the far-flung islands of the western Pacific. The objects exhibited are described and pictured in Kaeppler’s (1978) catalogue of the exhibition. Numerous artefacts featured feathers, often as the most obvious feature, and it is this collection of feather artefacts that has told us most of what we know about how Hawaiians integrated feathers into their material culture before any influence by Europeans. I examined most of these Hawaiian feather artefacts and was able to identify many of the birds from which the feathers came.The feathers of Hawaiian honeycreepers dominate among the objects, particularly those artefacts that appear to be of greatest cultural significance (e.g., idols, cloaks, and ka¯ hili ). Plate 9 depicts several examples of such artefacts.

Cloaks and capes Surely the most famous of the feathered objects in Hawaiian material culture are the spectacular feather capes, ‘ahu‘ula (⫽ red shoulder garment), which were large enough to drape in graceful and abundant folds from the shoulders to the ankles of ‘ali‘i, who often stood over 6 feet tall.These objects were worn only by the highest-ranking male ‘ali‘i and were symbols of power and status. Although women may have prepared feathers for use, ‘ahu‘ula were constructed only by high-ranking men (Kaeppler 1970). According to Kaeppler, these

280 Appendix 1: Honeycreepers in Hawaiian material culture cloaks were inherited from one chief to another, worn only on important occasions or in battle, and were taken as battle prizes by the conqueror. Judging from the early collections, most of the cloaks made before western contact contained primarily the red feathers of the ‘I‘iwi and yellow feathers ¯ ‘o¯. from either the Hawai‘i Mamo or Hawai‘i ‘O Kaeppler (1970) estimated that a single cloak could contain as many as a half million feathers.Although we will never be sure of the exact number, Kaeppler speculates that only about seven, the number illustrated by Sarah Stone (Force and Force 1968), ‘ahu‘ula were known to have been collected during Cook’s voyages, testament, perhaps, to their rarity and importance. Among the objects collected by Cook were about 20 capes (short cloaks draping only to about the waist). The feather capes show much greater variation than do the cloaks in terms of the types of feathers used in construction. Red feathers are almost invariably those of the ‘I‘iwi, but there is one cape (Kaeppler 1978, fig. 67), collected on Kaua‘i, the neckline of which is edged with patches of bird skin with the feathers still attached! Some of the patches of skin are ‘Apapane, others are ‘I‘iwi, and the patches of skin with yellow feathers are proba¯ ‘o¯‘a¯ ‘a¯ .The main body of bly thigh feathers of the ‘O the cape is decorated with chicken (⫽Red Junglefowl) feathers, and possibly some seabird feathers (e.g., Great Frigatebird and Red- or White-tailed tropicbirds). Many of the capes contain red ‘I‘iwi feathers and yellow feathers from either the Hawai‘i Mamo or one of the ‘o¯‘o¯ species, usually Moho nobilis from the Island of Hawai‘i. One very interesting cape (Kaeppler 1978, fig. 66) was edged on the top and sides with red (‘I‘iwi) and yellow ¯ ‘o¯) triangles, and with a row of mamo (Hawai‘i ‘O feathers at the bottom outer edges.The central part of the cape is ornamented with the red central rectrices of the Red-tailed Tropicbird, interspersed with some neck and back feathers of male frigatebirds. The tropicbird and frigatebird feathers have all been split in half down the central axis of the feather, which has the effect of making the feather spiral or curl. Besides being indicators of social status, anthropologists have speculated that cloaks and capes

worn during battle could have acted as shields, preventing or mitigating injuries from the clubs, stones and spears used in war. The heavy woven netting to which feathers were tied were made from the fibres of olona¯ , an endemic Hawaiian plant, made more available through intentional cultivation. These fibres are among the strongest plant fibres known (Wagner et al. 1990), and were used to make ropes prized by mountain climbers in the nineteenth and twentieth centuries before the advent of synthetics.

Feathered helmets High-ranking ‘ali‘i often wore mahiole, feathered helmets, during battle or important ceremonies. These objects varied tremendously in shape and style, but were constructed of olona¯ netting ornamented most often with red, yellow, and black feathers, although a small number feature green feathers. Again, the red feathers on mahiole I have examined are invariably from ‘I‘iwi, yellows from both mamo and ‘o¯‘o¯, and black feathers most likely (based on their colour and structure) from ‘o¯‘o¯, usually M. nobilis. These helmets, like the cloaks and capes, signified power and prestige, and were said to be worn at occasions of state and during battle. No doubt they enhanced status because of their value and because they increased physical stature by adding to height. However, they could also have afforded protection from head injuries during combat. Kaeppler (1985) hypothesises that ‘ahu‘ula and mahiole were worn exclusively in sacred or dangerous situations and in battle, and that these objects lent both physical and sacred protection. Holt (1985) includes photographs of one post-contact mahiole (p. 97) and one pre-contact mahiole (p. 99).

Ka¯hili Another type of artefact associated with the ‘ali‘i were ka¯hili, feathered standards.These were of generally two sizes, tall (ka¯hili ku¯) objects, as much as 20 feet in height, and shorter ones that could be held in the hand.The use and significance of these objects is not well understood because so few of them survive

Appendix 1: Honeycreepers in Hawaiian material culture 281

A1.1 A collection of standing kahili being curated at BPBM. Many of these contain small numbers of honeycreeper feathers.

from Cook’s first voyages, and early explorers took few notes on their use and appearance (Rose 1979; Rose et al. 1993). A passage by William Ellis (Ellis 1782: II: 156), third voyage, provides this description: They have also a kind of fly-flap, made of a bunch of feathers fixed to the end of a thin piece of smooth and polished wood: they are generally made of the tail feathers of the cock, but the better sort of people have them of the tropick birds feathers, or those belonging to a black and yellow bird called mo-ho. The handle is very frequently made of one of the bones of the arm or leg of those whom they have killed in battle, curiously inlaid with tortoise-shell: these they deem very valuable, and will not part with them under a great price.This ornament is common to the superiors of both sexes.

Ellis’s description does not do justice to the variety of forms of ka¯ hili, but he was correct in observing that the presence of human arm or leg bones in such objects was of great significance to their owners. Hawaiians believe that the arm and leg bones are places where mana or spiritual power resides, and objects containing these bones were of great spiritual and political importance. Although ka¯ hili were probably used to keep away insects and cool the ‘ali‘i, they also marked the presence of chiefly individuals and were believed to provide a kind of spiritual protection (Rose et al. 1993: 274). Rose et al. (1993) provide a detailed analysis of 75 ka¯ hili ku¯ in the collections of the BPBM. However, their sample contained only two objects collected from Cook’s voyages. Thus, their sample of ka¯ hili reflects materials and construction techniques typical of the nineteenth and early twentieth centuries. The ka¯ hili collected on Cook’s voyages and exhibited at Bishop Museum contained few honeycreeper feathers. One of two ka¯ hili ku¯ from these collections contained neck and back feathers of male Great Frigatebirds and flank feathers of Red-tailed Tropicbirds. The second (Kaeppler 1978, fig. 99) was ornamented with central tail feathers of Red-tailed Tropicbirds, neck feathers of roosters, and a small number of yellow mamo feathers.The kumu or handles of these ka¯ hili ku¯ consisted of central wooden shafts encircled by cylinders of human bone interspersed with rings of ‘ea or hawksbill turtle Eretmochelys imbricata shell, each ring made from a single scale. Other ka¯ hili collected on Cook’s voyages (I examined three) were made with the flight feathers, especially the two central tail feathers, of ‘o¯‘o¯, probably M. nobilis. It is interesting to note that these feathers were split in half all the way down the central shaft, which curls the feather. This technique of feather splitting was also used in one of the feather capes mentioned above.

Feather images A small number of akua hulu manu, or religious idols, were collected on Cook’s voyages, and these are thought to be images of various gods. They are constructed on a base of woven ‘ie‘ie aerial

282 Appendix 1: Honeycreepers in Hawaiian material culture roots covered with a tight fitting net of olona¯. Most are covered primarily with red ‘I‘iwi feathers, sometimes underlain with yellow or black feathers, which give depth and texture to the appearance of the object or serve to demarcate facial features. One object (Kaeppler 1978, figs 56 and 57) has a small patch of ‘Apapane feathers at the centre of the forehead and a few more scattered throughout. This image’s eyebrows consist of black feathers that I was unable to identify. The eyes of these images are made with pearl oyster shells and carved spheres of wood representing pupils. The teeth are from domestic dogs Canis familiaris. A few are crowned with human hair. Most have rather frightful expressions. Perhaps I should include in this category a very unusual object regarded by many to have sacred ceremonial significance: the feathered temple shown in Kaeppler’s (1978) fig. 60. The framework of this object is of ‘ie‘ie aerial roots, onto which red ‘I‘iwi feathers have been lashed. The edges of the temple are of yellow feathers from ¯ ‘o¯. The ‘door’ of the both mamo and Hawai’i ‘O temple is bordered by hawksbill turtle shell. The temple resembles a house and may have signified the dwelling place of a god or gods.

Feather lei Many lei (feathered ornaments for head or neck) were collected on Cook’s voyages of discovery, but few survive that can be definitely attributed to these collections. The feather lei I examined from Cook’s voyages were all made using feathers of Hawaiian honeycreepers. Solid-coloured lei, usually yellow or red, were less common than those made in alternating bands of colour. The principle colours used in lei were yellow and red, followed by black and green. It is among the feather lei that we most frequently find green feathers, and to date I have been unable to identify the honeycreepers from which these originated. H. D. Pratt (pers. comm.) considers the colour of some of these ¯ ‘u¯ feathers. green feathers close to that of some ‘O The use of modern molecular techniques could possibly solve these identification problems.The red feathers, as one might expect, are mostly ‘I‘iwi feathers, with an occasional ‘Apapane here and

¯ ‘o¯ and Hawai‘i there. The yellows are Hawai‘i ‘O Mamo. Many pre- and post-contact lei are pictured in Holt (1985: 48-63).

Other objects A number of other feather artefacts were collected on Cook’s voyages.These include feathered helmet bands, which are similar to lei, except that some were very thick, being made of feathers lashed to a net backing of ‘ie‘ie or olona¯. Hawaiians made a number of kinds of musical instruments, among which were ipu or gourd rattles used by dancers of hula.The handles of ipu were decorated with feathers from a variety of birds, including some honeycreepers. A few feathered aprons (Brigham called them ‘mats’) were collected during the Cook voyages (Rose 1978). Little is known of the function of these objects, though Kaeppler (1978), noting that the aprons have ‘loops and lacings reminiscent of Hawaiian ankle ornaments’, speculates that they may have been worn across the chest or stomach for protection during battle.

Post-contact change in feather artefacts Among the Polynesian civilisations, Hawai‘i’s social organisation was unique, and arguably the most complex ( Jennings 1979; Kirch 1985). Based upon archaeological, linguistic, and other evidence, anthropologists have described social and political change in Hawai‘i in considerable detail.They have determined that, at the time of Western contact, divisions of land and designations of land tenure in Hawai‘i were established by the ‘ali‘i. In other Polynesian societies, and presumably early Hawaiian society, rights to land derived from one’s genealogy. By the time Cook arrived, Hawai‘i had developed a stratified class society, and only the ‘ali‘i maintained genealogies (Kirch 1985). Kaeppler (1985) points out that ‘like other Polynesian societies, Hawai‘i had special forms of material culture that were used specifically by those of rank.’ Furthermore, she suggests, changes in forms of material culture accompanied changes in Hawaiian social relationships and organisation. One of the most

Appendix 1: Honeycreepers in Hawaiian material culture 283 dramatic, and certainly the most studied, periods of social and cultural change in Hawai‘i took place during the late eighteenth and early nineteenth centuries, with the early influence of continental cultures. Kaeppler (1985) has examined how Hawaiian feather artefacts reflect post-contact transformations in Hawaiian culture. Of interest to students of Hawaiian honeycreepers are the changes in form and colour of artefacts, particularly ‘ahu‘ula and ka¯ hili, large objects requiring many thousands of feathers for construction. Of the capes and cloaks assembled and catalogued by Kaeppler (1978) for the ‘Artificial Curiosities’ exhibit at BPBM, eight are pictured in the catalogue. Of these, only one is a cloak (Kaeppler 1978, fig. 65), the rest are capes. The cloak is red (‘I‘iwi) and yellow (‘o¯‘o¯ and mamo). Of the seven capes pictured, two (figs 71 and 74) are made entirely from honeycreeper and honeyeater feathers; the remainder consists largely of feathers from chickens and various seabirds. Of the 30 or so cloaks and capes known to have been collected on Cook’s voyages, at least 20 are capes.The cloaks, capes, and other objects ornamented entirely or mostly with feathers of honeycreepers and honeyeaters were possessed only by the most powerful ‘ali‘i, and were probably few in number (Kaeppler 1985). Changes in form (shape, colour) and function (sacred vs. political significance) of these objects began immediately after western contact. Feather artefacts, especially cloaks and capes, were given by the Hawaiians to Western explorers and their colleagues in great numbers. Based on our understanding of the cultural importance of these objects, we may conclude that with these gifts the Hawaiians expressed respect and perhaps even reverence for the recipients. History also suggests that, although they may not have appreciated the sacred and political importance of some feather artefacts, Westerners appreciated them because of their great beauty, craftsmanship, and the investment of effort they represented, including that required to catch the birds and fashion the objects. Although before Western contact feather objects of the kind possessed only by ‘ali‘i were relatively rare, in the late 1700s and early 1800s, many such objects were constructed and given as gifts. For Hawaiian hon-

eycreepers, this meant their feathers came into greater and greater demand. Coupled with the advent of firearms in Hawaiian culture, this must have tremendously increased the numbers of birds taken. Thus, the question arises, what role did Hawaiian feather work play in the decline and extinction of Hawaiian honeycreepers? The answer is not a simple one.

Impacts of feather collecting on Hawaiian honeycreepers How many feathers does it take to make an ‘ahu‘ula? I once obtained permission from the BPBM to closely examine the famous Ka¯mehameha cloak (Holt 1985: 130, 131), which was made entirely from yellow mamo feathers. My approach was to count the number of feathers in several areas of known size, and then, using the measurements of the size of the entire cloak, to extrapolate the number of feathers used. Next, I examined several mamo specimens to find out how many feathers each bird could yield. Because the yellow feathers used in the cloak were only undertail and uppertail coverts (ko‘omamo), this was relatively simple.A single Hawai’i Mamo can yield one to two dozen such feathers, depending on the stage of moult of the individual bird. In the end, I estimated that approximately 80 000 birds had to be captured to produce enough feathers for the Ka¯mehameha cloak. According to David Malo (1951: 77), ‘An ahu-ula made only of mamo feathers was called an alaneo and was reserved exclusively for the king of a whole island, alii ai moku; it was his kapa wai-kaua or battle cloak.’ Whether birds were killed after they were captured for their feathers is a controversial question. Catching honeycreepers in the first place must have been exceedingly difficult. Handy et al. (1972) state that birds were caught by smearing sticks with the sticky sap of the breadfruit and placing the sticks in trees where the birds were expected to perch.This was surely an ingenious technique, but one that would require knowledge of the birds’ habits and great patience as well. According to Malo (1951: 38), all of the honeycreepers whose feathers we know were used in feather artefacts were eaten by

284 Appendix 1: Honeycreepers in Hawaiian material culture the Hawaiians. He claims their flesh was ‘good eating’ or ‘delicious’. Nevertheless, the notion persists that feather collectors practised conservation by removing only a few feathers from each bird and then releasing it.The truth, I suggest, is somewhere in between.The observations of William Anderson quoted above indicate that these birds were killed in numbers. It would have been difficult indeed to resist the temptation to eat birds in an environment that was relatively protein poor. Additionally, unless the bird catcher was able to remove the very sticky breadfruit sap from the bird’s feet and feathers, there is little chance it could have survived. Rose et al. (1993: 298) also discuss this issue in some detail. If indeed many of the honeycreepers caught by pre-contact Hawaiians were killed, to what degree did feather collecting before 1778 contribute to the decline and extinction of these birds? Based on the distribution of honeycreepers at the time of contact and the data provided by records of fossil honeycreepers, I believe that pre-contact Hawaiians contributed in only a minor way to the decline of

the honeycreepers whose feathers they treasured. First, the songbirds whose feathers were used in greatest numbers (‘I‘iwi, mamo, and the ‘o¯‘o¯s) were still present in large numbers at the time of Western contact. It is my view that the impact of pre-contact feather collecting by Hawaiians was unlikely to have caused the extinction of any species of forest songbird or even contributed significantly to their declines.The addition of firearms to the toolbox of the feather collectors, combined with the greater demands for red and yellow feathers of certain forest birds, may have accelerated the late nineteenth-century declines of those species already hard hit by disease, predation, and habitat change. Though pre-contact Hawaiians may have contributed significantly to the decline of numerous seabird species and caused the extinction of many flightless land birds and some of the forest birds, these impacts were associated with the search for high protein foods, the development of agriculture, and the management of habitat for human use, not the acquisition of feathers.

Appendix 2: Scientific names and families of plants mentioned in the text ‘a‘ali‘i ‘a‘e ‘a¯kala ‘a¯kia ‘a¯koko ‘akulikuli ‘aheahea ‘a¯la‘a alani alena ‘aweoweo bananap banana pokaa beach naupaka black acaciaa blackberrya blue guma Brazilian peppera breadfruitp button sedge Christmasberrya coconutp eucalyptusa firetreea Formosan koaa fuchsiaa garden nasturtiuma geranium golden shower treea gorsea guavaa ha¯ha¯ ha¯ha¯ (Fig. 8.11) ha¯ha¯‘aiakamanu ha‘iwale

Dodonaea viscosa Sapindus saponaria Rubus hawaiensis Wikstroemia spp. Euphorbia lorifolia Sesuvium portulacastrum Chenopodium oahuense Pouteria sandwicensis Melicope spp. Boerhavia repens Chenopodium oahuense Musa spp. Passiflora molissima Scaevola sericea Acacia melanoxylon Rubus argutus Eucalyptus globulus Schinus terebinthefolius Artocarpus exilis Fimbristylis cymosa Schinus terebinthefolius Cocos nucifera Eucalyptus spp. Myrica faya Acacia confusa Fuchsia magellanica Tropaeolum majus Geranium Cassia fistula Ulex europea Psidium guajava Clermontia or Cyanea spp. Cyanea angustifolia Clermontia fauriei Cyrtandra sp.

Sapindaceae Sapindaceae Rosaceae Thymeliaceae Euphorbiaceae Aizoaceae Chenopodiaceae Sapotaceae Rutaceae Nyctaginaceae Chenopodiaceae Musaceae Passifloraceae Goodeniaceae Fabaceae Rosaceae Myrtaceae Anacardiaceae Moraceae Cyperaceae Anacardiaceae Arecaceae Myrtaceae Myricaceae Fabaceae Onagraceae Tropaeolaceae Geraniaceae Fabaceae Fabaceae Myrtaceae Campanulaceae Campanulaceae Campanulaceae Gesneriaceae

286 Appendix 2: Scientific names and families of plants halapepe hau hele ‘ula hau kuahiwi Hawaiian olive Hawaiian persimmon Hawaiian raspberry Hawaiian vetch ha¯wane haup? hapu‘u hinahina kahakai ho‘awa hoi kuahiwi huakehili ‘uka hulumoa ‘ie‘ie ihi ‘ili‘ahi ‘ilima ironwooda kahili gingera ka¯lia kanawao Kaua‘i koli‘i kauila ka¯wa‘u kawelu kipukai koa kokio kea kokio ke‘oke‘o ko¯lea lau nui (or simply ko¯lea) kopiko Koster’s cursea kuku‘i (candlenut)p kupala lama lapalapa lehua lehua mamo loulu loulu-lelo lychee maiapilo maile makaloa ma¯mane

Pleomele aurea Kokia drynarioides Hibiscadelphus giffardianus Nestegis sandwicensis Diospyros sandwicensis Rubus hawaiensis Vicia menziesii Pritchardia spp. (fruit) Hibiscus tiliaceus Cibotium spp. Nama sandwicensis Pittosporum spp. Smilax melastomifolia Scaevola kilaueae Korthalsella complanata Freycinetia arborea Portulaca lutea Santalum freycinetianum Sida fallax Casuarina equisetifolia Hedychium gardnerianum Elaeocarpus bifidus Broussaisia arguta Trematolobelia kauaiensis Alphitonia ponderosa Ilex anomala Eragrostis variabilis Heliotropium curassavicum Acacia koa Hibiscus waimeae Hibiscus arnottianus Myrsine lessertiana Psychotria spp. Clidemia hirta Aleurites moluccana Sicyos pachycarpus Diospyros sandwicensis Cheirodendron platyphyllum Metrosideros polymorpha (flowers) Metrosideros macropus Pritchardia spp. (tree) Pritchardia hillebrandii Litchi chinensis Capparis sandwichiana Alyxia oliviformis Cyperus laevigatus Sophora chrysophylla

Agavaceae Malvaceae Malvaceae Oleaceae Ebenaceae Rosaceae Fabaceae Arecaceae Malvaceae Dicksoniaceae Hydrophyllaceae Pittosporaceae Liliaceae Goodeniaceae Viscaceae Pandanaceae Portulacaceae Santalaceae Malvaceae Casuarinaceae Zingiberaceae Elaeocarpaceae Hydrangeaceae Campanulaceae Rhamnaceae Aquifoliaceae Poaceae Boraginaceae Fabaceae Malvaceae Malvaceae Myrsinaceae Rubiaceae Melastomaceae Euphorbiaceae Cucurbitaceae Ebenaceae Araliaceae Myrtaceae Myrtaceae Arecaceae Arecaceae Sapindaceae Capparaceae Apocynaceae Cyperaceae Fabaceae

Appendix 2: Scientific names and families of plants 287 manele manono Methley pluma miconiaa mountain apple (‘o¯hi‘a‘ai)p mountain sandalwood naio naupaka kahakai naupaka kuahiwi nohoanu nohu (puncture vine) nuku ‘i‘iwi ‘o¯ha¯ (Fig. 8.11) ‘o¯ha¯ ke¯ pau ‘o¯ha¯ (Species accounts, Fig. 8) ‘ohe (or ‘ohe‘ohe) ‘ohe naupaka ‘o¯helo ‘ai (or just ‘o¯helo) ‘o¯helo kau la‘au (or tree ‘o¯helo) ‘o¯hi‘a-ha¯ ‘o¯hi‘a-lehua or ‘o¯hi‘a o¯lapa olomea olona¯ olopua o¯puhe pa‘iniu pa¯pala paperbarka peacha pigweed pili (grass) pilo pilo (Maui) pinea pohaa po¯huehue (seaside morning-glory) puaokama pukiawe red sandalwood sandalwood sandbura sea purslane scheffleraa silk oaka slash pinea soapberry

Sapindus saponaria Hedyotis terminalis Prunus cerasifera ⫻ salicina Miconia calvescens Syzygium malaccense Santalum paniculatum Myoporum sandwicense Scaevola sericea Scaevola gaudichaudiana Geranium arboreum Tribulus cistoides Strongylodon ruber Clermontia clermontioides Clermontia hawaiiensis Clermontia grandiflora Tetraplasandra spp. Scaevola glabra Vaccinium dentatum Vaccinium calycinum Syzygium sandwicensis Metrosideros polymorpha Cheirodendron trigynum Perottetia sandwicensis Touchardia latifolia Nestegis sandwicensis Urera glabra Astelia spp. Charpentiera spp. Melaleuca quinquenervia Prunus persica Portulaca spp. Heteropogon contortus Coprosma spp. Coprosma ochracea Pinus spp. Physalis peruviana Ipomoea pescaprae Sicyos maximowiczii Styphelia tameiameae Santalum haleakalae Santalum spp. Cenchrus echinatus Sesuvium portulacastrum Brassaia actinophylla Grevillea robusta Pinus elliottii Sapindus saponaria

Sapindaceae Rubiaceae Rosaceae Melastomaceae Myrtaceae Santalaceae Myoporaceae Goodeniaceae Goodeniaceae Geraniaceae Zygophyllaceae Fabaceae Campanulaceae Campanulaceae Campanulaceae Araliaceae Goodeniaceae Ericaceae Ericaceae Myrtaceae Myrtaceae Araliaceae Celastraceae Urticaceae Oleaceae Urticaceae Liliaceae Amaranthaceae Myrtaceae Rosaceae Portulaccaceae Poaceae Rubiaceae Rubiaceae Pinaceae Solanaceae Convolvulaceae Cucurbitaceae Epacridaceae Santalaceae Santalaceae Poaceae Aizoaceae Araliaceae Proteaceae Pinaceae Sapindaceae

288 Appendix 2: Scientific names and families of plants strawberry guavaa sugia swamp mahoganya tagasastea tobaccoa tree heliotropea tree ohelo ‘uluhe (fern) wahine-noho-mauna wauke (paper mulberry)p wiliwili wiliwili haolea aAlien

introduced or cultivated species. Polynesian introduction.

pAboriginal

Psidium cattleianum Cryptomeria japonica Eucalyptus robusta Cytisus palmensis Nicotiana tabacum Tournefortia argentea Vaccinium calycinum Dicranopteris linearis Adenophorus tamariscinus Broussonetia papyrifera Erythrina sandwicensis Erythrina variegata

Myrtaceae Taxodiaceae Myrtaceae Fabaceae Solanaceae Boraginaceae Ericaceae Gleicheniaceae Grammitaceae Moraceae Fabaceae Fabaceae

Appendix 3: Scientific names, families, and subfamilies of non-Hawaiian birds mentioned in the text (excluding those in Tables 2.2 and 2.3) Auk, Great Bananaquit Bellbird, New Zealand berrypeckers Brambling Bunting,Tristan Bunting,Wilkins’s Canary, Island Chaffinch Chat,Yellow-breasted cowbirds Crossbill, Red crossbills Dove,Whistling Finch, Cocos Finch,Warbler flickers flowerpeckers Flycatcher, Galaápagos Flycatcher, Guam Flycatcher,Vermilion Frigatebird, Great fruit-doves Goldfinch, American Hawfinch holarctic creepers Hoopoe Huia Hummingbird, Broad-tailed Junglefowl, Red (chicken) Kite, Snail longbills lorikeets, brush-tongued Martin, Southern Mauritius Kestrel

Pinguinus impennis Coereba flaveola Anthornis melanura Melanocharis spp. Fringilla montifringilla Nesospiza acunhae Nesospiza wilkinsi Serinus canaria Fringilla coelebs Icteria virens Molothrus spp. Loxia curvirostra Loxia spp. P. layardi Pinaroloxias inornata Certhidea olivacea Colaptes spp. Dicaeum spp. Myiarchus magnirostris Myiagra freycineti Pyrocephala rubinus Fregata minor Ptilinopus spp. Carduelis tristis Coccothraustes coccothraustes Certhia spp. Upupa epops Heteralocha acutirostris Selasphorus platycercus Gallus gallus Rostrhamus sociabilis Toxorhamphus spp. Vini spp. Progne modesta Falco punctatus

Alcidae Coerebidae Meliphagidae Dicaeidae Fringillidae: Fringillinae Emberizidae Emberizidae Fringillidae: Carduelinae Fringillidae: Fringillinae Parulidae Icteridae Fringillidae: Carduelinae Fringillidae: Carduelinae Columbidae Emberizidae Emberizidae Picidae Dicaeidae Tyrannidae Monarchidae Tyrannidae Fregatidae Columbidae Fringillidae: Carduelinae Fringillidae: Carduelinae Certhiidae: Certhiinae Upupidae Callaeatidae Trochilidae Phasianidae Accipitridae: Accipitrinae Melanocharitidae:Toxorhamphini Psittacidae Hirundinidae Falconidae

290 Appendix 3: Scientific names, families, and subfamilies of non-Hawaiian birds meadowlarks mockingbirds (Galaápagos) mountain-tanagers nuthatches orioles (American) Parrotfinch, Pink-billed parrotfinches Pigeon, Passenger Ploughbill,Wattled redpolls Robin, Black (Chatham Is.) rosy-finches sapsuckers scythebills Siskin, Red Siskin, Saffron siskins spiderhunters Starling, European Thrush,Tristan treecreepers Tropicbird, Red-tailed Tropicbird,White-tailed Tui Vanga, Sickle-billed Vireo,Yellow-throated Warbler, Black-and-white Warbler,Yellow Yellowthroat, Common

Sturnella spp. Nesomimus spp. Buthraupis spp. Sitta spp. Icterus spp. Erythrura kleinschmidti Erythrura spp. Ectopistes migratorius Eulacestoma nigropectus Carduelis spp. Petroica traversi Leucosticte spp. Sphyrapicus spp. Campylorhamphus spp. Carduelis cucullata Carduelis siemiradzkii Carduelis spp. Arachnothera spp. Sturnus vulgaris Nesocichla eremita Climacteris spp. Phaethon rubricanda Phaethon lepturus Prosthemadera novaeseelandiae Falculea palliata Vireo flavifrons Mniotilta varia Dendroica petechia Geothlypis trichas

Icteridae Mimidae Thraupidae Sittidae Icteridae Estrildidae Estrildidae Columbidae Pachycephalidae Fringillidae: Carduelinae Petroicidae Fringillidae: Carduelinae Picidae Dendrocolaptidae Fringillidae: Carduelinae Fringillidae: Carduelinae Fringillidae: Carduelinae Nectariniidae Sturnidae Turdidae Climacteridae Phaethontidae Phaethontidae Meliphagidae Vangidae Vireonidae Parulidae Parulidae Parulidae

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Index Species accounts may be found on the page numbers set in bold type and illustrations on the pages set in italic. Plates appear between pages 184 and 185.

A ‘A‘ali‘i 208 adaptive radiation xxiv, 3, 17, 63, 69, plate 8 a‘e 19 ‘aheahea 133 Aidem, Joan 42 Aidemedia 225 A. chascax 225 A. lutetiae 226 A. zanclops 225 ‘a¯ kala 17, 132, 136 ‘Akeke‘e 230–231, plates 6, 8 ‘Akepa 226–229, plates 6, 8 ‘akialoas 75–249 bill evolution 71, classification 79, disease 166, ha¯ha¯ nectar in diet 137, in Maui bark-picking guild 133, insect diet 131, 134, 138, sympatric 66 ‘Akialoa, Hoopoe-billed 66, 249 ‘Akialoa, Kaua‘i x, 245–246, plate 3 ‘Akialoa, Lesser 248–249, plates 3, 8 ‘Akialoa, Maui-nui 247 ‘Akialoa, O‘ahu 246–247, plate 3 ‘Akiapo¯la¯‘au 6, 133, 254–257, plates 4, 8 ‘Akikiki 219–221, plates 5, 8 ‘Akohekohe 264–267, plates 7, 8 ‘a¯ koko 130 Alaka‘i Plateau 13, 14 Alaka‘i Swamp ix ‘Alala¯ 11, 19 alani 17 ‘alauahios 48, 190 classification 80 historical distribution 65 ‘Alauahio, Maui 20, 192–195, plates 5, 8 bill 92 ‘Alauahio, O‘ahu 191–192, plates 5, 8 Albatross, Laysan 200 allozymes 55, 57, 58 alphabet, Hawaiian xxvi Amadon, Dean xi, 37, 38, 48 ‘Amakihi, Greater 18, 46, 243–244, plates 4, 8 ‘Amakihi, Hawai‘i 17, 19, 20, 41, 94, 104, 118, 132, 134, 136, 144, 240–243, plates 3, 8 ‘Amakihi, Kaua‘i 235–237, plate 3 bill 94, feeding on kanawao 136, nectar-stealing 140 ‘Amakihi, Lesser 232, see ‘Anianiau

‘amakihis 32, 235–244 classification 79, double invasions 65, 66, feeding in eucalyptus trees 134, generalist diet 134, insect diet 131, island populations 65, nectar in diet 136, switching to insects 139, taxon cycle 65, temperature 149 ‘Amakihi O‘ahu 238–239, plate 3 American Museum of Natural History 37 amphibians 23 ‘Anianiau 57, 92, 136, 232–234, plates 4, 8 ants 23 ‘Apapane 20, 81, 260–263, plate 6 feathers in artefacts, 278, 280, 282, nest 151 Atkinson, Carter 184 avian malaria 28, 31, 160, 166, 167 avian pox 31, 160, 166, 167 ‘aweoweo 20 Axis axis 27 B Baldwin, Paul H. 37 Ballieu,T. 34, 204 banana flowers 138 banana poka 31, 135, 139, 140 Banko, Paul C. 184 Banko,Winston 38 Bates, Henry W. 109 bathing 110 bats 21 behaviour see individual species accounts Berger, Andrew J. 39, 184 Bernice Pauahi Bishop Museum 6, 35, 48 bibliography 291 bills, types of 85 crossed and twisted 90 morphologies 67 shape in cardueline and drepanidine finches 86 bill clapping 121 bioenergetics 148 biological species concept 77 birding 181 black acacia 31 blackberry 31 Bledsoe, A. H. 55 Bloxham, Andrew 33

336 Index blue gum 31, 147 Bock,Walter J. xi, 38, 39, 48 Bos taurus 25 Bowles, John ix, 37 breeding biology 55, 149, see also individual species accounts breeding in captivity 180 breeding season 150 Brighamia 18 bromeliads 18 Bruner, Philip L. ix, 184 Bryan,William Alanson 35, 157 Bulbul, Red-vented 28 Bush-Warbler, Japanese 28 butterflies 23 butterfly, Kamehameha 133 C calls 112 alarm/social 113 juvenile 113 Campanulaceae 17 Canary, Laysan 198 Canis familiaris 282 Cann, Rebecca L. 40, 184 Capra hircus 25 Cardinal, Northern 28 cardueline finches xxiv, 47, 49, 50, 52 adaptive radiation 69, bill 85, biochemical relationship with drepanidines 55, cranial features 51, cranial similarities with Hawaiian honeycreepers 83, flocking 123, migration 55, mobbing 127, myology 83, nest sanitations 55, plumage 53, response to ‘spishing’ 126, songs 54 Carothers, John H. 40 Carpenter, F. Lynn 40 Casey,Tonnie L. C. 38, 184 cats, feral 27, 127, 177 cattle 25 centipedes in diet 131 centrifugal speciation 64 character displacement 66, 146 and drepanidine finches 64, 65 Chloridops 210 C. kona 210–211 C. regiskongi 212–213 C. wahi 212 Chlorodrepanis 75, 235 Christmasberry 31 Ciridops 273 C. anna 273–275 C. tenax 275 cladistics xxviii Clermontia 18, 142, 144 C. fauriei 18 coconut flowers 135 Coerebidae 45, 48 coerebids 50 Collins, Mark S. 184 coloration, factors affecting 108 competitive exclusion, 91 comprehensive biologic species concept 78

Conant, Sheila xxiii, 38, 39, 40, 184, 278 concordance testing xxx consonants, Hawaiian xxvi contact notes 112 convergence xxix, 48, 53, 67, 109 Cook, James 24, 33, 157 coralbean 20, 142, 144 Cornell Laboratory of Ornithology 40 cranial anatomy 84 cranial fenestra 52, 84 cranial kinesis 84 cranial osteology 48, 51 Creeper, Hawai‘i 46, 181, 221–224, plates 5, 8 creepers, Holarctic 92 Creeper, Kaua‘i 219, see ‘Akikiki Creeper, Maui 193, see ‘Alauahio, Maui Creeper, Moloka‘i 196, see ‘Ka¯ ka¯wahie Creeper, Oahu 191, see ‘Alauahio, O’ahu Creeper, Olive-green 221, see ‘Amakihi, Greater crossbills 55, 90 crows 21 Crow, Hawaiian 11, 27, 126, 128 Culex quinquefasciatus 31, 166, 167 Cyanea 18 cytochrome b 56 D Darwin’s finches 69, 70 deer, axis 27 Delissea 18 Deppe, Ferdinand 33 Diamond Head 13 ‘dieback’ 10 diet see individual species accounts diphthongs, Hawaiian language xxvi diseases in birds 157 carriers 160, management of 176, and parasites 169 dispersal 15 display flights 122 displays, antagonistic 123, mating 122 DNA–DNA hybridisation technique 55, 58 DNA in honeycreeper systematics 58 DNA sequence data 60 Dole, Sanford B. 34 dominance hierarchy in nectarivors 138 double invasions 64, 65, 66 Dove, Spotted 28, 164, 166 Dove,Whistling 68 Drepanidae xxv Drepanididae xxiv, xxv, 44, 45 DNA sequence data 60 genera 73 Drepaniidae xxv Drepanis 77, 267 D. coccinea 269–273 D. funerea 268–269 D. pacifica 267–268 Dysmorodrepanis 217 D. munroi 217–218 Duck, Laysan 15

Index 337 E East Maui 11 Eddinger, Robert 39, 184 ‘Elepaio 36, 69 Emperor Chain 7, 62 Emperor Seamounts 62 Endangered Species Act 172 Engilis, Andrew 48 epiphytes 18 epizootic diseases in Hawaiian birds 166 Eretmochelys imbricata 281 eucalypts, planting of 31 Europeans, arrival of 24 evolutionary systematics xxviii extinct and threatened honeycreepers 173 eyeworm 169 F Fabaceae, ornithophily 144 Falco peregrinus 128 feather artefacts in Hawaiian culture 278 feather-collecting, impact on Hawaiian honeycreepers 283 feathered helmets 280 feather wear 108 feeding ‘guilds’ 129 feeding noises 121 feeding territories in nectarivores 138 feral ungulates 19, 24 management of 176 Finch, Cocos 68 Finch, Cone-billed 276–277 Finch, Laysan 15, 20, 46, 62, 78 198–200, plates 1, 8 Finch, Grosbeak 210 finches, drepanidine Hawaiian xxiii, xxiv, 44, 48, 50 bill 85 plumage colour 107 Finch, Kona 210 see Kona Grosbeak finches, emberizine 49 finches, fringilline 55 Finch, House 55 Finch, Makawehi 202–203 Finch, Maui-Nui 203 Finch, Nihoa 14, 46, 62, 78, 201–202, plates 1, 8 Finch, Ridge-billed 277 firetree 31 Fleischer, Robert C. xi, 58, 60, 184 flight 110 flock cohesion notes 114 flocking 123–124 flocks, mixed species 124–126, 128 conspecific 126 flowerpeckers and berrypeckers, bill 88 flowerpeckers, parallels 68 flycatchers, monarchine 21, 69 flycatchers, tyrant 69 folivory 133 forest, lowland dry 19 mamane-naio 19, 20, 27–28, 148 temperate mixed 19 fossil birds 41–43

‘founder effects’ 64 Freed, Leonard A. 40, 184 French Frigate Shoals 14 Frigatebird, Great 280 Fringillidae 44, 49, 50 Frohawk, F.W. 34 frugivory 130 fruit-doves 68 G Gadow, Hans 44 Gagne, Betsy Harrison 42, 184 Gagne,Wayne C. 42, 184 Galápagos Islands 3, 69, 70 gapers 50, 225–226 Gaper, Curve-billed 225 Gaper, Maui-nui 226 Gaper, Straight-billed 225 geckos and skinks 23 genetic engineering 181 Geometridae 131 geraniums, Hawaiian 142 Giffin, Jon G. 184 glottal stop xxvii goats 25 goldfinches 50 Goldfinch, American 104 Great Mahele 31 Grosbeak, King Kong 212–213 Grosbeak, Kona 19, 210–211, plate 1 Grosbeak, Mauka 276 Grosbeak, Parrot-billed 76 Grosbeak, Wahi 212 guava 132 guava strawberry 31 H ha¯ha¯ 18, 137 ha¯ ha¯‘aiakamanu 18, 132, 137 habitat see individual species accounts habitats and plant communities 15 Hakalau Forest National Wildlife Refuge 125, 167, 176 Haleakala¯ 9, 11, 12, 20, 175 Ha¯na Rainforest Project 38, 39 Hanawi Natural Area Reserve 127, 175 Harrier,Wood 21 hapu‘u 17, 153, 178 hau 24 hau hele ‘ula 142, 143 hau kuahiwi 143 Hawaii Audubon Society 36 Hawaii Volcanoes NP 16, 17, 19, 20, 167, 177 Hawaii, University of 39, 40, 41, 184 Hawaiian Bird Survey 36 Hawaiian Endangered Bird Conservation Program 180 Hawk, Hawaiian 21 hawk-moths 23 head-scratching 111 Hemignathus 74, 75, 234, 250 H. (Akialoa) ellisianus 246–247 H. (Akialoa) lanaiensis 247 H. (Akialoa) obscurus 248–249

338 Index H. (Akialoa) stejnegeri 245–246 H. (Akialoa) upupirostris 249 H. (Chlorodrepanis) flavus 238–239 H. (Chlorodrepanis) kauaiensis 235–237 H. (Chlorodrepanis) virens 240–243 H. (Hemignathus) affinis 253–254 H. (Hemignathus) hanapepe 250–251 H. (Hemignathus) lucidus 252 H. (Hemignathus) munroi 254–257 H. (Viridonia) sagittirostris 243–244 Henshaw, Henry W. 35, 157 Herpestes auropunctatus 27 heterobills, 66, 71, 80, 95, 131, 138, 250–256 Hibiscadelphus 20 adaptations for pollination 142 extinction and loss of pollinators 143 Himatione 260 H. freethii 263–264 H. sanguinea 260–263 ho‘awa 19 hoi kuahiwi 18 homeothermy 148 homoplasy xix, 67 honeybees 23, 31 Honeycreeper, Black-faced 187 see Po’o-uli Honeycreeper, Crested 264 see ‘Akohekohe Honeycreeper, Laysan 263–264, plate 6 honeycreepers, Neotropical 49, 50 plumage 53 song flights 54 Honeyeater Laysan 263 see Laysan Honeycreeper honeyeaters 21 adaptive radiation 69 similarity with Hawaiian honeycreepers 67, 68 tongue shape 70 Honolulu 13, 31 Hookbill, Lana¯‘i 217–218, plates 2, 8 hoopoe 67 Hosmer Grove 31, 139, 242 Howarth, Frank 42 Huala¯lai volcano 9, 10 hummingbirds 145 Hui Manu 28 Hwamei 28 hybridisation and distinctive coloration 109 I, J ibis 21 ‘ie‘ie 18, 19, 130, 143 ‘ili‘ahi 19 ‘ilima 20, 133 ‘Iniki, Hurricane 9, 147, 183 insectivory 131 Institute of Pacific Islands of Forestry 41 International Biological Program 39 interorbital septum 48, 52, 84 Ipomoea indica 20 I. pes-caprae 20 ‘I‘iwi 269–273, plates 7, 8 tongue 102, feathers in artefacts 278, 280, 282, 283, feeding 98, odour 46, pollinating mamane 144, relationships 57, bill 98

Jacobi, James D. 38, 184 James, Helen xi, 6, 42, 75, 184 Jeffrey, Jack 184 Jungle-fowl, Red 280 K Ka‘ala, Mount 13 kahili (feathered standards) 278, 279, 280, 281 kahili ginger 31 Kaho‘olawe 11,12 Ka¯ ka¯wahie 196–197, plates 5, 8 Kamakou Preserve 160 kanawao 17, 130, 131 Kaneohe Bay 13 Kaua‘i 8, 13, 63 koli‘i Kaua‘i 98 kauila 19 Kaupo Gap 12 kawau 17 kawelu 20, 200 Kealakekua Bay 33 Keauhou Bird Conservation Center 180 Kepler, Cameron and Angela Kay 184 Keulemans, J. G. 34 Kilauea Caldera 177 Kilauea volcano 9, 10 Kipahulu Valley Expedition 38 Kipuka Pua‘ulu 19 kipukas 11 Knudsen,Valdemar 34 Ko‘olau Gap 12 Ko‘olau Range 13 koa 16, 17, 129 koa-finches 17 diet 129, double invasions and character displacement 66, sexual selection 66 Koa-Finch, Greater 207–209, plates 1, 8+46 Koa-Finch Lesser 209–210, plate 1 Koa-Finch Yellow-headed 209 Kohala Mountain 25 Kohala volcano 9 Koke‘e 19 kokio ke‘oke‘o 139, 140 ko¯lea lau nui 17, 131 Kona coast 11 Kona region 11 ‘kona weather’ 9 kopiko 19 Koster’s curse 31 kuku‘i 24 Kure and Midway atolls 7 Kure Atoll, discovery 62 L Lahaina 12 lama 20 La¯na‘i 11, 12, 36 language, peculiarities and conventions of Hawaiian xxvi, xxvii Laughing-thrush, Melodious 28 lapalapa 17 Lasiurus cinereus semotus 21

Index 339 lateral flange condition 84 Laysan 8, 14,20, 62 lehua blossoms 98 lehua mamo 16 lei (necklaces) 278 Leiothrix, Red-billed 28 lek behaviour 123 Lepidoptera 23 Lepson, Jaan K. 40 lifespan 155 lingual wings 46, 70, 100 lizards 21 Lo‘ihi Seamount 7 Lobelia 18 lobeliods 17, 18, 98, 130, 137, 139 co-evolution with birds 141, 143 longevity and survival rates 155 Loope, Lloyd L. 184 lorikeets 137 loulu 17, 20 loulu-lelo 274 Lousiana State University Museum of Natural Science 40 Loxioides 203 L. bailleui 204–207 Loxops 73, 226 L. caeruleirostris 230–231 L. coccineus 226–229 M MacMillen, Richard E. 40 Magumma 77, 231 M. parva 232–234 Mahukona volcano 9 maiapilo flower 136 maile 18 Malvaceae 20 ornithophily 143 ma¯ mane 19, 20, 129, 142, 144 mammalian predators, management of 177 Mamo, Black 268–269, plate 7 Mamo, Hawai‘i 267–268, plates 7, 8 mamos, 267–268 diet 136 taxon cycle 65 manele 19 Mannikin, Nutmeg 28, 164, 166 manono 17 mating displays 122 Maui 11, 125 Maui-nui 11 Mauna Kea 7, 9, 10, 27 Mauna Loa 7, 10, 147 177 Melamprosops 187 M. phaeosoma 187–190 Melamprosops odour 46 tongue 103 meliphagids 141 mice 27 Miconia calvescens 32 Midway Atoll 7, 15, 179 millipedes in diet 131

Mills Collection 34 Mills, J. D. 34 Millerbird, Laysan 15 mimicry, Batesian 109 mimicry, vocal 120 mints 18, 139 mites in diet 131 mitochondrial DNA 58 Mo‘omomi 13, 42 moanalos 21, 23, 25, 42 mobbing 47, 127 mockingbirds 70 Moho nobilis, 280 Moloka‘i 11, 12, 13 mongoose, Indian 27 Morin, Marie 184 mosquitoes 23 and spread of disease 160, 166, 178 biting an ‘Apanane 161 mouflon, impact of 27 moults and plumage, terminology used xxvi Mountainspring, S. 89, 184 mountain-tanagers, bills 87 Munro, George C. 34, 36, 157 Mus musculus 27 musculature 52, 83 Myrtaceae 16 N naio 19, 20 nasal operculum 52, 67 National Park Service 41 naupaka beach 20, 141 naupaka kahakai 20 Necker Island 63 nectarivory 135–141 nectar-stealing 32, 139 Nene 27 nest ‘helpers’ 154 nestlings 153 nests 151, see also individual species accounts nest sanitation 153 Newton, Alfred 34 Ni‘ihau 8, 13 niche partitioning 131, 132, 138 Nihoa 8, 14, 20, 63 nohoanu 142 nohu 20, 64 Northwestern Hawaiian Islands 8, 13, 35 nukupu‘u 138, 166 Nukupu‘u, Kaua‘i 250–251, plates 4, 8 Nukupu‘u, Maui 253–254, plate 4 Nukupu‘u, O‘ahu 252, plate 4 nuthatches 68, 92 Nuttall,Thomas 33 O O‘ahu 13 odour, drepanidine 44, 46, 70, 127 oha kepau 137 ‘oha 18, 137, 269 ‘ohe 17

340 Index ohelo kau la‘au 17, 130 ‘ohe naupaka 140, 141, 144 ohi‘a 20, 99, 133, 135, 138 ‘o¯hi‘a-ha¯ 16, 17 ‘ohi‘a-lehua blossom 135 ‘okina xxv ‘Ola‘a Forest, epizootic disease 167 ‘Ola‘a Tract 17 ‘o¯lapa 17, 130, 131 olive, Hawaiian 19 ‘Oloku‘i 12, 13 olona, 280 olopua 19 Olson, Storrs xi, 6, 42, 184 omnivores 134 ‘o¯‘o¯s 49, 70 opuhe 130 Oreomystis 74, 218 O. bairdi 219–221 O. mana 221–224 Oreomystis, bill 93 ornithophily 142, 143, 144 Orthiospiza 275 O. howarthi 276 ‘O‘u 18, 213–216, plates 2, 8 Ovis aries 27 O. musimon 27 Owl, Short-eared 46, 127 P pa‘iniu 18 pair-bonding 150 palatine process of the premaxilla 51, 84 Palila 19, 27, 179, 204–207, plates 1, 8 diet 129, odour 46, sonogram of song 115 Palmer, Henry 34, 157 Palmcreeper, Kaua‘i 275 Palmeria 264 P. dolei 264–267 panhandling 134 pa¯ pala 19 paperbark 31 paper-mulberry 24 parallelism 67 homoplasy and convergence 67 parks, refuges, and preserves 174 Paroreomyza 74, 190 P. flammea 196–197 P. maculata 191–192 P. montana 192–195 Paroreomyza, bill 93 Parrotbill, Maui 257–260, plates 2, 8 parrot-finches 70 parrots, convergence 68 parsimony xix Parulidae 50 PAUP method 57, 60, 61 PCR technique 57 Pearl and Hermes Reef 15, 20 Pearl Harbor 13 Penguin Bank 11

pepper, Brazilian 31 persimmon, Hawaiian 20 Perkins, R. C. L. 25, 34, 157 Peterson, Roger Tory ix, 38 Pheasant, Kalij 28 philopatry 156 phylogenetic species concept 77 phylogenetic systematics xxviii phylogenetic trees xxix, 58–61 pigs 23, 24, 25, 176, 178 pilo 17, 131 pine, slash 242 plantaris 83 Plasmodium relictum 166 plate tectonics theory 62 Ploughbill,Wattled 68 plumage maturation 104 Polipoli Springs 31 pollination 17 and extinction of mamos and ‘akialoas 145 avoiding 139, 145 pollinators, birds as 23 Polynesia 3, 7 adaptive radiation 70 Polynesians 23 Po‘o-uli 187–190, plate 2 Poxvirus avium 166 Pratt,Thane E. 184 predator defence 46 predators 24 Hawaiian honeycreepers response to 126 mobbing 127 Pritchardia remota 20 Pseudonestor 76, 257 P. xanthophrys 257–260 Psittirostra 73, 213 P. psittacea 213–216 Pterylosis, plumage and coloration 103 Pu‘u Wa‘a Wa‘a 10 pukiawe 17 Punchbowl Crater 13 puncture vine 20, 64 R rabbits 14 Rail, Laysan 15 Raikow, Robert J. 39 rainforest 12, 16 ‘rain shadows’ 9 Ralph, C. J. 41, 184 Ralph, Carol Pearson 41 Ramsey, Fred L. 184 rat, black 27 rat, roof 27 raiding a nest 164 rat, Norway 27 rats 23, 27 management of 177 Rattus exulans 23 R. norvegicus 27 R. rattus 27

Index 341 redpolls 50 Rhodacanthis 207 R. flaviceps 209–210 R. palmeri 207–209 Richards, Lawrence P. 37 Richardson Frank, ix, 37 Riper III, Charles van 39, 184 Rollandia 18 roosting 111 rosy-finches 53 Rothschild,Walter 34 Royal Hawaiian Agricultural Society 28 S sandalwood 19 sandalwood trade 24 San Diego, Zoological Society of 180 Scaevola 141, 142 Scott, Michael J. 40, 184 scythebills, bill similarities 67 seabirds 24 Sea Eagle,White-tailed 21 seamounts 7 seed-eaters 129 Serinus 73 Shallenberger, Robert 38, 40, 184 sheep, impact of 27 Shovelbill, Kiwi 69, 224 Shovelbill, Pololei 69, 224 sickle-bills 94 Sincock, John L. 38, 41, 184 siskins 50 site tenacity 156 skeletons 51, 83 skinks and geckos 23 Smithsonian Institution 6, 39 Molecular Genetis Laboratory 184 snails in diet 131 snakes 183 snake brown tree 182 soapberry 19 solitaires 21 songs, 114–121 advertising 114 whisper 119 Solitaire, Green 243 South Kona 16 spiderhunters, bill similarities 67 spiders in diet 131 ‘spishing’ 126 Stenogyne 18, 142, 143 stilt-owls 21 sunbathing 111 sunbirds, tongue shape 70 Sus scrofa 24 Sushkin, P. P. 49 swamp mahogany 31 Sykes, Paul W. 184 sylviine warblers 21 synapomorphy 44 systematics xxviii

T tagasaste 135 Tanager Expedition 14, 36 tanagers 4, 49, 50 genetic connections with Hawaiian honeycreepers 55, parallels 68, plumage similarities 53, songs 54, in West Indies, Galápagos and Cocos Islands 55 taxon cycle 64, 65 Telespiza 197 T. cantans 198–200 T. persecutrix 202–203 T. ultima 201–202 T. ypsilon 203 territoriality 55,150 Thraupidae 50, 52 thraupine honeycreepers, jaw muscles 52 todies 69 tongue 100 drepanidine tubular 4, 56, 71, 102 of drepanidine and other finches 47 of Hawaiian honeycreepers 101 seed-cup 100 warbler-like 100 tourism 181 Townsend, John Kirk 33 tradewinds 8 translocation 178 trap-line feeding 139 tree fern 17, 178, 153 treecreepers, Australian 92 tree heliotrope 133 Trematolobelia 18 Tristan da Cunha, 70 Tropicbird, Red-tailed 280 turtle, hawksbill 281 U,V ‘Ula-‘ai-ha¯ wane 17, 273–275, plate 2 United States Exploring Expedition 33 UPGMA numerical program 57 Usnea lichen in nests 152 Vancouver, George 24, 278 Vanessa tameamea 133 Vangulifer 224 V. mirandus 224 V. neophasis 224 Vanga, Sickle-billed 67 Vestiaria 77 vetch, Hawaiian 142 Viridonia 75, 243 vocalisations 54, 112–121 Volcano Village 19 vowels, Hawaiian xxvi W Wai‘ale‘ale 13, 14 Wai‘anae Range 13 Waimea Canyon 13, 26 Waimea Plains, effect of cattle 25

342 Index warblers 56 Ward,William V. 40 Warner, Richard E. 40, 184 wauke 24 West Maui 11, 12 West Moloka‘i 11, 13 West Nile virus 182 wiliwili 20, 142, 144 Wilson, Scott 25, 34, 157 wing flutter 121 White-eye, Japanese 28, 126

wood warblers 50, 68 diet 134 woodpeckers 5 Woodside, David W. 37, 38, 184 X,Y, Z Xestospiza 276 X. conica 276–277 X. fastigialis 277 Zeigler, Alan 3, 6, 42, 184 Zusi, Richard L. 39, 52