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Contents

List of colour plates xix List of abbreviations xx Plan of the book, names and terms xxiii Honeycreeper Topography xxxi Frequently mispronounced words of Hawaiian origin xxxii

PART I General chapters 1 The Hawaiian honeycreepers: evolutionary triumph 3 and ecological tragedy 2 The honeycreepers’ world 7 3 Discovery and research: historical perspectives 33 4 Origin and evolution 44 5 Classification 72 6 Form and function 83 7 Behaviour 110 8 Ecology and breeding biology 129 9 Diseases and parasites 157 10 Status, conservation, and the future 172

PART II Species accounts 11 Genus Melamprosops 187 Po‘o-uli Melamprosops phaeosoma 187 Genus Paroreomyza 190 O‘ahu ‘Alauahio Paroreomyza maculata 191 Maui ‘Alauahio P.montana 192 Kakawahie P.flammea 196 Genus Telespiza 197 Laysan Telespiza cantans 198 T.ultima 201 Makawehi Finch T.persecutrix 202 Maui-Nui Finch T.ypsilon 203 Genus Loxioides 203 Loxioides bailleui 204 Genus Rhodacanthis 207 Greater Koa-Finch Rhodacanthis palmeri 207 Lesser Koa-Finch R. flaviceps 209 Genus Chloridops 210 Kona Grosbeak Chloridops kona 210 Wahi Grosbeak C. wahi 212 Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xvi

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King Kong Grosbeak C. regiskongi 212 Genus Psittirostra 213 ‘O‘u Psittirostra psittacea 213 Genus Dysmorodrepanis 217 Lana‘i Hookbill Dysmorodrepanis munroi 217 Genus Oreomystis 218 ‘Akikiki Oreomystis bairdi 219 Hawai‘i Creeper O. mana 221 Genus Vangulifer 224 Kiwi Shovelbill Vangulifer mirandus 224 Pololei Shovelbill V.neophasis 224 Genus Aidemedia 225 Straight-billed Gaper Aidemedia chascax 225 Curve-billed Gaper A. zanclops 225 Maui-nui Gaper A. lutetiae 226 Genus Loxops 226 ‘Akepa Loxops coccineus 226 ‘Akeke‘e L. caeruleirostris 230 Genus Magumma 231 ‘Anianiau Magumma parva 232 Genus Hemignathus 234 Subgenus Chlorodrepanis 235 Kaua‘i ‘Amakihi Hemignathus (Chlorodrepanis) kauaiensis 235 O‘ahu ‘Amakihi H. (C.) flavus 238 Hawai‘i ‘Amakihi H. (C.) virens 240 Subgenus Viridonia 243 Greater ‘Amakihi H. (Viridonia) sagittirostris 243 Subgenus Akialoa 244 Kaua‘i ‘Akialoa H. (Akialoa) stejnegeri 245 O‘ahu ‘Akialoa H. (A.) ellisianus 246 Maui-nui ‘Akialoa H. (A.) lanaiensis 247 Lesser ‘Akialoa H. (A.) obscurus 248 Hoopoe-billed ‘Akialoa H. (A.) upupirostris 249 Subgenus Hemignathus 250 Kaua‘i Nukupu‘u H. (Hemignathus) hanapepe 250 O‘ahu Nukupu‘u H. (H.) lucidus 252 Maui Nukupu‘u H. (H.) affinis 253 ‘Akiapola‘au H. (H.) munroi 254 Genus Pseudonestor 257 Maui Parrotbill Pseudonestor xanthophrys 257 Genus Himatione 260 ‘Apapane Himatione sanguinea 260 Laysan Honeycreeper H. freethii 263 Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xvii

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Genus Palmeria 264 ‘Akohekohe Palmeria dolei 264 Genus Drepanis 267 Hawai‘i Mamo Drepanis pacifica 267 Black Mamo D. funerea 268 ‘I‘iwi D. coccinea 269 Genus Ciridops 273 ‘Ula-‘ai-hawane Ciridops anna 273 Kaua‘i Palmcreeper C. tenax 275 GENERA INSERTAE SEDIS 275 Genus Orthiospiza 275 Mauka Grosbeak Orthiospiza howarthi 275 Genus Xestospiza 276 Cone-billed Finch Xestospiza conica 276 Ridge-billed Finch X. fastigialis 277

Appendix 1: Honeycreepers in Hawaiian material culture by Sheila Conant 278 Appendix 2: Scientific names and families of plants mentioned in the text 285 Appendix 3: Scientific names, families, and subfamilies of non-Hawaiian 289 mentioned in the text

Bibliography 291

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(1991), while those of historically known genera Organisation are based on my earlier work (Pratt 1979a) ampli- The text is divided into two main sections. Part I is fied from more recent sources. Species accounts an overview of evolution begin with the full English and scientific names, a and natural history in 10 chapters. Part II comprises selective synonymy of scientific names (major ref- accounts of each species in systematic order with erences only), and other vernacular names, both diagnostic descriptions of the genera. The nine English and Hawaiian. Etymology of the current colour plates depict all historically known honey- vernacular and scientific names is given where creeper species and representative artefacts made known. Systematics discusses briefly the species’ from honeycreeper feathers.The birds are shown in evolution, relationships, and taxonomic history characteristic poses and in native plants upon which cross-referenced to discussions in Part I.Any recog- they are known to feed or perch. Almost all were nised subspecies are discussed here. The Descrip- painted directly from specimens, but a few that are tion includes an overall length measurement (cm) not represented in American collections were based taken mostly from Pratt et al. (1987) and intended on specimen photographs or colour lithographs in a only to establish relative sizes among species. Col- copy of Rothschild (1893–1900) in the John S. oration is described in general terms meant to sup- McIlhenny Collection of the Louisiana State Uni- plement the colour plates. Unlike previous volumes versity Library. At the back are three appendixes: in this series, detailed colour descriptions are not Appendix 1, contributed by Sheila Conant, dis- given, nor are the usual linear measurements, both cusses the importance of honeycreepers in Hawaiian of which are readily available in the recently pub- culture; Appendix 2 is a tabular listing of the scien- lished BNA accounts. Voice is described, begin- tific names and families of plants mentioned in the ning with calls and ending with primary songs. text, following Wagner et al. (1990); and Field identification compares the species with Appendix 3 lists scientific names and families of others that might cause confusion for observers. birds that occur outside Hawai‘i that are mentioned Even extinct species may include this heading to in the text, taxonomy following Sibley and Monroe help prevent mistaken ‘rediscoveries’ (see Pratt and (1990) or AOU (1998). Native and introduced Pyle 2000). Distribution, habitat, and status Hawaiian birds other than honeycreepers are listed in references a range map where feasible, but maps are two tables in Chapter 2.And finally,a Bibliography not given for species known from subfossil remains lists all the publications known to the author that or from only a few historical specimens. Current deal with Hawaiian honeycreepers in any substantive ranges are shown in solid tone, historic ranges in way,plus literature cited that would not otherwise be cross-hatching. For species that are not mapped, included. range may be discussed in detail, emphasising the In Part II, the diagnoses of genera known localities where the birds were last found. Diet and only prehistorically are based on James and Olson foraging, Breeding biology, and Life cycle Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxiv

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and demography complete each species account. group of birds, so placing them in a family of their For many poorly known species, sections may be own was the only reasonable course. Now that deleted or combined to save space and avoid con- most authors accept that the Hawaiian honeycreep- stant repetition of ‘no information’. ers are a sister group of the cardueline (see When using the Bibliography, one should Chapter 4), maintaining them in a family of their understand several conventions.With a few excep- own is less tenable. Raikow (1985), however, pres- tions, tabular references such as field checklists, ents a cogent analysis of the relationships as cur- museum catalogues, world checklists, and other rently understood, and recommends maintenance simple lists are excluded. Only the most recent of the family Drepanididae. Koblik (1994), virtually example is given for some lists, such as the US alone among recent authors, agrees with Raikow. Federal list of Endangered Species (USFWS Whether the Hawaiian honeycreepers are a sepa- 1999a), which are updated periodically.Periodicals rate family or a subfamily or tribe within the are abbreviated following modern conventions but Fringillidae, they are widely recognised as a distinc- substantive names are not abbreviated, i.e. Journal of tive, monophyletic, geographically isolated group the Ni‘ihau Ornithological Society would be abbrevi- that has undergone a spectacular burst of adaptive ated J. Ni‘ihau Ornithol. Soc. Authors’ given names radiation. If included in the Fringillidae, they are listed as initials only. References are cited in clearly stand apart in their degree of variation from the text by use of the ‘name/date’ system. Page the other members of the family, all of which are numbers within references are given only for clearly ‘finches’ in the general sense (see discussion direct quotations. Usually only the author’s last of names below).Their notoriety is such that pack- name is given, but in cases that would be confus- aging them as merely a subgroup of a well-known ing, initials are given as well. For example, the Bak- and widespread family of birds does a disservice to ers, Paul and Helen, are co-authors of two papers both groups of birds as well as to the naturalists, published in 2000; on one he is the senior author, ornithologists, evolutionists, birders, and conserva- while she is the senior on the other.To distinguish tionists who might benefit from studying them. them, they had to be cited as ‘P.and H. Baker’ and They clearly are a ‘family’ in a colloquial sense, and ‘H. and P. Baker’, respectively. Both my colleague their ranking in the taxonomic hierarchy is cur- Thane K. Pratt (no relation) and I are senior rently moot. Consequently, this volume in the authors of numerous publications listed. I use his series ‘ families of the world’ is devoted to them initials with his citations in cases where doing oth- unapologetically and without prejudice as to erwise would cause undue confusion, and use just whether they are a ‘real’ family or not. ‘Pratt’ for my own. Honeycreepers or finches? Terminology In this volume, ‘Hawaiian honeycreeper’ and ‘hon- eycreeper’ are used as the general term for members The family concept of the Drepanidinae (or Drepanididae or Drepani- Previous volumes in this series have dealt with bird dini or Drepaninini), and ‘Hawaiian finch’ refers families whose categorical status has not been specifically to those honeycreepers that retain finch- equivocal.The Hawaiian honeycreepers are an ex- like characteristics. Recently, a few authors have ception: although long regarded as a full-fledged rejected the name ‘Hawaiian honeycreeper’ as a family, Drepanididae, classifications since the 1970s group name, preferring to refer to these birds sim- have increasingly moved the group further down ply as ‘finches’ or ‘Hawaiian finches’, because not all the taxonomic hierarchy as a result of our increas- members of the group feed on honey (ϭnectar), ing knowledge of origins and relationships. Earlier many of them do not creep, and they are related to authors could only speculate about the nearest rel- a group of finches. In my opinion, calling these birds atives of this diverse but obviously interrelated finches serves only to obfuscate. Certainly no one Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxv

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using the term ‘honeycreeper’ in this context thinks highly varied Hawaiian honeycreepers simply as of it literally, any more than they do for such other ‘finches’. A few writers have attempted to compro- bird group names as nuthatch, roller, courser, thrasher, mise by using terms such as ‘honeycreeper-finches’ oystercatcher, spiderhunter, nutcracker, warbler, car- or ‘finch-honeycreepers’, but such constructs are dinal, bishop, sandpiper, or shearwater. These are cumbersome, unnecessary, and, in my opinion, not simply useful epithets with a long tradition of Eng- particularly helpful. The word ‘finch’ has a rather lish usage whose literal meaning is now secondary. specific general meaning in English that has nothing Furthermore, few such designations apply to every to do with classification. The new shorter Oxford Eng- member of a group. Many warblers do not warble, lish dictionary (Brown 1993) defines it as ‘Any of some flycatchers do not catch flies, relatively few numerous small birds, typically having babblers babble, and not all mockingbirds mock. short, stout beaks adapted for eating, belonging Denying the longstanding name ‘honeycreeper’ on to the family Fringillidae and certain related fami- these grounds is pure pedantry. lies.’This definition would certainly be misapplied When Hawaiian honeycreepers were first dis- to an ‘amakihi or ‘I‘iwi. The dictionary’s ‘related covered by European scientists, the family designa- families’ include Emberizidae, Cardinalidae, Passeri- tion of many of them was far from obvious, so they dae, Ploceidae, and Estrildidae, all of which include were distributed among a variety of avian families. birds that could be called ‘finches’(see Zusi 1978 for Eventually, the interrelationship of all the thin- that specific usage). The term ‘Hawaiian finches’ billed birds was recognised, but those with finchlike could even be misinterpreted to refer to members of bills continued to be regarded as true finches until these families introduced to the islands.A Hawaiian the end of the nineteenth century (see Chapter 3). honeycreeper by this other name would sing as By the time their relationship to the thin-billed sweetly, but its distinctiveness would be lost in (and mostly nectarivorous) birds was established, nomenclatural ambiguity. Recently, the nickname the name ‘honeycreeper’ was well entrenched. ‘dreps’ has come into use (Pratt 2002b), and it Furthermore, the finchlike honeycreepers were appears occasionally in these pages. regarded as the result of evolutionary convergence Readers who consult the historical literature will with ‘true’ finches until the late 1970s, when the find the basic name of the honeycreeper taxon cardueline finch connection first became widely variously written as Drepaniidae, Drepanidae, and accepted. Pratt (1979a) used the term ‘Hawaiian Drepanididae. Mayr (1943) and several major refer- finches’ specifically for the finchlike honeycreepers ences (Amadon 1950; Baldwin 1953) used Drepani- to separate them as a group from the rest of the idae, but Drepanididae is now considered the honeycreepers, and most writers since, e.g. Schluter correct form. In references and quotations from his- (1988), have used the term with that restricted torical texts, I have avoided the use of ‘[sic]’ after meaning. earlier name variants because these forms were not Olson and James (1982b), however, used the term incorrect when they were used. ‘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 Names of honeycreeper 1985). Their usage is not without merit because it calls attention to relationships that differ strongly subgroups from the long-held belief that these birds descended Virtually every author since the beginning of the from a thin-billed ancestor, and also calls attention twentieth century has found it convenient to subdi- to the fact that a disproportionate number of the vide the Hawaiian honeycreepers into smaller species species known from prehistoric remains are finch- groups for discussion purposes.Some have recognised like. However, I agree with Amadon (1986) that tra- two groups, some three. Whether these should be dition and accurate communication far outweigh recognised taxonomically has been controversial, and any perceived instructive value in referring to the is addressed philosophically in Chapter 5. However, I, Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxvi

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too,find the need to be able to refer to certain group- as English loan-words. As such, they can be (and ings. In my 1979 dissertation, I recognised three often are) written without the orthographic con- subfamilies that I now regard as representing evolu- ventions (see below) of the Hawaiian language. tionary grades rather than clades. These correspond I have chosen to write bird, plant, and place names more or less to the two subdivisions of Perkins (1903) of Hawaiian origin with the proper orthography to and Amadon (1950) except that their ‘green birds’ aid the reader with pronunciation. The ultimate group is split into finchlike and thin-billed divisions. authority on the Hawaiian language, and the source These groupings are still convenient: of much of the etymology presented herein, is the Pukui and Elbert (1971) Hawaiian dictionary. 1. The ‘red and black’ birds of Perkins and We will never know what the people who Amadon, which I also refer to as the nectari- drove them to called the species of vores (they could also be called drepanidines, Hawaiian honeycreepers known only from prehis- but for obvious reasons to do so would lead to toric remains. Most of their names were probably confusion). long forgotten by the time of Cook’s arrival. Nor 2. The finchlike honeycreepers, also called did James and Olson (1991), the describers, create drepanidine finches, Hawaiian finches, and English counterparts for their Latin names, but psittirostrines (after the genus Psittirostra). Ziegler (2002), a close colleague of Olson and 3. The thin-billed ‘green birds’, which can be James, proposed several. English names matter in called the hemignathine group (after the genus this case, mainly to emphasise the fact that these Hemignathus). newly described species are not ancient fossil forms like dinosaurs, but were part of the same Some genera, e.g. Melamprosops, Paroreomyza, Ore- modern avifauna as the historically known spe- omystis, and Pseudonestor, do not fit comfortably into cies. The names used in this book were chosen in any of these broad categories, but that does not collaboration with Helen James (pers. comm.) destroy the usefulness of the groupings for making because I thought it presumptuous to name these generalisations. birds on my own.

English names of Hawaiian Peculiarities of the Hawaiian honeycreepers language Part of the charm and mystique of Hawaiian hon- Hawaiian has a limited alphabet, with only 12 con- eycreepers, for English speakers at least, stems from sonants and 5 vowels.Vowels are pronounced as in their exotic-sounding names, nearly all derived Romance languages such as Spanish or Italian. from Hawaiian. From the earliest days of European Diphthongs are pronounced with more separation discovery, English speakers adopted the Hawaiian of the merged sounds than in English, and are names for native birds in the islands, perhaps always stressed on the first member of the pair. because none of the familiar European bird names Most (ei, eu, ou, ai, ae, ao) are pronounced as one seemed appropriate. As pointed out by Olson and would expect, but au always sounds like the ow of James (1995), these vernacular names are the only how rather than the au of fault. ones that have remained stable through two cen- The consonants are pronounced straightforwardly turies of taxonomic twists and turns, and one can- as in English, with the exception of w, which can not understand the classic honeycreeper literature be pronounced as either w or v. Initially or after u without knowing them. Most species have no other or o, it is w; after e or i, it is a soft v; after a it is correct name in English, so the names can be correctly either way,but in a given word usually is pronounced regarded, as many already have been in a variety of one way or the other by custom (‘I‘iwi ϭ ee- dictionaries, including the Oxford English dictionary, EE-vee; Kakawahie ϭ KAH-KAH-wah-HEE-eh; Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxvii

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Kaho‘olawe ϭ KAH-ho-oh-LAH-veh; Hawai‘i ϭ correctly. The next section, ‘Frequently mispro- Ha-WYE-ee or Ha-VYE-ee). These examples also nounced words of Hawaiian origin’, lists pronunci- include the ‘okina, written as an opening quote mark ations of honeycreeper names of Hawaiian origin (‘),which functions as an additional consonant.It rep- and a selection of other frequently mispronounced resents the stoppage of sound between syllables of the Hawaiian place names and terms. English interjection ‘uh-oh’ or some local British pronunciations of the tt in ‘little’ (‘li‘l’). This ‘glottal Language conventions stop’ had no equivalent letter in the Roman alphabet, and is often misinterpreted as a diacritical mark or a The interplay of English and Hawaiian sometimes mark of punctuation.Writing Hawaiian names with- produces some linguistic tangles. One involves the out the glottal stop makes them difficult to read and fact that Hawaiian has no plural form for nouns, pronounce, just as if a letter has been deleted (Pukui plurality being indicated by context. Depending on and Elbert 1971). An initial ‘okina, as in ‘amakihi, is personal preference more than anything else, vari- heard only in a word sequence such as ‘Kaua‘i ‘Amak- ous modern scientific writers may or may not plu- ihi’ (all Hawaiian words end in vowels, so the initial ralise Hawaiian-derived names. In English, hunters glottal stop breaks the vowel glide between words). have traditionally dropped plurals, as in ‘We shot The ‘okina is essential to avoid ambiguity and to des- three quail, two turkey,and some grouse’, and mod- ignate diphthongs. For example, the Hawaiian name ern birders, especially in Hawai‘i, have adopted this for Bulwer’s Petrel is ou, but one of the honeycreep- lingo both in speaking and writing. For this book, ers is called ‘O‘u . Only the ‘okina distinguishes a I have taken the position that bird names of Hawai- diphthong from the two-syllable pronunciation. ian origin are now English loan-words and can In general, Hawaiian words are accented or be pluralised. However, I restrict the use of plurals stressed on the next-to-last and alternating preced- to enumerating kinds rather than individuals. For ing syllables, with the final stress the strongest example, there are several ‘amakihis (Kaua‘i ‘Amak- (‘amakihi ϭ AH-mah-KEE-hee). Words of five ihi, O‘ahu ‘Amakihi, Common ‘Amakihi, and syllables are stressed on the first and fourth Greater ‘Amakihi), but one might correctly report (‘akialoa ϭ AH-kee-ah-LO-ah;‘alauahio ϭ AH-lau- ‘I saw only three ‘amakihi on the ‘Aiea Trail’. ah-HEE-oh). However, many words have irregular The previous example also demonstrates the stresses indicated by a horizontal line, called a widespread practice in ornithological and birding macron, over the vowel. But this stress is not quite literature of capitalising the specific English names the same as an irregular accent in, say, Spanish, of birds. I follow that practice herein because I because it is superimposed over the stress that would believe it reduces confusion. Hawaiian bird names, be present normally. English speakers can approxi- including those of honeycreepers, are standardised mate the effect by thinking of a syllable with a by the American Ornithologists’ Union (AOU macron as if it were a separate word (‘akepa ϭ AH, 1998), albeit without ‘okina and macrons, so they KEH-pah, not AH-kep-ah). Many words with take on the mantle of proper names that can be macrons are actually phrases written as one word. capitalised. For those outside the AOU area, I use For example Haleakala was originally a phrase (hale the names of Sibley and Monroe (1990). a ka la,‘house of the sun’) and is pronounced HAH- A final convention involves tense. Throughout lay-a-kah-LAH. Several names of Hawaiian honey- this book I often use the present tense to describe creepers have similar etymologies. Stressed syllables actions of extinct birds, especially when comparing (and stressed monosyllabic words such as la, above) them to living species or when they are part of a list originated as repeated vowels with a glottal stop that includes both extinct and living species. It sim- between (Ziegler 2002) that in casual pronunciation ply reads better and avoids circumlocution. I use became slurred together (as the syllable po in ‘Aki- past tense when the subject is something that obvi- apola‘au, which originated as po‘o (ϭhead). Know- ously no longer occurs or when present tense ing that helps to pronounce words with macrons would be very misleading. Bear in mind that several Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxviii

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honeycreepers are currently in that limbo of not taxonomy (Chapter 5).The latter is included within having been observed for a considerable time but systematics and involves only the naming of taxa not ‘officially’ declared extinct. Using present tense (defined above) and grouping them with related in those cases at least gives them the benefit of the taxa in a hierarchy.Taxonomy is as old as mankind, doubt. 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 Terminology of moults and explanation here of evolutionary philosophies and plumages their terminology may be of some help. I follow the now widely used terminology of Humphrey and Parkes (1959), with the names of Evolutionary systematics moults and plumages capitalised. Nestlings begin with a coat of down, which is replaced before fledg- When Charles Darwin first proposed the principle ing by the Juvenal plumage. Most of the Juvenal of evolution, many of the details, especially those plumage is worn for a relatively short time, and is dealing with genetics and the process of speciation, replaced in the First Prebasic moult by the First Basic remained to be worked out.The earliest hypothes- plumage. The flight feathers from the Juvenal is of Hawaiian honeycreeper evolution (Perkins plumage, however, are retained for a full year.These 1903) was developed well before these phenomena feathers are usually distinguishable from the rectri- were thoroughly understood. Eventually, a consen- ces and remiges of adults, allowing first-year birds to sus called the ‘modern synthesis’, as elucidated pri- be distinguished. In birds that have two distinctive marily in the works of Ernst Mayr (1943, 1963, plumages each year, Basic plumage is worn during 1969), developed. Mayrian evolutionary systematics the ‘off-season’, and the Alternate plumage during the dominated studies of honeycreepers from the 1940s breeding season. Hawaiian honeycreepers do not to the 1970s. 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 Phylogenetic systematics differs progressively as a bird ages, as in several The last quarter of the twentieth century witnessed Hawaiian honeycreepers with delayed plumage a sea change in systematic philosophy, from evolu- maturation, it is referred to as First Basic, Second tionary methods to phylogenetic systematics, which Basic, etc.When a bird reaches a stage in which one grew mainly from the work of the German evolu- plumage looks the same as the next, it is said to be tionist Willi Hennig (1966). Phylogenetic sys- in Definitive plumage.The term immature is used for tematics did not gain wide acceptance among birds whose exact age is not known but which have ornithologists until relatively recently, and the dif- not reached the Definitive stage.The term subadult ferent approaches of these philosophies are still may be used for an immature that is approaching being sorted out.The two schools treat some of the Definitive. same terms differently,and phylogenetic systematics has its own esoteric jargon that may not be famil- iar to the lay reader. Systematic philosophy and jargon The techniques of phylogenetic systematics are Systematics, the science of evolutionary history, called cladistics. An excellent short summary of relationships, classification, and naming of organ- cladistic methods in the Hawaiian context is given isms has a vocabulary all its own. Some familiar by Funk (1995), but for a more detailed discussion words have specialised meanings in this context, see Forey et al. (1992) and references cited therein. and many terms are used solely by systematists. Lay- In cladistics, groupings are defined by one or more men often confuse systematics (Chapter 4) with shared derived character states or synapomorphies.In Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxix

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other words, all the groupings that share a uniquely razor is a guideline, not a law.Funk (1995: 33) states derived condition are presumed to have descended ‘This principle does not preclude the possibility of from a common ancestor that had that condition. convergent or parallel evolution; it simply states that Such a monophyletic group of taxa is called a clade. when there is no reason to think otherwise, two A clade may or may not correspond to a taxon, but characters that appear to be the same are treated all taxa are clades in phylogenetic classifications. In as homologous.’ In other words, convergence is contrast to evolutionary systematics, clades are never assumed. Nowadays, cladograms are con- defined solely on the basis of synapomorphies, structed using computer programs, especially one whereas in evolutionary systematics groupings are known as PAUP*, an acronym for Phylogenetic allowed based on characters that are primitive, i.e. Analysis Using Parsimony (Swofford 1999). holdovers from an ancestral evolutionary stage. In phylogenetic systematics, such paraphyletic groups may be called evolutionary grades, but they are not Phylogenetic trees recognised taxonomically. Shared primitive charac- A hypothetical reconstruction of evolutionary history ters are called symplesiomorphies. A character that is is called a phylogeny.Traditionally,a phylogeny is illus- derived (an apomorphy) at one level becomes prim- trated in the form of a tree or dendrogram, something itive (a plesiomorphy) at a higher level. Uniquely like a family tree in human genealogy, using specia- derived character states that are found in only one tion events (often hypothetical) as branching points taxon are called autapomorphies. Groups that are or simply showing a pattern that does not define the each other’s closest relatives are called sister groups, branching points at all. In evolutionary dendrograms, and if they are named, sister taxa.A series of changes taxa are grouped on the basis of overall similarities, in the same character, each derived from the other, and a group is considered monophyletic as long as all is called a transformational series. Such series are the of its members share a common ancestor, even if it phylogenetic equivalent of Bock’s (1970) ‘micro- does not include all the descendants of that ancestor. evolutionary sequences’ among Hawaiian honey- If a dendrogram is based on numerical data,it is called creepers. For a transformational series to be used in a phenogram (Sneath and Sokal 1973). A dendrogram reconstructing evolution, one must determine the that is constructed using cladistics is called a cladogram. direction or polarity of the changes.The most com- Its branching points are based on transformational mon method for determining polarity is called out- series of synapomorphies. In phylogenetic systemat- group comparison. If a character occurs in two or ics, to be monophyletic a group must include all the more forms or states within a group, the state found descendants of a common ancestor. Groups that do in the sister group and/or more distantly related not include all descendants are termed paraphyletic.In taxa is assumed to be the plesiomorphous or prim- this book, the term monophyletic is used in the lat- itive condition. ter sense. In both systems, any taxon that can be An important element of cladistics is the concept shown to be polyphyletic, i.e. comprising descendants of parsimony. The rule of parsimony, or ‘Occam’s of two or more different ancestors, is assumed to be razor’ as it is sometimes called (referring to William erroneously constituted. of Occam, a medieval Englishman who first stated Phylogenetic trees are never intended to repre- the principle), basically means that one never sent observed fact.They are hypotheses about how assumes a complex or convoluted explanation if a evolution may have proceeded. In the earliest sys- simpler one is available. Convergence, now often tematic studies of Hawaiian honeycreepers, includ- called homoplasy when it occurs among members of ing Perkins (1903), Bryan and Greenway (1944), a closely related group, is relatively rare in nature Amadon (1950), Baldwin (1953), and Richards and because it usually requires a hypothesis of more Bock (1973), relationships were usually hypothe- complex evolutionary patterns than the alternative sised based on similarities and differences, with the that similar things are derived from a common phylogeny developed later.This methodology rep- ancestor. Nevertheless, it does happen; Occam’s resents the evolutionary school of systematics, but Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxx

xxx Plan of the book, names and terms

in the context of phylogenetic reconstruction is should still be regarded as tentative. Despite the better referred to as the eclectic school (Raikow enthusiasm of the practitioners of these revolution- 1985). In it, a hypothetical phylogeny attempts to ary techniques, who tend to dismiss contrary evi- express both character transformations and geneal- dence from phenotypic characters without serious ogy. In phylogenetic studies, the phylogeny, which comment, I believe, like Raikow (1986), that con- is purely genealogical, comes first and is then used cordance testing with traditional methods is the to determine relatedness. My own early work (Pratt only reasonable way to evaluate these hypotheses. 1979a) was a product of its times in that I used The earliest phylogenies, such as that of Perkins cladistics to develop a phylogeny,but allowed eclec- (1903), were actually drawn to resemble trees, but tic thinking to influence my classification. More more recently a horizontal arrangement, with the recently, classifications have been based primarily tree lying on its side, has become the standard. To on phylogenetic principles, but the most widely facilitate comparisons of various trees in this book, followed one (AOU 1998) still retains elements of I have redrawn those published elsewhere in a com- earlier eclectic thinking.The classification proposed mon style using the program MacClade (Maddison in this book (see Chapter 5) is based solely on phy- and Maddison 2000). In no case is the tree topology, logenetic patterns (Chapter 4). i.e. the branching sequence, changed, but other The majority of phylogenetic trees that have information, such as branch lengths used in some been published since 1979 have been based on bio- trees, may be lost through this simplification. For the chemical or genetic studies.The fact that phyloge- same reasons, I have converted the taxonomy of the nies developed by these techniques seem to change originals to modern equivalents and have used ver- in fundamental ways from one study to the next, nacular names because they are the only ones that even in the same laboratory, indicates that results remain comparable across various studies. Honey Creepers_FM.qxd 11/19/2004 8:01 AM Page xxxi

Honeycreeper Topography

nape forehead upper back

lower back tertials malar area rump wing bars primary coverts secondary coverts uppertail coverts scapulars median coverts lesser coverts auricular (cheek)

eye-ring

rectrices lore (tail feathers) primaries tomium (flight feathers) vent chin secondaries throat flank breast thigh belly

hallux toe claw

postocular stripe side of neck crown superciliary (eyebrow) back forehead secondaries lore maxilla (upper mandible) primaries culmen tail tomium

gonys (lower) mandible

undertail coverts breast flank tarsus belly