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EXTINCT BIRDS
Julian P. Hume
CHRISTOPHER HELM
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Contents
Foreword7
Preface to the Second Edition 9
Acknowledgements11
Introduction13
Glossary17
Extinct birds 19
Struthionidae (Ostriches) 19 Bucerotidae (Hornbills) 242
Casuariidae (Emus) 20 Picidae (Woodpeckers) 243
Apteryigidae (Kiwis) 25 Acanthisittidae (New Zealand wrens) 250
Aepyornithidae (Elephant birds) 26 Pittidae (Pittas) 255
Dinornithidae (Giant moas) 30 Furnariidae (Ovenbirds) 256
Emeidae (Lesser moas) 33 Grallaridae (Antpittas and ant-thrushes) 258
Megalapterygidae (Upland Moa) 38 Maluridae (Glasswrens) 259
Sylviornithidae (Scrubfowls) 40 Tyrannidae (Tyrant flycatchers) 260
Numididae (Guineafowl) 41 Turnagridae (Piopios) 262
Phasianidae (Grouse and quails) 42 Meliphagidae (Honeyeaters) 265
Anatidae (Ducks, geese and swans) 47 Dasyornithidae (Bristlebirds) 266
Spheniscidae (Penguins) 65 Acanthizidae (Australian warblers) 266
Procellariidae (Petrels) 67 Callaeidae (Wattlebirds) 267
Hydrobatidae (Storm-petrels) 73 Notiomystidae (Stitchbird) 271
Podicipedidae (Grebes) 73 Cinclosomatidae (Quail-thrush) 271
Threskiornithidae (Ibises) 76 Cracticidae (Currawongs and butcherbirds) 272
Ardeidae (Herons, egrets and bitterns) 79 Campephagidae (Cuckoo-shrikes) 273
Sulidae (Boobies) 84 Oriolidae (Orioles) 275
Phalacrocoracidae (Cormorants) 85 Rhipiduridae (Fan-tailed flycatchers) 276
Anhingidae (Darters) 87 Monarchidae (Monarchs) 277
Accipitridae (Eagles, kites and hawks) 88 Corvidae (Crows) 281
Falconidae (Falcons and caracaras) 93 Petroicidae (Australasian robins) 283
Otididae (Bustards) 97 Paridae (Tits and Chicadees) 284
Rhynochetidae (Kagus) 97 Mohoidae (Mohos) 285
Aptornithidae (Adzebills) 98 Hirundinidae (Swallows and martins) 290
Rallidae (Rails and coots) 99 Macrosphenidae (Crombecs) 291
Turnicidae (Buttonquails) 141 Phylloscopidae (Leaf warblers) 291
Haematopodidae (Oystercatchers) 142 Acrocephalidae (Reed warblers) 292
Charadriidae (Plovers) 144 Pycnonotidae (Bulbuls) 297
Scolopacidae (Snipes and sandpipers) 145 Sylviidae (Sylvia warblers) 298
Laridae (Gulls) 152 Locustellidae (Grasshopper warblers) 299
Alcidae (Auks) 152 Cisticolidae (Cisticolas and allies) 301
Columbidae (Pigeons and doves) 155 Zosteropidae (White-eyes) 301
Strigopidae (Kakas, keas and kakapos) 189 Regulidae (Kinglets) 305
Psittacidae (Parrots and macaws) 191 Troglodytidae (Wrens) 306
Cuculidae (Cuckoos) 217 Mimidae (Mockingbirds and thrashers) 309
Tytonidae (Barn owls) 221 Sturnidae (Starlings) 310
Strigidae (True owls) 224 Turdidae (Thrushes) 319
Caprimulgidae (Nightjars) 234 Ploceidae (Fodies) 326
Aegothelidae (Owlet-nightjars) 235 Estrildidae (Waxbills, munias and allies) 327
Apodidae (Swifts) 236 Fringillidae (Finches) 328
Trochilidae (Hummingbirds) 236 Parulidae (New World warblers) 357
Brachypteraciidae (Ground-rollers) 239 Icteridae (New World orioles) 360
Halcyonidae (Kingfishers) 240 Emberizidae (Buntings and New World sparrows) 362
Upupidae (Hoopoes) 241 Thraupidae (Tanagers) 367
5
References516
Index595
6
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Foreword
by Errol Fuller
To many people, the word extinct conjures up visions of gigantic creatures living eons ago, beasts separated
from us not just by their size and terrifying power, but by unimaginable gulfs of time. Their connection with
our own lives seems so remote that it is easy to reduce them to figures of fantasy, rather than as creatures
that were once of flesh and blood. We may hear the story of the dinosaurs or of the mammoths with awe
and wonder, but these tales hardly impact on our own lives.
But in recent years the idea of extinction and extinct creatures has begun to assume a new dimension.
The frightening speed with which the natural world is being destroyed forces us to consider extinction in a
much more personal way. No longer can it be reduced to a matter of great lumbering animals created in
celluloid to excite, alarm or entertain us. The concept of the extinction of species is now at our doorstep,
rather than confined to a distant past to which we need pay little real attention.
With this awareness has come a growing interest in species that have become extinct in comparatively
recent times, an interest that leads to the realisation that such species have an intrinsic fascination all of their
own. Many of those that have vanished in what might loosely be termed ‘recent historical times’ have left
behind traces and records of their interaction with humanity. They seem almost close enough to touch, yet
will now remain forever just out of reach. Not only do they retain the elusive and intriguing quality of
mystery, but many were remarkable creatures in their own right. Among birds are many species that have
fascinating stories attached to their names, or were weird and wonderful in appearance. There is the
Passenger Pigeon, for instance, and the Huia, and the Great Auk. And, of course, there is the Dodo.
But perhaps it is not just the interesting and curious nature of many species that makes recently extinct
birds so fascinating. Perhaps the very idea of a bird becoming extinct is something that is particularly star-
tling. After all, birds are such symbols of life, movement, vitality and freedom. Their capacity for flight and
their very familiarity – the fact that they live so noticeably all around us – make it doubly difficult to come to
terms with the idea that a species should no longer exist.
Anyone who finds the subject intriguing (and perhaps many who don’t yet realise that they do) will
discover that this second edition puts all the relevant information concerning recently extinct birds at their
fingertips – and brings it right up to date. This is not just another annotated list; it is a comprehensive
account of what is known. A book of this kind cannot be compiled after just a few months routine research.
It is a labour of love, and represents Julian Hume’s lifetime study. It combines great knowledge and experi-
ence with an enormous passion for the subject, and scholarship and practical experience in the field are
evident throughout. For years Julian has travelled the world seeking out snippets of information or gaining
insights in the actual terrains that extinct birds once inhabited. Like me, he cannot resist drawing and
painting those things that interest him, and his charming line drawings give the book an added dimension.
Doubtless, this volume will be compared with the book on extinct birds that I wrote several years ago,
but in reality any such comparison is of little purpose. My book is something of a romantic ramble through
the subject, whereas the present work is a tremendous compendium of information. No stone has been left
unturned in the pursuit of facts – some obscure, some easier to come by – and the result of all this effort is
presented in a form that is crystal clear and easily accessible. Anyone coming to this work may rest assured;
Julian Hume has ruthlessly uncovered whatever information exists, and set it down in this truly encyclo-
paedic volume.
Errol Fuller
It has been just five years since the publication of the first edition of Extinct Birds, yet already significant
developments have taken place. The inclusion of taxa known from the fossil record and birds not recorded
in recent years increases the number of extinct species by 48 and subspecies by seven. This makes dismal
reading, but is partially offset by the rediscovery over this period of two full species and 21 subspecies.
Greenway (1967) noted cheerfully in his preface to the second edition of Extinct and Vanishing Birds of the
World that six forms of bird hitherto thought extinct had been rediscovered, including the now probably
extinct Eskimo Curlew Numenius borealis. He then lists one species and four subspecies as ‘almost certainly
extinct’, of which the Aleutian Canada Goose Branta canadensis leucopareia and Anjouan Scops Owl Otus
capnodes have since been rediscovered. Greenway also included the Huia Heteralocha acutirostris in this cate-
gory, despite the fact that it had not been recorded since 1907. The problem here, of course, is this: when is a
bird species considered extinct? The IUCN (International Union for the Conservation of Nature) and
BirdLife International have defining categories, but these can be idiosyncratic. For example, the Pink-
headed Duck Rhodonessa caryophyllacea, which could still be caught in numbers until at least 1927 – ten live
birds arrived together in that year at Foxwarren Park, Surrey, England, alone – was last recorded in 1949.
There have been no confirmed reports since, yet it is listed as possibly extant. By contrast, the elusive Po’ouli
Melamprosops phaeosoma, which occurred in the extensive forested slopes of Maui in the Hawaiian Islands,
was last seen in 2004 and is now considered certainly extinct.
There is no doubt that pronouncing a species extinct prematurely can have detrimental effects. If a bird is
considered gone, all hope for it is then lost and it no longer receives protective legislation. But even when a
conservation project receives copious financial support, it may still not be enough to save a species. For
example, a long-term government-funded project to save the Orange-bellied Parrot Neophema chrysogaster, a
bird that has a precarious annual migration from southern mainland Australia to its breeding grounds in
southwest Tasmania, appears to have been in vain. The wild population is down to fewer than 50 birds and
continues to decline. Around 300 individuals are kept in captivity to safeguard the future of the species, but
this parrot is almost certainly destined to become Extinct in the wild; In 2015, BirdLife International predicted
that this would happen within three to five years. The Orange-bellied parrot will no doubt join a list of birds
that have already suffered the same fate, birds such as the Alagoas Curassow Mitu mitu, Guam Rail
Gallirallus owstoni, Socorro Dove Zenaida graysoni, Guam Kingfisher Todiramphus cinnamominus and
Hawaiian Crow Corvus hawaiiensis.
In their work on human-caused bird extinctions, Milberg & Tyrberg (1993) and Tyrberg (2009) showed
that the majority of documented cases after 1600 AD occurred on islands. But times have changed, and this
is certainly not the case today. Human interference on continental land masses has destroyed and frag-
mented natural habitats, thus creating isolated ‘islands’ in a sea of urban, industrial and agricultural
development. Combined with ever-increasing anthropogenic encroachment and pressure, these remnants
are under serious threat along with the species that they support. This is creating a new phase in the already
gloomy record of bird extinctions, and one that mirrors the catastrophic effects that have already beset
many island birds.
Furthermore, the sad predicament facing conservationists today is that with so many birds now on the
verge of extinction, choices have to be made as to which can or cannot be realistically saved. As many as 60
species have populations numbering fewer than 50 individuals, some with less than ten, and the number of
species included in this category is increasing rapidly. Many international conservation organisations are
working hard to implement protective measures to reverse this trend, and the success stories are testament
to their efforts. But the fight to prevent more species ending up in this book should not be just left to conser-
vationists; it should be of paramount importance in all government agendas. Surely it must be worth the
effort!
9
I have been most fortunate to have worked and corresponded over the years with an array of scientists,
artists and colleagues interested in extinct birds. In particular, Storrs Olson and Errol Fuller have been inspi-
rational, and our endless discussions have provided me with so many insights. I am forever grateful for
that. I especially thank Jim Martin from Bloomsbury. Jim has worked closely and patiently with me on 1st
and 2nd editions of this project; his inspired editorial skills and background knowledge kept things moving
forwards, often when seemingly impassable obstacles loomed ahead. Both books are much improved by his
efforts. I also thank Katie Read, whose editorial skills helped see this second edition through. I thank the
following friends and colleagues for correspondence and discussions, and for those who have provided
access to museum specimens, rare books and manuscripts in their care, in particular Robert P ryfls-Jones,
who has been an inspiration throughout my later career. I am also extremely appreciative to colleagues and
to the many readers of the book, who provided details that helped improve the factual content of the text.
I especially thank Michael Walters for his contribution to the first edition of this book.
In the United Kingdom I thank Lorna Steel, Anthony Cheke, Carl Jones, Glyn Young, Nigel Redman,
Dave Martill, Dave Loydell, Bob Loveridge, Mike Barker, Darren Naish, Stig Walsh, the late Cyril Walker,
the late Colin Harrison, Nick Arnold, Graham Cowles, Sandra Chapman, Phil Rainbow, Florence Gallien, Jo
Cooper, Mark Adams, Douglas Russell, Alison Harding, Effie Warr, Judith Magee, Paul Cooper, Mike
Brooke, Ray Symonds, Mathew Lowe, Clem Fisher, Tony Parker, Alan K. Jones, Malgosia Nowak-Kemp,
Richard Hing, Andrew Kitchener, Alan Knox and Nigel Collar. In the United States I thank Helen James,
Dave and Lida Burney, Dick and Karen Carruthers, Dave Staudt, Paul Sweet, Carla Dove, Gary Graves,
Pamela Rasmussen, Guy Tudor, Albert Gilbert, Donald Drake, Michael Walther, Johanna Humphrey, Joel
Cracraft and Dave Steadman. In France I thank Cécile Mourer-Chauviré, Roger Bour, Christian Jouanin,
Jean–François Voisin, Daniel Goujet, Ronan Allain, Claire Sagne, Virginie Bouetel and Christophe Thébaud.
In Réunion I thank Sonia Ribes, Jean-Michel Probst, Pierre Brial, Dominic Strasberg, and Auguste and
Christine de Villèle. In Mauritius I thank the late France Staub, the late Claude Michel, the late Jean-Michel
Vincent, the late Alfred N orth-Coombes, John Mauremootoo, the late Clare Mauremootoo, whose passing
was so premature, Anwar Janoo, Alan Grihault, Christian Foo Kune, Vikash Tatayah and all of the staff both
past and present of the Mauritius Wildlife Foundation, Vincent Florens, Claudia Baider, Gérard Bax, Pierre
and Elizabeth Baissac, Mike and Marie-Josée Martial-Craig, Jean-Marie Huron, Michele Sabelli, Vikash
Rupear and the staff of the Mauritius Institute. In Rodrigues I thank Aurel Andre, Richard Payendee,
Arnaud Meunier, Alfred Begué and Sweety Sham Yu. In the Netherlands I thank Perry Moree, Pieter and
Els Floore, René Floore, Kenneth Rijsdijk, Marijke Besselink, Perry de Louw, Hanneke Meijer, Tamara
Vernimmen, John de Vos, René Dekker, Leon Claessens, Ria Winters, Hein van Grouw, Menno Leenstra,
Justin Jansen, and all of the Dutch team who have been working on the Mare aux Songes project over the
years. In Austria I thank Ursula Göhlich, Anita Gamauf and Hans-Martin Berg. In Germany I thank Sylke
Frahnet, Pascal Eckhoff, Melanie Nayyal and Michael Stiller. In Spain I thank Josep Alcover Tomas and Juan
Carlos Rando. In Italy I thank Carlo Violani. In Japan I thank the late Hiroyuki Morioka, Satoshi Yamagishi
and Hiroto Kawabata. In Australia and New Zealand I thank Steve Bourne, Liz Reed, Kathryn Medlock,
Tammy Gordon, David Maynard, Paul Schofield, Alan Tennyson, Ross Galbreath, Jason Frogatt, Matt
Rayner, Murray Williams, Jamie Wood, Chad, Ondrea and Pearl Richards, Nick Porch and Joe Forshaw, and
in particular Trevor Worthy, whose achievements have been a source of motivation. In the Soviet Union I
thank Vladimir Loskot and Vera Richter. I thank in particular the Biodiversity Heritage Library resource
www.biodiversitylibrary.org, which has made research for this book so much easier.
For support and friendship I particularly thank Owen Griffiths, who has done so much in forging the
conservation movement in the Mascarenes and Madagascar, and whose foresight into rewilding
11
programmes have now achieved international recognition, and Greg Middleton, whose knowledge of caves
in the Mascarenes is second to none, and who has provided so much support during field work.
The following funding bodies have provided support for much of my field work: I thank the Percy
Sladen Centenary Fund, Gen Foundation, Natural History Museum Special Fund, Dodo Research
Programme, François Leguat Ltd, Mauritius Wildlife Foundation, and University of Portsmouth.
Finally I thank my family. To my mother, brothers and sister, my late Nan, and to my father, who would
have been so proud of this project, to my daughters, Jade, Jasmin and Jeradine, and to my partner, Lindsey
Hubbard, whose love and support helped see me through.
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Introduction
Extinction is a natural process that has existed for as long as life itself. It has been estimated that 99% of all
species that have ever existed have disappeared. In general, the process of extinction is comparatively slow,
and a species’s ‘lifespan’, from origin to extinction, averages between 1 and 10 million years. In addition to
these background, gradual extinctions, many mass extinctions have taken place; the most famous of these
saw the seemingly abrupt demise of the dinosaurs at the end of the Cretaceous. Mass extinctions appear
catastrophic and sudden when viewed in the context of geological time, but in fact probably took thousands
of years. As these events happened millions of years ago, they are often regarded as something from a
bygone age, far removed from our present world. Yet despite its role as a vital natural process, the scientific
concept of extinction is a relatively recent phenomenon. Before humans evolved on the planet, mass extinc-
tion events were caused by events such as extra-terrestrial impacts, volcanism, and changes of climate and
sea-level. However, we now recognise a new agent of mass destruction – ourselves. In the last millennium,
the impact of humans on the natural world has accelerated out of control, at a rate unprecedented in the
Earth’s long history. There is hardly any part of the Earth’s biosphere that has been spared from some kind
of human influence, either direct or indirect.
No vertebrate group has suffered more than the birds. They have colonised all of the world’s land-masses
including the most remote archipelagos. They are often specialised to feed on certain foods and live in
certain habitats, and they frequently become secondarily flightless. These are essentially the reasons why
birds are so vulnerable to human interference. Birds are reasonably large, conspicuous and sought-after for
food, feathers or as museum specimens; hence there is a wealth of literature that documents the discovery,
and later, the disappearance, of many species. But their demise tells one important and fundamental story,
one that epitomises what can go wrong when humans interfere with the natural world.
Avian extinction was first covered in a rather eccentric book entitled Extinct Birds, published in 1907 by
Walter Rothschild, founder of the Tring Museum and an avid and passionate collector of anything rare and
extinct. The volume was exquisitely illustrated by the top artists of the day, including John Gerard
Keulemans and George Lodge. The Rothschild family was in the banking business, which allowed Walter
to indulge both scientifically and financially in his hobby. This included sending collectors to many parts of
the world to bring back specimens for his personal collection. Without the efforts of Rothschild and his
collectors, we would be unaware of many of the avian extinctions that have taken place, especially on
islands. Fifty years later, the subject was tackled again by James Cowan Greenway Jr., when he produced
Extinct and Vanishing Birds of the World in 1958 (followed by a second edition in 1967). Greenway spent years
compiling material for his now-classic work, which included endangered, extinct, and hypothetical species.
The book was illustrated by David Morrison Reid-Henry, who produced (in my opinion) the best ink draw-
ings of birds ever produced. In 1987 (and in an expanded edition in 2000), Errol Fuller brought the subject
up to date with his lavishly illustrated volumes and dramatic and moving prose. Fuller’s work did much to
raise public awareness of avian extinction, and the books continue to be cited by academics as an important
scientific source. All of these works concentrated on species known to have become extinct since 1600, an
arbitrary date often cited by recent authors based on the time when written documentation began in earnest.
The sheer magnitude of bird extinctions that have taken place makes shocking reading. Many more
recently extinct species known from subfossil remains await description, and a number of critically endan-
gered species will probably disappear within the next decade or two. For those doubting that a
human-induced mass extinction event is taking place, you need look no further. The facts presented within
these pages speak for themselves.
13
This book
In keeping with tradition, this book is also entitled Extinct Birds, but it has a major difference from its pred-
ecessors; it describes all species and subspecies known from skins, written accounts and illustrations, and
also includes the recent palaeontological record. It is thus the first fully comprehensive treatment of the
subject ever undertaken. The fossil record shows a truer extent of bird extinctions, yet it is often ignored by
compilers, giving a great underestimation of the actual numbers involved. The fossil record is, of course,
incomplete, and an underestimate of the actual level of extinction; no doubt there are many avian taxa that
have vanished without trace, or have yet to be found as subfossils.
The species order follows the recommendations of the International Ornithological Committee (IOC),
but some bird families are in a constant state of flux, so the taxonomic arrangement is not intended to be
definitive. Nomenclature of recently extinct species broadly follows Howard & Moore Complete Checklist of
Birds of the World (Dickinson & Remsen 2013; Dickinson & Christidis 2014), though I have amended to
follow a common- sense approach where appropriate, especially for common names of long-extinct
subfossil-only species.
There has been an ongoing major taxonomic revision of bird orders and family placements (Dickinson &
Christidis 2013; Dickinson & Christidis 2014; del Hoyo & Collar 2014; Del Hoyo et al. 2017), which were not
all fully available in time for incorporation into this volume. This issue will be addressed in a future edition.
Extinct birds
This section includes all species known to have disappeared in the last 700 years that are represented by
museum specimens, including skins, skeletons, subfossil remains and eggs, and also those species known
only from credible accounts and illustrations. These generally have a documented history, and their declines
have been noted over time. The areas of former distribution have generally been drastically altered beyond
recognition, or have been overrun by exotic species. These birds have been sought for over a number of
years without success. It can be stated with absolute certainty that they once existed and are now either
extinct or probably extinct.
This section, I hope, is as exhaustive as possible, although there are of course some birds that are
presumed extinct pending further ornithological expeditions to far-flung places, and a number that are
known to have existed based on vague travellers’ tales; perhaps subfossil discoveries will fill in some of the
blanks over the years to come. I have tried to be as comprehensive as possible but acquiring detailed infor-
mation can be fraught with difficulties, especially when dealing with obscure scientific references, the
location of museum specimens, and the sometimes spectacularly confused literature. I would be pleased to
hear (care of the publishers) of any omissions or inaccuracies that we can rectify in forthcoming editions of
this book.
Hypothetical birds
The second section covers species that are known from a unique type or a few specimens, or from accounts
and illustrations, the validity of which remain in doubt. These birds may represent valid taxa, in which case
they may be now extinct, but equally they may be aberrant, hybrid or vagrant individuals.
These ‘hypothetical’ species often stem from the travelogues and systematic works of the 17th, 18th and
early 19th centuries. These contain illustrated and written accounts of a number of species that have never
been satisfactorily matched with any extant species. Some of the descriptions are so meagre as to be virtu-
ally impossible to identify. Others represent plumage variations, aberrant specimens, hybrids, artefacts, or
even d reamed-up birds. However, some may (and in some cases almost certainly do) represent extinct
species. Those that are probably trustworthy are given full accounts in this section. Those that are doubtful
and invalid are briefly discussed in Appendix 2, with full references for those requiring further
information.
14
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Introduction
THE APPENDICES
Appendix 1: Data-deficient taxa
The first appendix looks at those species and subspecies that are considered ‘data deficient’; that is, they
have not been recorded reliably for many years, but occur (or occurred) in extremely remote areas that
remain poorly known ornithologically (in some cases the breeding grounds may be completely unknown),
or are extremely skulking or cryptic. While not formally treated as extinct, these taxa may now be gone;
however, they may equally well survive (though doubtless in tiny numbers). I hope that this book will
inspire the next generation of field ornithologists to attempt to establish whether populations of these birds
cling on.
15
The world of zoological nomenclature is riddled with often obscure terms and abbreviations. This is espe-
cially apparent in the lists of synonymy which accompany each species entry. While we have tried to keep
this to a minimum, some of these terms do inevitably crop up. This list of the more commonly used terms
should help the reader interpret the text.
aff. From Latin affinis, meaning a species related to but not identical with the named species.
alien An alien, exotic or non-native species, occurring outside its normal distribution, which
has arrived there usually by human agency, either accidentally or deliberately.
allopatric A term used to describe two or more populations that are geographically separate.
carpometacarpus A bone in the wing that is made up of fused metacarpal bones and some carpal bones
(finger and wrist bones).
cf. From Latin confer, meaning to compare with.
conspecific Of or belonging to the same species.
contra From Latin. In contrast or opposition to, or against.
co-type See syntype
emendation An improvement or correction of something that is erroneous or faulty.
et al. From Latin et alii, when referring to more than two authors in a citation.
ex Meaning examined or taken from an often unpublished source.
exotic An alien, exotic or non-native species, occurring outside its normal distribution, which
has arrived there usually by human agency, either accidentally or deliberately.
family A taxonomic group containing one or more genera.
fide From Latin, meaning on the authority of, with reference to a publication or to a cited
published statement.
form An individual or group of individuals that vary from other members of the same taxon
in minor ways.
genus A term applied to a taxonomic category ranking below a family and above a species
(plural genera) and generally consisting of a group of species exhibiting similar characteristics.
holotype One particular specimen of an organism to which the scientific name is specifically
attached; usually shortened to ‘type’.
Holocene A geological age that began around 10,000 years ago at the end of the Pleistocene.
ibid. From Latin ibidem, meaning the same place. This is a term used to provide a reference
for a source that was previously cited. It has a very similar meaning to idem.
id. From Latin idem, meaning the same. This is used to replace the name of a repeated
author, thus removing the need to repeat the full citation.
introduced An alien, exotic or non-native species, occurring outside its normal distribution, which
has arrived there usually by human agency, either accidentally or deliberately.
lapsus An involuntary mistake made while writing.
mandible The lower jaw or bill in birds.
maxilla The upper jaw or bill in birds. Sometimes called the rostrum or upper mandible.
monospecific A genus containing only one species.
monotypic A species without subspecies.
MS Abbreviation of manuscript. MSS is plural.
nec. Latin, meaning ‘and not (of), nor (of)’.
17
nomen nudum The term is used to indicate a designated name which may well have originally been
intended to be a scientific name, but is not accepted as one because it has not been
published in the correct manner with an adequate description.
nomen dubium This term is used to indicate a taxonomic name that cannot be assigned with certainty
to any taxonomic group because the description is inadequate and the original spec-
imen may no longer exist.
neotype A specimen which has been selected as the type for naming a species or subspecies,
when the holotype (see type) is lost or has been destroyed.
nominate If a species is split into two or more races or subspecies, the first one to be identified or
described becomes the nominate race or subspecies.
partim Latin meaning in part. May be used in citations to indicate scientific names that are
vaguely synonymous.
phenotype The observable physical characteristics of an organism, such as morphology and
biochemistry, as determined by both genetic make-up and influence of environmental
factors.
Pleistocene A geological age that spanned the period from 1.8 million years to around 10,000 years
ago.
polytypic A species with two or more subspecies.
postcranial Referring to all parts of the skeleton other than the cranium (skull).
race Another word for ‘subspecies’.
scientific name The recognised name given to an organism, consisting of a genus and species. This is
also called the binomial name.
sic From Latin, generally placed inside square brackets [sic]. It is added after a quote or
reprinted text, and indicates to the reader that the passage appears exactly as in the
original source, which may include original errors.
skin A specimen that has been collected and skinned, and then the body stuffed. The skin
may be prepared flat for ease of storage (e.g. in a museum), or mounted (mounted skin)
in a life-like pose.
species A group of organisms that share common characteristics and are capable of breeding to
produce fertile offspring. A species is given a two part (binomial) name.
sp. An abbreviation for species, referring to an undescribed species.
ssp. An abbreviation for subspecies, referring to an undescribed subspecies.
subfossil Remains in which the fossilisation process is not complete. The material described as
subfossil in this book is primarily due to it being of a relatively young age.
subspecies Taxonomic rank below species; two subspecies differ from each other but can still inter-
breed, though they usually do not due to geographic isolation. Subspecies are given
three-part (trinomial) names.
sympatric A term used to describe two or more species or populations that occur in the same
geographical area.
synonym A scientific name that is or was used for a taxon that also goes by another scientific
name.
synonymy A chronological list or record of the scientific names that have been applied to a species.
syntype An additional type specimen or specimens from which a holotype has not been desig-
nated. The syntypes collectively represent the type name.
tarsometatarsus The bone in the lower part of a bird’s leg made up of fused metatarsal and tarsal bones
(foot bones).
taxon (plural taxa) A taxonomic unit in the Linnaean system of biological classification e.g. order, family,
species, subspecies.
teste From Latin, meaning by the evidence of.
tibiotarsus A long bone situated between the femur and tarsometarsus made up of the tibia fused
with proximal bones of the tarsus.
type One particular specimen of an organism to which the scientific name is usually
attached.
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The accounts
Struthionidae (Ostriches)
The Ostrich Struthio camelus is the world’s largest living bird and is presently found throughout s ub-Saharan
Africa. It was formerly much more widely distributed, occurring eastwards to the Middle East. There are
five recognised subspecies. Three of the four extant races are not considered threatened, but the North
African Ostrich S. c. camelus has declined drastically and is considered to be in serious danger of extinction.
Over-hunting is the primary reason for the Ostrich’s decline.
The Syrian Ostrich Struthio camelus syriacus once inhabited open semi-desert and desert plains in a contin-
uous range across the Arabian Peninsula and the Middle East. Its last stronghold was north to 33°N, east to
Kuwait, to Jordan, and the Syrian Desert south into the Arabian Peninsula (Jennings 1986); it also may have
occurred in southern Israel and the Sinai (Shirihai 1996).
Rothschild (1919) described the Syrian Ostrich at a meeting of the British Ornithologists’ Club on May
14, 1919, at a time when the ostrich was apparently still common, and hunting had little effect on numbers.
Arabs originally hunted Syrian ostriches on horseback using guns and falcons (mind-boggling in itself), but
this technique was extremely difficult as the ostriches were wary and could not be approached closely;
Cheesman (1923) reported that a skilled Sleb (nomadic tribes from these deserts) hunter considered himself
lucky if he killed more than two or three a year. Potts (2001) suggested that the Syrian Ostrich was not
hunted for its meat but primarily for its feathers, and in certain areas for its fat. The eggs were prized by the
Arabs, who considered them a delicacy. The shells were also used for decoration, and were frequently to be
found hanging in mosques. Until the early 20th century, hunting pressure did not appear to have any
marked effect on ostrich numbers. Meinertzhagen (1954) said that the bird was common in the deserts until
1914, but precision rifles arrived in Arabia during the First World War, and later motor vehicles, so the birds,
which must have suddenly become rather easy targets, were doomed.
The last record was of an individual killed and eaten some time between 1940 and 1945 by Arabs near the
oil pipeline in the area of Al Jubail, in north-east Saudi Arabia (Greenway 1967); another two birds were
supposedly shot in 1948, and there is a dubious second-hand record of a dead female found in 1966 at the
Wadi al-Hasa, near Ghor el Safi in Jordan (Shirihai 1996). The carcass was washed away by the floodwaters
of the Jordan, so no specimen was available to confirm the observation.
Meinertzhagen (1954) claimed to have seen a dozen or so Syrian Ostriches in captivity in Amman and
Damascus in 1920, and wanted to buy some to bring back to London Zoo, but funds could not be raised.
This was a pity, for this subspecies might easily have survived and increased in captivity.
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Habits
Burckhart in Prater (1921) recorded the breeding habits:
This bird breeds in the middle of winter and lays from twelve to twenty one eggs. The nest is generally made at
the foot of some isolated hill. The eggs are placed close together in a circle, half buried in sand, to protect them
from rain, and a narrow trench is drawn around, whereby the water runs off. At ten or twelve feet from this
circle, the female places two or three other eggs which she does not hatch, but leaves for the young ones to feed
upon immediately after they are hatched. The parent birds sit on the eggs in turn; and while one is so employed,
the other stands keeping watch on the summit of the adjacent hill.
In historical times, the Syrian Ostrich was most numerous in Al Jawf Province in northwest Saudi Arabia,
near the Jordanian border (Robinson & Matthee 1999). The birds were associated with herds of the now
extinct Saudi Gazelle Gazella saudiya and Arabian Oryx Oryx leucoryx, which was extinct in the wild (but has
been reintroduced in a handful of places across the region).
Casuariidae (Emus)
The Emu Dromaius novaehollandiae is the second largest living bird and is confined to Australia, where it has
been divided into three subspecies. It was once persecuted for crop damage and hunted for food, but
although it has declined in some areas, it remains generally unthreatened. However, the populations that
once occurred on Tasmania, King and Kangaroo Islands were all exterminated in the 19th century.
The history of the two extinct emus of King and Kangaroo Islands is exceedingly confused (see Pfennigwerth
2010 for a comprehensive review). Both species were discovered on the voyage of Captain Matthew Flinders
in 1802, but they were only mentioned in passing and not considered different from those of the mainland
(Flinders 1814). Later the same year, Captain Nicholas Baudin visited the islands, and the naturalists on his
expedition, Péron, Leschenault, Bailly and Le Sueur, made observations on the flora and fauna (Péron 1809;
Horner 1987; Baudin 2004). The expedition collected at least three live adult emus on King Island, and two
or three adults on Kangaroo Island (Jouanin 1959; Baudin 2004). Remarkably, after a voyage of 15 months or
so, two of the emus, a possible male and a female, arrived alive at the Jardin des Plantes, Paris (Jouanin
1959; Pfennigwerth 2010); the female died in April 1822 and is preserved as the unique Paris skin. The male
– the last of his kind – died in May 1822, and is now the Paris skeleton. A skeleton in Florence, also believed
to be a male, was lost for many years until rediscovered in 1900 (Giglioli 1901). Contrary to previous reports,
this skeleton is a composite from two different individuals, and does not represent the bones of the Paris
skin. Newton (1900), Giglioli (1907) and Mathews (1910b) believed that a fourth specimen existed in the
Liverpool Museum, but it appears that this may have been a juvenile mainland emu.
Confusion over the provenance of the island emus began almost as soon as they were discovered
(Pfennigwerth 2010). Brasil (1914) argued that because the Baudin expedition visited King Island from
10–23 December 1802, when the weather was very bad, the expedition members remained in camp and did
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variation in Dromaius novaehollandiae, so Heupink et al. recommend that the King Island Emu deserves
subspecific status only. However, del Hoyo & Collar (2014) retain it as a full species.
Habits
Péron (in Milne-Edwards & Oustalet 1879; Pfennigwerth 2010) gained a considerable amount of informa-
tion on the habits of the King Island Emu from Daniel Cooper, who was evidently a careful observer:
The bird was generally solitary, but gathered in flocks of ten to twenty at breeding time, then paired and
wandered off in pairs. In plumage it was black in the adult, greyish in the young. The sexes were alike, but it was
suggested that the male was perhaps “brighter”, and was also very slightly larger than the female. There was no
seasonal variation in plumage. The largest birds were up to four and a half feet long and the heaviest weighed
from 45 to 50 pounds. They fed on berries, grass and occasionally seaweeds, and their feeding times were mainly
in the mornings and evenings. They were seen to run quite quickly, but as they were fat they ran much less
quickly than the birds of the mainland. They swam well, but only when obliged to do so. They frequented lagoons,
and tended to seek shade rather than open country. The nest was usually situated near water and under the
shade of a bush. It was constructed on the ground of sticks, and lined with dead leaves and moss; it was oval in
shape and not very deep. Seven, eight or nine eggs were laid, always on 25 to 26 July, a remarkable instance of
breeding synchronisation in a bird in which it is difficult to see the selective advantage. Incubation was by the
female, possibly also by the male as the latter apparently developed a brood patch. Neither bird left the nest, the
non-incubating parent remaining nearby. The chicks were striped and left the nest two to three days after
hatching.
The botanist Robert Brown, who accompanied the Flinders Expedition, examined the dung of a King Island
Emu and noted that it comprised mainly of the berries of Prickly Cedar Leptecophylla juniperina (Vallance et
al. 2001).
As with the King Island Emu, it was Flinders (1814), while visiting Kangaroo Island in 1802, who made the
first mention of Kangaroo Island Emus. The Baudin expedition visited the island later the same year and
collected some specimens, but unlike the King Island birds, their present whereabouts is unknown. Flinders
had found the island uninhabited, but commented on the extraordinary evidence of previous conflagra-
tions, particularly fire damage, and these may have resulted in the extermination of the birds. It seems clear
from the extracts quoted here (from Flinders’s narrative) that he did not collect any emus:
Not less than thirty emus or cassowaries were seen at different times; but it so happened that they were fired at
only once, and that ineffectually. They were most commonly found near the longest of the small beaches to the
eastward of Kanguroo Head . . . where some little drainings of water oozed from the rocks . . . The bay is perfectly
sheltered from all southern winds; and as there were several spots clear of wood near the beach, it is probably
[there] that the kanguroos, and perhaps cassowaries, might be numerous.
Specimens
The first subfossil remains of Kangaroo Island Emu were discovered in 1903 at The Brecknells, an area of
sandy hills in Cape Gantheaume, and others have subsequently been found (Parker 1984a). Unlike the
conclusions of Giglioli (1901) and Morgan & Sutton (1928), who did not notice any differences between the
King and Kangaroo Island birds, Parker considered the Kangaroo Island Emu to be quite distinct. Compared
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with the King Island birds, the tibiotarsus and tarsometatarsus averaged greater in total length, and the
Kangaroo Island birds also exhibited no apparent sexual dimorphism.
An adult skin in the Geneva Museum was presented in 1827 by M. Moricaud. This was believed to come
from Kangaroo Island. But according to Jouanin (1959) it is a female Australian mainland Emu, possibly the
bird brought back by Le Naturaliste (see King Island Emu, above); this bird died in Paris Zoo in 1809. Thus
no skin specimens of the Kangaroo Island Emu are known to exist; an illustration (plate 36) accompanying
the account of Péron (1807), which was executed by Charles-Alexandre Le Sueur, artist on Baudin’s voyage
(see Milne-Edwards & Oustalet 1899), remains the only unequivocal depiction.
A unique egg
An egg, now in Tring, was collected by Dr John Dunmore Lang, a minister of the Church of Scotland and
principal of the Australian College at Sydney, apparently in 1830. It is accompanied by a note that reads:
Dr. Lang of the Sydney College brought this egg from Kangaroo Island. It was found hanging from the beam in a
Sealer’s hut at Nepean Harbour and had recently been brought in from the bush.
The egg subsequently passed to the collector George Dawson Rowley, and when his collection was sold at
Stevens Sale Rooms in November 1934, it was acquired by George E. Mason, who gave an account of its
history (Mason 1936).
If Mason’s information on the egg is correct, the bird must have died out between 1830 and 1836, before
formal white settlement in 1836, having been rare for some years before. The settlement comprised colonists
of the South Australian Company, but sealers lived on the island prior to this; they certainly collected eggs
and probably killed birds for food. Mainland emus were introduced in 1826 and later, so hybridisation may
have occurred as well. Between 1802 and 1830, the only recorded sighting seems to have been from one
Captain George Sutherland, who saw some in the south of the island near Hawk’s Nest Station in 1819
(Condon 1967), at a time when the island appears to have been covered with thick impenetrable scrub.
Although Tasmanian Emus had been recorded by Gunn (1852) in 1837, who presented two specimens to
Tring, it was not until the early 20th century that they were regarded as specifically distinct (Le Souëf, 1904a,
b). Gunn thought that Tasmanian Emus were smaller than mainland ones, but bones found subsequently
indicate that there the differences were slight. Le Souëf (1904b) examined early references to emus in
Tasmania, noting that they were formerly plentiful, and were often recorded by early settlers. He mentioned
that in 1803, the Rev. R. Knockwood recorded a bird with six young in his diary, and John Meredith informed
Le Souëf that he had seen a pair with six young in 1851. Mr. Ransom of Killymoon hunted emus with dogs
in about 1840, and apparently Captain Hepburn of Roy’s Hill or St. Paul’s Plains (or both) found a nest with
eight or nine eggs, which were subsequently hatched by a turkey. From these other birds were bred,
allowing Hepburn to build up a colony. Colonel William Vincent Legge (1907), who was married to a
Tasmanian girl, recorded some specimens captured in 1845:
One of the shepherds of the estate, H. Wyburn, was resident at the bogs, and used not infrequently to bring
eggs to the house, and about the year 1845 succeeded in capturing two young birds, which were conveyed to
Rockfort and reared in the goose-yard. They lived about the homestead for several years, and were tame and
mischievous, coming to the open ‘French’ windows of the dining-room to be fed, thrusting their heads into
the room at times.
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Extinction
During the early years of the 19th century, Tasmanian emus were both hunted for food and persecuted as a
pest. The extensive use of fire to clear open grassland and scrub for agriculture had made them very scarce
by the 1830s (Gould 1848; Dove 1924). Dove (1926) further supplied important information about the prob-
able ultimate reason for the emu’s extinction. Soldiers were employed during the 1830s to shoot emus in
order to provide provisions for the troops and prisoners. They were paid 1s 6d per pound of flesh by the
government, and a single private assisted by two prisoners could average 1,000 pounds of emu-flesh per
month. It is unlikely that the species could tolerate such intense slaughter long-term, and by around 1845
the last unequivocal records were made of wild birds (Gunn 1852; Dickison 1926; Dove 1926). Furthermore,
by the early 1850s, Australian mainland emus had been imported from Victoria, and these may have hybrid-
ised with any remaining native birds.
Reports were made of captive emus, especially of a pair kept by the Van Diemen’s Land Company in a
park at Circular Head, which bred and reared six young in 1845 (Dove 1924, 1926; Dickison 1926), but these
may also refer to imported birds. A pair of emus were said to have been given to a Baron von Steiglitz, of
which one survived until 1873, and an emu that lived in the city park in Launceston, Tasmania, in the 1870s
was supposed to be the last of the Tasmanian species. The specimen was not preserved, however, so it was
almost certainly one of the mainland imports, and von Steiglitz’s birds probably were, too.
A leg of a Tasmanian Emu is now in my possession, and so far as I can judge from it, as a very imperfect spec-
imen, there are differences in the arrangement and size of the scales, which may justify the separation of the
Tasmanian emu from that of New Holland. Still further research and examination are necessary.
This leg was lost for about 70 years, but was re-examined by Scott (1923), who agreed with Gunn that the
scales were different from those of the Australian mainland species. Another skin specimen held in
Frankfurt was thought to have been a Tasmanian Emu, but the provenance is in error (Steinbacher 1959).
A possible subspecies (or maybe a separate species) may have once occurred on Flinders Island. The
Australian ornithologist Dominic Serventy was informed by a resident of the island, Frank Jackson, that
fragments of emu eggshell had been found (Serventy 1967; Hume & Walters 2012). If they did once occur
there no other evidence is known; this was confirmed by a palaeontological survey in 2015 conducted by
the author, palaeontologist Lorna Steel and cave expert Greg Middleton (Hume et al. in press).
Habits
Little is known about the habits of the Tasmanian Emu. Legge (1907) stated that the bird inhabited ‘remote
hill marshes and upland plains, and also open country on the littoral of the north and east coasts’. His
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father-in-law, Major W. Gray, owned an upland moor called Kearney’s Bogs, where emus occurred regu-
larly in the 1840s. Gunn (1852) recorded some remarks made by a Mr J. Hepburn to J. Milligan:
Mr. J. Hepburn informs me [Milligan] that the booming noise is not peculiar to the female and that the male
bird does, though not frequently, make the same sound. The Tasmanian Emus share the toils of incubation
between the sexes, but upon the mother devolves the care of bringing up the young brood, to which the male
parent, for the most part, displays an unnatural and most bitter antipathy.
Apterygidae (Kiwis)
The kiwi is the smallest living ratite and restricted to New Zealand, where it is nowadays divided into five
species. The male incubates the pale green or ivory white egg for a period of up to 76 days, one of the
longest of any bird. The egg itself is the largest compared with the size of the adult bird, and accounts for
approximately 26% of the body weight. Kiwis have suffered from o ver-hunting, habitat loss and predation
from introduced mammals, and some species are in critical danger of extinction.
The Little Spotted Kiwi is the smallest kiwi species. It is divided into three subspecies (Mathews 1935b);
these once occurred on South Island (A. o. owenii and A. o. occidentalis) and North Island (A. o. iredalei),
though these are considered to represent a single species by some authorities (e.g. Herbert & Daugherty
2002).
The South Island subspecies were distributed from Marlborough south to southern Fiordland, whereas
the North Island form is known from just one specimen, taken in 1875 from the Tararua Range. It was
collected by Morgan Carkeek, of the Survey Department, in snow-grass at an elevation of 3,000 feet (914m),
on Mount Hector at the head of the Hutt River (Buller 1876). Carkeek noted that at a higher elevation the
bird was comparatively abundant, but he also observed it below the snow-line, where it frequented ‘mossy
places in the bush free from undergrowth’. The South Island Little Spotted Kiwi was considered common in
the 1870s (Sales 2005), but numbers plummeted as demand for specimens from European museums
increased. The birds were regularly hunted by prospectors and farmers for food, and the flightless, terres-
trial kiwis were extremely vulnerable to predation by feral dogs and cats; in addition, the kiwi generally
lays only one egg, so reproductive rates are low. During the 1890s and early 1900s, and in an effort to save
the species, some South Island birds were captured and released on off-shore islands (BirdLife International
2011). They have also been released on mainland North Island, and their future appears to be safe.
The North Island subspecies was less fortunate, and has not been seen since 1875. The combined effects
of habitat change, predation by introduced mammals, and o ver-hunting extirpated the population before
protective measures could be put in place. The race is now considered to be extinct.
Habits
Little was recorded about the North Island race in life, but on Kapiti Island Little Spotted Kiwis prefer short
flax or seral scrub and mature forest. They nest in an excavated burrow, in a hollow log, under tree roots or
in dense vegetation, where one or sometimes two eggs are laid. They are entirely nocturnal and forage by
scratching with their strong feet and claws and probing in leaf litter and soft soils for insects, worms and
deep-burrowing larvae. They also occasionally take fruit. Their eyesight is very poor so they rely on an
25
acute sense of smell to locate prey items. The call is somewhat similar to the Brown Kiwi Apteryx australis,
but is more tremulous.
The Giant Elephant Bird was easily the heaviest bird known, heavier even than the largest moa, Dinornis
robustus. It was approximately 3m tall, and may have weighed up to 438kg, nearly half a metric ton (Amadon
1947). However, Worthy & Holdaway (2002) suggest this is a vast overestimation based on measurements
on the robustness of the femur, and that in keeping with the estimated weights of the largest moas, A.
maximus was more likely to have weighed around 275kg. Nonetheless, this was still heavier than D. robustus.
The Giant Elephant Bird was flightless with only rudimentary wings; the sternum had no keel for the
attachment of wing muscles. The tibiotarsus was longer than the tarsometatarsus, which suggests that the
birds were not agile but best described as ponderous browsers and grazers. The bill was long and deep with
large narial openings, so the birds probably had a w ell-developed sense of smell. The presence of pits and
depressions on the frontal bone of the cranium suggest that the elephant bird had a small crest (Andrews
1896c), as did some species of moa with a similar cranial characteristic (Parker 1892). The egg was the largest
known, with a volume the equivalent of 160 chicken eggs (Hawkins & Goodman 2003).
Extinction
A number of hypotheses have been proposed for the extinction of the elephant birds. It is possible that intro-
duced domestic fowl brought diseases, against which the elephant birds were completely defenceless
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(MacPhee & Marx 1997), but this seems unlikely. They appear to have been vulnerable to over-hunting and
perhaps egg harvesting; knife cut-marks are present on some of the remains and eggs have been found in
association with archaeological midden sites. This may have been coupled with habitat modification and
perhaps a natural aridification of the island caused by climate change (Burney et al. 1997; Tovondrafale et al.
2014). Either way, the entire elephant bird family is now extinct. Unlike the moas of New Zealand, at least one
species may have survived until the 17th century, a time when Europeans became established on the island.
The first French governor of Madagascar, Étienne de Flacourt (Flacourt 1658), made the following report:
The ‘vouropatra’ a large bird which haunts the Ampatres and lays eggs like the ostriches; so that the people of
these places may not take it, it seeks the loneliest places.
The name Vorompatra (or Vouropatra) is an ancient Malagasy name for the bird, and translates as ‘bird of
the Ampatres’. The Ampatres are today known as the Androy region of southern Madagascar (Pearson &
Godden 2002). The species Flacourt was referring to cannot now be ascertained. No more trustworthy
reports were made, so the last of the elephant birds must have died out shortly after Flacourt’s account.
Habits
Although the elephant birds disappeared before anything was recorded about their habits, it is probable
that they were forest inhabitants, perhaps ecologically similar to the cassowaries Casuarius sp. of northern
Australia and New Guinea. Elephant birds may have been important seed-dispersers of forest trees and
shrubs (see Dransfield & Beentje 1995). An isotopic study of Aepyornis eggshell and bone has shown that
these birds generally fed on C3 plants (trees, shrubs and forbs, or herbaceous flowering plants), and also on
succulents (CAM plants), but not on C4 plants (tropical grasses) (Tovondrafale et al. 2014). This strongly
indicates that elephant birds were browsers and not grazers. Elephant birds also had a strong conical bill,
which may have been an adaptation for foraging underground tubercles or for breaking hard fruits
(Goodman & Jungers 2014). ‘Nothing is known about their breeding habits, but a histological analysis by
Ricqlès et al. (2016) has shown that Aepyornis maximus exhibited lines of arrested growth in certain bones,
indicating that the young took several years to reach maturity. Nothing is known about their breeding
habits. Although eggs and egg shell fragments are abundant in certain localities, fecundity may have been
low and perhaps only one or two eggs were laid each year. The huge numbers of shell fragments that have
been found in sand dune deposits in the arid south-west (Grandidier 1868) suggest that this area may have
27
been an important breeding ground for the birds, but eggshell fragments have also been found elsewhere
on Madagascar.
Only two predators were large enough to take elephant bird eggs and young. Juvenile elephant birds
may have provided food for the Malagasy Crowned Eagle (see p. 91), whereas eggs and chicks may
have provided food for the largest Madagascan carnivore, the now-
extinct Giant Fossa Cryptoprocta
spelea.
Hildebrandt’s Elephant Bird was described from a tarsometatarsus, but a nearly complete skeleton was
subsequently found, which is now in Paris. Other mounted specimens are in Antananarivo, Madagascar
and Tring. It was the smallest member of the family, and was approximately the size of a small cassowary
Casuarius sp. at about 1.0–1.2m, but it could probably reach up to 1.5m when browsing. It was a robust
species, and may have weighed up to 65kg.
This dubious species is known from a single femur, and differed only in its less robust size. If more speci-
mens become available, it is likely to fit well within the individual variation of Hildebrandt’s Elephant Bird
A. hildebrandti.
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This species is known from a number of elements including the femur, tarsometatarsus, cranium,
mandible, coraco-scapula, humerus, tibiotarsus and pelvis. It was substantially larger than Hildebrandt’s
Elephant Bird Aepyornis hildebrandti, rivalling some of the larger moas in size and weight, but smaller
than A. maximus.
Considering the discovery of extreme sexual dimorphism in the moas in which the female could be up to
1.5 times larger than the male (Bunce et al. 2003), the smaller specimens described as new species – i.e. A.
hildebrandti, A. medius and A. gracilis – might all represent males, or clinal variation of A. maximus.
The genus Mullerornis comprised three or four species, all of which were smaller than their closest rela-
tives, the elephant birds of the genus Aepyornis. Milne-Edwards & Grandidier (1894) and Lamberton
(1934) separated them on size and certain characteristics of the bones, comparing them in size with the
cassowaries Casuarius spp. One species, M. betsilei, is known from Pleistocene deposits found only in the
centre of the island. Lamberton (1934) provided an exhaustive study of the osteology of these birds, and
described a new species, M. grandis, which was the largest of the genus, but there are doubts concerning
its validity.
Mullerornis differed in being slightly more gracile in the dimensions of the bones compared with the
smallest members of Aepyornis, especially A. hildebrandti (Andrews 1904, Lamberton 1934), but all have
been described from rather inadequate p ost-cranial remains, the differences between which are poorly
defined. Like the moas of New Zealand, this was often based on size alone. Mullerornis rudis was described
from a tibiotarsus and tarsometatarsus and compared with M. betsilei, the smallest and most gracile of the
genus, it was more massive, with a remarkable widening of the distal end of the tarsometatarsus and large
tendinal openings (Milne-Edwards & Grandidier 1894). A subfossil bone possibly belonging to a species of
Mullerornis has been radiocarbon-dated to c.1260 bp, suggesting that the bird was still extant at the end of
the first millennium (Burney et al. 2004). It presumably died out for the same reasons as Aepyornis.
This species was originally known from a tibiotarsus found in coastal deposits in the south-west of
Madagascar. Remains were later found in Ambolistra, Belo and Ampoza in the south-east, and Antsirabe
and Ampasambazimba in the centre of the island (Lamberton 1934), which include a partially complete
individual. The Coastal Elephant Bird M. agilis was smaller than the Ostrich Struthio camelus, being slightly
larger than the Australian mainland Emu Dromaius novaehollandiae. The legs of the M. agilis were massive,
but less so than the Robust Elephant Bird M. rudis, and the proportions of the limbs differed from those of
the Emu. Although the femur and tibiotarsus were approximately equal in total length, the tarsometatarsus
was short and broad in M. agilis (and is long and slender in the Emu), suggesting that like Aepyornis,
Mullerornis were not agile birds. The sternum was completely flat without a keel, and wings were probably
absent.
The cranium, although differing in certain characters, was similar to Aepyornis, but the bill has yet to
be found. Lamberton’s monograph on Mullerornis defines distinct osteological differences between this
29
genus and Aepyornis, but the differences are less striking between the various species of Mullerornis. If
more material becomes available, M. agilis may well turn out to be referable to individual variation
in M. rudis.
Habits
Being smaller than Aepyornis, the Mullerornis elephant birds doubtless had a different ecology, and perhaps
utilised different food sources. E
ggshells referable to Mullerornis have been found in a number of localities,
but not in huge quantities (as is the case for Aepyornis); the eggs differed in being much thinner and in
having a smoother surface. There is no indication of clutch size, but it was probably small.
The subfossil remains of the genus Dinornis have been previously divided into as many as 40 different
species from both the North and South Islands of New Zealand, the majority of which were separated by
size alone. However, using genetic sex determination in ancient bones, Bunce et al. (2003) showed that only
two species of Dinornis actually existed, and that the size differences were attributed to sexual dimorphism
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Dinornithidae
in which the female was approximately 1.5 times the height and 2.8 times the weight of the largest males.
Not only did Dinornis exhibit the greatest sexual size dimorphism in any bird, it also varied in size
depending on altitudinal, temporal and biogeographical factors. The two species thought to have existed
are D. novaezealandiae of North Island and D. robustus of South Island (Bunce et al. 2009); even with this
present understanding, it appears that there are some distinct lineages within the Dinornis complex that
await further analysis (Baker et al. 2005).
The extinction of the moas can be almost certainly attributed to the actions of humans. The proto-Maori
arrived sometime before 1300ad, but most if not all genera were extinct prior to the European discovery of
New Zealand in 1642 by Abel Tasman (Gill & Martinson 1991; Vickers-Rich et al. 1995). Holdaway & Jacomb
(2000) have postulated that the disappearance of all moas may have occurred in less than a century.
Furthermore, Holdaway et al. (2014) has shown that only a small population of Polynesians, with numbers
no greater than 2,000, was necessary to completely exterminate moas in the habitable areas of eastern South
Island. The rapidity of their demise can only be explained by human over-hunting and habitat alteration;
one of the few documented cases of the ‘blitzkrieg effect’ in which hunting of adult birds far exceeds fecun-
dity (Martin 1967; Diamond 2000). The birds were slow-growing with low rates of reproduction, not
reaching sexual maturity until at least 10 years old of age (Turvey et al. 2005). Archaeological evidence from
Maori middens has shown that the sudden onslaught of hunting and e gg-collecting proved disastrous to a
family of birds that had hitherto evolved in the absence of mammalian predators and had lost the necessity
for fast reproductive rates. When humans arrived on New Zealand the moas were doomed.
Habits
The North Island Giant Moa favoured wet, lowland, podocarp and Nothofagus beech forests, was rare or
absent in drier areas, and an infrequent visitor to the subalpine zone (Worthy & Holdaway 1993). An
abundance of Dinornis and other moa remains, including stomach contents and coprolites (preserved
droppings) (Burrows et al. 1981; Horrocks et al. 2004), provides a large amount of information about their
probable habits. The North Island Giant Moa foraged by snipping and plucking twigs, leaves and berries
from a diverse range of low trees, shrubs and small plants up to 3m off the ground, with its broad flat-
tened bill. Preserved stomach contents found at Pyramid Valley contained stems and twigs of Olearia
and Coprosma (Gill & Martinson 1991). The gizzard contained large numbers of stones to aid in the
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breakdown of food. Hartree (1999) discovered a number of probable nesting sites used by at least four
species of moa. They were all under the shelter of limestone or sandstone rocks, and the birds probably
used fallen logs and dense vegetation as well. The nest was a small scrape in the ground (Owen 1879;
Hartree 1999) lined with moa-snipped twigs (Hutton & Coughtrey 1875), or soft twigs, feathers and
grasses (Hartree 1999). According to Hartree, it appears that only one egg was laid, and the nest was
only used once. All of the nest localities were found between sea level and 820m in altitude. As females
accounted for most of the moa bones preserved in swamps and deposited in museums, Huynen et al.
(2010) suggest that the females went out foraging and the smaller males incubated the eggs, as seen in
the incubation behaviour of kiwis Apteryx sp. Huynen et al. have also shown that the moas that exhibit
extreme sexual dimorphism (i.e. Dinornis, Euryapteryx and Emeus), laid eggs with the thinnest shells of all
known birds, and thus the small size of the male may have been an adaptation to prevent damage to the
eggs while incubating.
Moa egg shell fragments are up to 2mm thick, much thinner than that of an ostrich (Gill & Martinson
1991). Eggs attributed to this species are the largest moa eggs known. All moa species were K-strategists –
species with low fecundity and slow maturation rates (Turvey et al. 2005), and were probably long-lived
birds, possibly reaching 50 years in age (Tennyson & Martinson 2007). Tennyson & Martinson further
suggest that only one or two eggs were laid each breeding season, and as they were large relative to those of
other birds, the incubation was probably long, being anything up to two months; the chicks were well
developed on hatching and able to feed immediately.
Using mitochondrial DNA, McCallum et al. (2013) identified distinct populations within species, espe-
cially in the widespread Little Bush Moa Anomalopteryx didiformis and Coastal Moa Euryapteryx curtus; the
latter species had distinct populations on North and South Islands. By contrast, in North Island Giant Moa
Dinornis novaezealandiae and South Island Giant Moa D. robustus, genetic diversity was most prevalent in the
east of the North Island and the west of the South Island respectively. Where these regions are characterised
by rich mountainous terrains and able to support long-term diverse habitats, McCallum et al. suggest that in
the North Island, the distinct populations of D. novaezealandiae and Mantell’s Moa Pachyornis geranoides
were highly structured and appear to have occupied the same geographic location for hundreds of thou-
sands of years. Separate populations of these two species occurred on the east and west coasts and were
divided by mountain ranges in the centre of the island.
This was the largest species of moa. It was taller than the Giant Elephant Bird Aepyornis maximus of
Madagascar, but it was much less massive. Most abundant in eastern South Island, this moa appeared to
prefer the drier, rain-shadow forested areas (Worthy & Holdaway 1993), where it occupied relatively open
shrubland or forest margins (Worthy 1989). This habitat was in natural decline during the Holocene, so the
birds were forced into less suitable habitat and thus were more susceptible to human hunting and habitat
destruction.
Two dessicated specimens of South Island Giant Moa found in Otago have the large scales of the lower
leg preserved, and show that it had prominent foot pads (Tennyson & Martinson 2007).
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Habits
Many birds need to ingest grit or stones to provide the necessary grinding action to allow the mastication of
tough vegetable matter. Moas were no exception and gizzard stones have been found associated with a
number of moa species. The stones were usually quartz pebbles ranging in size from 10mm to large stones
exceeding 110mm in length (Wood 2007). Individual Dinornis gizzards could sometimes contain several
kilograms of stones (Worthy & Holdaway 2002).
Analysis of gizzard contents have shown that the South Island Giant Moa fed on twigs and berries, plus
occasional herbs and grasses (Tennyson & Martinson 2007).
Fossil moa footprints have been found on numerous occasions, and stride and inferred speed can be
extrapolated as long as the hip height of the bird can be determined. Alexander (1983), Duncan & Holdaway
(1989), and Worthy & Holdaway (2002) concluded that moas had an average speed between 3.8 and
4.34km/h, depending on the size of the species involved. They were slow-moving animals. Evidence from
trackways, which are likely to have been made by moas, suggest that these w ell-worn trails were made by
the birds moving through dense vegetation (Horn 1989), but this interpretation is still open to debate.
Adult moas probably had one natural predator, the huge Haast’s Eagle Harpagornis moorei (see p. 88). The
presence of puncture marks in the pelvises of some moa specimens (Worthy & Holdaway 2002) suggest that
this eagle pounced on the birds from behind, inflicting serious wounds to the body, and perhaps the coup de
grâce was a bite to the back of the neck.
However, Rawlence et al. (2011) suggest that not all puncture marks are derived from Haast’s eagle. The
endemic Kea Nestor notabilis, a parrot noted for its ability to attack live and dead sheep in order to reach fat
deposits beneath the skin, would have also had the ability to scavenge dead or dying moas, especially those
trapped in swamps. The long and powerful beak of the kea could just as easily puncture a moa’s pelvis.
Anomalornis gracilis Hutton 1897, p. 546, pl. 47, fig. A (Opito, near Mercury Bay, Coromandel). Junior secondary homonym of
Dinornis gracilis Owen 1854
Anomalornis didiformis (Owen); Hutton 1897, p. 547
Anomalornis owenii (Haast); Hutton 1897, p. 549. Unjustified emendation
Anomalopteryx parvus (Owen); Oliver 1930, p. 45
Pachyornis owenii (Haast); Archer 1941, p. 44. Unjustified emendation
Specimens Subfossil remains are in Auckland, Canterbury, Manawatu, Wanganui and Wellington, New Zealand; and
London. Mummified remains are in Invercargill, New Zealand.
Status Extinct. Known only from subfossil remains. Date of extinction c.1400.
Range North, South and Stewart Islands, New Zealand.
This was one of the smallest species of moa, standing approximately 50–90cm tall and weighing 26–64kg
(Bunce et al. 2009). It was a slender species with a stout, rather robust bill. The jaw musculature was large,
and this species could place considerable pressure when biting. With overlapping cutting edges on the
mandible, it could snip stems, twigs and leaves with great force (Worthy & Holdaway 2002). This species
once occurred on both North and South Islands, being more numerous on North Island. On South Island its
remains have been found in north-western and southern parts of the island. It preferred high-rainfall
Nothofagus beech forests.
Using molecular evidence derived from moa coprolites, Wood et al. (2012) showed the food preferences
of the Little Bush Moa consisted mainly of fibrous material taken from trees and shrubs in the forest under-
storey (confirming the work of previous authors). The Little Bush Moa was restricted to lowland
closed-canopy forest, and Wood et al. noted further that its feeding ecology was shared by at least two other
species of moas, the Upland Moa (see p. 38) and South Island Giant Moa (see p. 32); both of these latter
species also grazed on herbs in unforested vegetation communities (Horrocks et al. 2004; Wood et al. 2008).
The Coastal Moa was another small species that exhibited extreme sexual dimorphism (Worthy 1987).
Males stood about 51cm tall and weighed approximately 12–20kg. The females were about 103cm tall and
weighed up to 109kg (Bunce et al. 2009). It had a blunt bill that was relatively weakly constructed (Worthy &
Holdaway 2002) so it probably foraged on softer plant material, including fruits and berries, and may have
eaten insects. It was a lowland inhabitant and occupied dry r ain-shadow forest, coastal dunelands, open
grasslands, and shrublands. Unlike all other moas, the Coastal Moa was found on both North and South
Islands, but the populations were undifferentiated. Furthermore, remains of ths bird are most numerous in
coastal deposits, which suggests that there may have been population interchange comparatively recently,
via Pleistocene land-bridges (Bunce et al. 2009).
In Euryapetryx, Emeus and probably all of the smaller genera of moas, birds exhibited tracheal elongation
in which the trachea was up to 1m in length and formed a large loop in the body cavity (Worthy & Holdaway
2002). This characteristic is also found in some other bird families, notably swans (Anatidae), cranes
(Gruidae) and guineafowl (Numididae), all of which are extremely vocal species. Worthy & Holdaway
speculate that the moas were capable of deep, resonating calls that could travel for long distances, and the
vocalisations may have been used to signify territories, attract mates, or to make contact calls in dense
vegetation.
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Emeidae
Gill & Martinson (1991) note that two eggs attributed to the Coastal Moa were discovered by a Mr L. J.
Mathews in 1900 at Doubtless Bay, Northland. Both eggs were in a bank where a stream cut through a sand
dune. One was unbroken and measured 120mm x 91mm, with a capacity of around 500ml, and estimated to
weigh 500g when fresh. This is the smallest known moa egg. By comparison, another egg discovered in the
1850s at Kaikoura, South Island and attributed to Dinornis is the largest ever found. It measures 240mm x
178mm with a capacity of almost 4 litres, the equivalent of 60 hen’s eggs, and would have weighed around
4kg when fresh (Gill & Martinson 1991). Subfossil remains and eggs of the Coastal Moa are common in
archaeological deposits; thousands of the birds and their eggs were eaten by the Maori (Tennyson &
Martinson 2007). As this species was extremely vocal, Tennyson & Martinson suggest that it was a flocking
species, which would have made it easy to locate and kill in large numbers.
The nomenclature of this species is extremely confusing. Archey (1941) and Oliver (1949) referred this
species to the genus Euryapteryx, without actually examining the bones. Worthy (2005b) resolved the
taxonomy and Euryapteryx geranoides is now synonymised under Pachyornis geranoides. Two other supposed
species, Pachyornis mappini and P. septentrionalis, are now known to be males and females of the same
species, both of which are also now synonymised under P. geranoides.
Mantell’s Moa was another extraordinary bird with massive short and robust legs, a blunt bill and squat
body. It also exhibited extreme sexual dimorphism. The males stood about 1m tall and weighed between 17
and 30kg, whereas the females were taller and may have reached between 40 and 100kg. It was the stoutest
member of the genus (Worthy & Holdaway 2002). Subfossil remains show that this species was restricted
mainly to the eastern lowlands of South Island during the Holocene (Worthy & Holdaway 1993).
The bill was short and blunt in this species, and the jaw was comparatively weak. This suggests that the
bird plucked foliage, rather than shearing it, and fed on softer leaves than other moa species (Gill &
Martinson 1991). Gill & Martinson note that plant remains found in the preserved gizzard of one individual
have been identified as the leaves, seeds and fruit of Matai Podocarpus spicatus, Pohuehue Muchlenbeckia
australis and Bush Lawyer Rubus sp. The gizzard stones measured 2–28 mm in diameter, with a total weight
of 253g.
Status Extinct. Known only from subfossil remains. Date of extinction c.1350–1450.
Range Western South Island, New Zealand.
The Crested Moa was a massive, squat bird, but smaller than the H eavy-footed Moa P. elephantopus and
Mantell’s Moa P. geranoides. It stood around 1.2m high and weighed around 75kg. The frontal region of the
cranium exhibited small depressions, interpreted as feather pits (Worthy & Holdaway 2002), so it is likely
that this was a crested species. It was the dominant species in fossil localities in both the far north and far
south of South Island, as well as in the cold alpine terrain. It was primarily an occupant of montane to subal-
pine areas (Worthy & Holdaway 1993).
Tennyson & Martinson (2007) suggest that the crest may have been used for sexual and territorial
displays.
The Heavy-footed Moa was widespread in the lowlands east of the Alps in the South Island and occu-
pied coastal shrublands, dunelands, grasslands and forest margins. These specialised birds appear to have
undergone a natural decline; evidence from the Punakaiki fossil locality has shown that P. elephantopus and
P. geranoides were replaced by Anomalopteryx didiformis and Dinornis robustus during the Holocene, in this
area at least (Worthy & Holdaway 1993). Once humans arrived on New Zealand, these slow, ponderous
birds would have been easily slaughtered, and the subfossil remains found in Polynesian middens on the
Waitaki and Rakaia Rivers suggest it was regularly hunted (Worthy & Holdaway 2002).
An intact pure white egg was discovered in the 1870s in central Otago; it contained an embryo, and may
be referable to this species, or perhaps to P. geranoides (see Hector 1872; Worthy & Holdaway 2002). In color-
ation at least, white eggs were laid by this genus, whereas greenish-blue ones were laid by Megalapteryx
didinus (see below).
If the egg does belong to this species, it is the second largest known, measuring 226mm x 155mm
(Tennyson & Martinson 2007).
The Eastern Moa was a medium-sized species reaching around 73cm tall in the male, and up to 1m tall in
the female. Males weighed around 36kg, and females up to 79kg (Bunce et al. 2009). Surviving feathers and
dessicated skin attributed to this species show that it was brownish beige in coloration, the feathers hair-like,
becoming shorter toward the head. The head itself may have been devoid of feathers. The legs were
extremely powerful with short, robust tarsi, and the feet were disproportionately large. It was probably a
rather ponderous, slow-moving species.
The Eastern Moa occurred in the lowlands and its remains were found in what are perceived to have
been dry rain-shadow forests and shrublands (Worthy & Holdaway 2002). Due to the lack of mitochondrial
DNA diversity in this species, Bunce et al. (2009) suggest that it went through a natural genetic bottleneck
followed by a rapid expansion during the Holocene, when its favoured habitat expanded. This would
account for the abundant subfossil remains, but with restricted genetic diversity. Analysis of gizzard
contents have shown that the Eastern Moa fed on fruit, seeds and leaves (Gill & Martinson 1991).
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The Upland Moa, as its name suggests, preferred montane and subalpine habitats, but it was also found in
the lowlands close to the coast where suitable habitat existed. However, its remains are most numerous in
the upland alpine zone (Gill & Martinson 1991; Worthy & Holdaway 1993). The Upland Moa male was
approximately 65cm tall and 28kg, whereas the female was up to 95cm tall and weighed up to 80kg. It was a
gracile, agile species with large, slender toes, which would have enabled it to manoeuvre easily in its
montane habitat. A preserved dessicated leg shows that the tarsi were feathered, which was another adap-
tation for habitation at high altitudes, especially with frequent snows (Worthy & Holdaway 2002). Very few
eggs have been found in association with the moas, but one exception was the discovery of a g reenish-blue
Upland Moa egg.
Claims by Europeans to have seen living moas in Fiordland, South Island, are all open to alternative
explanation. The prime candidate for a late-surviving moa is the Upland Moa. However, not a single shred
of physical evidence exists, and a thorough analysis of all of the reports and archaeological evidence has
shown that most were probably derived from confusion with the extant South Island Takahe Notornis hoch-
stetteri, or entirely mistaken (Anderson 1989). Like all moas, it appears that the Upland Moa disappeared
within a century or so after Polynesian occupation of the island, and at least two centuries before Europeans
arrived in New Zealand.
The fossil record has shown that this species, as well as some other moas, became smaller over time, and
that the Upland Moa that survived into the Holocene was not only the smallest of the genus (Worthy &
Holdaway 1993), but along with the Little Bush Moa Anomalopteryx didiformis (see p. 33) was the smallest of
all moa species. There were also size differences between h igh-altitude populations where the specimens
were large, compared with the smaller lowland forms. These discrepancies are the primary reason why so
many different species were originally described, especially since size was the main character used.
Megapodiidae (Megapodes)
The megapodes are chicken-like terrestrial birds that incubate their eggs by burying them in mounds of
decaying vegetation, or in volcanic soils or hot sand. The family is divided into seven genera, distributed in
Australia and New Guinea, and islands of Indonesia, the Bay of Bengal and the western Pacific.
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Megapodiidae
The terrestrial habits of megapodes have made them particularly vulnerable to hunting, over-harvesting of
eggs and predation by introduced animals, leading to drastically reduced populations, particularly on
oceanic islands. The fossil record shows that megapodes once occurred on many islands in the Pacific,
notably Fiji, Tonga and New Caledonia, with undescribed species from the Bismarcks, Solomons, Vanuatu
and Loyalty Islands; all have disappeared due to anthropogenic activities (Steadman 1999, 2006a).
The Consumed Megapode was a large species, larger than any extant megapode, but smaller than the
Pile-builder Megapode M. molistructor of New Caledonia (see below). However, it still retained the ability
to fly and seems to have occurred throughout Tonga and perhaps Fiji (Steadman 2006a). It was part of the
M. freycinet superspecies group and was broadly sympatric with the smaller M. freycinet in Tonga. It appears
to have died out within a couple of centuries of human occupation of the islands.
The Pile-builder Megapode was the largest member of its genus, with very robust tarsi (Balouet & Olson
1989), but it was dwarfed by the New Caledonian Giant Scrubfowl Sylviornis neocaledoniae (Steadman 1999;
see below).
A dubious species tentatively assigned to Megapodius (Gray 1862b), M. andersoni (see p. 371), may repre-
sent another extinct megapode on New Caledonia, but the available evidence is too vague to make any
further assumptions (Balouet & Olson 1989). The Pile-builder Megapode was almost certainly hunted to
extinction by humans.
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This megapode was the same size or slightly larger than the extant O range-footed Scrubfowl Megapodius
reinwardt, but with reduced wings and robust legs. Based on these skeletal characters, Worthy (2000)
suggested that it was probably flightless or almost so, unusual among the smaller megapodes, which would
have made it particularly vulnerable to hunting and introduced predators.
Sylviornithidae (Scrubfowls)
The relationships of the New Caledonian Giant Scrubfowl Sylviornis have been debated, but in general this
bird was considered to be a giant megapode (e.g. Olson & Balouet 1999; Worthy 2000) or a megapode-like
galliform, a fact that led Mourer-Chauviré & Balouet (2005) to place it in its own family, Sylviornithidae,
based mainly on the very distinct anatomy of the skull. Recent work has shown it to be a stem galliform,
distant from the megapodes, and a sister taxon to the Noble Scrubfowl Megavitiornis altirostris (Worthy et al.
2016), which is now placed in the same family.
The giant, flightless New Caledonian Giant Scrubfowl reached the extreme in insular adaptations, being
unlike any other flightless bird in its morphology and probable ecology. It was about 1.2–1.6m tall and may
have weighed up to 40kg (Steadman 1999). The bill was deep and laterally compressed, and some well–
preserved jaws exhibit a distinct knob, somewhat reminiscent of a cassowary Casuarius sp. The wings were
reduced to stumps, the legs were short but massive, with strong toes and long claws. It was once thought
that like other megapodes, Sylviornis was a mound-builder, and that giant mound structures known as
tumuli were its nesting mounds
(Green & Mitchell 1983). However,
Worthy et al. (2016) have shown that
Sylviornis had none of the morpho-
logical adaptations seen in
megapodes and that its claw
morphology was similar to that of
chickens Gallus gallus. Sylviornis
almost certainly did not use ecto-
thermic incubation, or build mounds;
the tumuli of New Caledonia are of
anthropogenic origin. In life,
Sylviornis may have been herbivo-
rous, feeding on low vegetation and
possibly digging for roots and tubers
New Caledonian Giant Scrubfowl with its specialised bill and feet, but
Sylviornis neocaledoniae Mourer-
Chauviré & Balouet (2005)
argue that the jaws are indicative of
another feeding strategy, perhaps an
invertebrate predator.
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Numididae
The bird is remembered in oral tradition on New Caledonia; it was said to be aggressive, could move
rapidly using its wings for balance, was reddish in coloration with a bony, solid casque in the shape of a star
on the head, and laid a single egg from November to April, but it did not incubate it (see M ourer-Chauviré
& Balouet 2005). When humans first arrived on New Caledonia, Sylviornis would have made easy prey, and
its eggs and young were particularly susceptible to introduced predators.
The possibility that at least three species of megapodes once occurred on the island of Viti Levu, Fiji, is quite
remarkable, with each differing in size and ecological niche. The Noble Megapode was a flightless species
with small wings and robust legs, the largest on Fiji, and exhibited considerable variation in skeletal
morphology (Worthy 2000), probably due to sexual dimorphism and a ge-related characters. It had a mark-
edly deep but narrow bill, which probably delivered a strong bite, but it lacked the necessary characters of
predatory birds. Worthy suggests that it fed on large fruits and was also quite capable of cracking open the
toughest seeds. Being totally terrestrial, it would have been quickly exterminated by humans and their
associated introduced animals.
Numididae (Guineafowl)
Guineafowl are m
edium-sized terrestrial birds restricted to Africa, with speckled plumage and a horny
casque on the head. The family contains four genera. The birds have long been domesticated and trans-
ported around the world. The guineafowl are generally not threatened, but continued habitat destruction
may impact on populations in the future.
The Helmeted Guineafowl Numida meleagris has been divided into more than 30 subspecies, most of which
are probably invalid. The species is extremely variable and characters used to separate some populations
are inadequate. It is widely hunted for food and domestic birds have been reared for centuries in many
countries across the world, including many parts of Africa. The Moroccan Guineafowl was a distinct
subspecies that, having suffered from severe habitat destruction and over-hunting, was last recorded with
certainty in the 1950s. Reports of birds in the 1970s were probably feral domestic birds, and rumours that a
captive population survived until the end of the 1980s have not been substantiated (Martinez 1994). It is
likely that this subspecies is now extinct.
Hartert (1919b) noted from illustrative evidence that the Moroccan Guineafowl was known to the
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Romans 2,000 years ago, and that the bird was transported to Italy via Greece and was known as the
‘Numidian Hen’. It was reported as having once occurred in Morocco and Tunisia.
The Heath Hen was formerly considered a full species, but is now generally regarded as the nominate race
of the Prairie Chicken Tympanuchus cupido. Three Prairie Chicken subspecies survive today: Greater Prairie
Chicken T. c. pinnatus, Lesser Prairie Chicken T. c. pallidicinctus, and Attwater’s Prairie Chicken T. c. attwa-
teri; all are greatly reduced in range, and threatened with extinction.
Original range
The original range of the Heath Hen is not known with certainty, for it probably began to decline soon after
European colonisation of North America. It is known to have occurred from Massachusetts to the Potomac
River, perhaps north to Maine and New Hampshire, and possibly south to the Carolinas. The subspecies
was only recognised in 1885 (Brewster 1885), which hitherto had been lumped together with the western
Prairie Chickens, all termed the Pinnated Grouse. However, by this time, the Heath Hen was already
confined to Martha’s Vineyard, an island in Buzzard’s Bay off the Massachusetts coast, the last stronghold
of the bird. The problems of range determination are also compounded by lack of specimens. Only six are
known from the mainland, while the rest were all collected on Martha’s Vineyard, so the original distribu-
tion can never be established. Gross (1928) suggested that the bird may not originally have been native to
this island but were introduced there. It had, however, been present since at least 1824. The six mainland
specimens of Heath Hen differ from those of Martha’s Vineyard, resembling the Greater Prairie Chicken
(Poole 1949), whereas six specimens in the Museum of Comparative Zoology, allegedly collected from
Martha’s Vineyard, also resemble the Greater Prairie Chicken. It is probable that collecting data is erro-
neous, unrecorded introductions of western birds took place, and that individual variation was also great.
Other distributional reports for Maine and New Hampshire are questionable. The former existence of the
bird in Maine rests solely on the authority of Audubon (1834), stating that they occurred on Mount Desert
Island and on Mar’s Hill near Houlton, but the Spruce Grouse Canchites canadensis had been called the
‘heath hen’ by residents (Palmer 1949), so Audubon may have been confused. Prairie Chickens were intro-
duced to Maine in the 1870s and it was probably these that were protected by law in 1876 and 1878 (Grinnell
1888; Palmer 1949). A similar confusion of names may have occurred in New Hampshire. A single eight-
eenth-century record for ‘Grouse’ (Belknap 1792) may refer to the Heath Hen, but Greenway (1967)
considered the record to be unconvincing.
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Phasianidae
In Massachusetts, the Heath Hen was evidently common and occurred at forest edges near the coast, and
in the valley of the Connecticut River (Wood in Forbush 1912). Wood resided at what is now the city of Lynn
in Massachusetts, and reported that the bird was common, its flesh was red and it was evidently hunted for
food. Prior to 1840, the bird was presumably still common at least in the eastern parts of Massachusetts, as
Nuttall (1840) recounted how Lieutenant Governor Winthrop was begged by his servants not to have Heath
Hen served more often than ‘a few times’ a week. Just 15 years later, they were no longer seen on the main-
land of Massachusetts; the bird had already gone by 1830 in the western part of the state, disappearing from
New England about the same time (Greenway 1967). A bill to protect them was passed in 1846 by the State
of Rhode Island, but no other evidence exists to show the Heath Hen occurred there.
In Connecticut the records are also meagre. Thomas Pennant (1785) described specimens in Blackburn’s
Cabinet that were collected in Connecticut, and as late as 1840, Thomas Nuttall said they were still to be
found on the plains in Westford, but they had disappeared from the plains in the Connecticut Valley near
Springfield about 1812 or 1813 (Forbush 1927).
As early as 1791, a law was introduced by Cornelius J. Bogert of New York to protect the Heath Hen on
Long Island. This bill protected the bird during the closed season, but seems to have been largely unen-
forced. The bird continued to decline in New York State and was almost if not completely extinct there by
1844 (Giraud 1844; De Kay 1844). The latter author said that it was still found on a few islands off
Massachusetts, in the mountains of Pennsylvania, and on Schooley’s Mountain in New Jersey. It was very
popular as a game bird and commanded high prices in markets.
It is known that a Heath Hen or Prairie Chicken was common on the plains of New Jersey before 1850
(e.g. Wilson & Bonaparte 1832) but it is not clear which subspecies was involved. The preserved skins
resemble the western race, but these may have been imported. The Heath Hen probably disappeared from
Pennsylvania by the 1870s (Greenway 1967), but the exact date is unknown; the last supposed sighting in
the east were birds observed near Broadhead’s Creek, Monroe County in 1869 (Weygandt 1906; Greenway
1967). It must have occurred in Chesapeake Bay, as a specimen catalogued in the United States National
Museum in Chicago was collected there, but this was destroyed in a fire (Greenway 1967).
The record of the bird occurring in Carolina is based entirely on the observations of Catesby (1743),
though he never saw the bird on either of his visits there (1710–19 and 1722–26), nor did he record any
person who had. Ironically, Catesby did not see the Heath Hen until he returned to England, where he
observed captive birds in 1743 at Cheswick, the seat of the Earl of Wilmington. The Earl’s birds were brought
from America but their exact origin had not been recorded. Further supposed occurrences in Maryland,
Delaware and Virginia are discussed in detail by Gross (1928), but no specimen exists.
Habits
The Heath Hen, like most gallinaceous birds, fed on a wide variety of food. Gross (1928) lists the following
as being among the main food items taken on Martha’s Vineyard: Bayberry Myrica carolinensis, Bearberry
Arctostaphylos uva-ursi, Partridge-berry Mitchella repens, and Blueberries Vaccinium vacillans, V.
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pennsylvanicum and Gaylussacia baccata; they also took rose hips, strawberries, acorns, leaves, buds and
insects. The nest was built on the ground, and usually composed of leaves, grasses and twigs. Unlike its
western cousins, which place the nest in an open site, the Heath Hen apparently always selected a
well-concealed site among dense vegetation (Gross 1928). The nests were so difficult to find that very few
were ever discovered, and there are only about two clutches and a couple of odd eggs in any collection. The
eggs were deep olive-buff, considerably darker in colour than those of the Prairie Chickens. The breeding
season was during June and July; the incubation and fledging periods do not appear to have been recorded.
New Mexico S
harp-tailed Grouse
Tympanuchus phasianellus hueyi Dickerman & Hubbard
Tympanuchus phasianellus hueyi Dickerman & Hubbard 1994, p. 133 (Folsom, Union County, New Mexico)
Specimens Specimens are in Berkeley, California; New Mexico; and New York.
Status Extinct. Last recorded in 1954.
Range New Mexico and probably Colorado.
Description 41–46cm (16–18in). Plumage generally white mottled with dark and light brown, lighter on underparts with
uniform V-shaped markings; tail with two central deck feathers; bill dark brown; iris dark orange; legs and feet orange-brown.
Males had yellow comb and violet display patch on neck; irregularly marked on deck feathers; female smaller with indistinct
comb, and more regular horizontal markings on deck feathers. New Mexican birds differed in being more tawny brown on
upperparts; intermediate in hue of underparts and ventral markings; and presence of throat markings compared with T. p.
jamesi and T. p. columbianus.
The Sharp-tailed Grouse T. phasianellus is a sedentary species occurring in the prairie regions of North
America and Canada. It has been divided into seven subspecies, which are not under immediate threat, but
are generally declining primarily due to habitat loss and o ver-hunting (Silvy & Hagen 2004). The New
Mexico population was restricted to the high mesas in Colfax and Union counties, New Mexico, and prob-
ably adjacent Las Animas County, Colorado. It had probably been declining for some time due to habitat
destruction and also increased aridity (Dickerman & Hubbard 1994). It was apparently common prior to
1926 (Ligon 1961), but the last records were in 1952 on the Sewell Ranch, Colfax County (Merrill 1967). The
subspecies T. p. jamesi was introduced to Sewell Ranch in 1952, and may have hybridised with the last
pure-bred New Mexican birds, but these possible hybrids have also now disappeared (Dickerman &
Hubbard 1994). Wetmore (1936) referred a subfossil bone collected about 255km to the s outh-west and
dated around 1300 ad to this form, indicating that it was much more widely distributed in the past.
Habits
The New Mexico Sharp-tailed Grouse occupied the high mesas between altitudes of 2,438–2,743m on grass-
lands mixed with dense stands of oaks, conifers and other woody growth on the steeper slopes (Dickerman
& Hubbard 1994). Grain crops were grown in large areas, and it appears that the birds were reliant on
spillage during the winter months (Ligon 1961), as habitat loss may have adversely impacted their natural
feeding grounds.
There has been some doubt as to the validity of this subspecies, but McGowan (1994) considers it a valid
taxon. The Black Francolin F. francolinus is a widespread species occurring in Cyprus and southern Turkey
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through the Middle East and Transcaucasia to Sikkim and Bangladesh. It is however decreasing in numbers
over much of its range due to over-hunting and habitat alteration. The Lake Amik Gölü population has
disappeared as a result of the almost complete drainage of its habitat in the 1970s, and has not been reliably
reported for some time. It may now be extinct, but if it does still survive the numbers must be extremely
small.
Habits
Little was recorded about the Amik Gölü birds, but the Black Francolin in general is a secretive and wary
bird, which never strays far from thick undercover, and occurs in grassy areas, savannas, and also crop
fields. It forages for grain, grass seeds, fallen fruit and insects on the ground, and will only fly when forced.
The last known specimens of the New Zealand Quail were taken at Blueskin Bay in 1867 or 1868, but the
bird was believed to have survived in parts of South Island as late as 1875 (Gill & Martinson 2010). The
extinction appears to have been extraordinarily rapid, for the bird was still common in 1861, at least in
places such as the open down of Waikonaiti, north of Dunedin, and in Nelson Province in the n orth-west of
South Island (Greenway 1967). Buller (1887–88) recounted how in 1848 Sir Edward Stafford held a shooting
party on his estate about 30 miles from Nelson and in a few hours 29 brace were bagged. Shooting was
banned the following year in the hope of preserving the birds, but the year after not a single quail could be
found.
The bird was still numerous in parts of South Island in 1861 and possibly lingered on till after 1870, but at
this time other quails were being imported for release, and confusions exist as to which species is involved.
Brown Quails Coturnix ypsilophora were not imported before 1866, but pheasants were imported as early as
1842, and it is thought that diseases brought with them may have been responsible for the extermination of
the New Zealand Quail (Greenway 1967), though this cannot be proven. It was probably a combination of
factors that led to the quail’s extinction. These included the widespread burning of lowland tussock grass,
which was the favourite haunt of the quail, plus imported rats, cats, ferrets and other predators, but these
had no effect on the numbers of the imported Brown Quail. However, the Australian species may have been
better adapted to cope with mammalian predators, which abound in Australia, than the New Zealand bird,
which evolved in a mammal-free environment.
Habits
The New Zealand Quail inhabited open grasslands and marshes, feeding mainly on seeds. On South Island
they occurred in open fernlands, and in dry grasses intermixed with low shrubby plants on Long Island in
Queen Charlotte Sound (Fleming 1982). The nest was a depression in the ground, lined with grass. The eggs
were 10–12 in number; pale brown blotched with darker brown. Incubation was recorded to last about 21
days. The call was a low purring sound, which had been likened more to an insect than a bird call. It was
something like twit, twit, twit twee-twit, repeated several times (Potts 1870; Greenway 1967).
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The extinction of Canary Island birds, around 13% of the avifauna, can be blamed on the negative effects of
deforestation, over-hunting and introduction of alien species by both the original inhabitants and later
European invaders (Illera et al. 2016); all extinct birds nested on the ground, and four out of the five were
flightless or poorly volant.
The Canary Islands are occupied by
two extant races of Common Quail, the
migratory Coturnix coturnix coturnix
which is found on the eastern islands,
and C. c. confisa, a permanent resident of
the western islands at a higher altitude.
The Canary Islands Quail once occurred
on a number of islands in the group,
probably including Gran Canaria and
Lanzarote, though no remains have
been found on these latter islands yet. It
appears to have been quite common
prior to the arrival of humans, but disap-
peared shortly after settlement, probably
due to predation by cats and perhaps
rats. It differed from the Common Quail
in having smaller wings but larger feet,
which suggests that it was evolving Canary Islands Quail
Coturnix gomerae
towards flightlessness.
The Himalayan Mountain Quail was described from captive specimens in the menagerie of the Earl of
Derby (J. E. Gray 1846), after which it was observed and collected for a period of just 30 years, and never
reliably reported again. Very few birds were taken (localities and dates are assembled in Collar et al. 2001)
and it is doubtful that hunting was the primary cause of its disappearance. Fuller (1987, 2000) considered it
significant that all the known records came from a very brief time period and believed that unusual weather
conditions brought them temporarily to the Himalayas from their usual habitat further north. As this bird
was so cryptic, it may have been overlooked elsewhere. Blanford (1898) believed that the long, soft plumage
was an indication that it came from a cold climate. There are more recent accounts that suggest that the
Himalayan Mountain Quail still exists (see Rahmani 2012), but none have been confirmed.
Habits
The Mountain Quail occurred in the altitude range of 1,650–2,400m, occupied long grass and brushwood on
steep slopes and was extremely cryptic. Birds could only be made to fly when flushed by a dog or trodden
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on, and when doing so made a shrill whistling note, but their flight was slow and heavy (Hutton in Hume &
Marshall 1879). It was encountered only in winter, and the authors suggested that it was a migrant from ‘the
better-wooded, south-eastern portions of Chinese Tibet’. Nothing was ever recorded about the quail’s
breeding habits but a yearling male in moult collected in June suggests it probably bred in September. The
little information known is derived only from those who shot them. Hutton (in Hume & Marshall 1879)
stated:
Two or three coveys came in November 1867, and some remained as late as June 1868, when this present spec-
imen was shot. They have not been seen since, and I never saw them before.
Finn (1911) gave the best summary of the species:
It seems to me to be a migratory bird, arriving in winter, although its small wings look ill-adapted for a journey
of any length. It goes in single pairs or coveys, and keeps close to the cover in grass jungle or brushwood, being
almost impossible to flush without a dog. Its flight is heavy, slow, and short; its food, grass seeds. The call is a
shrill whistle.
47
This goose was considerably smaller than the South Island Goose Cnemiornis calcitrans. The first bones were
found at Kaiiwi in 1886, but Forbes gave no adequate description, and his types were thought to be lost.
This led to Oliver (1955) renaming the species and designating neotypes. However, the Forbes types subse-
quently turned up in the Natural History Museum, London, and Dawson (1958) therefore restored the
original, revalidated name. It was, he says, ‘a most elegantly moulded goose’. There is one possible histor-
ical account (see Worthy & Holdaway 2002), quoted from C ockburn-Hood in 1875, which states that a
goose, with rufous plumage and unable to fly, could be caught with dogs and was esteemed for food. The
reasons for its extinction are not known, but there is no doubt that a large, flightless goose would have been
extremely vulnerable to over-hunting.
The Bering Cackling Goose was a small dark form of the Cackling Goose B. hutchinsii, which formerly
occurred on the Komandorskie and probably the Kurile Islands (Bergman 1935), the westernmost range of
the species. They may have migrated at least to Japan, but data is lacking. The birds were doubtfully distinct
from the Aleutian subspecies, B. h. leucopareia, and are generally considered conspecific with it. The popula-
tion was exterminated due to o ver-hunting, and predation by the Arctic Fox Vulpes lagopus, some time in the
early 20th century, with the last sighting occurring in either 1914 or 1929. Little more is known about the
Bering Cackling Goose.
The Aleutian subspecies almost suffered a similar fate after Arctic Foxes were introduced to the Aleutians
by Russian fur traders between 1836 and 1930. B. h. leucopareia was thought to have become extinct, but a
small population was discovered on Buldir Island in 1962 (Palmer 1976). The geese made a remarkable
comeback once the Arctic Foxes were controlled; this taxon is no longer considered endangered.
Subfossil remains of a giant goose were found in a lava tube above Kailua in the North Kona District on
Hawaii (Olson & James 1982). The remains were fragmentary so its taxonomic status could not be deter-
mined. More material was found subsequently in another lava tube in North Kohola, which showed that
this goose, now termed the Giant Hawaiian Goose, was larger than any other Hawaiian anatid, living or
extinct. It was also flightless, but the wings were not as reduced as those of the m
oa-nalos (see p. 63) (Olson
& James 1992).
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Although it remains undescribed, the Giant Hawaiian Goose is now known from a number of fossil
localities; mitochondrial DNA analysis has shown that it is derived from the Canada Goose Branta
canadensis, and closely related to the Nene B. sandvicensis and the Greater Hawaiian Goose B. hylobadistes
(Paxinos et al. 2002) (see below).
The Giant Hawaiian Goose was massive, averaging 1.4 times the weight of the largest race of Canada
Goose, B. c. maxima, and 4.5 times the weight of the Nene (Paxinos et al. 2002). Its skull was extremely robust
and the bill robust and deep. The leg bones were massive and the wings were disproportionately small.
As the island of Hawaii is less than 0.5 million years old and the genetic differences were slight between
the Giant Hawaiian Goose and its close Branta relatives, Paxinos et al. suggest that gigantism, flightlessness
and a robust cranium evolved within that relatively brief period. Evolutionary trends towards gigantism
can occur comparatively rapidly, a scenario mirrored by the rapid evolution of the world’s largest eagle,
Haast’s Eagle Aquila moorei (see p. 88). The lack of terrestrial, herbivorous mammals must have been an
important factor in Hawaiian anatid evolution. As a consequence, selective pressures resulted in their rapid
gigantism and switch from wetland habitats to the vacant niche of terrestrial herbivory (James & Burney
1997; Paxinos et al. 2002).
Being large and flightless, the Giant Hawaiian Goose was probably hunted to extinction soon after
Polynesians settled on the islands. Carbon dating of bones indicates that it may have survived until at least
the 16th century (Paxinos et al. 2002).
This was a large, robust-limbed, small-winged goose, closely related to the extant Nene Branta sandvicensis
and the undescribed Giant Hawaiian Goose Branta sp. (Paxinos et al. 2002)(see above). It occurred on the
southern slopes of Mount Haleakala. Similar, a s-yet undescribed species occurred on other islands of the
Hawaiian Archipelago (Olson & James 1991). Uniquely, subfossil remains show that this species was in the
evolutionary process of losing its flying ability, with some specimens volant, others flightless (Olson &
James 1991; Paxinos et al. 2002). Being terrestrial and at best a poor flier, it no doubt represented easy prey to
the Polynesians and to introduced predators.
There is some confusion over the status of the Cygnus swans of New Zealand and the Chatham Islands.
A supposed endemic New Zealand Black Swan Cygnus sumnerensis, related to the extant Australian
Black Swan Cygnus atratus, was described from subfossil remains on New Zealand, on the basis of its
larger size (Forbes 1892d). Worthy & Holdaway (2002) have shown that these subfossil remains are
undifferentiated from that of the Australian Black Swan, and not considerably larger as proposed by
Forbes (1892d). Worthy & Holdaway therefore suggest that a resident population of Australian Black
Swan C. atratus – and not a different species – was exterminated by Polynesian colonists at some time in
the past.
However, subfossil remains from the Chatham Islands are distinct. These birds had a larger and stouter
bill, and comparatively shorter wings compared with subfossil remains found on mainland New Zealand
(Milliner 1999). This suggests that the population of black swans once found on the Chatham Islands prob-
ably represent yet another now-extinct species from these islands.
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Geese were mentioned by the Dutch on Mauritius during the 17th century (Cheke & Hume 2008), but
without providing any details. The English mariner Marshall in 1668 (in Kahn 1927) gave the best
description:
Here are many geese, the halfe of their wings towards the end, are black, and the other halfe white. They are not
large but fat and good [to eat].
A carpometacarpus from Mauritius was described by Newton & Gadow (1893) as belonging to a Comb
Duck Sarkidiornis, larger than Knob-billed Duck S. melanotus. However, Andrews (1897), and more recently
Cowles (1987), showed that this specimen was actually referable to the genus Alopochen, and was closely
related to the Egyptian Goose Alopochen aegyptiacus. A few more subfossil elements have since been found,
which indicate that the Mauritian goose was smaller than Egyptian Goose, but had more robust legs. This
may have been due to the goose becoming more terrestrial, a fact supported by an account from the log of
the President in 1681 (Barnwell 1950–54):
Up a little within the woods are several ponds and lakes of water with great numbers of flamingoes and gray teal
and geese; but for the geese these are most in the woods or dry ponds.
Habits
Little was recorded about the habits of the Mauritius Sheldgoose. An account written by Johannes Pretorius
on Mauritius in 1666–69 (in Hume & Winters 2015) provides the only information:
Geese are also here in abundance. They are a little larger than ducks, very tame and stupid, seldom in the water,
eating grass, sometimes 40 or 50 or even 100 together. When they are being shot, the ones that are not hit by the
hail stay put and do not fly away. They usually keep to the north side of the island, far away from where the
people live, except in the dry season when they are forced to drink on the other side of the island, and sometimes
near the lodge.
The Mauritius Sheldgoose probably disappeared due to over-hunting and perhaps predation of eggs
and chicks by introduced mammalian predators, especially cats.
Analysis of subfossil remains collected by Kervazo in 1974 (Cowles 1994) clearly show that a derivative of
the Egyptian Goose Alopochen aegyptiacus, closely related to the Mauritian species, once occurred on
Réunion Island. Dubois (1674) in 1671–72 gave the best account:
Wild geese, slightly smaller than the European geese. They have the same feathering, but with the bill and feet
red. They are very good [to eat].
Nothing more can be said about them other than that they differed from Egyptian Goose by having more
robust leg bones and a shorter, deeper bill (Mourer-Chauviré et al. 1999). Over-hunting appears to be the
primary cause of extinction, but no doubt introduced predators would have taken eggs and chicks.
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The Swiss palaeontologist, Charles Immanuel Forsyth Major, spent two years in Madagascar from July 1894
to August 1896, and amassed a vast amount of vertebrate remains, including much from the extinct mega-
fauna (Jenkins & Carleton 2005). He sent the material to the then British Museum of Natural History,
London, which included a series of subfossil remains belonging to an anseriform. These were collected at an
ancient lake site at Sirabé (now Antsirabe) in central Madagascar (Rothschild 1907a). Andrews (1897a) iden-
tified some of them as being clearly referable to Alopochen, but they differed from the only surviving member
of the genus, the Egyptian Goose A. aegyptiaca. He named the new species Alopochen sirabensis after the
collecting locality.
The Antsirabe Sheldgoose was one of the most common species found at this ancient lake; Goodman &
Jungers (2014) suggest that it was a gregarious species, feeding at the edges of aquatic habitats on herba-
ceous plants, seeds, invertebrates and small vertebrates. Two radiocarbon dates from A. sirabensis bones
collected at Antsirabe give 19,250 and 17,100 years BP (mean calibrated dates of 22,860 and 20,170) (Crowley
2010), but this duck may have survived until at least the 15th or 16th centuries; however, it had certainly
disappeared before the arrival of Europeans (Goodman & Rakotozafy 1997). The reason for its extinction is
not clear, but a natural aridification of the island and over-hunting by humans may have been responsible
(Young et al. 1996; Crowley 2010; Goodman & Jungers 2014).
until at least the 14th or 15th centuries, but probably succumbed to a natural aridification of the island and
from over-hunting by the Malagasy people (Young et al. 1996; Crowley 2010; Goodman & Jungers 2014).
The Crested Shelduck was always considered a rare species, even at the time it was first collected, and is
known from just three specimens. The first was taken in April 1877 near Vladivostok, Russia, the last two in
1913 or 1914 and 1916 in Korea (Greenway 1967), but 19th-century Japanese descriptions and paintings of it
suggest it was previously more widespread (see Nowak 1983). Supposed recent sightings of the Crested
Shelduck (Sok 1984) have not been confirmed (see Collar et al. 2001 for a full review). If it does survive, the
population of this enigmatic and elusive bird must be tiny.
Habits
Little is known about the Crested Shelduck’s habits. It was mainly recorded in pairs or small flocks in
coastal localities, and it also occurred on forest islands within rivers. The structure of the bill infers that it
was a grazing species, not a filter feeder (Nowak 1983).
Milliner (1999) discovered abundant subfossil remains of a Tadorna shelduck in cave and dune deposits on
Chatham Island, which differed from the New Zealand endemic Paradise Shelduck T. variegata in having a
larger, more robust skull and jaws and slightly smaller wing elements. This suggests that it was a sedentary
species, with reduced flying capabilities. The Chatham Islands avifauna suffered extremely high extinction
rates after humans colonised the islands, especially amongst wildfowl (Worthy & Holdaway 2002; Young et
al. 1997), so it is likely that the Chatham Islands Shelduck was rapidly exterminated through over-hunting
by Polynesian settlers.
This duck was related to, but was considerably larger than, the Pink-eared Duck Malacorhynchus membrana-
ceus of Australia and Tasmania. From a study of its anatomy (Worthy & Holdaway 2002), the New Zealand
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Pink-eared Duck was a filter feeder, dabbling on the surface of open water, and not a diving species.
Over-hunting coupled with a specialised diet probably contributed to its extinction. Henry Travers, the
New Zealand naturalist, reported seeing a ‘peculiar Teal’ with ‘bright scarlet markings on the wings’ on
Chatham Island in November 1864 (Tennyson & Martinson 2007). There is a possibility that this was one of
the last surviving New Zealand Pink-eared Ducks.
Finsch’s Duck is closely related to the Australian Wood Duck Chenonetta jubata, and probably derived from
it (Worthy & Olson 2002). It is one of the most common species to be found in Quaternary fossil deposits on
both eastern districts of North and South Island (Worthy & Holdaway 2002). Hamilton (1893) suggested
that the large numbers of bones found together might be indicative of the depredations of the predatory
Haast’s Eagle:
To account for the great number of such bones it has been suggested that perhaps the eyrie of the Harpagornis
was on the rock above the chasm, and that these are the relics of his foraging expeditions.
This duck was about the size of the Mallard Anas platyrhynchos at 56–65cm (22–26in) in total length, but with
comparatively reduced wings, a short bill and robust legs, which indicate that it had reduced powers of
flight and was more terrestrial. The bill was short and stout and adapted for grazing (Worthy & Holdaway
2002). Worthy & Holdaway noted that due to the abundance of Chenonetta subfossil remains from different
localities, a rate of evolutionary change toward flightlessness could be measured; mean wing-length
declined 10% between 10,000–12,000 years ago and around 2,000 years ago, while mean leg-length remained
constant, showing that w ing-reduction can occur in just a few thousand years. This phenomenon is also
exhibited in subfossil remains of the Greater Hawaiian Goose Branta hylobadistes (see p. 49).
Subfossil remains of Finsch’s Duck have been found in Maori middens, and being terrestrial and poorly
volant, thus easy to catch, there is no doubt that over-hunting plus probable predation by the Pacific Rat
Rattus exulans were responsible for its extinction.
Ducks of the genus Anas are adept at colonising remote island archipelagos, and in the absence of terrestrial
mammalian predators, often become flightless. An undescribed species of Anas, which had a more robust
cranium and longer and narrower bill than its probable closest relative, the New Zealand Brown Teal Anas
chlorotis, once occurred on the Chatham Islands (Milliner 1999). It seems to have disappeared after the
arrival of Polynesians, who no doubt hunted it to extinction.
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Only two specimens of this duck are known, the co-types collected in 1874 (Streets 1876) which were appar-
ently juveniles (Todd 1996). Since then Coues’s Gadwall has never been seen, although an expedition from
the B. P. Bishop Museum in Hawaii visited the island in 1924, as did the collector Rollo Beck for the American
Museum of Natural History at around the same time (Greenway 1967). The cause of extinction is not known,
but it is probable that it was a combination of over-hunting and predation by introduced animals.
The population is believed to have resulted from accidental visits by the Common Gadwall Anas streptera,
which remained to breed and evolved in isolation. However, some authorities consider it a poorly differen-
tiated subspecies. The Common Gadwall certainly occurs throughout the general Pacific area. They are
vagrant to the Hawaiian Islands, and occasional elsewhere (Pratt et al. 1987). One W. G. Anderson of
Honolulu, who was born in around 1895 on Fanning Island and spent his youth both there and on
Washington Island, stated that there were many migrant ducks in his day, but no resident ones, and hunting
them was regular sport (Wetmore 1925a). This suggests that the species, if indeed it was a resident one,
must have been extinct by then.
Habits
Coues’s Gadwall inhabited freshwater lakes and peat bogs on Washington Island, an uplifted atoll in which
the lagoon has lost much of its salt water and subsequently partially filled with fresh water (Greenway
1967). It had an increased number of bill lamellae compared with nominate A. s. streptera, which suggests a
specialised diet (Todd 1996). Nothing was ever recorded of the bird’s ecology.
The islands of Amsterdam and St Paul are situated more than 3,000km from any continent. They were often
visited by early mariners and their commensal animals, so that all native land birds were exterminated
before being scientifically described. The first possible account of Amsterdam Island Duck was by William
de Vlaming, who in 1696 thought he saw two f our-footed animals in the reeds. He probably saw terrestrial
birds, as there are no known native land mammals, but equally may have seen introduced rats.
Both islands were visited in February 1793 by the British Embassy to China under Lord Macartney
(Barrow 1807). John Barrow, in his account of the voyage, mentions a small brown duck on St. Paul, which
was probably distinct from that of Amsterdam Island (see p. 54). Amsterdam Island was not visited by a
naturalist until 1874, by which time it had already been well-used by sealers and whalers. Several wrecks
had occurred on its shores, and by 1874 the island was infested with rats. Unsurprisingly, no land birds
were found (Vélain 1877). In 1955–56, Patrice Paulian collected subfossil bones of a small duck on the island,
as well as the remains of a ‘mummified’ rail which immediately crumbled to dust (Jouanin & Paulian 1960).
The duck bones were examined and found to be most similar to the Garganey Anas querquedula; but Jouanin
& Paulian concluded that they must be vagrants.
The Brown Pintail Anas georgica occurs over much of South America. There are three recognised races: the
nominate georgica, confined to the island of South Georgia, spinicauda, occurring over much of the mainland,
and the isolated niceforoi in central Colombia. Nicéforo’s Pintail was only discovered in 1946 and last seen in
1952, just six years later. It was sedentary, and restricted to Andean lakes and marshes between 975m and
3050m in central Columbia (Todd 1996). It was well known to local hunters, who, along with habitat modi-
fication, may have been responsible for eliminating the species. Nothing was recorded about its habits.
The three types, a male and two females, were collected in 1885. Arundel was the first naturalist to visit the
island, and reported that there were no ducks initially, but later they appeared and were fairly numerous.
Phillips (1922–26) considered that A. a. modesta could not be distinguished from nominate Northern Pintail
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A. a. acuta, which occurs in the Hawaiian Islands and perhaps also in the Marshall Islands. Lister (1891)
discussed the birds of the Phoenix Islands, and listed the duck as the only land bird. As the population has
now gone, it is impossible to confirm the validity of this bird.
This was a local representative of the Sunda Teal Anas gibberifrons, a species which ranges from the Andaman
Islands to Australia, New Zealand, New Caledonia and Macquarie Island. Details about its ecology are
unknown. Bradley & Wolffe (1956), who published on the birds of Rennell Island, mention the teal only
taxonomically, and give no indication as to whether or not it was observed. It was restricted to the only
lagoon on Rennell Island (Todd 1996), and became extinct due to the introduction of Tilapia fish (Carboneras
1992), which presumably destroyed its food supply.
A small duck was reported from Mauritius and Réunion during the 17th century, which appears to have
died out around 1700 on Mauritius and 1710 on Réunion (Cheke & Hume 2008). The discovery of subfossil
remains (Newton & Gadow 1893) confirm that the species was a member of the genus Anas, related to the
Sunda Teal Anas gibberifrons complex, but it was larger than its closest relative on Madagascar, the
Madagascar Teal A. bernieri, and smaller than the other Madagascan Anas, Meller’s Duck A. melleri. It
appears that the Mascarene Teal was quite capable of flying between Mauritius and Réunion, and there
might have been some seasonal movement (Hume et al. 2004).
A report from the log of the ship President from 1681 (Barnwell 1950–54) (see Alopochen mauritiana, p. 46),
while visiting Mauritius, mentions that a ‘gray teal’ occurred on inland lakes in great numbers. They must
have disappeared rapidly after this date, as they were not mentioned again after 1700. Similarly, they
occurred in great numbers on Réunion, but were last seen a decade later. Over-hunting appears to be the
primary cause of extinction, but introduced cats would have been formidable predators.
Fossil remains of a pochard Aythya sp., a genus present in Madagascar, South-east Asia and Australia, have
been found on Réunion (Mourer-Chauviré et al. 1999). The Madagascar Teal Aythya innotata is most similar
to the Réunion Aythya subfossil material, so it may well have been this species or a closely related one that
once occurred on Réunion. The only historical account is an ambiguous mention of ‘ducks and teal’ in 1710,
which may indicate this species.
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The pelvis of this bird was erroneously described by Milne-Edwards (1896) as belonging to a
Diaphorapteryx rail; however, Oliver associated a skull with it, describing it as a ‘stoutly built duck’. A
recent molecular study has shown that the Chatham Islands Duck is a basal member of a subclade of
three New Zealand teals, the volant Brown Teal Anas chlorotis, and flightless Auckland Island Teal A.
aucklandica and Campbell Island Teal A. nesiotis (Mitchell et al. 2014). The Chatham Islands species was
the largest member of its genus, weighing around 1.5kg or even more, and it was completely flightless
(Mitchell et al. 2013; Williams 2015a, 2015b), with a heavy cranium and well-developed salt glands
(Worthy & Holdaway 2002). As the Chatham Islands Duck was incapable of flight, it would have made
it vulnerable to overhunting by humans and predation from introduced mammals, especially the Pacific
Rat Rattus exulans. This duck appears to have become extinct at some point between the 13th and 15th
centuries (Mitchell et al. 2013).
An additional undescribed member of this subclade once occurred on Macquarie Island, and is known
only from subfossil remains (Tennyson & Scofield 2013; Mitchell et al. 2013).
Habits
The Chatham Islands Duck possessed a large carpal knob, which was used in territorial disputes, similar to
those of modern-day steamer ducks (Williams 2015b). This, and the possession of salt glands, has been
interpreted as evidence for year-round territoriality, and consequently a limited distribution, perhaps
restricted to the shore-line of the Chatham Island lagoon and coastal regions (Worthy & Holdaway 2002;
Williams 2015a, 2015b). It fed in a marine or marine-influenced environment, or on a diet of highly saline
foods (Williams 2015a), possibly including seasonally available plant and invertebrate food items.
Much has been written about this duck, but the information is fragmentary and even contradictory. For a
bird which was known, and known to be rare, for so long, it is a great pity that so few studies were ever
carried out, particularly as the Pink-headed Duck appears to have been unusual in many ways.
The great 19th-century Indian ornithologist, Allan Octavian Hume, made exhaustive enquiries into the
Pink-headed Duck’s distribution in the Indian region. He stated that there were no records of the duck in
Ceylon (Sri Lanka), anywhere south of Mysore, or in Konkan, Gujarat, Kutch, Kathiawar (parts of modern
Gujarat), Sind, Rajputana (a state corresponding roughly to modern Rajasthan), Central Indian Agency
(north-west Madhya Pradesh) or the Central Provinces (central India) (Hume & Marshall 1879–81). There
were a few records from the North West Provinces (a region roughly corresponding to modern Uttar
Pradesh and Uttarakhand). Hume saw or heard of it only once in 20 years, but one was shot near Fatehpur
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Extinction
The birds were shot frequently by ‘sportsmen’
in the last 20 or so years of the 19th century, but
the species was always considered rare; hardly
anywhere did it occur in large numbers. They
were not particularly good to eat, but were
often shot nevertheless; however, Jerdon (1864)
claimed that they were good eating. That it survived for so long is probably due to the fact that the centre of
its distribution was a large plain crossed by deep, crocodile-infested rivers, heavily inundated by floods and
inhabited sparsely by humans, but commonly by tigers (Baker 1908). This area has become more populated
in the 20th century and has been drained and cultivated. Habitat destruction appears to be the main reason
for the Pink-headed duck’s extinction, but over-hunting is also likely to have been a factor (Collar et al.
2001). Bucknill (1924) reported that the species appeared to have virtually disappeared by the time of
writing, and inferred that it may have been partly migratory, for at Lucknow it was only a very rare winter
visitor in cold weather, and its distribution may have been to some extent dependent on the rains. Bucknill
spoke of a number of known hunters in the provinces of Bihar and Orissa who had been attending big duck
shoots in Bengal for more than twenty years and had never seen a Pink-headed Duck, though they were
always on the look-out for it. He quotes a number of other accounts of the bird, and gives his reasons for its
extinction as follows:
It is non-migratory and confined to India, and consequently was shot at all the year by those to whom a close
season is a dead letter. There are probably many more fowling pieces in use among the ryots than there were forty
years ago. Then too, vast areas of swampy ground have been brought under cultivation within the last half-
century . . . it [probably] moults in the autumn and for a short period completely loses its power of flight – a
circumstance which would be another serious disability to its economy.
The last wild bird was trapped in June 1935 in the Darbhangra District, Bihar and given to Charles. M.
Inglis, the naturalist and curator of the Darjeeling Museum (Mehta 1960; Ripley 1962). An accepted obser-
vation of a wild bird was made in 1939 in the Burdwan district, and the last stems from Manroopa Lake,
Bakhtiarpur, Bihar in 1948–49 (Collar et al. 2001; Rhamani 2012). This latter record is based on a letter (Raj in
Singh 1966), but there are some doubts as to its authenticity. The letter states:
This is to inform the Bombay Natural History Society that, in the year 1947 on the 27th January in the
afternoon, I shot a Pink-headed Duck at Manroopa Lake in Khagaria subdivision, Dist. Monghyr, in my
estate area known as Bahadurpur Estate. The said duck, six in number, took off from Manroopa jheel and
came over me and I shot only one. Sir Hugh Dow, Governor of Bihar, and Mr. E. O. Lee, i.c.s., Member of
the Board of revenue, Bihar, were in the shoot. The latter gave me a letter of testimony which read that the
Pink-headed Duck is rarely found in India and he congratulated me, but this letter is misplaced at present
. . . Again in 1948–49 I saw some Pink-headed Duck on Lake Manroopa, about 5 to 8 of them, but they did
not come over me so I could not shoot.
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An editorial response to the claim mentioned that the letter of testimony was not traced, Mr. E. O. Lee had
died, and that Sir Hugh Dow remembered the shoot but not the shooting of a Pink-headed Duck; however,
Dow saw no reason to doubt the story. Unfortunately, the shot specimen was not preserved, which would
have proven the record beyond doubt. Other observations continue to be made, but none have offered any
absolutely confirming evidence. While considered extinct (Ripley (1950, 1952) thought it extinct since 1950,
with a spate of observations made in that year all referrable to the Red-crested Pochard Netta rufina), this
duck may just cling on.
Captive birds
A number of Pink-headed Ducks were kept in captivity. The first in Europe arrived at London Zoo on
January 12, 1874, with further arrivals in 1887, 1892, and 1897 (Prestwich 1974); one died at the Berlin Zoo in
1908, and an individual was received in 1912 by a G. D. Tilley at Darien, Connecticut, in the United States,
but it only survived for a few days (Philipps 1922–26). Calcutta Zoo kept Pink-headed Ducks on a number
of occasions, and a pair of males was present in 1923 (Prestwich 1974), and one died around 1930 (Ali 1960).
The most important collections of live birds were made by Alfred Ezra, an avid aviculturist. Ezra estab-
lished a menagerie at his home at Foxwarren Park, Cobham, Surrey from 1919 until the 1950s, and kept an
impressive collection of birds and animals, including Pink-headed Ducks (Ezra 1925, 1942; Delecaour 1956).
Ezra received five Pink-headed Ducks (three drakes and two ducks) in 1925 from his brother, Sir David
Ezra, who had offered rewards to Indian bird-catchers (Ezra 1925). Two died almost immediately, but by
1926 the others had settled well (Ezra 1926). Another 10 mostly male birds arrived in 1929 (Ezra 1930), from
which two pairs were sent to his friend, Jean Delacour, a renowned ornithologist (Prestwich 1974; Fuller
2013). They were received at Delacour’s residence, the Chateau de Clères, Normandy, where Delacour had
assembled a vast collection of animals and birds, including many rare species, housed in a zoo in the
grounds of the castle (Delacour 1941). The last of Delacour’s Pink-headed Ducks died in 1936 (Todd 1996).
Delacour remarked to Todd that ‘they were very stupid birds because they refused to breed’.
Ezra’s live birds were collected in southern Goalpara in Assam, India, and in eastern Rangpur in northern
Bangladesh, near the confluence of the Tista (Teesta) and Brahmaputra Rivers (Ripley 1962), but they have
not been recorded again in these regions. Ezra (1942) noted that the birds were long-lived, some living for
more than 10 years, but made no attempt to nest, despite being kept under seemingly ideal conditions. Any
chance of breeding Pink-headed Ducks in captivity ended during the winter of 1932, when the last female at
Foxwarren Park succumbed (Prestwich 1974). The last of the male Foxwarren Park Pink-headed Ducks
appear to have died by 1936 (Ali 1960), but Ali reports a letter by a Mr H. G. Deignan stating that Sir David
Ezra had a living male in his aviaries in Calcutta in 1945. If correct, this is the last record of a captive bird.
Habits
The ducks occurred by small ponds surrounded by bushes and high grass, and in colder weather resorted
to rivers. Hume & Marshall (1879–81) stated that they inhabited mainly lakes and swamps, and were never
seen on running water. Simson (1884) noted that the birds were non-migratory, living in small flocks of
8–12, which he believed to be family groups, and that they did not associate with other duck species.
Blanford (1898) said that the duck was found throughout the year in small flocks of 4–10, or occasionally up
to 40 in the cool season, and in pairs from April to September. F. A. Shillingford (in Hume & Marshall 1879–
81) noted that the gizzard of a bird he examined was found to contain ‘half digested water weeds and
various kinds of small shells’, while two stomachs examined by other hunters contained small shellfish, a
crab and small shells (see Collar et al. 2001). The breeding season commenced in April. The birds began to
build their nests in May, the eggs being laid in June and July. One nest was concealed in the middle of big
tufts of high grass not more than 500 yards (457m) from the water, and was formed of grass with a few
feathers, but, unlike most duck’s nests, was not lined. The eggs were unique among ducks, being quite
round and china white (interestingly, Baker (1908) wondered if the eggs Shillingford sent to Hume – the
only ones known – were atypical, since they were unlike normal duck eggs). The clutch size was 5–10. Both
male and female were known to have been put up from the nest simultaneously, but whether the drake
incubated or merely remained nearby is uncertain. Like many other bird species, the female would try to
lead predators away from the nest by feigning injury.
J. C. Parker (in Hume & Marshall 1879–81) reported that the flight of the duck was ‘very powerful and
rapid’, and Shillingford (in Hume & Marshall 1879–81) noted that the call was like that of a drake Mallard,
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‘with a slight musical ring about it’. Delacour & Scott (1954–64) described the breeding display of presum-
ably captive Pink-headed Ducks:
The voice of the male is a whizzy [wheezy] whistle recalling that of the Mallards, but lower and weaker. The
female has a low quack. The males display in company, again like Mallards, but much more simply; they puff out
their short feathers with the neck shortened and resting on the back, then stretch the neck upwards and utter
their call. The females show in a rudimentary way the posture of the dabbling ducks.
Finn (1915) stated that they made a characteristic wing-whistle when flying in an aviary.
Relationships
The relationships of the Pink-headed Duck have proved difficult to determine. Hume (in Hume & Marshall
1879–81) thought it was closely related to Anas, and that he would have merged the two but for the differing
eggs. However, Johnsgard (1961) believed the species to be closely related to the Netta and Aythya pochards,
based on examination of museum skins and feather protein analysis (Woolfenden 1961; Brush 1976), which
has proved to be correct. Rhodonessa has a sister relationship with the Red-headed Pochard Netta rufina
(Livezey 1998a; Thomas & James 2016). Thomas & James have further shown that the unusual pink feathers
are pigmented with carotenoids, which is extremely rare in waterfowl. Only the aptly named Pink-eared
Duck Malacorhynchus membranaceus from Australia exhibits similar pink plumage, but it is only distantly
related to the Pink-headed Duck, indicating multiple origins of these plumage carotenoids.
The Labrador Duck seems to have been always reported as rare, and perhaps was a species in natural
decline. It was said to be uncommon even in the 17th century, rare in the 19th, and had virtually disap-
peared by 1871 when one was shot near Great Manan Island, New Brunswick. The last definite record is of
one shot on Long Island in 1875; however, Gregg (1879) records one taken in Elmira, Chemung County,
New York on 12 December 1878, but as this specimen has disappeared its identity cannot be confirmed.
There were, however, conflicting reports of its abundance. Chilton (1997) succinctly summarised most of
what is known about the Labrador Duck, citing several sources that claimed that the bird was common, and
remarking that so much about the duck is speculative.
The known winter range was along the Atlantic coast of North America from Nova Scotia south to
Chesapeake Bay. Many birds were sold in the markets of Baltimore, Philadelphia, New York and Boston.
Wintering birds also occurred in New Jersey and New England, while a few individuals were seen in the
Bay of Fundy and around Grand Manan Island in New Brunswick. The reasons for this duck’s extinction
are not fully understood. Despite being occasionally shot it was little persecuted, and not considered good
eating.
Habits
Where exactly these birds bred is unknown, and there is no reliable information on the nest and eggs.
Supposed eggs of this species have been shown to belong to R
ed-breasted Merganser Mergus serrator and
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The Auckland Islands were discovered in 18 August 1806 by Captain Abraham Bristow and named after the
English politician Lord Auckland. The merganser, which was confined to the islands, was first recorded by
Jacquinot in 1840 (Kear & Scarlett 1970). Despite subfossil remains attributed to a Mergus sp. being found on
the New Zealand mainland, suggesting that the Auckland species once had a wider distribution, the skel-
etal remains differ, so the Auckland Islands Merganser appears to have been endemic to the Auckland
Islands (Murray et al. 2014). There is an unconfirmed report (see Westerkov 1960) of the bird having been
seen at Campbell Island to the south of the Auckland Group in the early 19th century by the Antarctic
explorer Robert McCormick (McCormick 1842; Young et al. 1997), who had previously seen the merganser
in the Auckland Islands in 1840 (McCormick 1884), but this was in error (Williams & Weller 1974; Giller et al.
2010). Waite (1909) searched for it unsuccessfully, and commented:
A sharp look out was kept along the coasts of the Auckland Islands for this bird, but it was not recorded as
having been seen. This is less remarkable when, as Captain Bollons later told me, he had not seen the bird on the
coast, but had found it occasionally at the head of the estuaries, and especially on the inland watercourses.
61
Another search was made between 1972–73 (Williams & Weller 1974), and this was also unsuccessful. Pigs
were imported to the islands in 1807 (Waite 1909), sheep and goats later (Chapman 1891), and mice were
inadvertently introduced with shipwrecks. Rats are believed also to have been introduced, though there are
none there now. Sealing and whaling stations were established in the early 19th century, though whaling
was abandoned in 1852. Wild dogs and rabbits were also introduced at some time. These factors must have
impacted on the merganser’s numbers.
The Auckland Island Merganser seems to have been already greatly reduced in its range by the early
Maoris, as a result of hunting for the pot. The species, in common with many wildfowl on remote islands
with no mammalian predators, was ridiculously tame. The reason for its final extinction is unknown, but
Kear & Scarlett (1970) thought that it may have been due to very large numbers having been shot during
1901 and 1902. The birds must have already been scarce due to introduced predators and habitat destruc-
tion. Waite (1909) emphasised that there was little doubt that the pigs had caused havoc among
ground-nesting birds. Williams (2012) has suggested that at the time of European colonisation, the Auckland
Islands population probably only numbered 20–30 pairs, with around 10 non-breeding birds. Their scarcity
was due to the lack of suitable habitat on the Aucklands, so it is no wonder that the birds succumbed so
quickly to extinction.
Habits
The merganser inhabited fresh and brackish waters of creeks and upper estuaries, occasionally wandering
into salt-water harbours. Isotopic analyses have shown that it fed predominantly in freshwater, and it was
observed foraging at the heads of estuaries, on the island’s watercourses, and in deep freshwater pools
(Williams 2012; Williams et al. 2012). Hutton (1901; see Williams et al. 2012) claimed to have taken a 90mm
Koaro Galaxias brevipinnis, a freshwater fish, from the bill of a merganser, while the stomach of the only
known preserved (and complete) specimen at Tring contained macerated fish bones, the mandibles of a
polychaete worm and an unidentified gastropod (Hume & Walters 2012; Williams et al. 2012). It possibly fed
on fresh-water shrimps, but this is unconfirmed. Little was ever recorded of its breeding. Kear & Scarlett
(1970) believed that the pair-bond may have been a long one. Four 7–10 day old ducklings were found in the
month of December; the clutch size numbered at least four. Adult birds in pairs were recorded in October,
November, January, May and July. Krone (in Williams 2012) provided a clue to the choice of nest site. He
implied that the nest was rarely seen, occurring on a steep cliff facing north or north-east, and that this duck
nested alongside cormorants and penguins. The eggs were never described. Unlike other mergansers that
dive when threatened, the Auckland Island birds were said to conceal themselves among rocks (Todd 1996).
Milliner (1999) reported the existence of a flightless merganser from the Chatham Islands, which has proved
to be a distinct, almost flightless species (Williams et al. 2014). It differed from the Auckland Islands species,
Mergus australis, in being smaller, approximately the same size as the Red-breasted Merganser M. serrator, in
having a shorter bill, and with the sternum, wing elements and pelvis even more reduced. Most of the
subfossil remains were found in the Te Ana a Moe cave near the island’s large and extensive saltwater Te
Whanga Lagoon (Worthy & Holdaway 2002; Williams et al. 2014). Isotopic analyses of subfossil remains
have shown that the Chatham Islands Merganser was a marine species (Williams et al. 2012), unlike other
mergansers, as the cranium has conspicuous salt-glands; this is indicative of a diet of marine prey from the
large Chatham Island lagoons. Although direct archaeological evidence is presently lacking (Williams et al.
2015), the arrival of humans with their commensal mammals, especially the Pacific Rat Rattus exulans,
would have rapidly exterminated the weakly volant Chatham Islands Merganser due to over-hunting and
rat predation of its eggs and chicks.
Murray et al. (2014) further demonstrated that mainland New Zealand also once harboured a merganser
population; and as it differed in its smaller size and other characters from the Chatham and Auckland
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Islands birds, they consider it to be distinct. It remains undescribed at present, but no doubt it will add to
the ever-increasing number of extinct birds from the New Zealand region.
Until recently, the validity of the New Zealand Musk Duck was surrounded by doubt. Harrison & Walker
(1970) concluded that this duck was not distinguishable from the Musk Duck Biziura lobata. However, Olson
(1977a) considered the evidence and suggested that the New Zealand form was larger, and therefore the
name should stand, pending further analysis. This was confirmed after detailed studies of all available
subfossil remains (Worthy 2002). The New Zealand Musk Duck had more robust legs and comparatively
reduced wings compared to its almost entirely aquatic Australian counterpart. The birds may have had
similar habits, preferring deep water in large, permanent swamps, lakes or estuaries, and were originally
more widespread on both North and South Islands (Lalas et al. 2014). The New Zealand Musk Duck may
have preferred larger prey than other duck species, including crayfish, large insects, molluscs and fish
(Tennyson & Martinson 2007). Like all extinct New Zealand anatids, this species probably succumbed to
over-hunting.
New Zealand S
tiff-tailed Duck Oxyura vantetsi Worthy
Oxyura vantetsi Worthy 2005a, p. 263 (Lake Poukawa, North Island, New Zealand)
Specimens Subfossil remains are in Wellington, New Zealand.
Status Known from subfossil remains only. Extinct since the 16th century.
Range North and South Islands, New Zealand.
The New Zealand Stiff-tailed Duck was a diminutive version of the Australian B lue-billed Duck Oxyura
australis, being about 10% smaller in size (Worthy 2005a). Otherwise it was osteologically similar. There was
no indication that the duck had reduced flight capabilities, so it probably migrated from one lacustrine
environment to another depending on seasonality or food requirements. It probably disappeared due to
over-hunting by the Maoris, some time in the 16th century.
The Hawaiian Islands, due to their isolation and absence of terrestrial mammals, witnessed an extraordi-
nary radiation of ducks and geese, which often reached great size and evolved specialised adaptations.
Most remarkable were the m oa-nalos, which superficially resembled geese, but actually evolved from
dabbling ducks (Sorenson et al. 1999). So different were these g oose-like ducks from other anseriforms that
Olson & James (1991) termed the name ‘moa-nalo’ for them, meaning ‘lost fowl’.
They had extremely large lamellae (tooth-like projections on their bills), huge robust legs and dispropor-
tionately small wings. From coprolite (fossilised dung) analysis, James & Burney (1997) suggested that
these birds were specialised for hindgut fermentation of plant fibres. Being flightless, they probably quickly
fell prey to Polynesian hunters, and eggs and chicks may have been vulnerable to the introduced Pacific Rat
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Rattus exulans. The moa-nalo Thambetochen chauliodous from Molokai and Maui was the first species discov-
ered, and appears to have been a lowland species.
This moa-nalo from Oahu was smaller and less robust than T. chauliodous, with a slightly longer, less
decurved bill. Subfossil remains were first discovered at Barbers Point, a lowland site situated on the Ewa
Plain in south-western Oahu (Holocene in age), and at Ulupau Head, a Pleistocene lake deposit in the crater
of Ulupau Head, Mokapu Peninsula, on the south-east coast (Olson & James 1991); the Oahu Moa-nalo, like
the Maui Nui species, seems to have been a lowland bird. Olson & James note that the differences between
the two are comparatively slight, which is surprising since the islands were never connected by a land-
bridge. What is certain is that both were to suffer the same fate and disappear for the same reasons: a result
of over-hunting, habitat loss and nest-predation by introduced mammals.
The Stumbling M oa-nalo was so named because of its propensity to fall into cave holes (Olson & James
1991), and thus become preserved in the fossil record. This species differed from Thambetochen in jaw
morphology, these being shorter and deeper with smaller tooth- like projections. It appears that the
Stumbling Moa-nalo was restricted to a montane habitat, and lived sympatrically with Thambetochen chauli-
odous on Maui. The mountain ranges of the Hawaiian Islands provided the last refuge for many n ow-extinct
birds; the Stumbling Moa-nalo may have survived longer
than other flightless anseriforms. Regardless, it almost
certainly disappeared for the same reasons, a combination Kauai Turtle-jawed Goose
of over-hunting with predation or competition by intro- Chelychelynochen quassus
duced exotic animals.
flightless. It would have been analogous to large terrestrial grazing mammals or giant tortoises on other
islands, and cropped tough vegetation with its specialised jaws. It has been suggested that some of the
Hawaiian plant genera, particularly the endemic lobelias of the genus Cyanea, evolved heterophyllic foliage
(different shaped leaves in the juvenile plant, which are seemingly inedible to herbivores) as a defence
against browsing by the large array of endemic geese on the islands (James & Burney 1997).
One of the most extraordinary bird species discovered in recent years was the Kaua’i Mole Duck Talpanas
lippa, a flightless, almost blind species, with highly developed tactile and olfactory senses. It was probably
analogous to the New Zealand kiwi Apteryx sp. (Iwaniuk et al. 2009). The Kaua’i Mole Duck was presum-
ably nocturnal, and used its broad, sensitive bill to forage for invertebrates on the forest floor. Like the Kiwi,
it was vulnerable to hunting and introduced predators, and would have disappeared soon after the human
settlement of the Hawaiian Islands.
Spheniscidae (Penguins)
Penguins are a group of flightless, aquatic birds almost exclusively confined to the Southern Hemisphere
and divided into six genera. They occur in Antarctica and some temperate regions, with some species
endemic to islands and island groups. They are vulnerable during the breeding season, when they form
rookeries on land. They have been over-hunted throughout their ranges, and as a consequence a number of
island species are threatened with extinction.
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The subfossil bones of this penguin are considered generically distinct, most resembling Megadyptes and
Eudyptes. The genus Eudyptes consists of about six species of small crested penguins, some of which breed
on islands in the southern Atlantic and southern Indian Oceans, but it is mainly concentrated in the New
Zealand region. Megadyptes is monotypic with M. antipodes, the Yellow-crowned Penguin, confined to New
Zealand and its offshore islands. It has been suggested that Tasidyptes hunteri may not have been native to
Hunter Island, but could have wandered there from an a s-yet unknown breeding locality (van Tets &
O’Connor 1983).
The Hunter Island Penguin may have disappeared as recently as c.1800 due to o ver-
hunting by
Tasmanian Aborigines, who themselves were wiped out by European settlers just 75 years later.
The Chatham Islands once harboured two species of penguin, the Chatham subspecies of the Fairy Penguin,
Eudyptula minor chathamensis, which is still extant, and an undescribed species of Eudyptes, known from
sand-dune subfossil remains collected on Mangere Island (Tennyson & Milliner 1994). The bones have been
variously referred to the Erect-crested Penguin E. sclateri and the Fiordland Crested Penguin E. pachyrhyn-
chus (Sutton 1979), but they differ from both of these species in having a deep but long, thin bill. This
penguin may have survived until as recently as 1872; Travers & Travers (1872) held a captive bird some time
between 1869 and 1872, and stated:
I obtained and brought to New Zealand a live specimen of this bird, which had come on shore to moult. I believe
it to have been a young bird. It remained for nearly three weeks without food, but on reaching New Zealand it
was fed partly on fish and partly on raw meat. It became very tame, following like a dog anyone who fed it. It was
unable to take its own food, which had to be placed in the gullet.
It became very fat and appeared to thrive, but, unfortunately, I
was unable to get fish for several days, owing to stormy weather,
during which it was fed on meat. It died somewhat suddenly,
which I attribute to the nature of the food, as, on being opened,
it presented no appearance of disease. It used its flippers in
climbing, and by their aid was able to travel up very steep
places if at all rough. Nothing could be more quaint than the
habits and appearance of this bird as it wandered about the
garden, or followed those it knew. Though generally considered
stupid, no doubt from its appearance, it was extremely cunning.
When placed at night in an enclosure with some poultry it
became master of the situation, its harsh cry and powerful beak
striking terror into the other occupants.
The extinction of this penguin was probably due to
over-hunting by early settlers and whalers.
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Procellariidae
The discovery of this extinct taxon was the result of mitochondrial DNA analysis, which showed that a
distinct penguin once occurred on coastal South Island, New Zealand. It was approximately 10% smaller
than the extant Yellow-eyed Penguin Megadyptes antipodes and differed in other characters of the skeleton.
Boessenkool et al. (2009) have shown that the highly endangered population of Yellow-eyed Penguin is a
comparatively recent arrival to New Zealand from sub-Antarctica (including the Aucklands), which only
became established after the extinction of the Waitaha Penguin; it exploited the suddenly vacant ecological
niche. Remains of the Waitaha Penguin are abundant in Maori midden deposits, so it probably became
extinct due to over-hunting soon after the Maori became established on New Zealand.
Procellariidae (Petrels)
The family Procellariidae are a diverse group of tube-nosed seabirds, which have been divided into 14
genera. They are distributed throughout the world’s oceans, and breed on mainland coastal areas, island
archipelagos and individual islands. They are often restricted to small breeding areas, where they show
strong site fidelity. Some genera, like the Pterodroma petrels, are nocturnal when breeding and nest in
burrows. They have been heavily exploited as a food source, and continue to suffer from habitat loss, pollu-
tion, and predation from introduced mammals. Some species are in critical danger of extinction.
Surprisingly little is known of the Jamaican Petrel, or Blue Mountain Duck as it was locally known. William
Thomas March, who had evidently seen the living bird, sent an account of its habits to Carte (1866), which
the latter quotes:
It is a night bird, living in burrows in the marly clefts of the mountains at the east and north-east end of the
island. The burrows form a gallery 6 to 10 feet long, terminating in a chamber sufficiently commodious to
accommodate the pair; from this they sally forth at night, flying over the sea in search of food (fishes) returning
before dawn. It is often seen on moonlight nights and at sunrise running about the neighbourhood of its domi-
cile, and sometimes crossing the road regardless of the labourers going to their work. I know nothing of its
nidification.
Little more is known about the bird. Bancroft’s nomem nudum Procellaria jamaicensis very probably refers to
this species, but he gives no description other than that the specimen he saw, brought by a Captain Pearce,
was dug out of its hole on the summit of the Blue Mountain Peak of Jamaica by a terrier:
. . . although not rare, it is with difficulty found, since it burrows only in crevices on the tops of our highest
mountains, scarcely accessible. The individual now sent was hunted by a terrier dog from a hole on the summit
of the Blue Mountain Peak, on 17th March last, and, as I am told, uttered the most piteous cries, like those of a
child, while being dragged forth. These birds are found in some number on that spot, and individuals have some-
times gone hither to hunt them. They probably resort thither chiefly in their breeding season, and are very
seldom seen flying except in the evening, when it is supposed that they proceed to sea. As they frequent this
island, and have not been observed elsewhere, the species, if new, might be called Proc. Jamaicensis.
Browne’s account is hardly more informative:
This bird is observed in all parts of the sea; it is rather smaller than a pigeon, of a dark or blackish colour, and flies
so close to the surface of the water that it frequently lies hid between the waves for a considerable time.
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The Jamaican Petrel was last recorded with certainty in 1880. The birds were formerly hunted with dogs
for food and this may have contributed to their extinction. A recent pelagic survey (Shirihai et al. 2010),
which searched the waters off Jamaica, Guadeloupe and Dominica intensively for this species, was
unsuccessful.
Specific status
The Jamaican Petrel has been considered to be a dark-phase or colour morph of the Black-capped Petrel or
Diablotin Pterodroma hasitata, a species that breeds on Hispaniola, formerly on Guadeloupe, and for a long
time was itself thought extinct. The Jamaican Petrel’s taxonomic status has still not been satisfactorily
resolved. Salvin (1896), for instance, points out that specimens are variable:
Some specimens are much greyer than others, both on the back, lesser wing-coverts, face, and under surface,
including the under tail coverts. Both forms are found in Jamaica, apparently living together.
March’s letter to Carte mentions that a Mr. [J?] Hill considered the dark Jamaican Petrel the same as the pale
Diablotin referred to by Atwood (1791) on Dominica, but the latter is much more likely to represent P. hasi-
tata. It is these accounts that have prompted modern writers to suggest that caribbaea and hasitata were dark
and pale phases of the same species. However, the available information suggests that it is just as likely that
the Jamaican Petrel was specifically distinct from the Diablotin.
The Providence Petrel Pterodroma solandri is a wide-ranging bird of the Pacific Ocean, but it breeds only on
Lord Howe Island and Philip Island (Marchant & Higgins 1990). A population formerly bred on Norfolk
Island, but no skin specimens are known to have been preserved. It was first mentioned by Lt. James King,
who was on Cook’s third voyage when Norfolk Island was discovered in 1777 (Iredale 1930), and appears
to have been wiped out there between 1788 and 1790 (Greenway 1967). The evidence for this is derived
from the diaries and letters of Lt. Ralph Clark, who speaks of the ‘Mount Pit birds’, which were extensively
harvested by the human population stationed on the island (see Whitley 1934). As there is a high degree of
island speciation among petrels, the Norfolk Island bird may have represented a distinct species or subspe-
cies related to P. solandri.
drop down out of the air as fast as people can take them up and kill them . . . They were at the end of May as plen-
tiful as if none had been caught, although for two months before there had not been less taken than from two to
three thousand birds every night.
By late September 1792, Collins (in Medway 2002) was told that the Mount Pitt Petrel was already in
serious decline, and no longer dependable as a food resource. Understandably, the petrel was unable to
sustain such huge losses, and became extinct less than 13 years after its discovery. The introduction of pred-
atory cats and rats would also have impacted on the population (Marchant & Higgins 1990), and these
vermin appear to limit the re-establishment of petrels on the island (Holdaway 1999).
A second seabird?
There may have also been a second species of petrel or shearwater on Norfolk Island. In a letter dated
February 11, 1791, addressed to a Captain Campbell (Whitley 1934), Clark refers to a second bird that bred
at a different time than the Mount Pitt petrels:
Everybody here owes their existence to the Mount Pit birds [Pterodroma solandri], before they entirely left us,
another bird came in and supplied their place, but was not attached to Mount Pit alone but was found in holes of
the ground in the day time, all over the island. They resemble the Mount Pit Bird in plumage, make, etc., all but
the feet, which are white, which the Mount Pit birds are not, they cut, when boiled, like mutton for which we
named them the flying sheep, they were also in abundance for six weeks, when their young took flight, and they
all left us.
Iredale and other writers have suggested that the unnamed species may have been the Flesh-footed
Shearwater Puffinus carneipes, which still breeds on Lord Howe Island, but has not occurred on Norfolk
Island in recent years.
This species was discovered in a flooded cave on Oahu and lava tubes in the North Kona district. It is a
small petrel, perhaps the smallest in its genus, with no obvious relatives among living species of Pterodroma
(Olson & James 1991). The only surviving Hawaiian petrel is the Bonin Petrel Pterodroma hypoleuca, which
was once found on the main islands, but now survives only on the islands of the north-west chain. The
Polynesian Rat Rattus exulans was responsible for the death of vast numbers of Bonin Petrel chicks and eggs
on Kure Atoll, in so much that no chicks survived during a breeding season in the 1960s (Seto et al. 1999).
Both of these Hawaiian petrels would have been subject to o ver-hunting by humans and predation by the
Polynesian Rat on the main islands, which no doubt led to their extermination.
Tennyson & Milliner (1994), Milliner (1999) and Cooper & Tennyson (2008) mentioned subfossil remains
referable to an extinct Pterodroma petrel collected on the Chatham Islands. The bird was larger than the
Chatham Petrel P. axillaris and the Black-winged Petrel P. nigripennis, but smaller than the Magenta Petrel
Pterodroma magenta, the three species that currently breed on the Chathams; it was also slightly larger than
the New Zealand breeding species, the Mottled Petrel P. inexpectata, which measures 33–35cm in length,
with a wingspan of 74–82cm. Using morphological and DNA evidence, Tennyson et al. (2015) proved
previous predictions correct and described a new species of gadfly petrel, P. imberi, from the subfossil
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remains, naming it in honour of Michael J. Imber, an expert on Pterodroma petrels. It was most closely related
to the Soft-plumaged Petrel P. mollis.
The subfossil remains of Imber’s Petrel were found in fossil dune sites alongside Magenta and Chatham
Petrels, and it was intermediate in size, which suggests that it had a different feeding ecology (Tennyson et
al. 2015). Tennyson & Milliner (1994) thought that the subfossil remains on Mangere were deposited before
1800, which suggests that Imber’s Petrel had probably disappeared before the arrival of Europeans.
Polynesian over-hunting caused their extirpation on Mangere, whereas introduced cats probably finished
them off on Pitt Island, possibly some time during the late 18th century (Tennyson & Millener 1994).
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Procellariidae
Bourne (1968) identified cranial and post-cranial remains of a large petrel discovered in caves on the Plaine
Corail, Rodrigues, which he referred to the Réunion Black Petrel Pseudobulweria atterrima, thus supposedly
confirming its presence on Rodrigues. Cowles (1987) referred post-cranial elements, also from Rodrigues, as
belonging to a new species of petrel from the island, and also supported Bourne’s (1968) identification of
some of the petrel material as a Pseudobulweria, thereby reinforcing the idea that the Réunion Black Petrel
once occurred on Rodrigues. However, both were incorrect in their interpretations. JPH has since found
large numbers of subfossil remains of an extinct petrel, almost certainly referable to Pterodroma, and larger
than any Indian Ocean species (Hume in prep.). It appears to have been restricted to Rodrigues. Tafforet
(1725–26) described a petrel nesting in holes in the centre of island, and this was the only time it was
mentioned. No doubt cats, introduced around 1745, and rats would have rapidly exterminated the birds.
The genus Bulweria has two extant but rather poorly known species, Bulwer’s Petrel B. bulweri, which
breeds in the North Atlantic, and Jouanin’s Petrel B. fallax from the north-west Indian Ocean. The Lesser St
Helena Petrel was a large species, about the size of Jouanin’s Petrel, at 31cm in total length. Olson (1975b),
after an examination of the bones, found that some of them resembled the Pterodroma gadfly petrels, whereas
others were like the related genus Bulweria, hence his name bifax – two-faced. Although similar in appear-
ance, the two genera differ in certain skeletal structures. On balance, Olson felt that the bird’s affinities were
more with Bulweria and he therefore assigned it to that genus.
The decline of the Lesser St Helena Petrel is shown by the diminishing presence of subfossil remains in
the more recent deposits, which Olson suggests is a possible result of some natural environmental influ-
ence. Any surviving population was probably extirpated after the arrival of Europeans and their
commensals in 1502.
This shearwater is smaller than the two other New Zealand endemic members of the genus, Fluttering
Shearwater P. gavia and Hutton’s Shearwater P. huttoni (Holdaway & Worthy 1994). Hutton’s Shearwater is
critically endangered today primarily due to predation by introduced animals (Cuthbert & Davis 2002),
while Fluttering Shearwater is not considered threatened. Scarlett’s Shearwater disappeared as a result of
the introduction of the Pacific Rat Rattus exulans, a lethal predator of seabird eggs and chicks, and habitat
modification and over-hunting by the Maoris. It had shorter wings than Hutton’s Shearwater, suggesting
that it was more sedentary (Tennyson & Martinson 2007).
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Four species of Puffinus shearwater once bred on the Canary Islands. The extant Manx Shearwater P. puffinus
and the Macaronesian Shearwater P. baroli have patchy distributions within the archipelego, whereas the
extinct Lava Shearwater P. olsoni and Dune Shearwater P. holeae (see below) were restricted to the eastern
Canary Islands of Lanzarote, Fuerteventura and some offshore islets (McMinn et al. 1990; Walker et al. 1990).
The Lava Shearwater was intermediate in size between the Manx Shearwater and Little Shearwater, but
smaller than the Dune Shearwater (Rando & Alcover 2008).
The occurrence of four sympatric species on the comparatively small Canary Islands is remarkable, but
might be explained by each species having separate reproduction times and choice of breeding site (Rando
& Alcover 2009). The Macaronesian Shearwater breeds on offshore islets and inaccessible cliffs, whereas the
Manx Shearwater breeds some distance inland and nests in Laurel forest on scarp slopes up to 1,000m in
altitude (Martín & Lorenzo 2001).
The numerous subfossil remains of the Lava Shearwater were discovered in lowland lava fields (McMinn
et al. 1990), where it probably nested in caves and lava fissures. This is in contrast with the Dune Shearwater,
which favoured sand dunes for breeding (Walker et al. 1990). The survival of the Manx and Little Shearwaters
and the extinction of the Lava and Dune Shearwaters was almost certainly a result of their breeding strategy,
as all species would have been hunted by humans for food, especially during the breeding season. The
Dune Shearwater became extinct soon after the arrival of people from northern Africa in the late Holocene,
around 3,300 years bp. This was probably due to the easy accessibility of the nesting sites (Rando & Alcover
2009). The nesting sites of the Lava Shearwater were much more difficult to locate, and radiocarbon dating
has shown that this species probably survived until at least the 15th century, after the arrival of Europeans
on the islands (Rando & Alcover 2008). However, it was rapidly exterminated due to o ver-hunting and
predation from introduced predators, especially rats.
Lava Shearwater
Puffinus olsoni
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Podicipedidae
Hydrobatidae (Storm-petrels)
The Hydrobatidae is divided into seven genera, which comprise some of the smallest of all seabirds. They
are cosmopolitan in distribution, being found in all of the world’s oceans. The storm-petrels are rather
similar in appearance, which makes identification difficult. Some species are very poorly known. They
exhibit strong breeding site fidelity, which has made them vulnerable to o
ver-hunting, habitat loss and
introduced predators.
The Guadalupe Petrel, discovered in 1887 and not seen alive since 1911, was related to the widespread
Leach’s Petrel Oceanodroma leucorhoa, and indeed was originally described as a race of it. It would probably
still be regarded as such but for the fact that a race of Leach’s Petrel also breeds on Guadalupe, and therefore
by definition the two are specifically distinct.
The Guadalupe Petrel was clearly a restricted and vulnerable species, as even on its only known breeding
station of Guadalupe it was confined to a ridge beneath the pine and oak cloud-forest on the top of Mount
Augusta, at the north end of the island. Greenway (1967) believed that imported domestic cats may have
been at least partly responsible for the bird’s extinction, but as Leach’s Petrel also bred on the island, there
may have been some measure of competition. In 1906, W. W. Brown (in Thayer & Bangs 1908) reported that
although the petrel was still abundant, cats did indeed prey on the colony, and summarised the situation as
follows:
The mortality among these birds from the depredations of the cats that overrun the island is appalling – wings
and feathers lie scattered in every direction around the burrows along the top of the pine ridge. The species,
however, is still breeding in large numbers in Guadaloupe [= Guadalupe], and sometimes at night the air
seemed to be fairly alive with petrels, their peculiar cries being heard on all sides.
Goats were also released on Guadalupe, and severe o
ver-grazing has been detrimental to all of the island’s
avifauna.
Habits
Anthony (1925) noted that the birds nested in burrows among the pines and oaks at about 760m (2,500ft)
above sea level. He explored the pine ridge in July 1922 but could find only a few very old burrows. It
bred earlier in the year than most of its relatives, from early March to mid-May, though a few eggs were
reported by Thayer & Bangs (1908) as late as June 17. The bird lined its burrows with leaves and pine
needles, and laid a single egg, white, with a wreath of minute reddish-brown or lavender dots at the
large end. Bryant (1887) described the call as here’s a letter, here’s a letter, with an answering call of for you,
for you.
Podicipedidae (Grebes)
The grebes are aquatic birds with torpedo-shaped bodies and lobed feet, occurring in both the Old and New
Worlds. The family is divided into six genera. Some species are widely distributed, whereas others are
restricted to single lakes. Two species have become flightless. The grebes with restricted distribution are
under extreme pressure from human habitat alteration.
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This grebe was first collected in 1929 and considered common (Delacour 1932), but from that point on the
species was in steady decline. The introduction of exotic fish, particularly Tilapia, may have depleted food
supplies. In addition, the increased use of gill-nets at Lake Alaotra undoubtedly killed many diving birds.
Severe deforestation and increased sediment input due to changes in agricultural practise have also lowered
the water quality of the lake. Since the 1940s and perhaps earlier, hybridisation with the African Little Grebe
has occurred; indeed, some consider the type specimen of the Lake Alaotra bird to be a hybrid (Voous &
Payne 1965).
Habits
Little was recorded about the habits of this bird, but see Safford & Hawkins (2013) for a comprehensive
overview. They were usually found in pairs, sometimes in association with Little Grebe T. ruficollis, a species
that colonised Madagascar as recently as 1945 (Milon 1946). Alaotra Grebes dived near lakeside aquatic
vegetation, feeding almost exclusively on small fish (Voous & Payne 1965). It appears that the species was
probably incapable of prolonged flight, so was restricted to Lake Alaotra and surrounding water bodies.
Well-grown young were collected in May and June (Fjeldså 2005), which suggests the breeding season
began early in the year.
This species was first discovered by Osbert Salvin in the nineteenth century (Salvin 1904), but he did not
consider the birds to be anything more than large specimens of the common Pied-billed Grebe Podilymbus
podiceps. The bird was not formally described until 1929 by Griscom (1932), who remarked:
This remarkable giant form is a very interesting illustration of variation correlated with isolation. Pied-billed
Grebes occur locally in parts of South America, and they all differ from North American birds in having slightly
longer, thicker bills, and darker underparts. These differences are carried to a remarkable extreme in gigas, which
is no more isolated than many another colony of this grebe, but those and other mutations have become enor-
mously developed and dominant, due to generations of inbreeding.
Prior to 1965, the grebe population was believed to be about 200 (Griscom 1932, Bowes & Bowes 1972),
but a decrease to 80 in 1965 resulted in strenuous efforts to conserve it (LaBastille 1974). In 1975, the popula-
tion rose to 200, but then crashed to about 50 birds in 1983 (LaBastille 1984). In 1987, Hunter (1988) conducted
studies on Lake Atitlán, and came to the conclusion that all the grebes present were Podilymbus podiceps. It is
not known when the smaller species first took up permanent residence on the lake, though it had been
known to winter there. Whether the common species had out-competed the larger flightless form or hybrid-
ised with it is not clear, but the introduction of exotic fish, drastic lake habitat modification and over-use of
gill nets are cited as the primary causes of extinction.
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Podicipedidae
Atitlán Grebe
Podilymbus gigas
Habits
Griscom (1932) found that the Atitlán Grebe was very wary, which he believed to be the result of it being
‘assiduously hunted by the natives’. It spent the day out in the lake far from shore, and could not be
approached within gun-shot range from a boat. The birds were strongly territorial, and breeding territories
were fiercely defended. The pairs displayed in formation with each other, which included ducking dives,
synchronised neck movements and bill-dipping (Fjeldså 2005). The breeding season began in March, with
egg-laying in April; clutch size was 1–5, and the chicks became independent after 10–12 weeks (LaBastille
1974). The Atitlán Grebe is believed to have fed mainly on the large native crab Potamocarcinus guatemalensis,
populations of which were devastated by the introduction of Largemouth Bass Micropterus salmoides to the
lake in 1960 (LaBastille 1990), but fish also made up a large part of the diet. The bird mainly called in the
early morning and early evening, and the usual contact call was a cuh-cuh, with a warning call poc (from
which it got its onomatopoeic local name). The pairs often called in duet, which consisted of a rapid pene-
trating chatter, and a loud, intense, undulating cut-cut-cuh-cuh-cuh, turning into a two note puk-puk, and
finally two or three braying notes eeyaw-eeyaw (Fjeldså 2005).
When the Colombian Grebe was first collected in 1945 by Borrero (Borrero 1947), it was reported as being
abundant. However, it had declined to about 300 birds by 1968. Only two records exist after that date; one
was seen in 1972, and the last definite sighting was of three birds in February 1977 (Llima & del Hoyo 1992).
An International Council for Bird Preservation expedition to Lake Tota in 1982 failed to find it (Varty et al.
1986).
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The direct reason for the bird’s decline and extinction appears to be habitat modification. Fjeldså (1984,
1993) suggests loss of habitat due to drainage and siltation, coupled with increased pollutants that destroyed
the lake’s Potamogeton pondweed, an alga perhaps vital to the grebe’s ecology. The introduction of exotic
Rainbow Trout Salmo gairdneri to Lake Tota in 1944 may also have adversely affected the grebe’s food
resources. Hunting in the breeding colonies may have also hastened its extinction (Fjeldså 2005).
Habits
The Colombian Grebe was an inhabitant of medium to large freshwater lakes with surrounding reeds, and
possibly fed by picking small arthropods from the weedy shallows. Borrero (1947) recorded that females
were ready to lay in August on Lake Tota; otherwise little more is known of the habits.
Relationships
A phylogenetic and population genetic study suggests that the Colombian Grebe derived from a colonisa-
tion event of South America by an ancestral group of grebes related to the North American race of
Black-necked (or Eared) Grebe Podiceps nigricollis californicus (Ogawa et al. 2015). The authors also demon-
strate that the Colombian Grebe was a recently established lineage clearly capable of rapid functional
response to a new habitat, and it was probably not genetically distinct enough from Black-necked Grebe to
warrant full species status.
Threskiornithidae (Ibises)
The ibis are medium-sized wading birds with long, decurved bills. They are wetland or forest birds, and
distributed widely in Africa, Eurasia, the Americas and several island groups. The family is divided into 13
genera. Most species are not threatened, but some have restricted ranges and are in critical danger of
extinction.
Until the discovery of subfossil remains, it was generally believed that a species of white dodo (see p. 423),
related to the more familiar Dodo Raphus cucullatus of Mauritius, once existed on the island of Réunion. The
basis for this belief arose entirely from early travellers’ accounts, and a series of white dodo paintings
executed in the 17th century (Hume & Cheke 2004). No specimens of the Réunion Ibis were collected. The
first descriptive account was made by Melet (1672) in 1671:
. . . [an]other sort of bird called solitaires which are very good [to eat] and the beauty of their plumage is most
fascinating for the diversity of bright colours that shine on their wings and around their neck.
Sieur Dubois, who visited Réunion in 1671–72, gave the best descriptive account:
These birds are so-called [solitaires] because they always go alone. They are as big as a large goose and have a
white plumage, black at the tip of the wings and tail. At the tail there are feathers approaching the tail of the
ostrich. They have a long neck and the beak made like that of the woodcocks, but bigger, and legs and feet like the
turkeys. This bird is caught by running after it, since it flies only very little.
Dubois’s account clearly describes a bird other than a dodo, and the discovery of skeletal remains revealed
that the ‘solitaire’ of his account (not to be confused with the Solitaire of Rodrigues; see p. 158) was in fact an
ibis (Mourer-Chauviré et al. 1995a, 1995b), a bird that better corresponds with the early descriptions.
Morphological details suggest that it was closely related to the Sacred Ibis T. aethiopicus of Africa or
Straw-necked Ibis T. spinicollis of Australia (Mourer-Chauviré et al. 1999).
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Threskiornithidae
Réunion Ibis
Threskiornis solitarius
Extinction
The introduction of predators such as cats is among the main causes for the extinction of the Réunion Ibis.
Overhunting was doubtless also important. In 1671, Melet (1672; see Cheke & Hume 2008) gave a good but
damning account of the slaughter of birds (presumably including the ibis) that took place on Réunion in the
early years of settlement:
There are birds in such great confusion and so tame that it is not necessary to go hunting with firearms, they can
so easily be killed with a little stick or rod. During the five or six days that we were allowed to go into the woods,
so many were killed that our General was constrained to forbid anyone going beyond a hundred paces from the
camp for fear the whole quarter would be destroyed, for one needed only to catch one bird alive and make it cry
out, to have in a moment whole flocks coming to perch on people, so that often without moving from one spot one
could kill hundreds.
The Réunion Ibis appears to have been a montane species, perhaps restricted in its distribution. Its range
swiftly contracted around the remote mountaintops as Réunion became more populated. The last definite
reference to them dates from 1708 (Cheke & Hume 2008), when Feuilley (1705) made the following remark:
The solitaires are the size of an average turkey cock, grey and white in colour. They inhabit the tops of moun-
tains. Their food is only worms and filth, taken on or in the soil.
Réunion’s endemic ibis became extinct soon after. Over-hunting and the introduction of predators such as
rats and cats are probable reasons for its extinction.
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The discovery of the subfossil remains of flightless ibises in different localities on Maui appears to represent
two species. One of these, the smaller, Apteribis brevis, was restricted to montane regions, occurring on the
southern Slopes of the Haleakala Volcano, including the Kipahulu and possibly the Lower Waihoi Valleys
(Olson & James 1991).
A second, larger and undescribed form occurred in the lowlands from sea level to at least 808m, and
perhaps up to 1,145m in altitude. The bill of the undescribed lowland bird was much longer than the
montane, A. brevis, being more similar to the lowland Molokai Flightless Ibis, A. glenos, but more decurved.
Skeletal remains of these ibises were found beautifully preserved in lava tube cave deposits, where a
roof-collapse had occurred. These acted as natural pit-fall traps into which the birds fell and were unable to
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Ardeidae
escape. Both A. brevis and the undescribed lowland form fared no better than their Molokai congener, and
probably disappeared rapidly after Polynesians arrived on the Hawaiian Islands.
The New Zealand Little Bittern was first obtained at Tauranga in 1836 by Rev. Stack, and never again
seen in North Island; this bird was presumably a wanderer. In 1861, Ellman listed a Little Bittern for
New Zealand:
A missionary, a good naturalist, once assured me he shot a diminutive bittern, resembling our own little bittern,
but in travelling he lost the skin.
Rev. W. Colenso (in Buller 1887–88) elaborated on this:
As far back as the year 1836 the Rev. Mr. Stack obtained at Tauranga a specimen of the Little Bittern, and sent it
to the late Gilbert Mair, Esq., J.P., who presented it to me. It was alive in my possession for some time, and I ulti-
mately sent the skin to the Linnean Society. None of the natives in the district knew the bird.
In March 1868 two specimens were taken at Kanieri River in Westland by a Mr. Shaw, and forwarded to the
Canterbury Museum. Only ten more have been found, nearly all from Westland. Docherty, who collected
some of the known specimens, sent Buller a note on the bird’s habits:
They are to be found on the s alt-water lagoons on the seashore, always hugging the timbered side of the same. I
have seen them in two positions, viz.: – standing on the bank of the lagoon, with their heads bent forward, studi-
ously watching the water; at other times I have seen them standing straight up, almost perpendicular; I should
say this is the proper position for the bird to be in when stuffed. When speaking of lagoons as the places where
they are to be found, I may mention that I caught one about two miles in the bush, on a bank of a creek; but the
creek led to a lagoon. They live on small fishes or the roots of reeds; I should say the latter, because at the very
place where I caught one I observed the reeds turned up and the roots gone. They are very solitary, and always
found alone, and they stand for hours in one place. I heard a person say that he had opened one and found a large
egg in it. They breed on the ground in very obscure places; I never heard their cry.
Buller also quoted Potts in an account of specimens in Canterbury Museum:
They were taken alive without any great difficulty, after which they were turned loose amongst the fowls in a
poultry-yard. They were found dead shortly afterwards – it is alleged, from exposure to the keen frosty night air,
being deprived of the accustomed protection afforded by the thickly-growing sedgy vegetation of their swampy
habitat. They had been observed standing motionless on a bare stem or stalk, from which they overlooked the
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The Mauritius Night Heron was placed in Butorides by Newton & Gadow, but Rothschild (1907a), while
naming the bird Ardea mauritiana, stated that it was probably a Nycticorax. This was confirmed by Cowles
(1987). This species was closely related to the Rodrigues Night Heron N. megacephala (see below), but was
less robust in the leg elements. Contrary to reports that this species was flightless, there is no evidence to
suggest that this was the case.
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No definite records of this bird exist except the account of Leguat (1708), who mentions ‘great flights of
bitterns’ while on Mauritius in 1693. Reasons for extinction are unknown, but no doubt introduced preda-
tors such as cats were primarily responsible.
Both Leguat (1708) and Tafforet (1725–26) spoke of ‘Bitterns’ that occurred on Rodrigues, and Hachisuka
correlated these accounts rather arbitrarily with the bones described by Milne-Edwards, though the latter
believed his bones to be identical with the bird described by Leguat (Milne-Edwards 1875). Hachisuka
(1953) stated erroneously that the bird was flightless and ignored Tafforet’s account:
There are not a few Bitterns which are birds which only fly a very little, and run uncommonly well when they are
chased. They are of the size of an Egret and something like them.
Günther & Newton (1879) suggested that this species probably had poor powers of flight and was becoming
behaviourally flightless. Cowles (1987) re-examined the skeletal remains and concluded that the wings
were not particularly short, but the legs had become longer and more robust, characteristics applicable to a
more terrestrial species. The beak was also comparatively large. According to Leguat, the Rodrigues Night
Heron readily fed on the endemic Phelsuma day-geckos, and he stated:
We had an abundance of Bitterns, as big and as good as capons; they are more familiar, and more easily to be
caught than the Woodhens [Rodrigues Rail Erythromachus leguati ... They [Rodrigues Day-gecko Phelsuma
edwardnewtoni] serve for Prey to some Birds, especially the Bitterns. When we beat ’em them down from the
Trees with a pole, these Birds would come and devour them before us, tho’ we did our utmost to hinder them; and
when we offer’d to oppose them, they came on still after their Prey, and still follow’d us when we endeavour’d to
defend them.
Nothing more is known about this heron. Pingré (1763) could not find the birds in 1761, so they probably
disappeared during the intervening years as a result of severe deforestation and introduced predators,
especially cats.
Dubois (1674) visited Réunion in 1672–73, and is the only person to record this species in life:
Bitterns or Great throats, large as big capons [domestic fowl], but very fat and good [to eat]. They have grey
plumage, each feather tipped with white, the neck and beak like a heron and the feet green, made like the feet of
‘Poullets d’Inde’ [domestic turkey Meleagris gallopavo]. They live on fish.
The description is reminiscent of juvenile Nycticorax. Unlike the other Mascarene night herons, the Réunion
species shows no reduction in wing elements, so was probably a capable flier ( Mourer-Chauviré et al. 1999).
The reason for its extinction is unknown, but it would have been subject to the same introduced predators
as its relatives on Mauritius and Rodrigues. Nothing else is known and the bird cannot be identified with
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any extant species. Hachisuka (1937b) said little more except to place the bird into his own (now invalid)
genus, Megaphoyx.
The range of the Rufous Night Heron Nycticorax caledonicus extends from Australia through Sulawesi, the
Palau Islands and New Caledonia to the Philippines. There was once an isolated and distinctive race on the
Ogasawara, or Bonin, Islands. The first specimen was collected in 1827 on Captain Beechey’s voyage to the
Pacific and the Bering Straits, and was described by Vigors (1839) as follows:
This species agrees in every respect with the Nyct. Caledonica, in its colours and the distribution of them; with
the exception of the colour of the bill, which is black in the latter bird. It differs essentially, however, in the shape
of the bill, which is much more solid and nearly straight, approaching in this respect to the bill of the Bitterns.
The proportions of the wing are also different, the length, from the carpal joint to the extremity of the largest quill
feather, being an inch less in our bird than in the allied species. Several of these birds were seen at the Bonin
Islands. Mr. Collie remarks that ‘three white tapering feathers formed a crested plume on these birds, but many
were shot without the crest. They frequented the rocks on the sea-shore. They had one caecum’.
Kittlitz (1832–1833) recorded it in 1828 and identified it with the Rufous Night Heron, while Vigors’s type
was placed in the collection of the Zoological Society and later transferred to the British Museum (see
Walden 1877; Seebohm 1890a), and then to Tring.
The Bonin Night Heron nested in low trees near the coast and frequented beaches and marshes. The last
specimen (now at Tring) was collected in 1889; subsequent Japanese expeditions failed to find this bird. The
cause of extinction is not known, but human interference is believed to have been responsible.
This large species of night heron occurred on the island of Niue in the south-west Pacific. Its closest relatives
were an undescribed subfossil species from Tonga, and the extant Rufous Night Heron N. caledonicus (which
occurs in New Caledonia). The Niue Night Heron was characterised by having a long, straight upper bill,
shorter wing elements (ulna, carpometacarpus) compared with the leg elements, and stouter legs (Steadman
et al. 2000).
Nothing is known about this species but Steadman et al. (2000) suggest that the Niue Night Heron may
have fed on the abundant crabs present on Niue, much as the extinct Bermuda Night Heron Nyctanassa
carcinocatactes (see p. 83) did on Bermuda (Olson & Wingate 2006). The reasons for its extinction are prob-
ably over-hunting by humans and predation by introduced mammals.
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Ascension lsland is a rather barren, inhospitable place, but it originally supported at least two endemic
terrestrial birds, the Ascension Rail Atlantisia elpenor (see p. 117) and a now-extinct species of night heron.
The Ascension Night Heron was described from comparatively ancient subfossil remains, discovered in
guano deposits and fumaroles (Olson 1977c). The presence of an endemic night heron on Ascension is
somewhat surprising, since the island’s harsh environment offers little in terms of the usual night heron
habitat. Despite this, Olson suggested that it was probably a resident species, as one individual represents a
juvenile bird, i.e. the population was breeding rather than vagrant. The proportions of its skeleton show that
it was smaller than the Black-crowned Night Heron N. nycticorax, the species it was most likely derived
from, with shorter wings and more robust legs, and it probably had poor flying ability (Bourne et al. 2003).
The bird may have survived by feeding on the eggs, young or waste and invertebrates associated with large
colonies of seabirds, or it may have foraged on the coasts.
The Ascension Night Heron possibly survived until the arrival of the first human colonists at the start of
the 16th century, but being terrestrial and a probable ground-nesting species, disappeared shortly after due
to the introduction of cats and rats.
Thévet
A reference by André Thévet in 1555 (see Gaskell 2000), in which he mentions a flightless bird called
‘Aponar’, has sometimes been considered referable to the Ascension Night Heron. Thévet stated that:
Furthermore in this island there is a certain kind of great birds that I have heard called Aponars; they have little
wings, and therefore they cannot fly. They are great and high like herons, the belly white and the back black as
coal, the bill like to a cormorant; when they are killed they cry like hogs.
The name Aponar is a derivation of the word ‘Apponath’, which had previously been used for the Great Auk
Pinguinus impennis by Jacques Cartier in 1534 (Fuller 1999). The auk never occurred anywhere near to
Ascension, but it would have been known to sailors that visited the island in the early days of colonisation.
There was clearly some confusion: perhaps the name was transferred from the auk to another bird that, on
the face of it, was broadly similar – a large, flightless, black-and-white bird with a large bill, the night heron
(in much the same way that usage of the word ‘penguin’ switched from the Great Auk to its Southern Ocean
equivalents during the late 16th century).
The Bermuda Night Heron was almost certainly a derivative of the Yellow-crowned Night Heron N. viol-
acea, a crustacean-feeding species that is a regular vagrant on Bermuda. Inferences made from early but
brief accounts suggest that a resident population once occurred there (Olson & Wingate 2006). The birds
appeared to be ridiculously tame and one report by Sylvanus Jourdan in 1610 (see Olson & Wingate 2006)
stated:
There are also great store and plenty of Herons and those so familiar and tame that wee beate them downe from
the trees with stones and staves: but such were young Herons: besides many white Herons, without so much as a
blacke or grey feather on them . . .
The accounts mention grey, white and black birds, coloration suitably appropriate to adult and juvenile
Nyctanassa night herons. The Bermuda birds differed from the Yellow-crowned Night Heron in having an
extremely short but heavy bill with a wide gape, a more robust skull, and shorter, more robust leg bones.
Olson & Wingate suggest that the Bermuda Night Heron was adapted to feed on large land crabs (including
one extinct Bermudan species), and that the specialised bill with wide gape was perfectly suited to feed on
this prey.
The night heron became extinct shortly after the colonisation of Bermuda by humans, and being terres-
trial, probably succumbed to introduced predators and perhaps a shortage of prey.
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Sulidae (Boobies)
The boobies generally occur in subtropical or tropical oceans, although the Morus gannets occur in temperate
zones. The Sulidae have been divided into three genera. They are not truly pelagic, and usually remain in
close proximity to coastal land masses.
Abbott’s Booby Papasula abbotti was originally placed in the genus Sula; however, it has been shown to be
morphologically distinct and warrants generic status (Olson & Warheit 1988). Abbott’s Booby breeds only
on Christmas Island in the Indian Ocean, where it is considered endangered, but vagrants are known to
occur in the Pacific Ocean (Pratt et al. 2009).
Abbott’s Booby Papasula abbotti is now confined to Christmas Island in the north-east Indian Ocean, but it
once had a much greater range. The type specimen of the species, a skin, was taken on Assumption Island,
in the south-west Indian Ocean (Ridgway 1893), and Steadman et al. (1988) have collected subfossil remains
from Tikopia, an island in the Solomon Islands in the Pacific. The Assumption population became extinct in
around 1909 as a direct result of habitat destruction during guano excavation (Stoddart 1981, Skerret et al.
2001). The relict Christmas Island birds are endangered.
The Ua Huka subspecies of Abbot’s Booby was larger in all elements than the nominate bird, and, pending
the discovery of more material, may represent a distinct
species (Steadman et al. 1988). Subfossil remains of the Ua
Huka Booby are from archaeological midden sites, which Ua Huka Booby
indicate that it was regularly hunted by Polynesians. Papasula abbotti costelloi
Abbott’s Booby today nests only in tall trees, so the Ua
Huka Booby would have also been affected by habitat
destruction. It probably disappeared rapidly after human
colonisation of the Marquesas Islands.
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Phalacrocoracidae (Cormorants)
Cormorants are long-necked, slender diving birds that occur on coasts and inland waterways around the
world. They are all piscivorous, and characteristically swim low in the water and periodically dry their
wings while on land. In general, they are not threatened but some are restricted in range.
Newton & Gadow (1893) erroneously described subfossil remains from the Mare aux Songes, Mauritius, as
belonging to an anhinga or darter. However, Olson (1975a) pointed out that these bones are referable to a
small cormorant (not an anhinga), closely related to the wide-ranging African Reed Cormorant Phalacrocorax
africanus. Further subfossil remains have since been recovered (JPH, in prep.), which indicate that the
Mascarene bird was smaller than the African mainland bird, though otherwise similar. A race, pictilis, occurs
on Madagascar, and this may have been related to the Mascarene population, but it is more likely that the
Mascarenes once harboured their own subspecies. Its disappearance was probably due to over-hunting and
perhaps predation by cats (Cheke 2013b).
Habits
The Mascarene Reed Cormorant was never recorded alive on Mauritius, but on Réunion it was mentioned
by two competent observers; Dubois in 1671–72 (Dubois 1674) and Feuilley (1705). The last account is the
most detailed:
The cormorants are the size of a duck and the same form, except their neck is a bit longer; [they] live in the meres
like flamingos and live in the same way. They are not eaten unless very young. They smell strongly of the marsh
and the wild.
Nothing else is known about the bird in life.
The explorer Georg Steller, who visited Bering Island on Bering’s i ll-fated expedition in 1741, was the only
naturalist to record the Spectacled Cormorant in life. Much of what we know about Steller and the
Spectacled Cormorant is due to the research of Leonard Stejneger (1925, 1936), who wrote several papers on
the bird and a biography of the naturalist, as well as translating and editing Steller’s diary for publication in
1925.
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Steller
Georg Wilhelm Steller was born in Windsheim, near Nuremberg, on 10 March 1709, and studied at
Wittenberg, Leipzig, Jena and Halle Universities (see Stejneger 1936). After graduating, he became an army
surgeon and left Germany for St Petersburg, where, soon after his arrival, he became physician to the
Archbishop of Novgorod. In 1737 he persuaded the Academy of Sciences to send him on Bering’s expedi-
tion to Kamchatka and the Bering Sea. The expedition encountered various disasters, including being
ship-wrecked, and ultimately the party became stranded on Bering Island on November 5, 1741, where
Bering himself died a month later. After building a new boat, Steller organised the successful return of the
survivors to Kamchatka.
It was during the expedition’s stay on Bering Island that the Spectacled Cormorant, so-called because of
the patches of bare skin or ‘spectacles’ around its eyes, was first described. It was certainly eaten by Steller
and his comrades; they used the Kamchatkan native cooking method of encasing the bird in clay with all its
feathers, and then baking it in a heated pit (Greenway 1967). Although cormorants are not generally consid-
ered palatable, Steller claimed that it was a juicy meal after this culinary treatment, and that one bird was
large enough to feed three starving men.
Steller was never to return to Europe. He spent four years wandering Siberia, studying plants on the
way. Worn out by the hardships suffered on his travels and by his heavy drinking, which grew progres-
sively worse, he was taken ill at Tyumen and died on 12 November 1746, aged only 37.
Pallas
For nearly a hundred years nothing more was heard about the Spectacled Cormorant until 1837 or 1839,
when Captain Belcher of The Sulphur visited Sitka in Alaska and was given a skin specimen by Governor
Kuprianoff, one of several in his possession. Three others were sent to St Petersburg shortly afterwards, one
of which was given to Leiden, and these apparently constituted the only ones known to Stejneger at the
time he was writing in 1899. The origin of these skins is uncertain, but Greenway’s (1958) statement that all
the known specimens were collected by Steller does not seem to be correct. Gould (1843) wrote up the
zoology of the Voyage of the Sulphur, and acquired a fifth skin, which is now at Tring. Stejneger (1925) refers
to a total of six specimens, but seven are now known, all apparently derived from Governor Kuprianoff.
Habits
At the time of Steller’s visit to Bering Island, the Spectacled Cormorant was considered a common bird,
occurring on its rocky shores. It seems, however, that it did not breed there, but only on the offshore islets,
Spectacled Cormorant
Phalacrocorax perspicillatus
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due to the large population of predatory Arctic Foxes Vulpes lagopus on the main island. These probably
constituted its only natural enemy. About this time or shortly afterwards the bird seems to have died out,
and there was no trace of it in 1882 when Stejneger (1883) visited Bering Island. He wrote:
The natives . . . remember very well the time when it was plentiful on the rocks, especially on the outlying islet
Are Kamen [= Riy Kamen]. About thirty years ago, they say, the last ones were seen, and the reason they give
why this bird has been exterminated here on the island is that it was killed in great numbers for food. They
unanimously assert that it has not been seen since, and only laughed when I offered a very high reward for a
specimen.
Stejneger found some subfossil bones of the bird on Bering Island in 1882, and Hartert (1920) said that
‘Sokolnikoff collected some sterna’. This suggests that the bird may once have been more extensive in range,
possibly occurring on other islands in the Commander Group and on the Aleutians. No eggs are known to
have been collected. It was described as a clumsy bird on land, possibly with reduced flying capabilities,
but not flightless. These attributes would have made it extremely vulnerable to human hunters and
predators.
Extinction
In 1826 the human population of Bering Island increased dramatically when a Russian–American Company
settled a number of Aleuts on it, and they apparently considered the cormorant a tasty delicacy. Fur trap-
pers also frequented the islands and no doubt killed the birds for food at every opportunity, so it is likely
that over-hunting was the primary cause of extinction. However, Hartert (1920) suggested disease as a
possible culprit:
As this cormorant is described as very stupid and was much persecuted, it is generally supposed that it was
exterminated by men. This may be, but it is perhaps as likely that it was finally wiped out by an epidemic disease,
such as took place in the winter of 1876–1877 when many thousands of Phalacrocorax pelagicus [Pelagic
Cormorant] were destroyed and masses of dead birds covered the beach all round the islands. The inhabitants of
the islands, however, asserted that it had been last seen about 1852 or so.
Anhingidae (Darters)
Darters or snakebirds are slimline, l ong-necked birds with a long, sharp bill and a wide distribution, occur-
ring in America, Africa, Asia and Australia. Four species are recognised; some populations of the Asian
species, the Oriental Darter Anhinga melanogaster, are considered near-threatened due to loss of habitat.
The Levant Darter is doubtfully distinct from the nominate race of African Darter A. r. rufa. The birds were
found in Amik Gölü (Lake Antioch) in south-central Turkey, and the marshes of the lower Euphrates and
Tigris rivers in southern Iraq. The Amik Gölü birds were partly migratory, wintering south to the Hula
valley in n orth-east Israel, with lesser numbers reaching the coastal plain of n
orth-west Israel. The popula-
tion in Iraq was more sedentary.
By the 1930s, the Amik Gölü birds had begun to gradually disappear due to the almost complete drainage
of the surrounding marshes, and the last birds wintering in the north of Israel were recorded in 1957. At
least 110 birds were counted in Iran in 1990 (Orta 1992), but no more information is available as to their
status. The Iraq population was considered comparatively stable, but the drainage of almost 90% of the
Mesopotamian marshes following the 1991 Gulf War has resulted in the probable extinction of the Levant
87
Darter. The marshes are now undergoing a re-irrigation programme but this has probably arrived too late to
the save the darters. Amik Gölü is now completely desiccated.
Little was recorded about the birds in life, other than they occurred in freshwater lakes characterised by
fringes of tall gasab reeds.
The former presence of an enormous eagle in New Zealand caused great interest among ornithologists at
the time of its discovery in the 1870s. Its massive size, and the fact that its bones occurred in swamps along
with those of moas, led to considerable speculation. Forbes (1892d) believed it to be a carrion-eater, relying
on dead moas; however, Owen (1879) was of the opinion that it actively preyed upon them. The eagle was
first thought of as a huge harrier that had followed flocks of moas across grasslands, but it is now well-
established that the pre‑human vegetation of New Zealand was forest or scrubland (Worthy & Holdaway
2002). On account of its supposedly short wing bones, various authors were led to believe it was a poor flier,
and Duff (1949) postulated that the eagle was evolving towards flightlessness.
Systematics
The systematic position of Haast’s Eagle has long been under debate. Holdaway (1991) analysed the meas-
urements of bones of moorei and assimilis (the two species originally described by Haast), which differed
only in size, and concluded that these represented males and females of one species: Haast’s Eagle was
sexually dimorphic. The two sexes, therefore, were different in their locomotion and predatory potential,
and may well have taken different prey; such sexual dimorphism (and partitioning of prey resources) is
relatively common in raptors. Holdaway suggested that Harpagornis, the monospecific genus into which the
bird was originally placed, was closely related to Aquila, and this now proved to be the case (Lerner &
Mindell 2005; Gill et al. 2010). Haast’s Eagle is now considered part of that widespread genus. DNA analysis
(Bunce et al. 2005) has shown that it formed a clade with two small forest eagles, Aquila morphnoides, the
Little Eagle, and A. pennatus, the Booted Eagle, both of which have weights of c.1kg and wingspans of
c.1.2m. The estimated weights and wingspans of male (11.5kg/2.14m) and female (over 14kg/2.43m)
Haast’s Eagle (Braithwaite 1992) mean it was 30% heavier than the world’s largest living eagle, the Harpy
Eagle Harpia harpyja of South America. Furthermore, it evolved from a small, lightweight forest raptor of
about 1kg to become the world’s largest eagle and, at up to 13 times heavier than its ancestor, the heaviest,
all occurring in a period of between 0.7 and 1.8 million years (Bunce et al. 2005) – a remarkable evolutionary
event.
Habits
Bones of Haast’s Eagle have been recovered from caves, swamps, estuarine deposits, river gravels, dunes
and Polynesian middens at more than 30 localities in South Island. Most sites are in lowland areas, in low
hills, or near the coast, suggesting that Haast’s Eagle was not a bird of high ground, with eastern South
Island being the centre of distribution. The wings were short and broad with a long tail, typical of many
forest eagles. It was a ‘perch and strike’ predator, flying at high speed from a tall vantage point before
crushing its prey with powerful talons. Haast’s Eagle was able to evolve immense size and strength due to
abundant large prey and the absence of competing mammals. It had exceptionally strong, stout lower legs
and feet, with incredibly long claws. The largest claw known was over 60mm long; it would have been
covered with a horny sheath that increased its length by at least another 30mm, making it overall the size of
an adult tiger’s (Tennyson & Martinson 2007). As the beak was long, Haast’s Eagle could have reached deep
inside carcasses for the vital organs, and also would have been well-suited for feeding on carrion. The
discovery of a number of moa pelvises punctured by the talons of this eagle is testament to the power and
strength of the bird. Apart from actively hunting large birds (the flightless Cnemiornis geese may also have
been important prey), it probably also took carrion, and was, until the arrival of humans, the top predator in
New Zealand.
A Maori legend of an extinct bird of prey, the Hokioi (Grey 1872), may well be referable to A. moorei, indi-
cating that it survived into relatively recent times. The explorer Charles Edward Douglas claimed that he
shot two raptors of immense size in the Landsborough River valley in the 1870s. If the story is true, it is
more likely that Douglas encountered Eyles’s Harrier Circus teauteensis (see p. 90). Haast’s Eagle is believed
to have become extinct between 800 and 400 years ago, primarily due to the extinction of large prey species,
and as a result of human-driven environmental change.
An almost complete subfossil sea-eagle referable to the genus Haliaeetus was discovered on Maui, along
with other remains on Molokai and Oahu (Olson & James 1982), but identification was difficult to deter-
mine due to its morphological similarity with the Bald Eagle H. leucocephalus and White-tailed Eagle H.
albicilla, of New and Old Worlds respectively. Molecular mtDNA techniques were used to help resolve this
issue, and it was found that the Hawaiian Eagle was more closely related to the White-tailed Eagle, but
differed insignificantly genetically (Fleischer et al. 2000); the authors concluded that the Hawaiian Eagle
represented a disjunct population of White-tailed Eagle that had colonised the Hawaiian Islands compara-
tively recently.
However, a recent, more refined mtDNA analysis concluded that the Hawaiian Eagle represents a
distinct (>3% divergent) mtDNA lineage related to White-tailed Eagle that diverged in the middle
Pleistocene, around 100,000 years ago (Hailer et al. 2015); the population may have been resident for at least
374,000 years, perhaps longer. The genetic difference is sufficient to warrant subspecific or even full species
status, despite there being no morphological distinctiveness in the subfossil remains (Hailer et al. 2015).
Once established, the Hawaiian Eagle became the largest terrestrial predator in the Hawaiian
89
Archipelago, a predator capable of taking the large, flightless moa-nalos and ibises. Fossil remains on Oahu
and Molokai were found in coastal deposits, whereas the almost complete skeleton was found at 1,463m on
the south slope of Mount Haleakala, Maui; the Hawaiian Eagle occurred from sea level up to montane
regions (Hailer et al. 2015). The eagle’s absence from the other Hawaiian Islands may well be an artefact of
an insufficient fossil record, rather than representing a true distribution within the archipelago.
The Hawaiian Eagle disappeared after the arrival of Polynesians on the island, due to deforestation and
the loss of large prey. It may have even nested on the ground in coastal regions, which would have made
eggs and chicks vulnerable to predation from introduced mammals such as pigs and rats.
The Hawaiian Harrier was smaller than any known extant harrier and exhibited Accipiter-like short, broad
wings, characteristics of a forest- inhabiting species (Olson & James 1991). Harriers are generally
long-winged, low-flying birds that inhabit open areas, which would have been much scarcer prior to defor-
estation by the Polynesians. The Hawaiian Harrier was probably a b ird-hunting specialist, and could easily
manoeuvre through thick forest to hunt, so it relied heavily on undisturbed forest and forest bird species.
Great changes were brought to the environment after the arrival of Polynesians to the Hawaiian Islands, so
the loss of habitat was probably the primary cause for its extinction.
There has been much confusion about the type material of this New Zealand harrier. Forbes (1892d)
described Circus hamiltoni (now considered to be a nomen nudum) and C. teauteensis from subfossil material
found at Te Aute swamp in North Island, but failed to cite specific specimens or describe its distinguishing
features, other than to say that they were both larger than the Swamp Harrier C. approximans. Scarlett (1953),
in his description of eylesi, using material found in Pyramid Valley, South Island, compared it with bones of
C. a. gouldi (the Australian and New Zealand race of Swamp Harrier), which were found in the same
deposits, and they proved to be consistently different. Various authors have disagreed over the status of C.
teauteensis since then. Worthy & Holdaway (2002) synonymised it with Scarlett’s eylesi, but noted that
Forbes’s material should be r e-examined to confirm this, while Tennyson & Martinson (2007) use teauteensis.
The latter name has been adopted by Gill et al. (2010).
Eyles’s Harrier was larger than the Swamp Harrier, with different proportions, so it had probably
evolved to feed on larger prey and with a different predatory strategy. Being about four times as heavy as
the Swamp Harrier, with a deeper bill and with the lower leg, feet and claws 50% larger, Eyles’s Harrier
could tackle small moa up to 40kg in weight, although New Zealand Pigeon Hemiphaga novaeseelandiae was
a favoured prey (Tennyson & Martinson 2007). Tennyson & Martinson suggest that the body proportions of
Eyles’s Harrier indicate fast-flapping flight, unique amongst harriers, making it an adept predator in a more
forested environment. It appears to have favoured dry forests and shrublands, ranging from the coast to the
subalpine zone. It disappeared for the same reasons as other extinct New Zealand raptors – prey extinction
and habitat alteration. It is possible that this bird survived into the post-European era.
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Accipitridae
Accipter hawks are generally f orest-adapted birds, with short, broad wings and long legs. New Caledonia
was once inhabited by four accipiters (Balouet & Olson 1989), of which only two survive today, the Brown
Goshawk A. fasciatus and the White-bellied Goshawk A. haplochrous. The Powerful Goshawk was the largest
species, with particularly strong and robust legs. Why this species and the Gracile Goshawk Accipiter
quartus (see below) disappeared while the other two accipiters survived is a mystery.
The Gracile Goshawk was the smallest of the four New Caledonia Accipiter hawks, with less robust leg
bones. The sympatry suggests each species occupied a different niche. Interestingly, fossils of the two extant
species, A. fasciatus and A. haplochrous, have not been found associated with the two extinct species, which
may suggest that they did not occur in the same habitat, or that the extant species may have arrived compar-
atively recently (Balouet & Olson 1989).
The Bermuda Hawk exhibited strong sexual dimorphism and had comparatively robust legs. It was almost
certainly a bird-hunting specialist, perhaps also taking bats and lizards (Olson 2008b). Although the remains
of this hawk have not been found in recent fossil beds, which may suggest a natural extinction event, an
account from 1603 by Diego Ramirez mentioned that on Bermuda ‘there are many very large dark herons,
many very handsome sparrow-hawks, so stupid that we even clubbed them’ (quoted in Olson 2008b). This
may represent an endemic hawk without fear of humans, i.e. Bermuda Hawk; all migrant birds of prey
would surely have been less easy to approach.
The discovery of a huge crowned eagle on Madagascar (Goodman 1994a), related to but larger than the
extant African Crowned Eagle Stephanoaetus coronatus, has led to much speculation about its size and
ecology. It has been suggested that reports of the Madagascar Crowned Eagle, together with discovery of
the giant eggs of Aepyornis, gave rise to the legend of the Roc, the mythical bird of Sinbad the Sailor’s tales
that could carry away elephants in its enormous talons.
The African Crowned Eagle is a forest specialist, capable of taking prey as large as an antelope weighing
up to 20kg, and also feeding on monkeys, birds and large lizards (Kemp 1994). It has short, broad wings and
incredibly powerful legs and feet, and dispatches its prey by ambush attack, crushing it with its feet. The
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Malagasy Crowned Eagle Stephanoaetus mahery catching a Ruffed Lemur Varecia variegata
Malagasy Crowned Eagle had similar hunting ability, and almost certainly preyed on the diverse lemur
fauna (Feduccia 1996) and pygmy hippopotamus of the island (Goodman 1994a, b), as well as large birds,
perhaps even the young of the extinct Giant Elephant Bird Aepyornis maximus. Studies of anti-predator
behaviour in two species of lemur, the Ring-tailed Lemur Lemur catta and Verreaux’s Sifaka Propithecus
verreauxi, when diurnal birds of prey fly overhead, may be a result of retained instinctive fears of large avian
predators, causing them to seek cover (Goodman 1994b). Goodman argues that extant raptors on
Madagascar only pose a marginal threat to lemurs (and perhaps only to juveniles), whereas large lemurs
would have been regular prey of the Malagasy Crowned Eagle (although this interpretation has been ques-
tioned by Csermely 1996). There was also pronounced reversed sexual dimorphism, where the female was
larger than the male (Goodman & Muldoon 2016); the sexes would have fed on different-sized ranges of
prey.
The Malagasy Crowned Eagle was the top predator in pristine Madagascar. Goodman & Muldoon (2016)
have shown that it occurred in at least two distinct bioclimatic zones of western Madagascar, one comprising
a mixture of closed-canopy subhumid forest, open savanna woodland and low-lying marshes, the other
more open dry forest. The Malagasy Crowned Eagle was subject to localised extinction events around 5,500
years ago due to a natural aridification of the island (Goodman & Jungers 2014; Goodman & Muldoon
2016), but the destruction of forests and hunting of prey species by the Malagasy would have played a
major role in the eagle’s complete disappearance, and it appears to have become extinct perhaps as recently
as the 16th century.
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Falconidae
The original specimens of this species were collected by Edward Palmer (Ridgway 1876b), who made the
following comments about the bird:
The ‘Calalie’ is abundant on every part of the island; and no bird could be a more persistent or more cruel enemy
of the poultry and domestic animals. It is continually on the watch, and in spite of every precaution often
snatches its prey from the very doors of the houses. The destruction of the wild goats is not so great, as these
animals are better able to protect themselves than the tame ones. No sooner is one kid born – while the mother is
in labour with the second – than the birds pounce on it; and should the old one be able to interfere, she is also
assaulted. No kid is safe from their attacks. Should a number be together, the birds unite their forces, and, with
great noise and flapping of their wings, generally manage to separate the weakest one and dispatch it. They
sometimes fasten upon the tongue when the poor creature opens its mouth to bleat, and have been known to tear
it out, leaving the animal to perish, if not otherwise destroyed. Sometimes the anus is the point of attack. The
birds are cruel in the extreme, and the torture sometimes inflicted upon the defenceless animals is painful to
witness . . . Even when food is plenty, they often attack living animals instead of contenting themselves with the
carcasses of those already dead, seeming to delight in killing. Should one of their number be disabled or wounded,
it is instantly dispatched by the rest.
Due to its depredations, it was systematically exterminated by the goat herders on the island, who poisoned
and shot it at every opportunity. It may have been tamer and therefore more vulnerable than mainland cara-
caras, which becomes shy when persecuted. Bryant (1887) noted that birds were shot when they came to the
few available water holes, and that during his stay they had become scarce; he only saw four birds in two
days on the island. When Palmer returned to Gaudalupe in 1889, he saw none. It was Rollo H. Beck, a field
collector, who unwittingly observed the last caracaras. In a letter sent to Robert Ridgway (1876b), Beck
described his actions, which were the last encounter with the Guadalupe Caracara:
Although I had no idea of it at the time it seems probable to me that I secured the last of the Guadalupe Caracaras
on Guadalupe Island on the afternoon of December 1, 1900. Of 11 birds that flew toward me 9 were secured. The
other two were shot at but got away. The 11 birds were all that were seen, but judging by their tameness and the
short time that I was on the island I assumed at the time that they must be abundant. All of the skins but one
went with my Galapagos material to Lord Rothschild in England. One skin which I kept for a while went finally
to the Thayer Museum.
Ironically, soon after the last Guadalupe Caracara was seen, the island was abandoned by humans. At least
six visits have been made by naturalists since 1900 (Greenway 1967); the bird has not been seen again.
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Habits
The nest was described by Swann (1925) as a ‘large affair of sticks on top of a pile of rubbish and cacti’, but
the birds were also reported to nest on cliffs. The eggs were whitish, blotched with reddish-brown.
According to a local informant of Palmer (in Ridgway 1876b), the clutch size was said to be three, and were
laid by 17 April. Palmer wrote:
Hundreds of birds have been destroyed by the inhabitants, both with poison and fire-arms, without noticeable
diminution of their numbers. They are said to lay three eggs, speckled like a gull’s. When surprised or wounded,
they emit a loud, harsh scream, something like that of the Bald Eagle. In fighting among themselves, they make a
curious gabbling noise; and under any special excitement the same sounds are given forth, with an odd motion of
the head, the neck being first stretched out to its full and then bent backward till the head almost rests upon the
back . . . They eat small birds, mice, shell-fish, worms, and insects. To procure the latter, they resort to plowed
fields, where they scratch the ground almost like domestic fowls.
They were known to feed on the carcasses of goats and of petrels, plus grasshoppers, beetles and other
insects.
Caracaras were an important component of the original West Indian avifauna. They diversified into a
number of distinct species, with some being large and almost flightless, while others differed little from
extant members of the group. Examination of the few available fragments shows that the Puerto Rican
Caracara was intermediate in size between the Crested Caracara Polyborus cheriway and the Southern
Caracara P. plancus. However, Olson & Hillgartner (1982) suggest that these cannot be differentiated from P.
cheriway; both Puerto Rican and Bahamas Caracaras (see below) might well be referable to Crested Caracara.
Crested Caracara occurs in Cuba, as well as the southern United States, Tobago and northern South
America. It would not be surprising if this species populated other West Indian islands. Fossil remains are
fragmentary, so how these populations differed is very difficult to say.
All birds of prey are persecuted by humans, and island species are particularly vulnerable. Perhaps all of
the larger West Indian islands once harboured a population of now-vanished caracaras.
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Falconidae
The Bahamas Caracara was smaller than both Crested Caracara Polyborus cheriway and the Southern
Caracara P. plancus, but with a more robust bill, comparatively reduced wings and more robust legs (Olson
& Hillgartner 1982; Suárez & Olson 2001b) (but see Puerto Rican Caracara above for taxonomic status).
Caracaras in general are opportunistic, terrestrial scavengers that occur in open country, particularly
savannas and prairies, which gives a good indication of the original habitat composition of the Bahamas.
The Bahamas Caracara may have survived into the Holocene and, as a consequence, would have been
affected by severe anthropogenic changes.
The most recently discovered of the diverse West Indian caracaras is the Jamaican Caracara (Olson 2008a). It
was a very large bird with much reduced wings, and was probably flightless, or almost so. It occurred in the
more arid areas of southern Jamaica, which would have provided an open habitat, more suited to a terres-
trial mode of life. It probably had habits similar to the modern Secretary Bird Sagittarius serpentarius, preying
on reptiles, small mammals and large insects by running after them. It is likely that introduced predators
and habitat modification by European colonists exterminated the Jamaican Caracara before it could be
recorded alive by naturalists.
Dubois (1674) was the only person to mention the Réunion Kestrel. He made the following comment:
The second ones [Réunion Kestrel] are named yellow-feet, with the size and shape of falcons. They do harm to
the fowls of the inhabitants and the game of the island.
This kestrel differed from the extant Mauritius Kestrel Falco punctatus in being larger, and without the short,
rounded wings typical of insular forest falcons, so it may have derived from a different ancestor
(Mourer-Chauviré et al. 1999). As it was renowned for taking the inhabitants’ domestic fowl, it would have
been persecuted accordingly, and disappeared as a result.
The Cuban Kestrel was a specialised small falcon. Compared with the American Kestrel Falco sparverius,
which has a distinct subspecies sparverioides on Cuba, and the migrant Merlin F. columbarius, it was interme-
diate in size, yet it had substantially longer legs, perhaps the longest of any Falco (Suárez & Olson 2001a).
This adaptation may have been a result of feeding strategy; Olson & Suárez hypothesise that the Cuban
Kestrel was a terrestrial hunter, chasing after its prey. It probably also nested on or near the ground, a poten-
tial hazard when faced with introduced predators and human-induced environmental changes. It perhaps
became extinct after settlement by Europeans at the beginning of the 17th century.
Cuban Kestrel
Falco kurochkini
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Otididae
Otididae (Bustards)
The bustards are an Old World family of generally large terrestrial birds, most often associated with dry,
open scrublands and steppes. The family is divided into 10 genera. Many species are regularly hunted;
coupled with habitat destruction, some species are at serious risk of extinction.
The Arabian Bustard Ardeotis arabs is distributed across North Africa, and divided into four subspecies:
nominate A. a. arabs from the Horn of Africa, Saudi Arabia and Yemen; A. a. butleri from southern Sudan; A.
a. stieberi from Mauritania and n
orth-east Sudan; and A. a. lynesi from western Morocco (Collar 1996). All
populations are declining due to habitat destruction and hunting. The population from western Morocco
was described in 1930 (Bannerman 1930b), but it declined over the next 60 years until it was last observed at
Lac Merzouga and Lac Tamezguidat between 1987 and 1993 (Fishpool & Evans 2001). It is now probably
extinct.
Habits
Little was recorded about the Moroccan Bustard, but the bird preferred s emi-desert, open grassy plains, and
open Acacia savanna. The breeding season was from April to June. Diet comprised a variety of large insects,
small vertebrates, seeds, and succulent parts of leaves.
Rhynochetidae (Kagus)
The Kagu is the only extant member of the family Rhynochetidae, and its relation-
ships are still unclear. The monospecific Sun Bittern Eurypyga helias of Central and
South America is considered to be a close relative (Fain & Houde 2004; Hackett et al.
2008), and perhaps the extinct adzebills of New Zealand (see p. 98), suggesting a
distinct Gondwanan lineage of birds. The Kagu is a terrestrial bird, bluish-grey in
colour with a large, erectile crest and red bill and legs. It is endemic to the mountain
forests of New Caledonia, where it feeds on invertebrates. The Kagu is highly endan-
gered due to predation from introduced mammals and deforestation.
The Kagu genus and family was long thought to be monospecific, but the discovery
of subfossil remains in 1980 showed that another species once occurred on New
Caledonia (Balouet & Olson 1989). This extinct species, the Lowland Kagu
Rhynochetos orarius, was larger than R. jubatus. Balouet & Olson suggest that as the Lowland Kagu
extinct lowland form has only been found in areas at or near the coast, the two Rhynochetos orarius
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species may not have been sympatric. With our present level of understanding, it is impossible to determine
if the two species were mutually exclusive or if there was a zone of overlap. The lowland areas of New
Caledonia would have suffered immediate degradation once human colonists arrived, and together with
over-hunting and predation pressure from introduced predators, the Lowland Kagu probably disappeared
soon after human contact. Conversely, the large size of the island and associated highland refugia allowed
R. jubatus to survive.
Aptornithidae (Adzebills)
The adzebills were an extraordinary family of birds whose affinities are ancient and remain obscure. They
were originally considered to be small moas, and later aberrant members of the Rallidae. Olson (1977)
believed that Aptornis was not a rail, but was more closely related to the New Caledonian Kagu Rhynochetus
jubatus, but this may be due to convergence. Weber & Hesse (1995), based on cranial osteology, suggested
that adzebills were a sister group to the Galloanserae, the gamebirds and waterfowl, but again this may be
due to convergence (Worthy & Holdaway 2002). Their affinities still remain unresolved.
The North Island Adzebill was first described from a tibiotarsus, collected in 1842 at Poverty Bay, North
Island, by Rev. Williams (Owen 1843). It was smaller than the South Island species, A. defossor (see below).
The discovery of more complete material showed that the North Island bird was an extremely large, flight-
less species, the size of a small moa at 70–100cm in height, with massive robust legs. The wings were
disproportionately small with a uniquely reduced carpometacarpus (Livezey 1994). The skull was extremely
large and thick-walled, the bill deep and robust, downward-curving with a pointed tip, and had thick
cutting edges. The neck was immensely strong and muscular. Stable isotope analysis (Worthy & Holdaway
2002) has shown that these birds
were almost certainly carnivorous.
They may have torn into decaying
wood for grubs and other inverte-
brates, or preyed on burrowing
birds and reptiles by digging them
out of their tunnels. Equally they
may have been opportunistic omni-
vores, using their strong jaws and
feet for digging for roots, tubers and
invertebrates.
Subfossil remains were recovered
from Holocene deposits in the more
arid lowland regions of North
Island, which suggests that they
were restricted to these areas. The
birds may have preferred shrub-
lands and grasslands rather than
subalpine shrubland or dense
forests (Worthy & Holdaway 2002).
Subfossil remains have been found
North Island Adzebill Aptornis otidiformis with a petrel chick
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Rallidae
in Maori middens, so the birds were certainly hunted as food. Being flightless and entirely terrestrial they
must have been easily caught.
Because of their restricted range and lowland distribution, Worthy & Holdaway (2002) suggest that the
adzebills disappeared soon after Maori occupation of the islands, perhaps 1,000 years ago, and prior to the
extinction of the moas. This may have been due to their extreme vulnerability, especially when o
ver-hunting
was combined with the introduction of dogs and the Pacific Rat Rattus exulans, which were probable preda-
tors of eggs and chicks.
The South Island Adzebill was discovered in a cave near Oamaru in 1863 by the Rev. Richard Taylor, and
other remains have been found in several localities in South Island. An almost complete skeleton in London
was collected at Oreti River, Southland. This species was larger than the North Island form; otherwise it was
morphologically similar. Although the external appearance of the adzebill is not known, a reconstruction by
Jeanette Winn in Worthy & Holdaway (2002) shows the bird to be extremely robust with a massive head,
neck and legs, but with the wings completely hidden by the plumage, as in the kiwis. Worthy & Holdaway
predict the body mass of the bird to have been around 18kg (ranging from 10–25kg), the size of a small moa.
It was probably a s low-moving, cumbersome species, whose only natural enemy as an adult was the giant
Haast’s Eagle Aquila moorei (see p. 88).
Adzebills appear to have been less widespread than the moas, so that the effects of hunting pressure and
predation would be far greater. As in the North Island species, the arrival of Polynesian colonisers and their
commensal mammals would have extirpated the population rapidly.
The Yellow Rail C. n. noveboracensis occurs generally in the northern United States and Canada with disjunct
populations in Florida and east California (Taylor & van Perlo 1998). Goldman’s Yellow Rail was restricted
to the Rio Lerma marshes at Lerma and San Pedro Techuchulco, central Mexico. It was known only from the
type collected July 11, 1904 until rediscovered by Dickerman (1971) in 1961. A few individuals were
collected, the last on July 22, 1964. The Rio Lerma marshes have now been almost completely ditched and
drained for agriculture, and the bird has not been recorded since. The race is considered to be probably
extinct (Howell & Webb 1995), but there have not been any serious ornithological surveys to ascertain its
continued presence, so there is a small possibility that it might still survive.
Habits
It was recorded in wet meadows with bunch grass, sedge and Typha marshes in undisturbed areas in vege-
tation less than 50cm tall. It occurred up to an altitude of 2,500m. The nominate subspecies feeds on a
diverse range of aquatic and terrestrial insects and arthropods.
Many naturalists, including members of the Whitney South Sea Expedition (an extensive series of explora-
tions mounted in the 1920s by the American Museum of Natural History, financed by Harry Payne
Whitney), searched for this rail without success. The last known specimen was collected some time prior to
1890 (Reichenow 1891a; Mlíkovský 2012), but there were reports of the bird from Taveuni in 1971 and from
Waisa, Vitu Levu in 1973 (Blackburn 1971; Holyoak 1979), although these remain unconfirmed. Introduced
cats and mongooses are thought mainly responsible for its extinction, although other ground-living birds
have apparently survived.
The closest relative of the Fiji Bar-winged Rail is found in the Solomon Islands (Peters 1932), and both
species may be relicts of a larger, more widespread group (Greenway 1967).
Habits
The only observations of its habits seem to have been made by Layard (1875), who stated:
Generally distributed; inhabits the thick Taro beds and swamps. It was very shy, and rarely seen, but is, I believe,
easily trapped by the natives, who call it “Mbidi.” It lays six eggs (in a nest made of sedges), of a
raspberry-and-cream-coloured ground, speckled chiefly at the obtuse end with light purplish and dark dry
blood-coloured spots . . . [3.8cm long x 2.8cm at the widest point] . . . They nest in November and December,
and, I think about March.
Mayr (1945) believed that the Fiji Bar-winged Rail was the mysterious ‘sasa’, a wingless bird known to the
Fijians, which bred in holes the ground in the mountains.
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Rallidae
Wake, Wilkes and Peale Islands form a low, scrub-covered atoll in the North Pacific. The Wake Island Rail
was discovered in 1892, but not described until 1903, when Rothschild (1903) received the first material. It
was flightless with robust legs, and a probable derivative of the volant Hypotaenidia philippensis group
(Taylor & van Perlo 1998; but see below). Reports before 1942 state that the bird was common, and it could
be seen in areas with Pandanus/Cordia scrub and low Sesuvium trees (Ripley 1977). Polynesians had intro-
duced the Pacific Rat Rattus exulans before the arrival of Europeans, so the rails were clearly able to live
alongside these rodents.
Extinction
In 1935, an airport and hotel were built on Peale Island, as one of a number of stopover points for long-
haul flights from San Francisco to Hong Kong (Krupnick 1997). The island was fortified in 1940–41
because of the threat of war with Japan, and it was during the period 1935–41 that hotel employees,
military personnel and tourists regularly encountered the rails and made notes or took photographs
(Olson & Rauzen 2011). The Japanese launched an air assault against Wake Island on 8 December 1941 as
part of a wave of coordinated attacks immediately following Pearl Harbor, and they were in control of
Wake by 23 December (Cressman 1992; Olson & Rauzen 2011). From this time onwards, Japanese troops
occupied the island under almost continuous siege, until their surrender on 7 September 1945. They
were exposed to horrific hardships, including continual heavy bombardment and starvation. Of around
4,400 troops stationed on Wake in 1941, only 1,242 were alive by the time Japan surrendered in 1945
(Cohen 1983). It is no surprise, therefore, that any likely food source would have been consumed. The
Wake Island Rail had evidently gone by the end of Japanese occupation. Bailey (in Greenway 1967)
reported the bird extinct on Wake, but thought it might survive on Wilkes and Peale, which was unlikely.
Peale is separated from Wake by a 100m channel, which appears to have restricted the rail to Wake and
Wilkes (which are much closer together).
Rails of the genera Hypotaenidia and Gallirallus were part of a widespread radiation that probably incor-
porated almost all habitable islands across the Pacific Ocean (see Steadman 2006a). Steadman estimated the
number of rails that might have once occurred in the Pacific, and suggested somewhere between a highly
conservative 442 to nearly 1,600; this may still not reflect the true diversity of the region’s rails before the
arrival of humans. The volant Buff-banded Rail Hypotaenidia philippensis is probably the founding species
for many Pacific island rallids, including the Wake Island Rail (Taylor & van Perlo 1998; Olson & Rauzen
2011), but it is possible that this may not have been the case. Kirchman (2009) suggested that H. philippensis
could have only recently radiated from the Philippines, and that its expansion may have been facilitated by
human habitat disturbance, especially in the clearing of lowland forests to promote agriculture and also the
elimination of flightless rallid competitors.
Habits
The Wake Island Rail was omnivorous, feeding on seeds, molluscs, insects, caterpillars and worms,
exposing them in loose soil with sideward thrusts of its bill (Wetmore, in Olson 1996b). It also took the soft
parts of hermit crabs and scavenged seabird corpses, even attacking sickly birds (Wetmore in Olson &
Rauzen 2011). Breeding was sporadic and influenced by the season. The nest was constructed on the ground
in saucer-like depressions under dense cover, with incubating birds covering themselves in loose vegeta-
tion. The rail had a peculiar breeding system, with up to three closely associated nests being tended by more
than two birds per nest, with young cared for by groups of adults until well after hatching (Olson & Rauzen
2011). The bird made a low chattering sound, and at close quarters a gentle cluck (Wetmore in Olson 1996b).
It was bold and inquisitive, but would scamper into cover when startled. Vaughn (1945; see Olson & Rauzen
2011) wrote the following account on the rail in April 1938:
They travel around with, and even eat from the same dish with, the small vegetation-eating rats that infest the
island. Long unacquainted with human beings and therefore unafraid, they have become quite friendly with our
101
men during the three years the Pacific Airways have occupied the island. They stand by dozens on the steps of
the hotel kitchen door, peering thru the screen at the staff and going crazy with delight when one of the Chinamore
kitchen-boys [natives of Guam] comes out with scraps for them. They walk over his shoes and jump high in the
air, just like young chickens at feeding time. During the heat of the day, they get under the hotel or go down into
the rat burrows to keep cool; at night, they go foraging abroad with the rats.
The remote and inhospitable subantarctic island of Macquarie was once home to a rail and a parakeet (see
p. 194). The Macquarie Island Rail was first discovered by sealers, who collected a female in March 1879;
this specimen was taken to Otago Museum. It was said at the time to be common on the south side of the
island, occupying tussock grassland (Hamilton 1894b), but very difficult to catch. Two others, possibly
collected at the same time, were later acquired by the Natural History Museum, London. These three appear
to be the only known specimens. Scott (1872) visited the island in 1870 and found the rails ‘not uncommon’.
He described two rails, a larger reddish one, and a smaller black one; the black individuals were probably
young birds. No one has seen the rails since 1879 (Vestjens 1963), and they had certainly gone by 1894
(Hamilton 1894b). It appears that introduced cats, rats and New Zealand Weka Gallirallus australis (Brothers
& Skira 1983), of which the latter two preyed on eggs and chicks, were responsible for the extinction of the
Macquarie Rail, hastened by the introduction of rabbits that reduced their habitat by o ver-grazing the
tussock grass. The rails had previously survived alongside cats for nearly a century, but cat numbers
increased after rabbit introduction, predating rails during the winter months when rabbits became scarce
(Garnett & Crowley 2000).
The type of this species was collected in 1840. Ernst Dieffenbach (who first discovered it) stated that it
was:
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. . . formerly very common, but since cats and dogs have been introduced it has become very scarce. The natives
call the bird Meriki, and catch it with nooses. I often heard its shrill voice in the bush, and after much trouble
obtained a living specimen.
Buller (1887–88) quoted a native correspondent named Roiri on the Chathams:
This bird, the Moeriki, disappeared in the third year after the occupation of the island by the Maoris. If the bird
still survives I will catch you some. It was a beautiful bird. I remember seeing it when I was a boy. The Maoris
called it a Popotai.
However, Roiri never found one for Buller, and it has not been seen since 1840 (Hutton 1872).
Dieffenbach’s Rail was entirely terrestrial and restricted in distributional range, so disappeared rapidly
after the arrival of Europeans. Bush fires, cats and rats were almost certainly responsible for its demise
(Greenway 1967).
Habits
Virtually nothing was recorded about the habits of Dieffenbach’s Rail. Only two accounts provide any
detail. An account from Pitt Island in the late 1830s may be referable to it (in Tennyson & Martinson
2007):
When gathering food. . . the subject that gave its sovereign least trouble was the dark-brown water-hen, of the
size of a barnyard fowl, which inhabited the skirts of the woods, and fed on the beach. It was unable to fly, and
made no attempt to escape when approached.
A second was recorded by White (1896):
The mehoriki was a very tame bird, but was only caught at certain seasons, being strictly preserved at others.
The eggs were never eaten if in the least turned (i.e., sat upon); children were always reproved for so doing. The
birds were caught by preparing large traps with wide wings to them, between which they were quickly driven.
These birds lived in and preferred the undergrowth of the bush, which afforded them concealment. (Rather diffi-
cult to round them up in such a place.) The flesh was said to be very delicate, and much relished by sick persons.
The mehoriki was a very watchful bird. No stranger could approach without it uttering its warning cry. (This is
a singular habit for a very tame bird.) In colour it was light-straw-coloured, and spotted like the New Zealand
bittern, but not so dull a grey as the latter. The eggs were spotted, and about the size of a medium or small hen’s
egg.
Description probably 23cm (9in). Generally black above with white spots or bars; abdomen, throat and eyebrow white;
hind neck ferruginous; breast grey; upper bill blood-red, lower bill b
lood-red tipped pale fuscous; iris red; legs and feet
flesh-coloured.
Although known only from Tahiti and hearsay reports from Mehetia, it is possible that the Tahiti R
ed-billed
Rail also occurred on a number of outlying islands (Greenway 1967). Johann Reinhold Forster, who was
naturalist on Cook’s second Pacific voyage, indicated that the bird was called ‘Oomnaa’ or ‘Eboonàa’ in
Tahiti and its neighbouring islands, and ‘Oomèia-Keteòw’ in Tonga-Tabu; yet he gave only Tahiti and the
neighbouring islands in the species’ range. It was abundant when first discovered on Tahiti around 1844 but
had disappeared by the end of the century, perhaps surviving on Mehetia, which is free of cats, until the
1930s (Taylor & van Perlo 1998). It may have been exterminated by cats and rats, but no more information is
available.
Forster’s plate
The Tahiti Red-billed Rail is known from Georg Forster’s plate no. 128, executed on Cook’s second Pacific
voyage. Georg accompanied his father J. R. Forster on the voyage, and was given the task of draughtsman
to record the natural history. The plate is inscribed “Rallus pacificus. Taheitee. Oomnaoe. Oomeea keto õw”.
It is life-sized. Much confusion has arisen from this illustration, as plates 127 and 128, which depict two
quite different rails, are both captioned ‘Rallus pacificus’. This led to pacificus being synonymised under
Rallus ecaudatus, now known as the Tongan Buff-banded Rail, Hypotaenidia philippensis ecaudatus (this
subspecies sometimes occurs in lists of extinct birds, but it is actually extant). The matter was clarified by
Lysaght (1953); plate 128 of Forster depicts Hypotaenidia pacifica.
In Rothschild (1907a), there is a figure of the Tahiti R ed-billed Rail, copied from the Georg Forster’s
unpublished painting by John Gerrard Keulemans, who was perhaps the greatest bird artist of them all. The
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bird is faithfully reproduced, except the legs are coloured red instead of being flesh-coloured (see Fuller
2000).
From June 9–July 11 1777, during Captain Cook’s third and final voyage to the Pacific, the two expedition
ships, HMS Resolution and HMS Discovery, anchored off Tongatapu in the southern Tonga Islands. William
Anderson, naturalist and ship’s surgeon on the voyage, noted that a number of Tongan birds had been
procured from the locals, which included pigeons, small rails and gallinules (Medway 2010). Some of these
specimens were purchased by Joseph Banks (Medway 1979), including a skin of the Tongatapu Rail
preserved in spirit (Medway 2009). John Latham (1785) had access to Bank’s specimens, and he described
the rail as:
Philippine Rail Var. B.
The head in this variety is paler, and the streak over the eye grey: the hind part of the neck transversely striated brown
and white: the middle of the back, and scapulars, white, with a very little mixture of brown on the first: wing coverts
olive brown, transversely blotched with white; second quills white on the inner webs, on the outer olive brown; the
greater quills olive brown, marked with large ferruginous spots; the first wholly white, the second white within: tail
even with the end of the quills, barred olive brown and white: all the under parts white: bill and legs pale yellow brown.
Inhabits Tongataboo. In the collection of Sir Joseph Banks.
Despite having distinct white underparts, as seen in the closely related Hypotaenidia pacificus (see above),
it was relegated to the synonymy of Hypotaenidia philippensis (Sharp 1894), and thus ignored by subsequent
authors (see Medway 2010). However, Medway considered the evidence and concluded that the Tongatapu
Rail was indeed a valid species, and almost certainly restricted to that island. It is not certain if the rail was
flightless, but like so many island rails, it was probably at best only poorly volant. Cook recorded in 1777
that the island was heavily populated and cultivated, and it had been for millennia (Steadman 2006), so it is
somewhat surprising that the Tongatapu Rail was still extant in the 1770s. The bird had survived hunting
by the Tongans, habitat destruction, and predation by introduced mammals, especially rats, for centuries.
However, Cook had introduced dogs from Fiji in 1773 on a previous visit to the island. The dogs were feral
in 1777, had multiplied to large numbers by 1793, and are known predators of rails (see Medway 2010). The
Togatapu Rail was not recorded again after its discovery in 1777, so presumably these feral dogs had hunted
them to extinction by the end of the 18th century (Medway 2010).
Dickinson & Remsen (2013) and del Hoyo & Collar (2014) have placed many of the Pacific extinct and extant
Gallirallus rails into the genus Hypotaenidia based on their distinct characters. The Pacific rails described
from subfossil remains and mentioned here were also probably members of Hypotaenidia, and almost
certainly derived from Buff-banded Rail Hypotaenidia philippensis; therefore I refer them all to Hypotaenidia,
until further evidence suggests otherwise.
The Rota Rail was a medium-sized species, overall smaller than the Buff-banded Rail Hypotaenidia philip-
pensis and Barred Rail H. torquatus, from which many of the Pacific island populations may have been
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derived (Kirchman & Steadman 2006a; but see Wake Island Rail). It was flightless, but the legs did not
exhibit the usual robustness, a character normally associated with flightlessness.
The Aguiguan Rail was a small to medium-sized species, smaller than the Rota Rail (above), but similar
to the Buff-banded Rail Hypotaenidia philippensis. The preserved sternum is fragmentary, and the leg
elements do not differ in size from the Buff-banded Rail, so its flying capabilities cannot be ascertained
(Kirchman & Steadman 2006a). Nearly all of the subfossil remains show charring from archaeological
fires, so this species probably constituted an important food source for humans when they first arrived
on the island.
This was a large rail, equal in size to the New Britain Rail Hypotaenidia insignis and Solomon Rail H. wood-
fordi (Kirchman & Steadman 2006a), with a total length of approximately 33cm. Subfossil remains are
fragmentary, but the bird appears to have been flightless. Along with H. insignis, the New Ireland Rail is
only the second known from the Bismarck Islands, but future palaeontological work will doubtless reveal
more species.
The following rail species from the Marquesas Islands, along with an undescribed species from Hiva Oa,
represent the only species known from eastern Polynesia; all were flightless, and all are now extinct. The
Tahuata Rail was a m
edium-sized, probably flightless species, equal in size to B
uff-banded Rail Hypotaenidia
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philippensis. The legs were robust, particularly in the tarsometatarsus (Kirchman & Steadman 2007). This
suggests that the Tahuata Rail was totally terrestrial, adapted to the forest floor of its island home, and
extremely vulnerable to extinction after the arrival of humans and their commensal animals.
This was a small to medium-sized flightless rail, equal in size to Buff-banded Rail Hypotaenidia philippensis
but with reduced wings, and with leg elements of a generally similar proportions, except that the shaft of
the tibiotarsus was extremely gracile, perhaps the thinnest relative to any rallid species, whether flightless
or volant (Kirchman & Steadman 2007). The reasons for this are unclear.
This was a small, flightless rail, with reduced wing elements and corresponding robust leg elements
(Kirchman & Steadman 2007), typical of flightless, terrestrial, island-inhabiting rails. Its remains have been
found in archaeological middens along with other vertebrate bones and shells, so it was regularly eaten by
the Polynesians. This was probably the primary reason for its disappearance.
The Huahine Rail was a medium-sized flightless rail, about the size of the volant Buff-banded Rail
Hypotaenidia philippensis, but with the wing elements greatly reduced relative to the leg elements, and with
stout legs (Kirchman & Steadman 2006b). This bird was one of only two Hypotaenidia rails known to occur in
the Society Islands (the other was the extinct Tahiti Red-billed Rail H. pacifica, which is only known from a
painting; see p. 103). Like most of the rails of eastern Polynesia, its remains were found in archaeological
kitchen middens, and it probably disappeared as a result of over-hunting and perhaps predation by intro-
duced animals.
The Niue Rail was a m edium-sized flightless rail, about the size of volant B
uff-banded Rail Hypotaenidia
philippensis, but it differed in a number of characters which suggests that it may not have been part of
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the same radiation as other Hypotaenidia rails throughout the Pacific (Steadman et al. 2000). Complete
pectoral material is lacking, but ratio measurements infer that it was probably flightless. The subfossil
remains were deposited in pre-human sediments to the island, but with the arrival of Polynesian colo-
nists, it would have succumbed quickly to over-
hunting and anthropogenic changes to the
environment.
The Mangaia Rail was a small species, about the size of Wake Island Rail Hypotaenidia wakensis (see p. 100) at
22–25cm in total length. It had a slender bill and strong, robust tarsometatarsi (Steadman 1986). It occurred
sympatrically on Mangaia with another extinct, flightless rail, the Mangaia Crake Porzana rua (see p. 120),
and it seems to have disappeared at some point after the arrival of Polynesians but before the first scientific
study in the 1970s (Holyoak 1974a). O ver-hunting, habitat alteration and introduction of mammalian pred-
ators appear to be the causes of its extinction.
This was a medium-sized, stocky rail which differed from its congeners in characters of the bill and femur,
and was more robust in the skull and legs compared with B uff-banded Rail Hypotaenidia philippensis
(Kirchman & Steadman 2005). The wing elements do not superficially indicate that this species was flight-
less, being approximately the same size as the B uff-banded Rail, but more robust. However, correlations
between other similar-sized but extant flightless species (e.g. Guam Rail Hypotaenidia owstoni) suggest that
the Eua Rail was flightless. It disappeared sometime after the arrival of Polynesians on Tonga 700 years ago,
but before the arrival of Europeans 200 years ago.
Tabuai is the largest of the islands that form the remote Austral Islands. The Tabuai Rail was another species
that approximated the size of volant B uff-banded Rail Hypotaenidia philippensis, with similar-sized leg
elements, although the tarsometatarsus at the distal end was broader, indicating a more terrestrial habit.
However, the wing elements relative to the leg elements were greatly reduced, so it was almost certainly a
flightless species (Worthy & Bollt 2011). This rail appears to have been part of the Hypotaenidia philippensis
radiation, but conceivably may have been from an earlier colonisation event, and Worthy & Bollt (2011)
suggest that it was perhaps a ‘sister species’ to the Mangaia and Niue rails.
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Meredith (1991), followed by Holdaway & Anderson (2001), reported the presence on Norfolk Island of a
flightless Hypotaenidia rail, closely related to but larger than B
uff-banded Rail Hypotaenidia philippensis, but
they did not describe it. Holdaway & Anderson (2001) indicate also that at least three other rallids were also
once present; Buff-banded Rail, a gallinule Porphyrio sp., and Spotless Crake Zaporria tabuensis (the latter, or
a race of it, may well be the Dark Rail Zapornia tenebrosus of Latham; see Ripley 1977 and p. 331). A rail, no.
79 in the ‘Sydney’ series, painted by John Hunter during the first European settlement of Norfolk Island in
1788 (Hindwood, 1965), has been cited as representing the Buff-banded Rail, but it differs from it in some
degrees, and therefore could equally be referable to the extinct Hypotaenidia species.
It appears that the rails were victims of o ver-hunting, both by Polynesian and European settlers, and
terrestrial rails would have been extremely vulnerable to predation by the introduced Pacific Rat Rattus
exulans. The sometimes horrific fate that was to befall its close congener, the critically endangered, flightless
Lord Howe Wood Rail Hypotaenidia sylvestris, and probably many other island rails, is summed up in the
account of Thomas Gilbert, Commander of the ship Charlotte, who stayed on Lord Howe Island in 1780 (see
Hindwood 1940):
Partridges [Lord Howe Woodhen] likewise in great plenty ran along the ground, very fat and exceedingly
well tasted. Several of these I knocked down, and their legs being broken, I placed them near me as I sat under a
tree. The pain they suffered caused them to make a doleful cry, which brought five or six dozen of the same kind to
them, and by that means I was able to take the nearly whole of them.
New Caledonian Wood Rail Gallirallus lafresnayanus Verreaux & Des Murs
Gallirallus lafresnayanus Verreaux & Des Murs 1860, p. 437 (New Caledonia)
Tricholimnas lafresnayanus Collar & Andrew 1988, P. 46; Hume & Walters 2012, p. 96
Specimens Specimens are in New York and Tring.
Status Extinct. The last specimens were seen around the end of the 19th century.
Range New Caledonia, Melanesia.
Description 46cm (18in). Predominantly rufous brown with slight tinge of olivaceous on lower part of back; top of the head
dull slate, supercilium and chin pale rufous grey; underparts dull slate; neck and upper breast tinged with rufous; wings with a
blunt spur, and sharp spine on shoulder; primaries black; underwing-coverts and axillaries black, transversely barred with
white; iris dark crimson; bill dark horn; legs
and feet brown.
New Caledonian Wood Rail
The few known specimens of New Gallirallus lafresnayanus
Caledonia Wood Rail were taken on
the east and west coasts (Fullagar et al.
1982), but the bird may also have
occurred in the hills up to 915m.
Greenway (1967) noted the steep and
rugged terrain of New Caledonia, and
believed it possible that the species
could still survive in remote parts
where dogs, cats and rats could not
reach. It probably occurred in the
forests of the interior, and in forested
river valleys near the coast, but it was
not a marsh bird (Layard & Layard
1882), a legend that Greenway (1967)
believed originated with the hydrog-
rapher Monsieur Bouquet de la Gry.
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There have been a number of reports that the rail still persists. Possible sightings are from the headwa-
ters of Rivière Blanche in the south of New Caledonia in the 1960s, and Moutn Panié in the north in the
1960s and in 1984 (Stokes 1979; Balouet 1986; Ekstrom et al. 2000). None have been confirmed, however.
A living specimen was exhibited by Sclater (1869b) at the Zoological Society in London. The bird had
been presented by Dr. George Bennett of Sydney. It had been sent to Bennett from New Caledonia, and
shipped back by him on the Parametta. It was probably the only living specimen to reach Europe. Its skin
was apparently not preserved. Extraordinarily, the bird laid an egg in 1871. This unique specimen is kept at
Tring.
Habits
Layard & Layard (1882) provided the only comments on the New Caledonia Wood Rail’s habits. It inhab-
ited the same type of woodland as the Kagu Rhynochetos jubatus, which occurs in evergreen forest (Ekstrom
et al. 2002). Although they never saw the wood rail in the wild, they kept at least one bird in captivity and
commented:
We have kept it in confinement, feeding it on Bulimi, raw meat, and garbage. It is nocturnal, and runs with
great rapidity. In walking it elevates the tail with the peculiar flip common to the Rails, and it can climb and
jump like a cat. If alarmed it will squeeze itself into the smallest holes and crevices and lie ‘perdue’ and motion-
less, feigning death for a long time. We have never seen it in its native haunts; all we have received have been
brought to us, after being snared or caught with dogs.
It was also believed to feed on insects.
The knock-on effect of the extinction of a single species can spell doom for smaller organisms such as
parasites that depend on it to survive. Many birds are hosts to specific parasites, and with the extinction of
the host the parasite will disappear as well. A phtilopterid louse, Rallicola piageti was a parasite that occurred
solely on the New Caledonian Wood Rail, and may now be extinct (Rózsa & Vas 2014). Other host-specific
lice and ticks that have also disappeared include Longimenopon dominicanum (host: Guadalupe Petrel
Oceanodroma macrodactyla; see p. 73); Acutifrons caracarensis (host: Guadalupe Caracara Caracara lutosa, see p.
93); Austromenopon confine (host: Slender-billed Curlew Numenius tenuirostris, see p. 389); Coloceras hemi-
phagae and C. restinctus (host: Norfolk Island Pigeon Hemiphaga spadicea, see p. 187); Psittacobrosus bechsteini
(host: Cuban Macaw Ara tricolor, see p. 208); Philopteroides xenicus (host: New Zealand Bushwren Xenicus
longipes, see p. 250) and the aptly named Rallicola extinctus (host: Huia Heteralocha acutirostris, see p. 267)
(Mey 1990, 2005; Durden & Keirans 1996; Buckley et al. 2012; Rózsa & Vas 2014). It is possible that thousands
of invertebrate species have disappeared as a direct result of the extinction of host or symbiotic birds.
The Bogotá Rail Rallus semiplumbeus is divided into two subspecies; the nominate is found on the Ubaté-
Bogotá plateau in Cundinamarca and Boyacá in the east Andes of Colombia, where it is considered
endangered (Taylor 1996; Taylor & van Perlo 1998), and the Peruvian subspecies peruvianus. This is known
only from the type, presumably from the Peruvian highlands and now lost, but it is distinct and should
perhaps be afforded specific status (Taylor & van Perlo 1998). The Peruvian Rail has not been recorded since
the type’s collection in 1886, and is now presumed to be extinct. Nothing was recorded about its habits.
There is an unconfirmed report of this species from Ecuador, but the subspecific identity is unknown
(Fjeldså & Krabbe 1990; Collar et al. 1992).
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The recent discovery of five extinct rails from Macaronesia highlights the necessity of obtaining a palae-
ontological record before any avian biogeographical conclusions can be made. All of these extinct
Madeiran and Azorean rails were derived from Water Rail Rallus aquaticus, but differed in their smaller
size, reduced wings, and shorter, but more robust limbs (Alcover et al. 2015). These Macaronesian Rallus
species probably disappeared quickly after the arrival of humans and the introduction of invasive
species to the islands.
The flightless Madeira Rail was the largest and heaviest of the Macaronesian rails, being overall smaller
and stouter than Water Rail, but with more reduced wings – the humerus was around 24% shorter (Alcover
et al. 2015). The proportions of the robust limbs (long femur – short and stout tarsometatarsus) suggest that
the Madeira Rail was a slow-moving bird.
The Porto Santo Rail was overall smaller than the Water Rail Rallus aquaticus, but it had relatively gracile
legs (Alcover et al. 2015); the wings were longer than in Maderia Rail R. lowei, but shorter than those of
Water Rail, which is indicative of flightlessness. Like the Madeira Rail, the proportions of the leg bones
suggest that this species was probably incapable of high speeds or rapid manoeuvrability (Alcover et al.
2015), which would have made it extremely vulnerable to predation by humans and introduced mamma-
lian predators.
The São Miguel Rail was endemic to São Miguel in the Azores and was a small, flightless species with
extremely reduced wings and short, robust limbs; the bill was more curved than in Water Rail Rallus aquat-
icus (Alcover et al. 2015). Water Rail feeds on aquatic vertebrates, invertebrates and water plants (Taylor &
van Perlo 1998), but in the absence of major wet environments on São Miguel, the resident rail probably had
a different feeding ecology. The endemic flightless rails of the Chatham Islands, Hypotaenidia dieffenbachii
(see p. 102) and Cabalus modestus (see p. 112) had modified, decurved bills that were used to extract inverte-
brates from loose sand and from leaf litter (Hume & Walters 2012). The proportions of the limbs in the São
Miguel Rail indicates a slow, ponderous gait, so it is likely that this species had a similar foraging strategy to
the Chatham rails.
Unlike the other Macaronesian rails, the Pico Rail Rallus montivagorum was volant, but with reduced flying
ability. It was slightly smaller than Water Rail Rallus aquaticus, with a reduced sternum and much shortened
tarsometatarsus (Alcover et al. 2015). During a period of low sea levels between 30,000 and 14,000 years ago
the islands of Pico and Faial were connected (Rijsdijk et al. 2014), so the rail may once have occurred on both
islands (Alcover et al. 2015). Radiocarbon14 dating has shown that the Pico Rail survived into the historical
period, at least until the arrival of the Portuguese, who colonised the islands in the first half of the 15th
century (Newitt 2005; Alcover et al. 2015).
This was the smallest of all Macaronesian rails, and the wings were relatively more reduced than in other
Azorean rails, with a corresponding reduction of the keel (Alcover et al. 2015), the attachment area for the
main flight muscles. Furthermore, the extremely reduced carpometacarpus was around 49% shorter than
that of the Water Rail Rallus aquaticus, which indicates that the São Jorge Rail had weakly developed primary
flight-feathers. The bill was elongate and slightly decurved, and Alcover et al. suggest that it was used to
probe in soft soil, mosses and deep leaf litter, as hypothesised for other long-billed, flightless rails such as
New Zealand Snipe-rail Capellirallus karamu (Worthy & Holdaway 2002) (see p. 130) and the Chatham
Islands rails (Hume & Walters 2012).
At least three other probably endemic Rallus species are known from fragmentary subfossil remains
found in the central Azores, from the islands of Graciosa, Terceira and Santa Maria Rail. The Terceira Rail is
also known from an exquisite silicified three-dimensional specimen of the trunk and legs, in which the
silicification process has preserved the feathers and skin. Alcover et al. (2015) refrained from describing
these taxa until better-preserved material becomes available.
The first specimens of this rail were collected by H. H. Travers on Mangere Island (Hutton 1872, 1874). They
consisted of an adult and an immature skin, plus an individual preserved whole in spirits. Mangere had
been completely deforested by settlers at the end of the 19th century, due to cutting and burning of the
forests and the ravages of introduced goats and rabbits. Cats had been introduced to control rabbits at an
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earlier date, so the Chatham Islands Rail stood little chance of survival, the last being taken between 1895
and 1900 (Greenway 1967). Oliver (1930) believed that collecting was largely responsible for the rail’s exter-
mination, but Greenway (1967) pointed out that it was only collected as subfossil bones on Chatham Island.
One adult and a downy chick are the only recent records for Pitt Island (Fleming 1939). Buller (1905) records
Travers collecting it on Mangere, and apparently Walter Hood informed him that it might still be found
there ‘although difficult to procure, owing to its semi-nocturnal habits’. Hutton (1873) remarked:
Both the birds obtained by Mr. H. Travers were full grown, one accompanied by her young one, and the other
containing well-developed ova; they were both exactly alike in colour and dimensions, in neither of which do
they show any approach to the colour and dimensions of R. dieffenbachii.
The Chatham Islands Rail was a specialised, distinctive rail, more so than Dieffenbach’s Rail, which
outwardly appears to have been a more recent arrival. However, Trewick (1997b) suggests that C. modestus
may have been derived from a more recent colonisation, evolving into a nocturnal, almost k iwi-like niche,
thus remaining in sympatry with its more generalist congener Hypotaenidia dieffenbachii. However, Holdaway
et al. (2001) question this scenario. Olson (1973b) suggested that the smaller C. modestus disappeared from
Chatham Island and Pitt Island due to competition with the larger H. dieffenbachii, but according to the fossil
record, they were sympatric on all three islands (Tennyson & Milliner 1994; Trewick 1997a).
Habits
Travers & Travers (1873) noted that the Chatham Islands Rail was nocturnal, and according to Hawkins (see
Forbes 1893b) it nested in holes in the ground and the young, when hatched, took refuge in fallen hollow
trees. Hutton (1873) said it inhabited rocky places on Mangere, as there is no sand on Mangere where the
birds could reside, and that Travers had informed him that both parents were with a chick on 5 January,
suggesting the breeding season was in November/December. The bird fed on small invertebrates, and
Buller (1905) found the stomach contained only the hard remains of beetles. Forbes (1905) believed that it
fed principally on sandhoppers, which on this island travelled a long way into the bush.
Lewin’s Rail Lewinia pectoralis is (or was) distributed from Flores in the Lesser Sundas through New Guinea
and eastern, south-eastern and south-western Australia to Tasmania (Taylor & van Perlo 1998). Cleland’s
Rail was an isolated population known from only three localities in south-west Australia, and not seen since
1932 (Whittell 1933; Taylor 1996). In general, Lewin’s Rail is considered common, as was Cleland’s Rail
prior to 1932, but extensive drainage and burning of its wetland habitat for agriculture and settlement
resulted in the latter’s rapid disappearance (Garnett & Crowley 2000). Seaches for it in the 1980s were
unsuccessful.
Habits
Cleland’s Rail was a bird of swamps and creeks with dense fringing cover (Whittell 1933). Like other races
of the species it was carnivorous, feeding on various invertebrates, probably frogs and tadpoles, and
possibly bird’s eggs.
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The White-throated Rail Dryolimnas cuvieri is an Indian Ocean island endemic; the nominate, volant subspe-
cies occurs on Madagascar, with a semi-flightless race D. c. abbotti on Assumption Island, and the flightless
D. c. aldabranus on the island of Aldabra (Taylor & van Perlo 1998). Another s emi-flightless species once
occurred on Mauritius (see below).
Assumption is a raised limestone reef island, with an area of 11.07km2, and situated 30km south-east of
the much larger atoll of Aldabra. It rises to a maximum height of about 6m; the limestone forms cliffs on the
northern part of the east coast, and much of the surface is covered in scrubby vegetation. Until June 1908 the
island was uninhabited, after which settlement began for the purposes of mining the large deposits of
guano. Between 1926 and 1945, 161,000 tons were exported, together with an unknown quantity prior to
1926 (Stoddart et al. 1970). The island’s ecosystem suffered irreparable damage, and a number of birds
which formerly bred there no longer do so. Of these, only the native rail was endemic and Ridgway (1894)
described it from a series of specimens collected by Dr W. L. Abbott in 1892. It was still very common when
Fryer (see Stoddart et al. 1970) visited the island in September 1908, soon after the settlement had been
founded. Nicoll (1908), who visited the island in 1906, noted that:
On first entering the belt of trees and low bushes which fringes the shore, we were greeted by a chorus of squeals
and grunts, as though a litter of pigs was hidden in the cover. This remarkable noise proceeded from a number of
rails, birds much like our water-rail but rather more stoutly built, and with wine-red breasts, barred on the
flanks and belly with black and white. These rails were very tame, and walked about close to us in a perfectly
unconcerned manner. We never saw one of them fly, or even try to do so: they trusted entirely to their legs when
pursued. In several patches of bush we came across family parties of them, and although the young were mostly
full-grown and feathered, we saw several which were still covered with black down. They were found on all parts
of the island, except on the summit of the sandy hill on the windward side. While uttering its remarkable note,
this rail stands quite still and puffs out all its feathers; from what I observed I should say that the skin of the
throat is also expanded. The notes are loud – a strange mixture of squealing, grunting and booming – and
during its song the bird appears to be gradually collapsing, until at the end it is once more of normal size. I have
heard our English water-rail utter a somewhat similar noise when near its nest, but its cries are never so loud as
those of the Assumption rail. We caught two of them alive and brought them safely to England, and they are at
the time I write living in the London Zoological Gardens.
Of the two rails in captivity, Meade-Waldo (1908) made the following comment:
The two rails we caught were easily kept: they would eat almost anything at once, but did not become, nor are
they yet (two years afterwards) any more familiar than they were on their native island. They have well devel-
oped wings, but do not appear to fly, although they can do so, as I have seen the individuals in the Western
Aviary, Zoological Gardens, perched on the highest perches. This rail is peculiar to the Island of Assumption,
and although closely resembling it, is quite distinct from the rail of Aldabra Island, only some 40 miles away.
But in spite of its abundance, Nicoll (1908) feared that introduced rats might be a serious threat to the rail:
Unfortunately, rats have been imported by some means into Assumption and now are very abundant. There is
little doubt that they devour many eggs of the rail and of other birds which nest near the ground, and should the
rats increase to any extent, there is a great danger of these interesting birds becoming extinct in the near future.
This proved to be the case. Some time between 1908 and the visit of V
esey-Fitzgerald (1941) in 1937, the rail
disappeared, undoubtedly as a result of a combination of habitat destruction and predation by rats. It was
probably also hunted for food by the settlers. Flightless representatives of White-throated Rail may have
once occurred on Astove and perhaps Cosmeledo (Gerlach 1997). Both populations were last recorded in
1908 and disappeared due to over-hunting.
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Habits
Other than Nicholl’s (1908) account, nothing more was recorded of its habits. The Assumption bird had
comparatively reduced wings and weak flight, with more robust legs than nominate D. c. cuvieri, so was
presumably more terrestrial.
Newton & Gadow (1893) referred subfossil remains of a rail discovered at the Mare aux Songes fossil locality
on Mauritius to the Madagascan Moorhen Gallinula chloropus pyrrhorrhoa, which is a resident species on the
island. Cowles (1987) re-examined the material and assigned it to the Madagascar White-throated Rail
Dryolimnas cuvieri cuvieri, which is a rare vagrant to the island. The type specimen of D. c. cuvieri was actu-
ally taken on Mauritius in January 1809 (Schlegel & Pollen 1868). The author examined the extensive
subfossil collections of Etienne Thirioux, who searched for subfossil remains in caves and boulder scree
around Le Pouce Mountain and the plains around the Moka Mountain Range in the centre of the island.
Among Thirioux’s specimens were more Dryolimnas bones including wing elements, which clearly shows
that a flightless derivative of Dryolimnas cuvieri once occurred on Mauritius (contra Cowles 1987; Cheke &
Hume 2008; Hume in prep.). Cheke (2013a) noted that an account written by Reyer Coenelisz during a visit
to Mauritius from April to September 1602 mentions large and small rails, and suggests that the large bird
refers to the Mauritius Red Rail Aphanapteryx bonasia (see p. 126) and the small bird to the new Dryolimnas.
However, such is the confusion with contemporary rail descriptions and subsequent misinterpretations by
more recent scholars (e.g. Frauenfeld 1868, Renshaw 1936, Hachisuka 1937a and 1953), the account is
tenuous, to say the least.
If Cheke is correct, the Mauritius White-throated Rail was extant when the Dutch first occupied the
island, but it would have quickly disappeared due to over-hunting and egg and chick predation by intro-
duced mammals.
The Réunion Rail was a large, robust bird, much larger than W hite-throated Rail Dryolimnas cuvieri, with
particularly robust legs. Wing elements are lacking, but the robust limbs suggest that it was flightless.
Dubois (1674), in 1671–72, mentions a ‘Râle des Bois’ (wood rail), which was probably referring to this
species (Mourer-Chauviré et al. 1999), indicating that it had survived at least until the 1670s; it was not
mentioned again. Like all of the other large bird species on Réunion, it disappeared due to o ver-hunting
and predation by introduced mammals, particularly cats.
St Helena is a rugged, volcanic island jutting abruptly out of the ocean, with steep sea cliffs and deeply
incised valleys. When discovered in 1502, much of it was thickly forested in the highlands, with more open
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Olson (1981) discovered the subfossil remains of a new species of rail in sand-dune deposits on Fernando de
Noronha, an island situated 345 km east of the eastern tip of Brazil. Most of the skeletal elements of several
different individuals were recovered. The rail was medium-sized, with reduction of the wings, but less so
than in other flightless species. Olson (1977b, 1981) noted that it differed morphologically from any of the
genera on mainland Brazil, and that it might be another form of Atlantisia. An account of the island was
written in 1503 during the fourth voyage of Amerigo Vespucci, the Florentine adventurer after whom the
Americas are probably named (Carleton & Olson 1999). Although the rail was not mentioned directly, a
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description in the account suggests that a flightless rail could have easily been so abundant as to have filled
a boat:
We found this island to be uninhabited, with many sources of fresh water and countless delightful trees, full of so
many sea and land birds that they were without number, and were so guileless that they let themselves be taken
in hand; and we took so many of them that we loaded a boat full of them.
The rail lived alongside the now-extinct Fernando de Noronha Giant Rat Noronhomys vespuccii, which
was equally abundant during Vespucci’s visit, but deforestation, over-hunting, and predation by intro-
duced Black Rat Rattus rattus, House Mouse Mus musculus and cats appear to have exterminated the rails
and native rat some time after 1503 (Carleton & Olson 1999).
Ascension is a barren, desolate island, which was discovered by the Portuguese in 1501. It appears that the
island was never forested (Osbeck 1771; Olson 1973a); the traveller Peter Mundy found only rushes and
grasses in 1656 (Temple & Anstey 1936), but reported goats, which had probably been released by the
Portuguese discoverers of the island. Rats were present in 1754 (La Caille 1763), which Osbeck (1771)
thought had escaped from the wreck of William Dampier’s ship, the Roebuck, in 1701, though it is probable
that they arrived earlier.
Peter Mundy
Clearly Ascension’s fauna and flora is impoverished, and it is surprising that any terrestrial bird species
could survive on such a place, but an endemic rail and a night heron (see p. 82) certainly did. Mundy landed
on Ascension on June 7, 1656
(Temple & Anstey 1936), and
made the only known descrip-
tion of the Ascension Rail from
life:
. . . much bigger than our ster-
lings [starlings] ore stares:
collour gray or dappled, white
and blacke feathers intermixed,
eies red like rubies, wings very
imperfitt, such as wherewith they
cannot raise themselves from the
ground. They were taken
running, in which they are
exceeding swift, helping them-
selves a little with their wings
. . . shortt billed, cloven footed,
thatt can neither fly nor
swymme. It was more than ordi-
nary dainety meatt, relishing like
a roasting pigge.
Subfossil remains of the rail Ascension Rail
have been found in the Mundia elpenor
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fumaroles situated in a particularly desolate lava field with old volcanic chimneys leading down into
underground caves. A considerable number of bones, mainly of seabirds with some rails, were found in
these caves (Ashmole 1963a, b), presumably of birds which had accidentally fallen in and been unable to
climb out.
The Ascension Rail was a m edium-sized, flightless rail. We know virtually nothing of its ecology. Olson
(1973a) thought it likely that it was a scavenger of seabird colonies, probably taking eggs and food dropped
by birds during the feeding of their young, and that it might have nested in small sheltered crannies in lava
flows, similarly to sheathbills Chionis sp. The reason for its extinction is not known but it seems likely to
have been due to the introduction of rats. Cats arrived in 1815; by 1823 they were abundant and had exter-
minated virtually every bird on the island. However, the rail had probably long disappeared before their
arrival.
The last known specimen of the Jamaican Wood Rail was taken in 1881. It is believed that the introduced
mongoose was largely responsible for its extinction (Bond 1936a), though it survived alongside this pred-
ator, as well as cats and rats, for generations.
Habits
Gosse (1847) gave the only detailed account about its habits:
It affects fresh water morasses, and secluded streams, rather than saline swamps, and is found even on the
mountain acclivities. I have shot it skulking among the aquatic weeds at Basin Spring. As it roams, it utters at
intervals of a few seconds, a cluck, like a hen. The remarkable thinness of body, common to all this tribe, beauti-
fully adapts them for making their way through close herbage. It is sometimes seen perched on a low tree by the
road-side, at which time it seems to have lost its usual shyness, and sits looking at the sportsmen until he nearly
comes up to it. Its flight is singularly ineffective; slow, heavy, and laboured; the head is projected, and the body
hangs down, as I have seen the body of some unwieldy Bombyx, distended with eggs: the feet are also pendent.
According to Bond (1936a), it was more of a ‘land rail’ than other West Indian species.
This crake was an inhabitant of reedlands and brush, and damp places within the original forest, but after
land clearance for the production of sugar cane, water shortage during dry periods forced the rails to
frequent household water tanks, where they were subject to predation by cats. Rats were also probably
serious predators. It was last recorded in 1925 (Momiyama 1930); although the last official sighting was in
1911 (Ingram 1911).
Habits
Momiyama (1930) gave the only account of the bird:
The Mamijiro Kuina ordinarily hides in the tako forest or in the luxuriant foliage of large bushes in the virgin
forest, but may be seen where dried grass has accumulated or where rain water is available for drinking, but in
time of drought this rainwater dries up, and as they need water, they may wander into the vicinity of water tanks
near houses. Recently both domestic and wild cats have taken to killing them. They have become pitifully scarce.
The last living form met a similar fate four or five years ago, and at present they are completely extinct.
In December 1827, Baron Friedrich Heinrich von Kittlitz called at the island of Kosrae, the easternmost of
the Caroline Islands, where he remained for about a month (Kittlitz 1858). Kosrae is a high volcanic island
rising to 656m and covered in thick impenetrable forest, which at that time of Kittlitz’s visit would have
included almost the whole island (except the very highest points of the mountains and the area around the
native villages).
While stationed on Kosrae, Kittlitz found a small blackish rail and collected the two known specimens.
By the time the next ornithological visitor landed on the island in 1880 (Finsch 1881), the rail could not be
found, although Finsch spent nine days searching for it. Coultas (see Greenway 1967), who was on the
Whitney South Seas Expedition, searched from January to June 1931 in the marshes at sea level, and also in
the mountains where it had originally been located. He reported:
In the olden days it was a sacred bird, but since the Christian missions have been established not much attention
has been paid to the older faiths . . . Several oldsters seemed to remember their forefathers speaking of this bird,
but none of them admitted having seen it, except an elderly deacon, a staunch pillar of the church, who claimed
to have had it pointed out to him twenty years previous to my visit.
With the help of a native from the Solomon Islands, Coultas set traps for the rail, but without success. It
presumably disappeared during the intervening years as a result of rat predation. It was not common even
in 1828 when Kittlitz discovered it (Kittlitz 1858), and during the next 50 years whaling ships regularly
visited the island for ship refurbishment; rats inevitably became established. They were found on the island
in great profusion in 1880 (Finsch 1881). The crake had previously been unmolested, because it was consid-
ered sacred by the islanders (Taylor 1996).
Flightlessness
There has been some debate concerning the flightlessness of this species. The wings are small and rounded,
the tail feathers soft and loose (see Hume in Fuller 2000 for an image of the skin), which led Stegmann (in
Baker 1951) to suggest it was flightless, a fact now confirmed (Steadman 1986). Ripley (1977), after examina-
tion of the specimens, concluded it was another dark endemic Zapornia, derived from Spotless Crake Z.
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tabuensis (Olson 1973b) or Henderson Crake Z. atra (Taylor & van Pedro 1998), but differing in being larger
in general proportions and with a larger, more robust bill.
Habits
This flightless rail was solitary, totally terrestrial, and occurred in the continually wet, shadowy places in
wet forests and swamps near sea level. Kittlitz described its call as ‘alluring and resounding’. Nothing more
is known about the bird.
As in Gallirallus and based on molecular phylogenetic studies, the genus Porzana has been recently split,
and the Pacific Porzana have been placed into Zapornia (Dickinson & Remsen 2013; del Hoyo & Collar 2014).
Many of the subfossil Pacific rails mentioned here were clearly derived from Zapornia crakes (e.g. Olson &
James 1991; Steadman 2006a); therefore I assign them to that genus as well.
This was a medium-sized, robust-limbed crake, larger in size than the Spotless Crake Z. tabuensis, but
with more reduced wing elements than any other Zapornia, except the extinct St Helena Crake Z. astricto-
carpus (Steadman 1986). It was most similar in size and skeletal morphology to the extant Henderson Island
Crake Z. atra, which was probably derived from Baillon’s Crake Z. pusilla (Olson 1973a); it is likely that the
Mangaia Crake had the same ancestral origins (Steadman 1986). The Mangaia Crake probably disappeared
shortly after the arrival of Polynesians, but a Zapornia species might have survived on Mangaia into the
historic period (see p. 126).
Considerable confusion existed about the number of species of Hawaiian crakes and their nomenclature
(see Olson 1999b). Two geographical subspecies are now recognised, Z. s. sandwichensis for the lighter
leeward population, and Z. s. millsi for the darker windward birds (see below). Although the last specimen
was taken in 1864, Henshaw (1902), without any corroboration, believed that one or other of the Hawaiian
Crake races might have lingered as late as 1893. Pease (1862) briefly summed up the situation in 1862:
There is a wingless bird of small size living in the island of Hawaii, which the natives call ‘Moho’, which is now
nearly extinct, having been killed off by the wild cats and dogs within late years; I have seen but a single
specimen.
Many naturalists have searched for it since, without success. The reason for its extinction is not known, but
it may have been due to rats. The mongoose, which has been blamed in the past, was, in Greenway’s (1958)
opinion, unlikely to have been the main culprit, as it was not imported to the island until 1883, by which
time the bird was already extinct, or almost so. Little is known of its habits, but it seems to have lived in
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harmony with the Pacific Rat Rattus exulans, and to have shared its burrows in times of danger (Perkins
1903). However, the imported Black Rat Rattus rattus would have been far less accommodating, and may
have been a major factor in the bird’s extinction. Hunting by Polynesians, deforestation and predation by
pigs, cats and dogs, plus introduced diseases from poultry, have also been cited as causes for its
disappearance.
Habits
This crake inhabited open grassy areas and low scrub, or clearings in rainforest. It occurred in the drier
leeward side of Hawaii, in any suitable forested areas. Subfossil remains have been found in Kiholo Bay in
the north to Manuka Bay in the south, and up to 500m above Makalawena (Olson 1999b). The Hawaiian
name of ‘Moho’ was supposedly in reference to a ‘bird that crows in the grass’.
Wilson and Evans (1890–99) provide a little more precious information about the birds’ habits:
Moreover, Hawelu told me that the mail-carrier had seen the bird cross his path within the last three years; on
the same authority, the Moho outruns any dog possessed by the natives, and it is possible to track it by its cry – a
whirring sound resembling the rising of a bevy of Quail, while its nest is made on the ground. The five specimens
were all procured by Hawelu in the scrub-covered lava-flats about five miles south of the Volcano House, but
more information than this I could not obtain, and my intelligent informant is now a leper on Molokai.
Five skins of Z. s. millsi are known, all from specimens taken on the windward side of Hawaii by a native for
the Hawaiian bird-collector and storekeeper James D. Mills (Olson 1999b). Rothschild had sent his collector
Henry Palmer to Hawaii, but Palmer failed to find the bird. Two specimens purchased by Rothschild were
found in Mills’s collection after his death (Manning 1982) and were bought for $260 by Frederic Turnill, the
Honolulu Port Surveyor, who resold them to Rothschild. Sanford Ballard Dole, a lawyer and amateur orni-
thologist, had acted as agent for Alexander Agassiz, the director of the Museum of Comparative Zoology at
Harvard University, who had wanted the specimens but had been unable to compete financially with
Rothschild. In a letter to Agassiz of August 1889, Dole reported a sighting of a crake, and hoped that they
were still extant (so that he could get a specimen!).
Habits
This crake occurred only on the windward side of Mount Kilauea, and probably ‘an area of about 40 miles
along this coast’ (Greenway 1967); also in the Ola’a area of Puna District. Rothschild (1893–1900) added:
It lived formerly in the country between Hilo and the volcano Kilauea, in places where thick grass, Vaccinum
and Dianella, forms the thickest cover possible. In former times the ‘Moho’ was a dainty on the tables of the
Hawaiian kings.
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Only two crake species are known historically from the Hawaiian Islands, the Hawaiian Crake Zapornia
sandwichensis (see above) and Laysan Crake Z. palmeri (see p. 124). The discovery of subfossil remains has
now shown that each of the main Hawaiian Islands had at least one species of crake, and up to three on the
larger islands (Olson & James 1991), and more await description. All of these island crakes disappeared due
to anthropogenic changes, a combination of deforestation, over-hunting and predation by introduced
animals. They appear to be derived from multiple invasions of Zapornia species and, as a result, were all
short-billed ‘crakes’ (Olson & James 1991).
Ziegler’s Crake was flightless and one of the smallest rails in the Hawaiian Islands, with reduced wings
and robust legs. It appears to have been quite variable in size, which may suggest sexual dimorphism. Its
wings were comparatively less reduced than its close relative Keplers’ Crake Z. keplerorum (see p. 123) on
Maui.
The tiny, flightless Molokai Crake was not only the smallest crake in the Hawaiian Islands, it was smaller
than any known extant member of the Rallidae (Olson & James 1991). A crake may have survived into
recent times on Molokai, as Perkins (1903) stated on the authority of a local resident, R. Meyer, that crakes
once lived there. Unfortunately no skin specimens were ever collected from the island, so nothing is known
about the bird’s appearance.
Molokai Crake Porzana menehune with Maui Nui Finch Telespiza ypsilon
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Keplers’ Crake appears to be descended from the same ancestral species as Molokai Z. menehune and
Ziegler’s Z. ziegleri crakes (Olson & James 1991). It was a small crake, larger than Z. menehune but smaller
than Z. ziegleri. It also differed from the latter in having a shorter, wider bill and more reduced wing
elements.
Ralphs’ Crake was sympatric with Ziegler’s Crake on Oahu, and presumably occupied a different niche. It
was flightless and one of the largest of the Hawaiian crakes. Its wings were not as reduced as the other
larger Hawaiian crake, Severns’s Crake Z. severnsi (Olson & James 1991). The subfossil remains of Ralphs’
Crake have been found only in coastal deposits, so it is possible that it was restricted to lowland habitats. If
this was the case, it would have disappeared extremely rapidly once humans colonised the island.
The flightless Severns’s Crake was sympatric with Keplers’ Crake on Maui, and along with Oahu’s Ralphs’
Crake, was the largest of the Hawaiian rallids. The pectoral girdle and wings were extremely reduced in
this bird, and the keel of the sternum was reduced to a flattened ridge (Olson & James 1991).
Easter Island or Rapa Nui is one of the remotest islands in the world; the nearest continental landmass is
Chile, situated over 3,500 km to the east. Polynesians colonised Easter Island in the first millennium and
had a thriving culture, producing the famous carved statues (moai). Trunks of the now-extinct giant palm,
Paschalococos disperta, were used to transport moai around the island (Rainbird 2002). However, over popu-
lation gradually destroyed the island’s natural resources, especially the endemic broadleaf and palm forests.
The loss of boat-building timber left the Rapa Nui people marooned and isolated from the outside world,
and left them with no ability to harvest marine life in offshore waters. The introduction of Pacific Rat Rattus
exulans, an ardent seed-predator, probably prevented regeneration of forest. By the time of European contact
in 1722, the Rapa Nui civilisation had collapsed, with the islanders numbering less than 3,000, and all of the
original forest had been destroyed (van Tilberg 1994). Diseases introduced by visiting mariners coupled
with raids by Peruvian slave-traders from 1862 decimated the remaining native population; by 1877 only
100 or so were left alive (Peiser 2005).
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During the Rapa Nui occupation, vast colonies of seabirds of at least 30 species were extirpated, and at
least five endemic land bird species became extinct. Steadman (2006) identified an extinct Zapornia crake
from archaeological remains at the coastal site of Ahu Naunau, Anakena, on the leeward coast. It was
smaller than the wide-ranging Spotless Crake Zapornia tabuenesis. The site also contained another large,
extinct rail (of which the material was too fragmentary to refer to any particular genus), two species of
extinct parrot, of which one was a very large species, and an extinct heron, more closely related to Egretta
than to other Pacific herons. The archaeological deposit has been dated, with 14C calibrated dates ranging
from 1000–1280 and 1220–1430 AD (Steadman 2006). The land birds became extinct at some point during
this period.
Laysan is a coral atoll of about 1.6 by 2.4km, with a maximum height of 7.5m. It forms part of the north-west
chain, a group of atolls and seamounts covering a distance of approximately 2,100km. Laysan was once
covered with a luxuriant growth of shrubby vegetation (Findlay 1886).
The loss of the Laysan Crake was a particularly sad case in the dismal history of anthropogenic bird
extinctions around the world, and is a classic example of the devastating short-sighted effects that humans
can have on insular oceanic island faunas.
Rabbits
In the 1890s, the crake was considered common and Laysan harboured a probable capacity of 2,000 birds. At
about the same time, a station was set up to extract guano, and the owners introduced rabbits and guinea
pigs to start a meat cannery. This was a complete failure. More disastrously, rabbits escaped, multiplied to
vast numbers, and ate the vegetation down to the ground. By 1915, the crakes were in serious decline, and
by 1923, almost extinct (Wetmore 1925b), disappearing completely just a month or so after (Hume in press).
However, in an attempt to save the bird, Laysan Crakes had been introduced to other islands. One Captain
Anderson introduced birds to Pearl and Hermes reef, east of Laysan, in 1929, but the island was swept by a
gale soon afterwards, and no birds were found in 1930 (Kaufmann in Fisher & Baldwin 1946). Crakes had
also been relocated on Lisianski, Eastern and Sand Islands (Bailey 1956), but died out on Lisianski due to the
introduction of rabbits. The populations on Eastern and Sand Islands in the Midway Atoll, however,
flourished.
The cruellest twist of fate then took place. In 1943, during US military construction on Midway, rats were
accidentally introduced and overran the islands (Fisher & Baldwin 1946): within two years the crakes were
gone. The last known specimen of the Laysan Crake was seen alive on Midway Island in June 1944 or
possibly June 1945 (Munro 1945; see Olson 1999 for a full summary). A successful eradication of rabbits on
Laysan by 1945 enabled the vegetation to recover remarkably quickly, and had crakes been returned there
they might well still be with us today.
There were attempts to translocate Laysan Crakes on several occasions. Palmer (in Rothschild 1893–
1900) sent live crakes to London in 1891, and they were also supposedly introduced to sugarcane fields on
Oahu in 1904 to aid in the control of insects, but after their release they were never seen again (Caum 1933).
A plan proposed by George Munro and Thomas M. Blackman to bring Laysan Crakes from Midway to
Oahu for breeding in the 1930s and 1940s failed due to negative politics and paperwork (Rauzon 2001; Pyle
& Pyle 2009).
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Habits
The Laysan Crake lived in thickets of Scaevola brush and coarse grass, though when common it also occurred
on the beach and near habitation. It fed on flies attracted to carcasses of seabirds. It was diurnal, active, fear-
less and inquisitive. It was said to run like a mouse. It would even walk right up to people to examine their
shoes. Sometimes it would take fright and rush away, but would be halted by the sight of an insect which it
would eat and then casually return to the scene of its fright. Bailey (1956) recalled how some crakes clam-
bered over his legs to reach the yolk of albatross eggs that he and a companion had been blowing. Bailey
was responsible for the trapping of many individuals for relocation on seemingly suitable islands.
Professor Hugo H. Schauinsland spent three months on Laysan in 1896 collecting specimens
(Schauinsland 1899), and the rails in particular made an impression:
The quaint rails were our steady companions when working. We could hardly open the door of our laboratory
without a few of these small fellows coming in with us, rummaging eagerly through our collections to enjoy the
innumerable flies which buzzed around them. It was utterly comical to observe them as they would interrupt
their hunting for a moment, in order to warble their most remarkable song, which had a certain similarity to the
clanging of the loudest sounding alarm clock. When we were skinning birds, they would even hop bravely right
up onto our preparation table, to pick a piece of suet or meat directly from our fingertips.
The nest was built on the ground among thick grass, or on tussocks. It was rather elaborately woven of
grass mixed with feathers, and had a domed top, though there seems to be doubt whether this was actually
constructed by the bird or ‘formed’ of matted grass. The nest was lined with down. It was more elaborate
than the nests of most crakes. William T. Brigham, curator of the Bernice P. Bishop Museum in Hawaii,
wrote to Agassiz on 20 June 1892 (Manning 1982) that he had kept a couple of these little crakes in his room
for a few days. He reported that they were omnivorous ‘taking seeds, figs, ants, spiders and even the flesh
of their kind’. Brigham implied surprise that they drank water readily although there was no standing
water on Laysan. They did not roost, and had a habit of standing on one leg.
Fisher (1903) described the call as a ‘plaintive, high-keyed little rattle, which resembles remotely an
alarm clock with a muffled bell or pebbles ricocheting on a glass roof’.
This flightless crake, along with the Laysan Crake Zapornia palmeri (see p. 124), was considered by Olson
(1973a) to be derived from Baillon’s Crake Z. pusilla. It had evolved for a long period in isolation, so had
characters not present in Baillon’s. Both Z. palmeri and Z. astrictocarpus had equally reduced wing elements,
while the latter had less robust pelvis and leg elements. More importantly, Z. astrictocarpus had evolved a
peculiar proximally fused carpometacarpus (wrist bone), unique among rallids, the purpose of which is
unclear. The crake was sympatric with the much larger St Helena Rail Atlantisia podarces (see p. 115), and
may have similarly utilised seabird colonies for food (Ashmole & Ashmole 2000), perhaps feeding on regur-
gitated fish remains or small insects associated with decaying bird corpses. There were probably few natural
predators, but the crake may have subject to predation from frigatebirds Fregata sp. while foraging among
the seabirds, and its eggs and young may have been eaten by the St Helena Rail (Olson 1973a; Rowland et al.
1998).
The St Helena Crake was abundant before the arrival of humans, but it no doubt succumbed to
over-hunting, deforestation and introduced animals within a few decades of human settlement.
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Description 15cm (6 in). Head, neck and undersides charcoal grey, paler on chin; back, wings and tail black; bill black; iris red;
legs and feet red.
Much confusion has arisen from the bird known as Miller’s Crake (Walters 1988). It appears in two illustra-
tions, one by Georg Forster from Tahiti, executed during Cook’s second voyage (1772–1775), the other by
Miller (1784), which was almost certainly a copy of Forster’s work (Sherborn & Iredale 1921; Lysaght 1956b).
There are also later descriptions by Latham and Gmelin.
Lysaght (1956b) believed Miller’s plate to depict the bird now known as Zapornia tabuensis, the Spotless
Crake, a widespread species occurring from Australia and New Guinea across the southern Pacific to the
Marquesas and south to New Zealand (Taylor & van Perlo 1998). However, Miller’s Crake is now known to
have occurred on Tahiti and possibly Mangaia (see below), and was perhaps related to Z. tabuensis, or to
Henderson Island Crake Z. atra.
If it was a flightless species, which appears to be the case, the introduction of predatory mammals would
have quickly eliminated the population. The Tahiti population probably became extinct soon after its
discovery in 1784.
Mangaia
Recent palaeontological work on Mangaia, the southernmost of the Cook Islands, has shown that two
populations of rails occurred sympatrically there, one related to Z. tabuensis (or indeed a race of it), and the
other the Mangaia Crake Zapornia rua (see p. 120) (Steadman 1985b). Intriguingly, Steadman questioned the
Mangaians about a mysterious small black rail they called the mo’o mo’o, which flew poorly and was rarely
seen or heard in the taro swamp in the centre of the island. This might refer to Z. tabuensis, although it is not
known on Mangaia, or it might represent a population of Miller’s Crake (or something like it) that survived
until relatively recently. Mangaia is considerably smaller than Tahiti, the largest island in eastern Polynesia;
if two small rails survived on tiny Mangaia there is no reason why two could not survive sympatrically on
Tahiti as well (Olson & Steadman 1987; Walters 1988).
Among the 17th century accounts from Mauritius are references to a bird known as Velt-hoender, Feldhüner
and Rebhüner (Strickland & Melville 1848), different from those used for the Dodo Raphus cucullatus, and
obviously referring to a different type of bird. In the account of Peter van den Broecke (1646), who visited
Mauritius in 1617, there is a crude image of a flightless bird beside a Dodo and domestic sheep, but with no
reference to it in the text. Thomas Herbert’s (1634) travels to Mauritius in 1629 depict a bird simply called a
‘hen’, and provides the first detailed account:
The hens in eating taste like parched [roast] pigs, if you see a flocke of twelve or twentie, shew them a red cloth,
and with their utmost silly fury they will altogether flie upon it, and if you strike downe one, the rest are as good
as caught, not budging an iot till they be all destroyed.
They were mentioned again by an anonymous Dutchman in 1631 (in Servaas 1887), who curiously called the
birds ‘soldiers’. Although his words cannot always be trusted, Cauche (1651) in 1638 described ‘red hens’
that could be caught with red cloth, while Mundy (in Temple 1914) in 1638 also described this phenomenon,
but refers to the bird as a ‘Mauritius Henne, of a yellowish Wheaten Coullour’. Marshall (in Khan, 1927),
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who was on Mauritius in 1668, referred to the same bird but calls it a ‘Dodo’, which has led to considerable
confusion about the true Dodo’s extinction date (Cheke & Hume 2008). Marshall’s account reads:
Here are also great plenty of Dodos or red hens which are larger a little than our English henns, have long beakes
and no, or very little Tayles. Their fethers are like down, and their wings so little that it is not able to support
their bodies; but they have long leggs and will runn very fast, and that a man shall not catch them, they will turn
so about in the trees. They are good meate when roasted, tasting something like a pig, and their skin like pig skin
when roosted [sic], being hard.
The final detailed account of live birds was penned by the preacher Johann Christian Hoffman (1680), who
was on Mauritius from 1673–75:
[there is also] a particular sort of bird known as toddaerschen which is the size of an ordinary hen. [To catch
them] you take a small stick in the right hand and wrap the left hand in a red rag, showing this to the birds,
which are generally in big flocks; these stupid animals precipitate themselves almost without hesitation on the
rag. I cannot truly say whether it is through hate or love of this colour. Once they are close enough, you can hit
them with the stick, and then have only to pick them up. Once you have taken one and are holding it in your
hand, all the others come running up as if to its aid and can be offered the same fate.
Illustrations
The Mauritius Red Rail was also accurately illustrated, with the two best images only coming to light in the
19th century. In a journal of the flagship Gelderland, which visited Mauritius in 1601, there is an ink study of
a recently killed bird by Joris Joostenszoon Laerle (Hume 2003), which is by the far the most accurate rendi-
tion, and the only one drawn on Mauritius. In 1868, Frauenfeld (1868) discovered hitherto unknown
paintings of Mascarene species, executed about 1610 and attributed to the Dutch artist George Hoefnagel. A
red rail arrived at the menagerie of Emperor Rudolph II of Austria, along with a Dodo (perhaps the first
complete specimen: Cheke & Hume 2008) and the first Mauritian Fruit Bat Pteropus niger (Cheke 2007). It is
not known if any of these animals were alive upon arrival, but they were all illustrated as stuffed individ-
uals; the red rail painting remains the only known colour depiction. A Red Rail is also supposedly illustrated
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in a painting by Roelandt Savery dated 1625 (Hachisuka 1953), but examination of the original by JPH
provides no support for this.
The discovery of subfossil remains in the Mare aux Songes in 1865 confirmed the rallid affinities of
Aphanapteryx (Milne-Edwards 1868, 1869), and further material obtained by Sauzier (Newton & Gadow
1893) showed that it was a flightless species. An almost complete, associated individual was also obtained
by Etienne Thirioux around the turn of the 20th century. The relationships of Aphanapteryx are difficult to
determine, as characters in the skeleton suggest affinities with Gallirallus, Dryolimnas, Atlantisia and Rallus
(Olson 1977b).
The Mauritius Red Rail appeared to be able to survive the onslaughts of human occupation and associ-
ated animals including monkeys, pigs and rats, right up until the end of the 17th century. They continued to
be hunted on a daily basis in the 1680s and early 1690s by Dutch settlers (Hume et al. 2004), but the introduc-
tion of cats around this time proved disastrous (Cheke & Hume 2008), and the rails, briefly mentioned by
Leguat as rare in 1693 (Leguat 1708), were not seen again.
Habits
Hardly anything is known of its habits, but a recently discovered document written by Johannes Pretorius
while on Mauritius in 1666–69 (Hume & Winters 2015) provides some important information about the
rail’s ecology:
The dodaers [Aphanapteryx bonasia] is a red bird, as big as a fowl, has short wings and cannot fly. It scratches in
the earth with its sharp claws like a fowl to find food such as worms under the fallen leaves. This bird is unbeliev-
ably stupid. When one waves a stocking cap and makes a sharp sound with the mouth, it immediately heads
towards that person, and if one carries a stick, all of them can be killed with it without any escaping. They are
fatty and greasy to eat. They have a long, sharp beak which is slightly curved at the end.
Pretorius’s account confirms that the rails were feeding on invertebrates amongst leaf litter, and Hachisuka
(1953) surmised that as the tip of the bill was very sharp and strong, it probably fed by crushing molluscs
and other shells, rather like an oystercatcher. Hachisuka was prone to rather over-zealous interpretation,
even describing the above-mentioned ‘Wheaten’ bird as a new species based solely on the account of
Mundy (Hachisuka 1937a). However, Hachisuka’s suggestion may well have some credence, as many shells
of some of the largest but now-extinct Mauritius land snails (e.g. Tropidophora carinata) have a puncture hole
on the ventral surface, suggesting bird predation (Owen Griffiths pers. comm.). The rails were probably
opportunist omnivores.
The Rodrigues Rail is distinct from the Mauritian Red Rail in a number of characteristics. They were first
reported by Leguat (1708) in 1691–93:
Our Wood-hens are fat all the year round, and of a most delicate taste. Their Colour is always of a bright Gray,
and there’s very little difference in the plumage between the two sexes. They hide their Nests so well, that we
cou’d not find ’em out, and consequently did not taste their Eggs. They have a Red List about their Eyes, their
Beaks are straight and pointed, near two Inches long, and red also. They cannot fly, their fat makes ’em too heavy
for it. If you offer them anything that’s red, they will fly at you to catch it out of your Hand, and in the heat of the
Combat, we had an opportunity to take them with ease.
They were also mentioned by Tafforet in 1725–26, who gave more details (from A. Newton’s 1875
translation):
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There is a sort of bird, of the size of a young hen, which has the beak and feet red. Its beak is a little like that of a
curlew, excepting that it is slightly thicker and not quite so long. Its plumage is spotted with white and grey.
They generally feed on the eggs of the land tortoises, which they find in the ground, which makes them so fat that
they often have difficulty in running. They are very good to eat, and their fat is of a yellowish red, which is excel-
lent for pains. They have small pinions [wings], without feathers, on which account they cannot fly; but, on the
other hand, they run very well. Their cry is a continual whistling. When they see anyone who pursues them they
produce another sort of noise, like that of a person who has the hiccups.
When Pingré (1763) visited Rodrigues to observe the Transit of Venus in 1761, he stated that the Rodrigues
Rail was extinct. Their rapid disappearance between 1726 and 1761 suggests that introduced cats were the
main culprits, but over-hunting by humans may have also contributed significantly to their demise.
Subfossil remains
Milne-Edwards (1874, 1875) possessed both the original accounts and the first subfossil remains, recently
discovered in caves on the Plaine Corail, Rodrigues. From these he determined that a flightless rail, related
to Aphanapteryx on Mauritius, had once occurred on Rodrigues. He proposed a new genus, Erythromachus,
for the Rodrigues bird, but as better material became available (Günther & Newton 1879), it was placed in
the genus Aphanapteryx. However, Olson (1977b) suggested that it should be retained in Erythromachus due
to distinctive differences in the skeleton, which was followed by Mourer-Chauviré et al. (1999). As in
Aphanapteryx, the bill is variable in curvature and pronounced sexual dimorphism in size is exhibited in the
skeleton.
De Booy’s Rail was a very large, heavy, flightless rail that may have persisted into comparatively recent
times. Its affinities are unclear. A report (in 1943) that a ‘flightless waterhen’ inhabited Virgin Gorda, was
now rare but once common, and which the older islanders recalled killing with sticks (Olson 1977b), may
have been referable to this rail. It was certainly hunted throughout its range, as large numbers of bones have
been found in archaeological middens, which suggest that it must have been an important food item for the
island’s p
re-Columbian inhabitants. Over-hunting was probably the main reason for its extinction.
This was a small, flightless rail, known only from fragmentary skeletal material from cave deposits near
Saint-Michel de l’Atalaye, Haiti. It was the smallest member of its genus (Olson 1974), and there was consid-
erable size variation in the specimens.
This species is known only from fragmentary skeletal remains from the Pi’o Domingo cave, near
Sumidero. It was a flightless rail, intermediate in size between Nesotrochis debooyi and N. steganinos, and
probably had a wider distribution on Cuba in the past (Olson 1977b). It probably disappeared due to
over-hunting.
The first subfossil remains of this bird were found in a limestone cave in Karamu, with subsequent material
discovered in several other localities on North Island (Olson 1975c). This flightless rail was one of the most
distinctive members of the Rallidae, with the longest bill and perhaps the smallest wings of any member of
the family (Olson 1977b). The bill was flexible, strongly decurved at the tip, and had sensory pits, all adap-
tations for probing (Olson 1975c). The legs were particularly robust, and the s nipe-rail may have filled a
kiwi-like role, probing the ground for invertebrates.
Hawkins’s Rail was a very large flightless rail, with a long, pointed decurved bill, tiny wings, and robust
legs with much shortened toes (Andrews 1896a; Olson 1977b). It is known from abundant subfossil remains
found in sand dunes on Chatham Island, and analysis of its anatomy suggests it is derived from a Gallirallus
ancestor (Olson 1975c).
Until recently, it was known only from the fossil record, but two accounts, one from Walter Rothschild’s
collector Sigvard Jacob Dannefaerd dated 1895,
and another by Alexander Shand in 1911, describe Hawkins’s Rail
Hawkins’s Rail in life. Both Dannefaerd and Diaphorapteryx hawkinsi
Shand questioned the oldest member of the
Moriori Polynesians, who recollected the rails
from his youth. The first account was reported by
Dannefaerd (from Cooper & Tennyson 2004):
Of the large rail that Forbes classed as
Aphanapteryx, the Moriori name for same is
“Mehonui” (nui = large). I hunted up the oldest
Moriori chief on the island and their tradition of the
bird is that it was larger than a goose and had a very
loud cry (like a Tue–ck). It always walked with the
head down, pecking in decayed wood on the ground.
They [the Moriori] would often catch them by
walking straight up to them from the front, but they
could not come over to them from the side. They [the
rails] used to sleep together in colonies and the
Morioris used to find out where a colony had their
sleeping place, then sneak quietly up, make a rush,
and kill the whole Colony. They described them as a
dull, brick red in colour.
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Shand (see White 1896; Cooper & Tennyson 2004) gave more details:
The Mehonui was usually captured on its sleeping place or nest, where several – six or eight – might be found
huddled together, as the Morioris declare, like pigs in a bed. Having by observation, found its sleeping place on
the ‘clears,’ the Morioris made long tracks leading up to it, carefully removing any sticks or obstructions which
might alarm the bird by cracking, and then, by making a stealthy rush, they pounced on and secured all in the
nest or sleeping place. This bird had a powerful strident call, which could be heard at great distances. Its neck
was said to be about as long as a man’s arm. The Mehonui was peculiar in this, that if any one approached it in
front it did not see him, and, approached thus quietly, was caught by the neck and strangled. It kept its head
continually on the ground looking for food, chiefly fernroot, which it burrowed for and dug out with its powerful
bill, making, it is said, a rooting like a pig; any one, however, coming from the side or behind was quickly
detected, and the bird made off. Its colour was a reddish-brown, something like the New Zealand Kaka.
A large, flightless rail that was seemingly so easy to catch must have been particularly vulnerable to
over-hunting. The introduction of non-native predators would no doubt have hastened its extinction.
The Viti Levu Rail was flightless, with much reduced wings, short robust legs and an elongate, slender bill.
The discovery of this species has shown that at least seven species of rail, in seven genera, were once sympa-
tric on Viti Levu (Worthy 2004). It probably disappeared due to the same factors that have exterminated
much of the oceanic island avifauna throughout the Pacific, a combination of o ver-hunting, habitat destruc-
tion and predation by introduced animals, particularly the Polynesian Rat Rattus exulans. Subfossil remains
of the rail have been found in the dry lowlands (Worthy 2004); this habitat would also have been particu-
larly susceptible to fire.
This was a large gallinule, a species originally placed in Tribonyx by Andrews (1897), of about the same size
as the Tasmanian Native Hen Tribonyx mortieri (43–51cm in total length), but wing bones have yet been
found so it is unknown whether it was similarly flightless. Tribonyx is confined to the Australian region;
therefore characters in the skeleton prompted Brodkorb (1965) to erect a new monotypic genus Hovacrex for
roberti. Olson (1977b) questioned the generic value of the characters, and noted that if more bones were
discovered, it might be possible to reconcile the Hova Gallinule with some other extant genus. Nothing
more is known about the bird or the reasons for its extinction.
Perhaps the most enigmatic of all rails, the ‘Oiseaux bleu’ once occurred on Réunion Island, but as yet not a
shred of physical evidence of any kind has been found to resolve its taxonomy. There can be no doubt that it
was a large, terrestrial Porphyrio derivative (Olson 1977b; Mourer-Chauviré et al. 1999), and possessed char-
acters – larger size and more robust legs – usually associated with reduced powers of flight. The gallinule
was mentioned on a number of occasions by trustworthy observers, and appears to have been restricted to
the mountains, particularly a montane, marshy plateau called the Plaine des Cafres. From the accounts it
can ascertained that the Réunion Blue Gallinule was considered good game, and although able to fly, could
easily be caught and killed with sticks.
Little is known about it. Dubois (1674), while staying on Réunion in 1671–72, gave the first description:
Oiseaux bleus, as large as solitaires. Their plumage is entirely blue, the beak and feet red, made like hen’s feet.
They do not fly, but they run extremely fast, such that a dog has difficulty catching them in a chase. They are
very good [to eat].
Feuilley (1705) in 1704 described them further:
The Oiseaux bleuff live in the plaines on top of the mountains, and especially on the Plaine des Cafres. They are
the size of a large capon, blue in colour. Those that are old are worth nothing to eat because they are so tough, but
when they are young they are excellent. Hunting them is not difficult because one kills them with sticks or with
stones.
De Villers (1701–10) in 1708 gave the only details about the nest:
One sees there [the Plaine des Cafres] a great numbers of oiseaux bleus which nest amongst grasses and
aquatic ferns.
Le Gentil (1727) in 1717, extended by ‘Père Brown’ (1773), confusingly referred to it as resembling a wood
pigeon (probably Alectroenas):
Towards the east of the island there is a little plateau up a high mountain called the Plaine des Cafres where one
finds a large blue bird whose colour is very striking. It resembles a wood-pigeon. It flies but rarely and always
barely above the ground, but it walks with surprising speed. The inhabitants have never called it anything other
than oiseau bleu; its flesh is quite good and keeps well.
The bird had probably disappeared by the early years of the 18th century, primarily due to over-hunting,
but the accidental introduction of rats in 1676 (Cheke & Hume 2008) would also have made the eggs and
chicks extremely vulnerable to predation.
Size
There has been some disagreement regarding the size of the Réunion Blue Gallinule (Cheke & Hume 2008),
as Dubois stated that they were ‘the size of solitaires’, and Feuilley that that they were ‘as large as a capon’.
However, the Réunion Solitaire Threskiornis solitarius (p. 67) would have been no more than 65cm in total
(the size of the Sacred Ibis Threskiornis aethiopicus), and capons (or domestic chickens) were approximately
the size of the ancestral Red Junglefowl Gallus gallus, again about 65cm in total length, so there is no real size
discrepancy in the accounts. The Réunion Blue Gallinule was approximately the size of a South Island
Takahe Porphyrio hochstetteri (see Milton 1951).
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The flightless North Island Takahe Porphyrio mantelli was originally described from a skull obtained by the
collector, Walter Mantell, from North Island in 1847. The extant South Island Takahe P. hochstetteri was
described later, and was considered a race of the North Island bird. Oliver (1930), Scarlett (1972) and others
noted that P. mantelli is larger than the population from South Island, but otherwise similar. However, DNA
evidence has shown that the North Island Takahe is genetically more similar to the Australian Purple
Gallinule P. melanotus than to the South Island bird; the two populations evolved independently, with those
on South Island from an earlier colonisation event (Trewick 1996, 1997b), although this is not supported by
their morphology (Livezey 1998b). Nothing is known about the habits of the North Island Takahe, but
presumably it had a similar ecology to the South Island species.
The North Island Takahe probably survived until the late 19th century. In 1894 a bird matching the
description of a takahe was caught in the Ruahine Mountains by the surveyor Morgan Carkeek (Philipps
1959). He took the specimen to the house of Roderick McDonald, where it was identified as the rarely seen
mohoau by a local Maori. Roderick’s son was present when the specimen was brought to the house, and he
informed Philipps that it resembled a Pukeko Porhyrio porphyrio and had blue feathers. The North Island
Takahe must have disappeared soon after due to anthropogenic changes and predation by introduced
mammals. The South Island Takahe was also thought to be extinct, but a tiny population was discovered in
Fiordland in 1948 (Oliver 1955).
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affects only the melanin and not the carotenoid pigments (reds and yellows), the Lord Howe Gallinule’s bill
and legs retained normal coloration. Unfortunately, these minor discrepancies in plumage detail, along
with uncertainty over provenance, has resulted in a wealth of confusing taxonomic literature, with taxa
being described from wholly inadequate evidence (see Rowley 1875; Mathews 1928).
Literature muddles
There is no doubt that there was a population of largely white gallinules on Lord Howe Island that disap-
peared during the early part of the 19th century. The Lord Howe Gallinule was confined to Lord Howe
Island, and there is no substantiating evidence to suggest that it occurred anywhere else (contra Rothschild
1907a; Pelzeln 1860b; Mathews 1928). It was said to have also inhabited Norfolk Island, but as the capable
naturalists the Forsters make no mention of a white gallinule when Norfolk Island was discovered it is
extremely doubtful that one ever occurred there historically (see Iredale 1910; Hindwood 1932). However,
subfossil remains of a gallinule have been found on Norfolk (see Norfolk Island Gallinule).
The Australasian race of Purple Gallinule, Porphryio porphyrio melanotus, has been a regular visitor to
Lord Howe Island over the last 130 years; relatively recently (in the 1980s) it established a breeding popula-
tion on the island (Hutton 1991). Hindwood (1965) considered that these blue birds from Australia may
have interbred with the resident Lord Howe Gallinules. However, the Lord Howe Gallinule differed geneti-
cally (see below) and was also flightless (or at least behaviourally so; van Grouw & Hume 2016), strongly
suggesting that no interbreeding occurred. A mtDNA analysis has now shown that P. albus is correctly given
full specific status, and it is actually most closely related to the Philippine Purple Gallinule race, P. p. pulver-
ulentus (Garcia-R & Trewick 2015); P. albus arose from a small number of migrant individuals of pulverulentus
during the late Pleistocene, around 500,000 years ago. But because the Philippine birds would have had to
migrate over other islands in order to reach Lord Howe, the authors remain cautious as to this conclusion.
Another interesting zoogeographical puzzle for future researchers to resolve.
Early encounters
The Lord Howe Gallinule was recorded over a period of just two years, from 1788–1790 (van Grouw &
Hume 2016). Only a few accounts are first-hand, and were written by visitors to the island either on the way
to or from Norfolk Island. David Blackburn, who was Master on the ship Supply, landed on Lord Howe 19
March 1788, and again on 16 May 1788 (Blackburn 1934), was the only person to mention the diet of the
Lord Howe Gallinule:
. . . on the shore we caught several sorts of birds. . . and a white fowl – something like a Guinea hen, with a very
strong thick & sharp pointed bill of a red colour – stout legs and claws – I believe they are carnivorous they hold
their food between the thumb or hind claw & the bottom of the foot & lift it to the mouth without stopping so
much as a parrot.
White (1790), who never personally visited Lord Howe but obtained information from others who had been
to the island (van Grouw & Hume 2016), described the ease with which the gallinules could be caught:
They . . . found . . . in great plenty, a kind of fowl, resembling much the Guinea fowl in shape and size, but
widely different in colour; they being in general all white, with a red fleshy substance rising, like a cock’s comb,
from the head, and not unlike a piece of sealing wax. These not being birds of flight, nor in the least wild, the
sailors’ availing themselves of their gentleness and inability to take wing from their pursuits, easily struck them
down with sticks.
Extinction
After being considered to be common when first encountered, the Lord Howe Gallinule disappeared very
quickly, possibly within a decade of discovery (van Grouw & Hume 2016). Lord Howe was not settled until
1834 (Hindwood 1940); Foulis (in Hindwood 1940) was on the island from 1844 until 1847 and undertook an
ornithological survey, the first in 63 years (van Grouw & Hume 2016). He made no mention of a white galli-
nule, which must have been long extinct by that time. Whalers and sealers had used the island for supplies,
and they probably hunted the bird to extinction. Rats and cats, the usual culprits in island extinctions of
birds, did not arrive until later (Hindwood 1932).
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Archaeological evidence has shown that Polynesians once occupied Norfolk Island for a period of two
centuries, after which they mysteriously abandoned the island (Kirch 1988). It was during the Polynesian
period that a number of bird extinctions took place. Holdaway & Anderson (2001) described subfossil
remains from a Polynesian midden site at Emily Bay, Norfolk Island, which mainly comprised seabirds, but
some terrestrial bird remains were present. Holdaway & Anderson (2001) discovered a few skeletal elements
of a Porphyrio swamphen, smaller than the race of Purple Swamphen Porphyrio porphyrio on New Zealand,
melanotus, and more similar to the small subspecies to the north of Norfolk Island rather than those of New
Zealand and Australia. Purple Swamphen is widespread in the Pacific (Taylor & van Perlo 1998), but it
appears to have only reached Norfolk Island infrequently (Schodde et al. 1983).
The presence of more than one individual differing from P. porphyrio suggests that a native population of
swamphens once occurred on the island, but over-hunting by the Polynesians exterminated them.
This was a very large, flightless gallinule, almost equalling the extant South Island Takahe Porphyrio hoch-
stetteri of New Zealand in size (around 63cm in total length), but characteristics of the bones suggest that it
was closer to a typical Porphyrio than to P. hochstetteri (Balouet & Olson 1989). The tarsi were longer but less
robust than in P. hochstetteri, and the bill was not as deep. There was also distinct body-size sexual dimor-
phism, the larger individuals probably being male. Although known only from subfossil remains, the New
Caledonia Gallinule may have survived into comparatively recent times. A passage in Verreaux & Des Murs
(1860) states that a bird the size of a turkey, called N’dino by the locals, inhabited marshy places. If this was a
reference to the gallinule, it must have become extinct shortly after.
Subfossil remains of gallinules have been collected on a number of Pacific oceanic islands, and many of
these, pending better material, await description. All these species suffered from the actions of human
beings and introduced predators, disappearing soon after colonisation by Polynesians. The Huahine
Gallinule was sympatric with the Huahine Rail Hypotaenidia storrsolsoni (see p. 107), and was a small species,
approximately the size of an American Purple Gallinule P. martinica, at about 27–36cm long (Kirchman &
Steadman 2006b). No wing elements have yet been discovered, so it is not certain that it was flightless, but
as almost all insular island gallinules have poor or no flying capabilities, it is most likely that the Huahine
Gallinule was a poor flier at best.
Until the discovery of the Huahine Gallinule (see above), the Marquesas species was the only known galli-
nule in eastern Polynesia (Steadman 1988); however, other species are waiting to be described. This suggests
that like Hypotaenidia rails, there was a radiation of Porphyrio gallinules throughout the region. This was a
small gallinule, found on two islands, Hiva Oa and Tahuata, which are separated by just 3km. The propor-
tions of the limbs are not sufficiently diagnostic to ascertain whether or not this bird was flightless, but it
probably had reduced flying capabilities.
There is evidence that the Marquesas Islands Gallinule may have survived until as late as 1937. A
painting by Paul Gaugin, executed in 1902, depicts a gallinule-like bird from the Marquesas in the lower
right corner being seized by a dog, which may be a representation of this species. It was known locally as
the Koau. Furthermore, the Norwegian explorer Thor Heyerdahl reported seeing a hen-sized flightless bird
in 1937 on Hiva Oa, moving rapidly between some tall ferns; he compared it to a rabbit moving through a
burrow (Heyerdahl, 1974). The bird must have died out soon after.
The New Zealand Flightless Gallinule was originally known only from incomplete postcranial material,
which made determination of its relationships problematic. The discovery of a skull and jaws showed that
the bird was closely related to the Australian Tribonyx gallinule group, and that has proved to be the case
(Olson 1975c, 1977b; Taylor & van Perlo 1998; Worthy & Holdaway 2002).
The gallinule had reduced wings with short, heavy tarsi. The skull was most similar to the extant but
equally flightless Tasmanian Native Hen Tribonyx mortierii. The wings were even more reduced than in T.
mortierii. Olson (1977b) hypothesised that the New Zealand species, along with the volant and flightless
Australian forms, were all descended from a flying ancestor.
Numerous fossil remains have been found at Lake Poukawa at Hawke’s Bay, which was originally
surrounded by forest, suggesting that the gallinule was a forest inhabitant (Tennyson & Martinson 2007).
Holdaway (1989) considered that it occurred in riparian swards (areas of open grassland kept short by
grazing moas), open forest glades, and always near water. Its diet comprised seeds, shoots and stem bases
of grasses and sedges, and it nested on the ground in a tussock or clump of sedge (Holdaway 1989).
Being flightless and easy to catch, this species was probably driven to extinction by over-hunting and
predation of its eggs and chicks by the Pacific Rat Rattus exulans, but despite this onslaught it may have
survived in remote areas until the 18th century (Tennyson & Martinson 2007).
middle; edge of outer primary white; bill and shield bright red with yellow tip; iris r eddish-brown with grey tinge; feet and legs
greenish-yellow.
The main island of Tristan da Cunha is a volcanic cone rising to approximately 2,130m, with an area of
41.4km2. Precipitous cliffs surround it on all sides except the n orth-west, where the settlement is situated.
There are several small streams, and freshwater lakes in the craters of the volcano.
Tristan da Cunha was discovered in 1506 by Portuguese sailors. The islands apparently remained undis-
turbed until 1643, when Dutch vessels brought back reports, and in 1656, Van Riebeck, the founder of Cape
Town, sent a ship to see if the island was suitable for occupation. It was not until 1810, however, that the first
permanent settlement was made.
Taxonomy
The Tristan Island Moorhen has been variously considered conspecific with or a subspecies of the closely
related (but extant) G. comeri of Gough Island, which lies 400km to the south-east of Tristan da Cunha
(Taylor & van Perlo 1998). Olson (1973a) suggests that the islands were populated by two different colonisa-
tion events, most likely by Common Moorhen G. chloropus, and each species evolved in parallel, being more
related to the volant G. chloropus than to each other.
The last recorded specimens of the Tristan Island Moorhen were received in London on May 25, 1861, of
which three were alive (Sclater 1861c). The Challenger Expedition found none in 1873 (Sclater 1881), though
the islanders stated it was rare (Beintema 1972). The species has never been seen since. Hunting by humans,
predation of eggs and young by introduced rats and wild pigs, which were found in the most excluded
areas (Carmichael 1818), and severe deforestation to promote grazing all contributed to the bird’s
extinction.
Habits
Little is known about the bird’s habits, but its ecology was probably similar to the Gough Island Moorhen
(Taylor & van Perlo 1998). Carmichael (1818) provided precious little detail:
The only land birds on the island are a species of thrush (Turdus Guianensis?), a bunting (Emberiza
Brasiliensis?), and the common m oor-hen (Fulica Chloropus [= Gallinula nesiotis]). These birds have spread
over the whole island, and are found on the table-land as well as on the low ground. The Fulica [Tristan Island
Moorhen] conceals itself in the wood, where it is occasionally run down by the dogs; the others fly about the
cantonment, and are so tame as to suffer themselves to be caught with a hand-net.
A Captain Patten (in Pike 1873) added little more:
There is a good deal of timber, though not high. The principal trees resemble the yew in foliage, with a wood like
maple, and burns well. Wild celery, dock, sorrel, and parsley are found. Gannets, penguins, albatrosses, Cape
cocks and hens, and a bird [Tristan Island Moorhen] something like a partridge, only it is black, and cannot fly,
are abundant.
Sclater (1881) gave the last account:
A third land bird which still exists on Tristan da Cunha, though no specimens were obtained by the Challenger
Expedition, is Gallinula nesiotis. The ‘Island-hen’ as the inhabitants call it, lives on the higher plateau, and is
‘scarcely able to fly.’
Similar remarks were made by other observers, e.g. Earle (1832) and Gurney (1853). No other details are
available.
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Description 25cm (10in): A small moorhen, with a red bill and bright yellow frontal shield. It had comparatively large eyes,
which suggests it was crepuscular or nocturnal (Greenway 1967). Adult head, neck and breast bluish; blackish on face, chin and
throat; upperparts dark olive tinged brown; underparts dark olive tinged green; lower rump, tail and u ndertail-coverts black;
bill reddish orange, more yellowish on frontal shield, eye ring red; iris brown-red; legs and feet light red; claws hornish-brown.
Immature bill r eddish-brown, tip darker brown; shield dirty yellowish; legs and feet reddish h orn-brown; claws h orn-brown.
Egg creamy white background with numerous purplish and reddish-brown spots, more concentrated at the larger end.
The Samoan Gallinule was first collected in 1869. Just a few years later, two birds were probably given to the
Challenger Expedition in 1873, and an egg was collected in October of the same year (Knox & Walters 1994).
The species probably survived until the end of the century; Reed (1980) reports that a local resident, Mr
Kurt Stoentzer of Apia, considered the gallinule extinct since 1907. The Whitney Expedition searched for it
in 1926 but found no trace. It was highly prized as food by the natives, and introduced cats and rats were
dangerous predators, despite the bird having survived alongside these enemies for years (Greenway 1967).
Bellingham & Davis (1988) thought they may have seen two birds on August 22, 1984 in upland forests on
Savai’i, which hints that it may still survive, although there has been no further confirmation. However,
Pratt & Mittermeier (2016) state that Bellingham & Davis’s observation was in error, and that they had misi-
dentified either adult or young Purple Gallinule Porphyrio porphyrio or Buff-banded Rail Hypotaenidia
philippensis. The Samoan Gallinule was only ever recorded in the lowlands, which made it extremely vulner-
able to the spread of introduced species; Pratt & Mittermeier, who made detailed surveys between 1977 and
2006, suggested that the Samoan Gallinule is extinct and has probably been so since the 1870s.
Taylor & van Perlo (1998) placed the Samoan Gallinule into Gallinula, but Olson (1973b, 1975d) gives
good reason to retain it in Pareudiastes, along with P. silvestris of Makira in the Solomon Islands (see below).
This assignation has proven to be correct (Dickinson & Remsen 2013; del Hoyo & Collar 2014).
Habits
Whitmee (1874), a missionary to Samoa, noted that the Samoans always referred to the gallinule as the ‘bird
that burrows like a rat and lives underground’. The burrow was described as being about three or four feet
in length, curving downwards and then rising to near the surface, where the nest was placed. However,
Whitmee obtained evidence that the Samoan Gallinule nested above ground, and that the nest was
constructed from a few twigs and a little grass, in which two eggs were laid. Taylor & van Perlo (1998)
suggest that it may have only temporarily used burrows for refuge.
Pritchard (1866) described a Samoan hunting technique for catching the birds:
The Punae (or ‘springer-up’) is another very remarkable bird, which is now also nearly extinct . . . The natives
state that it burrows in the ground; feeds on grubs, worms, and insects; runs very fast; and when first startled
from its burrow, make a long spring upwards from the ground, but having very small wings it cannot fly . . . It
is said to be excellent eating, and as such was very esteemed by the natives. In the days when the punae were
plentiful, regular hunting parties used to go after them; dogs were trained to find their burrows; and the bird
was taken by placing a net over the burrow, in which it became entangled when, emerging from its haunt, it
made the long spring with which it invariably started when disturbed.
Whitmee (1874) was informed that it occurred on both Upolu and Savai’i, but it was apparently more
common on the latter, and was ‘only ever recorded some distance inland amongst the mountains’. He gave
precious few details about its habits:
The Pareudiastes evidently feeds upon insects. The Samoans are very clever at taming and feeding birds; but I
believe they have never yet succeeded in keeping this bird alive more than a few days. The man who caught the
one I sent to Canon Tristram last year tried to feed it on vegetable food; but it died in a few days. Another lately
procured on Savaii died in the same way on a vegetable diet. But one caught two months ago, after being fed by
cramming with vegetable food for three or four days, was brought, apparently in a hopeless state of starvation, by
one of M. Godeffroy’s collectors. He offered it some insects, which it eats readily, and on which diet it soon recov-
ered its health. It was fed for a fortnight upon beetles, grubs, &c., and was thriving well; but unfortunately, it
escaped from its cage one day while it was being fed.
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Rallidae
The Makira Gallinule was obtained once (an adult male) during the Whitney South Seas Expedition (Mayr
1933a). It occurred in rather inhospitable country where few naturalists had searched for it. There have been
a number of recent reports, so it may still survive, but these lack confirmation. It was reported by the Oxford
University Expedition in 1953 (Galbraith & Galbraith 1962) to be ‘well-known to the natives, and appar-
ently not rare below Nagasi’, and local people apparently noted its presence in 1974 (Taylor & van Perlo
1998); Dutson (2011) mentions local reports in 2002. Rats and cats are the likely culprits for its disappear-
ance, as the interior of the island remains well forested.
Habits
Little was recorded about the biology of the bird. Mayr (1933a) reported that the natives called it Kia, and
said it lived in the dense undergrowth of mountain forests, flew little and was hunted by them with dogs. It
must have been very rare at this time, as the expedition failed to obtain a second specimen in spite of many
rewards offered. The type locality was the village Húnogáraha or Hanagáraha, which is at an altitude of
580m in the centre of the island, about 17km south of Wanoni Bay. The region was covered with primary
forest, native plantations and secondary growth, and had many brooks and creeks which cut into the moun-
tainsides, but no standing water. The Makira Gallinule is thought to have been a species confined to dense
undergrowth on steep slopes in mountain forests.
Mayr (1933a) considered the Makira Gallinule to be generically distinct with no close relatives, but most
similar to the Samoan Wood Rail Pareudiastes pacificus; Olson (1973b, 1975d), noting similarities in the bill,
frontal shield, and tarsometatarsus, referred Mayr’s Edithornis silvestris to Pareudiastes. It differed from P.
pacificus in having a larger bill, feet and frontal shield, and its legs and feet were proportionally longer.
Milne-Edwards (1867) described a large coot from a series of bones found at the Mare aux Songes marsh,
Mauritius. It was thought to be a derivative of the Common Coot Fulica atra (Olson 1977b) or Red-knobbed
Coot F. cristata (Mourer-Chauviré et al. 1999). More recently F. newtonii has also been identified from subfossil
material discovered on Réunion, which is inseparable from the Mauritius population.
Contrary to suggestions that the Mascarene Coot was flightless (e.g. Olson 1977b), Newton & Gadow
(1893) indicated there was only slight reduction in the sternal keel, and M ourer-Chauviré et al. (1999)
suggested that it still had the capability to fly between the two islands in comparatively recent times.
Habits
Dubois (1674) in 1671–72 gave the best account on the bird:
Water-hens, which are as big as chickens. They are completely black, and have a big white crest on the head.
The ‘big white crest’ refers to the frontal shield, a characteristic fleshy feature that occurs in a number of
rallid genera. The coot disappeared from Mauritius sometime after 1693, when Leguat (1708) stated they
were rare. The population on Réunion may have survived until the late 17th century, but François Martin in
1665 and 1667 (in Lougnon 1970) stated that even by the 1660s the coots were in serious decline:
[In 1665] The river basin [at Saint-Gilles] was covered in geese and w
ater-hens, and the depths full of fish . . .
the water-hens allowed one to approach almost [close enough] to catch them by hand; we sent them all on
board. [But by 1667] We saw neither geese nor w ater-hens on the Etang de St. Paul which was formerly covered
in them.
Both populations were presumably exterminated through over-hunting and introduced predators.
Subfossil remains of a giant, flightless coot were found on Chatham Island and North and South Islands,
New Zealand, in the late 19th century. They were originally recognised as distinct (Forbes 1892b; Hamilton
1893), but Scarlett (1955a) concluded that the two were identical. The coot was variously placed in different
genera, but apart from its great size, Olson (1975c) considered that it was not generically different from
typical coots of the genus Fulica, and that the Chatham Island birds differed only subspecifically in charac-
ters of the humerus and skull from those of New Zealand. Worthy & Holdaway (2002) r e-examined the
subfossil remains, however, and concluded that the differences are in fact enough to warrant full species
status.
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Turnicidae
The Chatham Island Coot, the largest coot known, was extremely large and heavy, weighing between
1.5–2kg – twice the weight of closely related coots. It had reduced wings and robust legs, but it was prob-
ably still able to fly (Holdaway 1989; Worthy & Holdaway 2002; Tennyson & Martinson 2007). The neck was
long, and it had disproportionally long legs and toes, which was an adaptation for walking on soft ground
(Tennyson & Martinson 2007). The Chatham Island Coot probably occurred in lakes, ponds, swamps and
along streams in forest, and fed on shoots and leaves of aquatic plants (Holdaway 1989) and perhaps on
eggs of seabirds and invertebrates (Tennyson & Martinson 2007). It differed from the New Zealand species
F. prisca by having salt-water excreting glands, indicating that these birds were adapted to the salt-water
lagoons on Chatham Island (Olson 1977b).
Like so many other rails restricted to oceanic islands, the arrival of humans and their commensal animals
would have rapidly exterminated the Chatham Island Coot, which had evolved in a terrestrial mammal-free
environment.
When Hamilton (1893) described subfossil remains of a coot from New Zealand, he was unaware that F.
chathamensis had already been described. In his description of Fulica prisca, he remarked that:
It must have been a bird nearly as large as the Notornis [= North Island Takehe, Porphyrio mantelli], but with
a small head, and a frontal shield like the pukeko and Notornis.
Rothschild (1907a) noted that the New Zealand species was smaller than the Chatham Island bird F. chatha-
mensis, with less reduced wings, while the proportions of the skeleton show that it was still capable of flight
(Worthy & Holdaway 2002; Tennyson & Martinson 2007). Olson (1977b) suggests that these coots were
derived from Eurasian Coot Fulica atra, first arriving in New Zealand whereupon it achieved great size,
then subsequently colonising the Chathams (and getting even bigger).
Subfossil remains are common in midden deposits, so the New Zealand Coot would have made easy
prey for the Polynesian colonists; their eggs and young would have also been susceptible to introduced
predators. It probably disappeared soon after permanent settlement of the islands.
The New Zealand and Chatham Island coots were nearly twice the weight of other, closely related coots,
and they had comparatively long necks and disproportionately long legs and toes, which was probably an
adaptation for walking on soft ground (Tennyson & Martinson 2007).
Turnicidae (Buttonquails)
The buttonquails are small terrestrial birds that superficially resemble quails, but they are unrelated. The
family is divided into two genera and they have a wide distribution in Africa, Asia, Europe and Australia,
including large islands and small island groups. Some species are threatened due to o ver-hunting and
deforestation.
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Description 19cm (7.5in). The Painted Buttonquail is g reyish-brown above with black, brown and white flecking; crown, face
and breast flecked with white; pale greyish below with white streaking; shoulder chestnut; bill greyish horn; iris red; legs and
feet yellow. Male slightly smaller and duller. Differed from Painted Buttonquail Turnix varius in having most feathers of rump
and upper sides almost entirely black, margined with whitish or buff, and in being slightly smaller.
The Painted Buttonquail T. varius occurs in Australia and a number of islands, and is considered unthreat-
ened throughout its range. A distinctive subspecies known only from the type specimen once occurred on
New Caledonia, and it (or a closely related taxon) has been commonly found as subfossils in a number of
cave sites. Debus (1996) has suggested that it should be given specific status. A cave containing numerous
subfossil remains deposited by Barn Owls Tyto alba has shown that the population, although numerous, has
been in decline for some time (Boyer et al. 2010), and the type specimen may have been taken just before the
species became extinct.
The buttonquail’s remains have also been collected from surface deposits in a number of cave sites
suggesting that it was once widespread in New Caledonia (Balouet & Olson 1989; Boyer et al. 2010; Anderson
et al. 2010). The reasons for its decline are unclear, but deforestation through burning and perhaps intro-
duced mammalian predators may have contributed to its disappearance.
Nothing is known about the biology of this particular race, but elsewhere the species occurs in grassy
forests and woodlands, foraging for invertebrates and seeds.
The Small Buttonquail Turnix sylvatica has an extremely wide distribution, occurring from n orth-western
Africa to South-east Asia, the Philippines and Indonesia, with nine subspecies currently recognised. One of
these, T. s. sylvatica, has often been listed as extinct, but it is still extant with a population of 5–10 pairs in
Spain and it still occurs on the north-west African coast. The Tawi-tawi birds endemic to Jolo and Tawi-tawi
Islands in the Sulu Archipelago have not been recorded since the 1950s, and may be extinct. The islands
have suffered severe deforestation and the bird’s terrestrial habits may have made it vulnerable to intro-
duced predators.
Habits
Little is known about its habits. Other Small Buttonquail races occur in dense thickets, scrub, open grass-
land and agricultural areas. These birds forage by gleaning and scratching on the ground predominantly for
grass seeds and various invertebrates. They are reluctant fliers, preferring to skulk, and are only flushed
with difficulty.
Haematopodidae (Oystercatchers)
The oystercatchers are generally all black or black and white coastal birds with long, red or orange bills and
are widely distributed around the world, being absent only from the north and south poles and some trop-
ical regions of Africa and Asia. There has been some decline in populations due to habitat interference, but
only one species, the Chatham Island Oystercatcher Haematopus chathamensis, is considered endangered.
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Haematopodidae
The Canary Islands Oystercatcher was one of several black species, as opposed to the more usual pied
plumage of oystercatchers. It was generally considered a subspecies of the African Black Oystercatcher H.
moquini, but Hockey (1982) recognised its distinctiveness, and gave it specific status.
It was first recorded by Webb & Berthelot (1832), in the years 1828–30, occurring on the shores of Graciosa
and on the deserted shores of Lanzarote and Fuerteventura. Meade-Waldo (1893), who made the bird known
to ornithologists and described it as uncommon, collected three individuals with Canon Tristram, and the
local collector Ramón Gómez got two in the same period, while Bannerman (1913a) obtained the last known
bird on June 3, 1913. However, he also records a possible sighting in the mid-1960s, but Lovegrove (1971)
could find no evidence for the bird’s continued existence in 1971. According to local fishermen and light-
house keepers (Hockey 1996), the Canary Island Oystercatcher had disappeared by around 1940. Being a
resident species made it vulnerable to hunting, egg harvesting, habitat disturbance and perhaps introduced
predators. However, Hockey (1982) looked at Middens and discovered that the size class of shellfish on
which the oystercatcher fed slowly grew smaller as humans ate their way through the stocks. So it appears
that the species died out largely due to food competition with local human inhabitants.
Habits
It was a resident species on Fuerteventura, Lanzarote, and some of the offshore islands such as Los Lobos,
Graciosa, Montaña Clara, Allegranza and possibly the Roque del Este. Its supposed occurrence on Tenerife
(Godman 1872) is incorrect (Bannerman 1963) and it probably never wandered from the eastern Canary
Islands. Bolle (1855) described the birds as running about busily on the sand just above the tideline, picking
up shells. They were reluctant to fly, preferring to run, even when hard-pressed.
Bannerman (1913a) stated that they avoided sandy beaches, probably because of human disturbance.
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eade-Waldo (1893) described the bird as uncommon, always seen in pairs and very tame – this habit may
M
well be the reason for its demise. However, the bird seen by Bannerman (1913a) in June 1913 was very wild
indeed. Its voice was much louder and stronger than that of the Common Oystercatcher and he described
the alarm call as peepe–peepe peepe–peepe. Its diet was like that of other oystercatches, comprising small
molluscs and crustaceans (Hockey 1996). The breeding season is unknown. The nest or eggs were never
recorded, but the late Wolfgang Makatsch of Germany (pers. comm.) claimed to have an egg in his private
collection. Meade-Waldo (1893) obtained females with well-developed eggs but never took shelled eggs
with his own hand. He claimed, however, that he had eggs sent to him, and it may be one of these that was
obtained by Makatsch.
Charadriidae (Plovers)
Plovers are m edium-sized shorebirds with short bills and long legs, distributed in both the Old and New
Worlds. They inhabit open grasslands as well as wetlands. These birds are divided into two genera. Some
species are showing marked declines due to habitat destruction.
Nearly all of the specimens of this bird were collected on Java, and it was believed to migrate between there
and Timor. A bird was said to have been collected on Timor in 1828 or 1829, and a specimen and clutch of
eggs were thought to have been taken on Sumatra; otherwise the bird has never been seen on either of these
islands. Mees (in King 1981) considered these early records open to doubt.
Another bird was collected near Jakarta, Java in 1872, and just before 1885 there was a report of several
perched on a ship’s railing off the north coast of the island. It was seen in the 1920s, but considered local and
uncommon, and the last record was in Meleman on the south coast of east Java in 1939 (Bock 1958). The
reasons for its disappearance are unclear, but over-hunting and habitat modification may be the primary
causes.
Habits
Collar et al. (1999) translated a draft manuscript by a Swiss-German plantation owner, Max E. G. Bartels,
who had recorded details in the early 1920s about the lapwing’s habits. The lapwings were restricted to the
extensive swamps of the Sedari estuary and its tributaries, and occasionally in the lowlands of the Tjitaroem
delta, and at Rawah Tangerang. Bartels found individual pairs inhabited a large area. He recorded:
As soon as an intruder enters a territory the birds are up and circling him, giving their characteristic persistent
calls, and only calming down when he has left the territory . . . During the rainy season the birds keep to areas in
the swamps which are relatively little-flooded, since despite their long legs they prefer not to walk in open water
like stilts . . . Even though they know perfectly well how to use their legs, I have never yet seen them walking
long distances, only flying. Their flight is not acrobatic like their European cousins; I usually see them in fast
direct flight. Their food consists mainly of water- or swamp-living insect larvae, water bugs, beetles, snails (the
shells are broken up by the birds’ strong digestive muscles) and seeds of aquatic plants, which are, however, only
used as a supplement.
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Scolopacidae
Madagascar Lapwing
Vanellus madagascariensis Goodman
Vanellus madagascariensis Goodman 1996, p. 609 (Ampoza, near
Ankazoabo, Madagascar)
Specimens Subfossil remains are in New York and Paris.
Status Extinct. Known from subfossil remains only. Disappeared around the 14th century.
Range Madagascar.
The Madgascar Lapwing is known from a pair of humeri found in two different fossil localities, one in
Ampoza near Akkazoabo, and one in Lambohararna, north of Toliara (Goodman 1996). Radiocarbon dates
have shown that this species died out around the 14th century due to a natural aridification of its habitat,
which resulted in a contraction of range. Goodman and Jungers (2014) suggest that the Madagascar
Lapwing inhabited a riverine habit, and fed and nested on sand bars in rivers and along river banks.
Nothing else is known about this species.
The Coenocorypha snipe are now confined to the New Zealand area, and are distinct from the more wide-
spread snipe of the genus Gallinago. Their taxonomy has been extremely confused, primarily due to a lack of
specimens and their misidentification, and erroneous locality data, particularly with specimens from the
Auckland Islands.
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These snipe are endemic to a single island or small island group, though none now occurs on New
Zealand itself. Undescribed subfossil remains have been found of a large species in New Caledonia (Balouet
& Olson 1989) and Norfolk Island, of which Holdaway et al. (2001) concluded that it was most similar to the
Auckland Islands species, C. aucklandica. Another species, C. miratropica, has been described from Fiji (see p.
147); this genus was much more widespread in the past.
In 1870, Captain Bennett, of the schooner Mary Ann, saw two snipe on Little Barrier Island. One of these
was captured alive but subsequently died in captivity, and was presented to Auckland Museum by Mr. T. B.
Hill. Snipe have not been seen on the island since, and the population is considered extinct. Worthy et al.
(2002) referred subfossil remains discovered on North Island to this species, and a bird was reputedly shot
on Browns Island, Hauraki Gulf, in 1820 (Miskelly 1987). On North Island, it appears to have been a victim
of over-hunting by Polynesians and predation by the introduced Pacific Rat Rattus exulans or cats (Turbott
1990).
Big South Cape Island seems to have been the last island on which this snipe occurred, though it had previ-
ously occurred on Jacky Lee Island, from which it had been first described by Rothschild on the basis of
three specimens sent him by H. H. Travers (Rothschild 1921a). It formerly occurred on South Island but is
only known from subfossil remains, so probably disappeared after Polynesians arrived on the island.
Guthrie-Smith visited Big South Cape Island in 1913 and found snipe. Ten years later, Weka Gallirallus
australis were plentiful on the island where there had been none before, and the snipe were confined to
higher parts of the island. Twenty years after that, Richdale found both Wekas and snipe rare. The snipe had
already been exterminated on Jacky Lee Island by introduced Wekas.
The calls of this bird are thought to have been responsible for the Maori legend of the Hakawai (Miskelly
1987), who was said to have been a descendant of Tangaroa, the god of the ocean, and Rehua, the star
Antares, guardian of the uppermost heaven. The snipe had a nocturnal flight display in which distinctive
vocals were made in combination with air reverberating through the tail feathers as the bird descended
(Higgins & Davies 1996). The places on Earth where the calls were heard gradually declined, mirroring the
decline of the snipe, until the last was heard in the 1960s. The South Island Snipe managed to cling to exist-
ence on Big South Cape Island until Black Rats Rattus rattus got a toehold on the island in 1964. Don Merton
(2004) recalled the last moments of the South Island Snipe. A rat plague had been reported by m utton-birders
on Big South Cape Island in March 1964, but only after five months could Merton get to the island. He
found that the rats had already devastated the island’s fauna, and it was already too late to save the last
population of Stewart Island Bush Wren Xenicus longipes variabilis and Greater S hort-tailed Bat Mystacina
robusta. He managed to capture two individuals of the South Island Snipe to try and relocate them to a
nearby rat-free island, but their feeding requirements were poorly known and both died two days later. The
snipe has not been seen since.
Habits
Guthrie-Smith (1936) stated that the snipe was very skulking and rarely flew, but trusted to concealment, a
habit that, sadly, would not have protected it from rats. It fed on red worms, for which it probed in the soil
and compacted leaf litter, and small pupae found on the surface of the ground. Guthrie-Smith found that
the nest was a very simple affair, little more than a depression in the conifer needles that littered the ground.
Two eggs were laid, very pale brown with dark and pale brown spots or blotches, all over but more
numerous round the large end. He observed five pairs and found that only one chick seemed to be
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successfully fledged. Unlike the Chatham Island Snipe Coenocorypha pusilla, where each parent will raise
one chick from a clutch of two, both South Island Snipe parents cared for a single chick (Miskelly & de
Lange 2006).
Guthrie-Smith described the birds when feeding:
Moving with a curious halting, hesitant gait, always they advanced rocking as if balanced on springs. Usually
the male, like the country Scotsman of half a century ago kirkward bent with his womenfolk, moves a foot or two
ahead. If lost to sight, however, for the briefest period, communication is kept up betwixt the pair by a low hoarse
double croak. This is uttered from time to time as the pair or trio irregularly progress, sometimes at a walk, some-
times at a trot, but always whether slow or fast – and if necessary they can dart and disappear like lightning
– probing, probing, quickly, eagerly, decidedly. The long bill is held well forward after the manner of the Kiwi – a
Lilliputian stride or two, five or six rapid spearings into the ground, a brief hesitation, a prolonged sniff, a deeper
and more assured perforation of the spongy soil, a quick little m ouse-like run, a pause, an advance, a downward
thrust of the beak, so they moved ahead. Each minute red worm hardly thicker than a pin could upon withdrawal
of the bill always be seen dangling at its extremity ere being passed downwards and swallowed.
The Chatham Islands are the only island group on which two species of Coenocorypha snipe lived sympatri-
cally. Forbes’s Snipe was larger than the extant Chatham Island Snipe C. pusilla, with a longer bill, although
the length was variable. The terrestrial Coenocorypha snipe cannot survive alongside introduced mamma-
lian predators, especially rats, so Forbes’s Snipe was no doubt a victim of these invaders, but why it should
have disappeared whilst C. pusilla survives is something of a mystery.
Forbes’s Snipe is known only from subfossil remains, but there is a likelihood that these cryptic, nocturnal
and elusive birds survived into historic times.
Meredith (1991) produced figures of subfossil remains of a Coenocorypha snipe discovered on Norfolk Island,
but referred them to the Auckland Island Snipe C. aucklandica. The material consists of a complete humerus
and two tibiotarsi. The Norfolk Island material is slightly larger than that of the Auckland bird, but Worthy
(2001) remarked that most snipe taxa have similar-sized humeri and tibiotarsi but differ in other ways,
including skeletal element proportions, plumage and behaviour. Therefore, it is likely that this isolated
population on Norfolk Island was specifically distinct. Shortly after the arrival of Polynesians and their
commensal mammals, especially the Pacific Rat Rattus exulans, a notorious bird predator, the Norfolk Island
Snipe disappeared.
The discovery of a Coenocorypha snipe on the tropical island of Vitu Levu is surprising as this genus is
usually restricted to southern temperate latitudes (Worthy 2003). It was much larger than any of the New
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Balouet & Olson (1989) referred two partial humeri and a partial coracoid to the genus Coenocorypha,
suggesting that there was once an endemic snipe on New Caledonia. This makes sense biogeographically,
as these skulking, elusive birds are capable island colonists, reaching a number of remote archipelagos
(Worthy 2001). The New Caledonia Snipe was larger than any of the Coenocorypha from New Zealand. The
subfossil remains were found in Gilles Cave, a fossil deposit situated 7km inland on the west coast (Balouet
& Olson 1989), which suggests that the snipe was a forest inhabitant. It is not known when this bird became
extinct, but all Coenocorypha snipe are extremely vulnerable to introduced mammalian predators, especially
rats, so it is likely that this bird disappeared rapidly once these predators arrive in New Caledonia.
Wetmore (1920) originally referred subfossil remains of this species to the snipe genus Gallinago, but Olson
(1976b) re-examined the material and concluded that they actually belonged to the woodcock genus
Scolopax. The Puerto Rico Woodcock S. anthonyi was smaller than the European Woodcock S. rusticola, but
larger in the humerus and tarsometatarsus than the American Woodcock S. minor. Osteologically, the Puerto
Rican Woodcock is more similar to Old World woodcocks than to the American species, but Olson suggests
that this is not indicative of any close relationship. There was also a size difference in some of the material,
which suggests sexual dimorphism.
The wings of the Puerto Rico Woodcock were reduced compared with other Scolopax woodcocks, so
perhaps it was more terrestrial. Wetmore (1920) suggested that the fossil localities where S. anthonyi was
discovered were originally forested, which is a habitat preferred by the genus. Nothing else is known of it,
or why it became extinct.
Range Canada from Yukon, North- West Territories, Nunavut, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, New
Brunswick, Prince Edward Island, Nova Scotia, Newfoundland and Labrador, and possibly northern Alaska. Wintered in
Uruguay, Paraguay and possibly Chile and southern Brazil, south to Patagonia.
Description 29–34cm (11.5–14in). Adult; upperparts generally mottled c innamon-brown and white; crown brown; generally
distinct supercilium; dark streaking on sides of face and neck; underparts light brown, with V and Y markings on breast and
flanks; belly r usty-yellow; wing-linings cinnamon in flight; bill blackish, with base fl
eshy-pink; iris brown; legs and feet dark
greyish or dark greenish slate.
In the early 1800s, millions of Eskimo Curlews were recorded following wintering migration routes
south-eastwards from Yukon and the North-West Territories, along the northern shoreline of Canada,
staging primarily in Labrador and Newfoundland, then flying south n on-stop over the Caribbean to South
America, where they wintered on the pampas of Argentina and Uruguay. The birds return migration north
to North America followed a different route over the Great Plains, through Texas and the m id-western
states (Environment Canada 2007). As befits a l ong-distance migrant, vagrants would occasionally end up a
long way off course, and there are records of Eskimo Curlews pitching up as far away as the British Isles.
As late as the 1860s, hundreds of thousands if not millions of birds were killed each year for the pot,
especially as the Passenger Pigeon Ectopistes migratorius (see p. 166) was by this time in serious decline and
hunters had turned their attention to other game, but by the 1890s curlew numbers had collapsed (Gill et al.
1998). The reasons for its disappearance in Canada and the United States include uncontrolled market
hunting (Gollop 1988), the almost complete destruction of prairies for agricultural development and the
loss of favoured burnt areas after the suppression of wildfires (Gill et al. 1998). The bird’s habit of occurring
in immense flocks, its lack of fear of humans, and their tendency to circle within gunshot range after
members of the flock were shot made them easy targets for hunters ( Gratto-Trevor 1999). The extinction of
the Rocky Mountain Grasshopper Melanoplus spretus in the 1900s, a primary food source on the return
migration (Gill et al. 1998), may have exacerbated the problem.
The Eskimo Curlew was seen only sporadically during the 20th century. It was photographed by Don
Bleitz on Galveston Island, Texas, in 1962, and a specimen obtained on Bermuda in 1963 (Bond 1965) is the
last known unequivocal record. Sightings of birds continue to be reported, including a generally accepted
account of 23 birds seen in 1981 in Texas, and there have been unconfirmed reports up to 2006. The bird has
not been recorded on its South American wintering grounds since 1939 (Environment Canada 2007). If it
still survives its numbers are extremely small.
Eskimo Curlew
Numenius borealis
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Habits
The Eskimo Curlew utilised a range of open habitats, preferring treeless tundra with dwarf vegetation,
grassy meadows and shorelines on the breeding grounds, whereas during the autumn migration they were
found on heathland, meadows, mudflats and salt marshes. In spring on migration north they were associ-
ated with m ixed-grass prairies, often in recently burned areas or those disturbed by grazing Bison Bison
bison (Gill et al. 1998). The birds were long-lived and monogamous, with a low reproductive rate.
Spring migration occurred in April to May, with birds arriving in the breeding areas in late May to June.
Nesting is thought to have occurred from June to August. However, the exact breeding range of the Eskimo
Curlew is open to debate. Only two confirmed areas are known, both in the North-West Territories of
Canada, at the base of Bathurst Peninsula and near Point Lake ( Gratto-Trevor 1999). Breeding is presumed
to have occurred between these two areas and also elsewhere (Gallop et al. 1986), but this has never been
confirmed. The nests were constructed from dried leaves or grass; four eggs were normally laid
(Environment Canada 2007), but no evidence of nesting has been recorded since 1866. The eggs were green
with brown blotches. The autumn migration began in late July through to October. In Argentina, the
wintering grounds were treeless grasslands interspersed with wetlands (COSEWIC 2009), with dry and wet
fields used in Paraguay (Gollop et al. 1986).
Eskimo Curlews fed by probing the ground with their long bills. They fed primarily on Crowberries
Empetrum nigrum during the autumn migration in Canada, and insects, snails and other invertebrates
during the rest of their migration (Gill et al. 1998). The call is poorly known, but reportedly included a
rippling tr–tr–tr, a soft whistle bee bee, and also clear whistling sounds.
voyage of Captain Cook in 1773, when the unique surviving skin specimen was taken (Zusi & Jehl 1970). It
was also illustrated by Forster’s son, Georg. The reasons for its extinction are unclear, but Cook had released
goats and pigs on the island, and these may have subsequently destroyed the sandpiper’s habitat. Greenway
(1967) thought that rats were more likely to have been the cause of its disappearance.
Two more individuals (now lost) were collected on nearby Moorea between September 30 and October
11, 1777 by William Anderson, the surgeon’s chief mate on Cook’s second and third voyages (Iredale 1938;
Stresemann 1950; Lysaght 1959). At least one of these birds was illustrated and, based on rather ill-defined
plumage differences, was described as a different species, P. ellisi (Sharpe 1906b; see below).
Habits
Virtually nothing was recorded about the W
hite-winged Sandpiper’s habits. Forster (in Lichtenstein 1844)
recorded that they occurred near small brooks, which suggests that they lived inland rather than on the
coasts.
This second species of Society Island sandpiper was decribed by Sharpe (1906b), based on an unpublished
plate by William Ellis in the Natural History Museum, London. Another illustration is attributed to John
Webber. There has been some doubt as to the validity of it and the supposed differences between it and the
Tahiti Sandpiper P. leucoptera (see p. 150) may be due to individual variation (Zusi & Jehl 1970). Walters
(1991) argues that in terms of biogeography and differences in plumage, P. ellisi warranted specific status,
but in the absence of any physical remains, the validity of P. ellisi of Moorea remains in doubt. The popula-
tion on Moorea, whatever its status, disappeared for the same reasons as those that beset the Tahiti birds – a
combination of habitat destruction and predation by introduced animals.
The type and only specimen of P. cancellata was observed and probably collected by William Anderson on
Captain Cook’s last voyage, around January 1778 (Stresemann 1950), on Christmas Island (now Kiritimati).
It may have occurred on other islands in the Kiribati Group. There is a painting of it by William Ellis, who
was assistant surgeon during the voyage. On the return of the fleet in 1780, the specimen passed into the
Banksian Collection, where Latham (1785) saw and described it:
7 inches and a half. Bill one inch, black: the feathers on the upper parts of the bird brown, edged with white:
under parts white, transversely barred with dusky: quills dusky, with the ends brown, and the margins and tips
very pale: tail the same, spotted on both webs with white: legs dusky. Inhabits Christmas Island. In the collection
of Sir Joseph Banks.
A number of authorities have synonymised the Kiritimati Sandpiper with the Tuamotu Sandpiper, but
work by Pierce & Blanvillain (2004) has shown that the two species are distinct, which makes sense on
zoogeographic grounds (Kiritimati is some 3,200km to the east of the Tuamotus).
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The Tuamotu Sandpiper is considered endangered with a declining population, mainly because of
predation by introduced rats (Pierce & Blanvillain 2004; BirdLife International 2011). The Kiritimati
Sandpiper was also susceptible to introduced predatory mammals, especially cats which had overrun the
island, and became extinct sometime around the 1850s (BirdLife International 2011).
Laridae (Gulls)
The gulls have a widespread distribution and are most often associated with coastal regions and
human-modified habitat. The taxonomy is confusing, with up to 11 genera being generally accepted. Some
species are restricted to isolated lake systems far from the coasts, or islands and island groups, where they
are vulnerable to habitat destruction.
Alcidae (Auks)
Auks are the Northern Hemisphere equivalent of penguins; the family is divided into 11 genera. Unlike
penguins, auks are volant birds (the only modern exception being the Great Auk Pinguinus impennis), and
spend most of their lives at sea, returning to land only to breed. Only the flightless Great Auk has become
extinct, but some species are in serious decline.
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The Great Auk was the largest and most penguin-like of all the auks; indeed, the name ‘penguin’ was origi-
nally applied to the Great Auk, before it became transferred to the Southern Hemisphere birds that we
recognise as ‘penguins’ today. The Great Auk was flightless and clumsy on land, but perfectly adapted for
life at sea, in which it used its wings to propel itself through the water with great speed and agility. DNA
analysis has shown that the extant Razorbill Alca torda is the Great Auk’s closest living relative, and both
have a sister group relationship with the Little Auk Alle alle (Moum et al. 2002).
Range
Much has been written about the extinction of Great Auk, but rather little is known about the bird itself (see
Fuller 1999). It once had a wide range in the North Atlantic, occurring along the coasts of Canada, the
north-eastern United States, Scandinavia, the west coast of Greenland, Iceland, the British Isles, France and
northern Spain, with stragglers, or more likely traded birds, reported in Florida (Montevecchi & Kirk 1996).
It also occurred in Sicily in the Mediterranean, as there are reports of the birds occupying Marausa Bay
(Crofford 1989). Remains have been found in archaeological sites in Gibraltar, in Pleistocene deposits in
Otranto on the s outh-eastern tip of Italy, and in Jersey, in the Channel Islands (Greenway 1967). The breeding
colonies appear to have been restricted to specific sites, but the known main Great Auk rookeries in rela-
tively recent times were Bird Rocks in the Gulf of St Lawrence, Funk Island off Newfoundland (which had
the largest colony), the Faroe Islands, Geirfuglasker and Eldey Island near Iceland, Papa Westray in the
Orkneys, and St Kilda Island off Scotland (Montevecchi & Kirk 1996; Fuller 1999; Cokinos 2000).
The Great Auk was hunted all over its range for centuries, as a food item and for its thick down. It was
only vulnerable when it came to shore to breed, as it was extremely difficult to catch when at sea (Greenway
1967). The populations were smaller in the eastern Atlantic, which was probably due to a longer period of
human exploitation, but it was still comparatively numerous in the more inaccessible breeding colonies up
to the 16th century, in particular on Funk Island, where it may have numbered 10,000 pairs when first
discovered by Europeans (Nettleship & Birkhead 1985). However, by the mid-16th century there had been a
major decrease in numbers, especially on the European side of the Atlantic. Concerns over its plight resulted
in the first official protection of the species in 1753, when Newfoundland unsuccessfully petitioned Britain
to stop the slaughter of sea birds. Penalties were put in place for killing adults or stealing eggs, but taking
birds for fish bait was still permitted. Britain finally imposed a ban on killing the birds in 1794 (Montevecchi
& Kirk 1996), but by this time the Great Auk was no longer a breeding species in Newfoundland. The vast
breeding population that once occurred on Funk Island was hunted to extinction, with the last birds disap-
pearing between 1785 and 1800 (Montevecchi & Kirk 1996).
The slaughter of the birds remained unabated, and by the turn of the 19th century a combination of
wasteful e gg-harvesting, when eggers only took eggs without embryos, discarding eggs with embryos
(Crofford 1989), and killing for food and for the skins, reduced the population to a few hundred. Only a few
reports are known after this time. Two birds were taken in the Orkneys in 1812, a unique winter-plumage
bird was taken in 1815 near Fiskaernes, Greenland, a bird one was captured on St Kilda in 1821, and one
was one taken at Waterford, on the south coast of Ireland in May 1834 (Greenway 1967). The last authentic
record for Great Britain was one taken in July 1840 on St. Kilda (Fuller 1999).
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Last stronghold
The Geirfuglasker, a volcanic island off the s outh-west coast of Iceland, was the last breeding stronghold. It
was generally inaccessible to humans, but even here auks were persecuted when possible, and the crew of
the Salamine, a privateer under British colours, spent a day in 1808 killing many birds and treading down
their eggs and young (Newton 1861). The Geirfuglasker became submerged after volcanic eruptions in 1830
(Fuller 1999), but approximately 50 individuals survived this natural disaster, and the birds continued to
breed on the nearby island of Eldey. Unlike the Geirfuglasker, Eldey was accessible on one side. Due to their
great rarity, the Great Auk was in high demand from collectors and institutions around the world. The
surviving auks on Eldey were much sought after, and birds were regularly collected (Crofford 1989; Fuller
1999).
On June 3, 1844, a group of fishermen, who had been commissioned to collect specimens, landed on
Eldey. They came across a breeding pair, with the female sitting on a single egg. The last of the Great Auks
were to meet a brutal end. Both birds were strangled and sold as museum skins and their carcasses
preserved in spirit, while the egg was apparently smashed (Newton 1865; Grieve 1885; Fuller 1999). This
has generally been accepted as the last definite record, but an individual sighted in December 1852 on the
Grand Banks of Newfoundland (Newton 1861) has now been accepted as authentic (BirdLife International
2011). The Great Auk was never seen again.
Habits
Little is known with certainty about the Great Auk’s habits, and much of the available data are based on
analogies with the Razorbill. They were colonial breeders, forming vast colonies, and it appears that the
breeding season began in May, and ended in mid-July (Greenway 1967). Only one egg was laid. Incubation
was undertaken by both sexes, with the combined incubation and fledging period lasting between 43 and
53 days; the fledging period of 10 days or so was extremely short (see Bengtson 1984). Great Auks spent the
rest of the year at sea, migrating south during the autumn and winter months.
The Great Auk was an adept swimmer and diver, and could roam up to 500km from the nearest shore,
reaching depths of up to 76m in search of food (Montevecchi & Kirk 1996). They may have fed in flocks,
probably on a diet exclusively of fish, particularly Menhaden Brevootia sp. and Capelin Mallotus villosus
(Olson et al. 1979), but the young may have been fed on plankton and crustaceans (Montevecchi & Kirk
Great Auk
Pinguinus impennis
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1996). The natural predators of the birds were the Polar Bear Ursus maritimus, Killer Whale Orcinus orca, and
White-tailed Eagle Haliaeetus albicilla (Crofford 1989; Montevecchi & Kirk 1996). The calls were similar to the
Razorbill only louder and deeper in tone, and included low croaking, gurgling noises and a hoarse scream
(Montevecchi & Kirk 1996).
Alfred Newton (1861), who along with his friend, John Wolley, had obtained information from local
Icelandic fisherman in 1860, perhaps left the best summary of its habits:
In former days, the G are-fowls were, in summer time, so constantly observed in the sea by the fishermen that
their appearance was thought little of . . . We were told by many people that they swam with their heads much
lifted up, but their necks drawn in; they never tried to flap along the water, but dived as soon as alarmed. On the
rocks they sat more upright than either Guillemots or R azor-bills, and their station was further removed from
the sea. They were easily frightened by noise, but not by what they saw. They sometimes uttered a few low
croaks. They have never been known to defend their eggs, but would bite fiercely if they had the chance when
caught. They walk or run with little, short steps, and go straight like a man. One has been known to drop two
fathoms off the rock into the water.
More has been written about the Dodo Raphus cucullatus of Mauritius than any other extinct bird species
(Fuller 2002; Hume 2006). But in reality, virtually nothing is known about it in life. Early Arab traders prob-
ably discovered Mauritius in the 13th century, followed by the Portuguese in 1516 (North-Coombes 1980),
but as far as is known, they never settled there and no written accounts exist. It was a Dutch VOC (Dutch
East India Company) fleet under V ice-
Admiral Wybrandt Warwijck that in September 1598 claimed
Mauritius for the Netherlands (Barnwell 1948; Moree 1998), at which point the documented history of the
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island began. This claim has often been attributed to Admiral Jacob van Neck, but the fleet was separated in
a storm around the Bay of Good Hope, and van Neck proceeded directly to the East Indies.
For the next 40 years, the island was regularly used as a refurbishment station for ships travelling to and
from the East Indies, but no permanent settlement was made. Due to the increasing presence of rival English
and French traders, the Dutch established a fort at Vieux Grand Port, south-east Mauritius in 1638, and
maintained an almost continuous settlement until 1710, after which time they abandoned the island (Moree
1998). It was during the Dutch period that the Dodo and most other large, terrestrial vertebrates became
extinct (Cheke & Hume 2008).
Early accounts
Van Neck returned to the Netherlands in 1599 (A True Report 1599), and the first account about the Dodo
was published:
There are also other birds there which are as big as our swan, with large heads, and on the head a veil as though
they had a small hood on their head; they have no wings but in their place there are three or four black quills, and
where there ought to be a tail; there are four or five small curled plumes of a greyish colour. We called these birds
Walghvogels [nauseous bird], partly because although we stewed them for a very long time, they were very
tough to eat, yet the stomach and the breast were extremely good, but also because we could get a profusion of the
turtledoves, which we thought had a rather better taste.
The journal was expanded in 1600 and 1601 (Het Tweede Boeck 1601) as more information from returning
mariners became available, and these editions illustrated the Dodo for the first time. Plagiarism was
standard practice, so the account of van Neck, although already second-hand, has been repeated more than
any other (Hume 2006). A few accounts are novel and genuine however. An anonymous Dutch sailor in
1631 (Servaas 1887; Winters et al. 2017) was the only person to mention the dodo’s diet:
These mayors [dodos] are superb and proud they displayed themselves to us with a stiff and stern face and wide
open mouth, very jaunty and audacious of gait, and would scarcely move a foot before us, their war weapon was
the mouth, with which they could bite fiercely, their food was raw fruit, they were also w
ell-adorned, but were
abundantly covered with fat, and so many of them were brought aboard, to the delight of us all.
The accounts of Herbert (1634), Mundy (in Temple 1914) plus other brief accounts (see Fuller 2002; Hume
2006) all sum up the presence of the Dodo on Mauritius until perhaps the 1640s, but for the next two decades
they were no longer mentioned. The Dutch abandoned the island between 1658 and 1664, and it was in
1662, after the loss of the ship Arnhem, that surviving mariners made landfall and lived on an almost starva-
tion diet until rescued. One party led by Volkert Evertszen waded across to an islet, whose location is still
disputed (Cheke & Hume 2008), and discovered a small population of Dodos, but even this account is open
to interpretation (see Fuller 2002).
The Dutch reoccupied Mauritius in 1664 and two commanders who were stationed there (Hubert Hugo
from 1673–1677, and Isaac Lamotius from 1677–1692) both list Dodos as being extant (Moree 1998; Hume et
al. 2004). However it has been suggested that a name change had occurred, and that their mention of Dodos
was in fact referable to the Mauritius Red Rail Aphanapteryx bonasia (Cheke 2006; see p. 126). Whoever actu-
ally saw the last Dodo, be it in 1662 or 1693, is of no real consequence, as by this time the bird was all but
extinct in any case. Direct hunting by humans has long been cited as the cause of the dodo’s extinction, but
this was almost certainly not the primary cause. The 1 7th-century human population on Mauritius was
small, never more than 100 people, occupying an island 61km long by 47km wide, and covering an area of
1,860km2 (Hume & Winters 2015). However, the introduction of Black Rats Rattus rattus, pigs, goats and
perhaps monkeys, all which would have been direct threats to eggs and chicks, and competitors for limited
food resources, are the likely culprits in the Dodo’s demise.
Exports
A very few Dodo physical remains, either live or dead, left Mauritius for Europe, India and possibly Japan,
but again details are contradictory. A foot was recorded by Clusius (1605), while another remained in the
collection of the Royal Society until at least the 1870s (Hume et al. 2006); a dessicated head (skin now
perished) is in Copenhagen. Herbert (1634) and Mundy (in Temple 1914) both saw live, captive Dodos in
the menagerie of the Mogul Emperor, Jahangir, in Surat, India, and a live specimen was exhibited in London
in 1638 (Strickland & Melville 1848). Tradescant (1656) mentions a complete bird, perhaps the same
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Affinities
The affinities of the Dodo have been investigated by a number of authors. Over the years, it has been consid-
ered a member of a large array of bird orders; the Dodo has been thought to be a diminutive ostrich, a rail,
and even a vulture. After examining a skull in Copenhagen, Professor J. T. Reinhardt proposed that the
Dodo was related to pigeons and doves (Reinhardt 1842). This claim was initially met with ridicule, but
after Strickland & Melville (1848) confirmed his theory after examining the Oxford Dodo head, the idea
became universally accepted (see Owen 1866a; Nowak-Kemp & Hume 2016b). DNA studies have now
concluded that the Dodo and closely related Rodrigues Solitaire Pezophaps solitaria (see p. 158) are a sister
clade, nested within the Columbidae and derived from the same common ancestor as the south-east Asian
Nicobar Pigeon Caloenas nicobarica (Shapiro et al. 2002; Peirara et al. 2007; Soares et al. 2016).
157
Although differing in their molecular clock dating, two previous studies suggested that the common
ancestor of the Dodo and Rodrigues Solitaire diverged from Caloenas around 42.6 mya (million years ago)
(Shapiro et al. 2002) or 33.6 mya (Peirara et al. 2007), and the dodo and solitaire lineages diverged from each
other 18–36 mya (Shapiro et al. 2002), or 15–30 mya (Peirara et al. 2007), in both cases long before Mauritius
and Rodrigues had emerged above the sea. A more recent study suggests this event took place more recently,
with the common ancestor of dodo/solitaire and Caloenas diverging around 18 mya, and the dodo/solitaire
divergence at around 13.1 mya (Soares et al. 2016). This strongly suggests that the volant common ancestor
of the Dodo and Rodrigues Solitaire island-hopped its way to Mauritius and Rodrigues from south-east
Asia over millions of years, and before the emergence of the Mascarene Islands, using now-submerged
oceanic islands that were exposed during periods of low sea levels. The islands would not only have been
more numerous en route, but would also have shortened the distance between land masses when crossing
open water (Hume 2011b).
Habits
Nothing can be stated with any certainty about the Dodo’s habits. In Clusius (1605), a large gizzard stone is
illustrated alongside the Dodo, and they were reported by early mariners as excellent for sharpening tools.
This suggests that hard seeds were an important part of the diet. A study using CT scanning of the Dodo
and Rodrigues Solitaire’s braincases showed that these birds had a brain completely in proportion to their
body size, and that they had a similar intelligence level to modern pigeons (Gold et al. 2016). Both birds had
large and well-differentiated olfactory bulbs, rather than larger optic lobes, which suggests that these
ground-dwellers relied on smell rather than sight to detect food. This included fallen fruit, land snails, and
perhaps marine invertebrates when foraging on the shorelines.
Cauche (1651) said that the Dodo laid a single white egg on a mass of grass in the forest, and the juvenile
call was like that of a young goose. However, he did not actually land on Mauritius and appears to have
confused his description with that of a Cassowary Casuarius sp. (Cheke & Hume 2008). Recent work at the
Mare aux Songes (de Boer et al. 2013; Rijkdijk et al. 2009, 2011, 2015) has shown that the Dodo occurred in
forest dominated by Tambalacoque Sideroxylon grandiflorum, Pandanus sp., and endemic palms.
Fat or thin?
The Dodo may also have exhibited seasonal fat cycles based on the austral winter and summer, the lean
form occurring during the non-breeding, hot rainy season from October to March, and the fat form from
March to September, the cool, dry season, but this is purely conjecture. Kitchener (1993) analysed biometrics
and osteology in order to reconstruct Dodo body mass, and concluded that in nature it was a relatively
slender bird, but still among the heaviest non-ratite birds known today.
Angst et al. (2011a) concluded that previous estimates of the Dodo’s body mass were too high, based on
the combined lengths of the femur, tibiotarsus, and tarsometatarsus. However, their methodology has been
questioned and considered unreliable (Louchart & M ourer-Chauviré 2011), which resulted in a highly
publicised counter-argument (Angst et al. 2011b). The issue of Dodo body mass still remains unresolved,
and some 300 years after its extinction, this emblematic bird continues to prompt heated debate.
The Solitaire or Solitary of Rodrigues was the closest relative of the Dodo Raphus cucullatus It was endemic
to Rodrigues Island, situated 650km to the south-east of Mauritius. Both species (often called didines)
derived from the same common ancestor, an ancestral pigeon closely related to the extant Nicobar Pigeon
Caloenas nicobarica (Shapiro et al. 2002). Each island provided different ecological constraints, and as a result
the morphology of the two species differed.
Ironically, the ecology of the more famous Dodo is virtually unknown, yet the same cannot be said about
the Solitaire. Observations on the bird made by François Leguat (1708) provide such detailed insights that,
remarkably, he inadvertently became the first person to describe territorial and sexual behaviour in birds
(Armstrong 1953; Livezey 1993). Leguat and his comrades remained on Rodrigues for two years from 1691–
93, finally making their way back to Europe after eight years. Leguat moved to London and was encouraged
by Sir Hans Sloane, founder of the British Museum, and others to write an account of his journey
(North-Coombes 1971). It is through Leguat’s work (see below) that we know so much about the biodiver-
sity of an oceanic island prior to human interference.
Tafforet
Julien Tafforet was a ship’s mate from Réunion, and was sent to Rodrigues to ascertain the suitability of the
island for settlement (North-Coombes 1971). He took a copy of Leguat’s book and, like Leguat, was
marooned on Rodrigues rather longer than he had hoped for. A storm blew up, and the waiting ship was
forced to return to Réunion, leaving Tafforet and six men onshore for 9 months in 1725–26. Tafforet wrote a
thorough account about the island and its fauna and flora. The document was only discovered in the latter
part of the 19th century ( Milne-Edwards 1874), and his description of the Solitaire adds more to our knowl-
edge of the bird:
The solitaire is a large bird, which weighs about forty or fifty pounds. They have a very big head, with a sort of
frontlet, as if of black velvet. Their feathers are neither feathers nor fur; they are of a light grey colour, with a little
black on their backs. Strutting proudly about, either alone or in pairs, they preen their plumage or fur with their
beak and keep themselves very clean. They have their toes furnished with very hard scales, and run with quick-
ness, mostly among the rocks, where a man, however agile, can hardly catch them. They have a very short beak,
of about an inch in length, which is sharp. They nevertheless do not attempt to hurt anyone, except when they
find someone before them, and when hardly pressed try to bite him. They have a small stump of a wing which has
a sort of bullet at its extremity, and serves as a defence. They do not fly at all, having no feathers to their wings
but they flap them and make a great noise with their wings when angry, and the noise is something like thunder
in the distance. They only lay, as I am led to suppose, but once in the year, and only one egg. Not that I have seen
their eggs, for I have not been able to discover where they lay. But I have never seen but one little one alone with
them, and if any one tried to approach it, they would bite him severely. These birds live on seeds and leaves of
trees, which they pick up on the ground. They have a gizzard larger than the fist, and what is surprising is that
there is found in it a stone of the size of a hen’s egg, of oval shape, a little flattened, although this animal cannot
swallow anything larger than a small cherry-stone. I have eaten them; they are tolerably well tasted.
Tafforet eventually returned to Réunion in 1726, and his report signalled the end for the Solitaire. It was still
considered common in 1733 (Gennes 1735), but must have disappeared rapidly after. Tafforet and Leguat
reported the huge numbers of giant tortoises on Rodrigues, perhaps the densest populations anywhere in
the world. This attracted much attention from passing ships, and a tortoise run was set up to provision
Réunion and the hospital on Mauritius with fresh meat (Cheke & Hume 2008). Goats were introduced some
time after 1726, as were cats, and the tortoise hunters indiscriminately harvested the animals; such was the
massacre of tortoises that the governor of Mauritius, Mahé de Labourdonnais, ordered controlled hunting.
This was impossible to enforce, and by time Pingré (1763) visited Rodrigues to observe the Transit of Venus
in 1761, most of the birds had already gone, or very nearly so. He inquired about the Solitaire and was told
that they still survived in remote places, but this was unlikely.
Subfossil material
The first subfossil bones of the Solitaire were discovered in 1786 in a cave deposit by Labistour, a resident on
the island (Hume et al. 2014). More bones were found in the same cave, which provided the material for
Strickland & Melville’s (1848) monograph on the didine birds of the Mascarenes. In 1864, Sir Edward
Newton obtained two bones from another cave and was given another one by a Captain Barclay (Newton
1865). Newton was sufficiently interested to persuade the resident magistrate on the island, Mr. Jenner, to
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make further searches, and the following year, 85 more specimens were obtained. This encouraged Newton
to fund an excavation, obtaining monies from the British Association (Hume et al. 2009), which resulted in a
haul of 2,000 Solitaire bones (Hume et al. 2014). Henry Slater obtained nearly as many bones as all of his
predecessors on the 1874 Transit of Venus expedition (Slater 1879b). Leguat’s account of the Solitaire was
regarded by some as being pure fiction (e.g. Atkinson 1922), but analysis of this material confirmed Leguat’s
testimony (North-Coombes 1991), and his observations can be correlated with characters of the anatomy
(Newton & Newton 1870).
Habits
Leguat (1708) recorded that the nest consisted of a pile of palm leaves on the ground, and in this a single egg
was laid. The pair would not permit any other individual within approximately 200 yards of the nest. The
territory was aggressively defended, the male attacking intruding males, and the female repelling intruding
females. If an intruding female appeared, the male would not attack, but would call his mate to deal with
her. This level of detailed observation of living animals was exceptional for this time. On the radius, ulna
and carpometacarpus (wrist), the Solitaire had a bony growth (‘musket ball’), and for periods of four or five
minutes at a time the bird would perform twenty or thirty rapid pirouettes, whirling its wings with a
rattling noise which could be heard about 200 yards away. This distance was probably the radius of the
territory. Many Solitaire pectoral bones have healed fractures, which not only indicate the extreme violence
used, but also the use of their wings as weapons, in a way similar to extant pigeons. Leguat indicated that
the wing-clattering was used to attract females as well as to defend territory. This defence of the territory
continued through incubation and until the young birds became independent. The incubation period was
seven weeks, and the chick was nurtured for several months.
A recent study by Hume & Steel (2013) has shown that the musket balls were present in both sexes, and
always initially developed from the processus extensorius of the carpometacarpus. The structure may have
grown in order to protect the bone at the site of impacts sustained when combat behaviour began in adult
life, creating a more effective weapon favoured by natural selection, or it may have developed under the
influence of hormones. Either way, once a territory was established there were frequent fights. The musket
ball continued to grow throughout adult life, with the oldest males and females exhibiting the greatest
development. The bony growth extended to the distal end of the radius and proximal wing phalanx, and
the structure would have been covered in a thick skin, making it even larger. The audible sound described
by Leguat (1708) and Tafforet (1725–26) was produced by either knocking the wings together or hitting the
sides of the body. No other bird known has evolved this particular type of sound or weapon.
Leguat’s remarkable observations show that the Solitaire had a complex social life. The diet comprised
fruit of Latan Palm Latania verschaffeltii, Screw Pine Pandanus heterocarpus, seeds and fallen leaves. Like the
Dodo, the Solitaire had a large, single gizzard stone, which was used to grind the hard seeds in its diet. After
the breeding season, it seems likely that the territory was temporarily abandoned, the chicks formed crêches
and during this period pairing occurred, which suggested that the Solitaire was strictly monogamous
(Armstrong 1953). However, Newton (1865) thought that in view of the size difference between the sexes
the bird was probably polygamous. Armstrong (1953) however pointed out that size sexual dimorphism
can occur without polygamy (e.g. in steamer ducks Tachyeres sp.), and that Leguat quite clearly indicated
that the Solitaire was monogamous, as is the situation with most pigeons and doves.
Although not officially described until 1904, Harcourt (1851) had noticed that the wood pigeon on Madeira
was larger than the nominate C. p. palumbus. In 1886, Hartwig (in Schmitz 1899) found the Madeira Wood
Pigeon nesting in the mountain woods, and considered it to be rare at that time. Since 1904, all attempts to
find it have failed, and there has been only one alleged sighting; this seems to have been a case of
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misidentification of the rare Trocaz Pigeon Columba trocaz, a closely related species. The reason for its extinc-
tion is unknown.
Habits
Little was recorded about the Madeira Wood Pigeon. It occurred in forests in the north of the island, and it
apparently bred year-round; a clutch of two eggs was recorded (Clarke 2006).
The Lord Howe Pigeon is known from three paintings; the first by George Raper of HMS Sirius (Hindwood
1964), painted in about 1790, another, by an anonymous artist and probably a copy of that by Raper, and a
third by John Hunter, also from around 1790. There are also a number of accounts. When Lord Howe Island
was first discovered, pigeons were numerous and ridiculously tame. They could be caught in the hand, and
made no attempt to escape when slaughtered. Thomas Gilbert (1789) wrote of them in 1780:
On entering the woods I was surprised to see large fat pigeons . . . sitting on low bushes and so insensible to fear,
as to be knocked down with little trouble.
Despite the images and numerous accounts, doubt had been expressed as to whether Raper’s painting was
actually of a bird from Lord Howe Island, or was actually a Columba vitiensis subspecies from another
locality (Mayr 1941a). Peters (1937 [1931–48]) placed godmanae in the synonymy of C .v. halmaheira (a race
occurring from the Moluccas through New Guinea to the Solomon Islands), believing the locality of Lord
Howe Island to be an error. Nevertheless, there is ample evidence that a large pigeon did occur on Lord
Howe Island, though, as Mayr observed, it might have been a Ducula rather than a member of the Columba
vitiensis group.
In 1788, ship’s surgeon Arthur Bowes (in Hindwood 1940) described his encounter with the birds:
The pigeons also were as tame as those previously described [Lord Howe Woodrail Tricholimnas sylvestris
and Lord Howe Swamphen Porphyrio albus] and would sit upon the branches of the trees till you might go
and take them off with your hands or if the branch was so high on what they sat, they would at all times sit till
you might knock them down with a short stick, many hundreds of all the sorts mentioned above, together with
Parrots and Parakeets, Magpies and other birds were caught and carried on board our ship and the Charlotte.
Like most pigeons, they were apparently good to eat and easily caught using sticks, stones, nooses or guns
(Higgins & Davies 1996). When the island was settled in 1834 they were doomed, despite there being only
16 residents. The pigeons lingered until at least until 1853, at which time they were becoming scarce, as
MacDonald (1853) noted:
. . . a very beautiful pigeon is occasionally to be seen on the flats, with metallic green tinting on the wings and
whitish-brown breast, deepening towards the neck.
The botanist John MacGillivray, who collected there later in the same year, was unable to find any speci-
mens (Hindwood 1938). Nothing was recorded about their habits, but they probably occurred throughout
the lowland forests, and foraged in trees and on the forest floor (Hutton 1991).
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Stejneger’s (1887a) original description represents merely a brief diagnosis, and little is known about this
pigeon or the background to its comparatively recent discovery. The type was collected by Mr. C. Tasaki,
one of Henry Pryer’s collectors. Pryer spent some years in Japan, and also visited the Bonin and Ryukyu
Islands and discovered several other new species in the area (Seebohm 1890b). He was apparently the first
to explore the Ryukyu Islands ornithologically, and both Seebohm (1887b, 1888) and Stejneger (1887c)
almost simultaneously wrote on the avifauna as a result of his researches, although Seebohm did not at first
realise that the Ryukyu Pigeon differed from the commoner Japanese Wood Pigeon Columba janthina, which
also occurs on the islands.
Kuroda (1925) regarded this bird as a race of Japanese Wood Pigeon, but at the time of writing did not
regard it as rare. The reason for the extinction of the Ryukyu Pigeon is unknown, though it may have been
due to destruction of its native forests; almost nothing was recorded about the bird’s habits, except that is
was a bird of dense, subtropical woodland. The Daito Islands were completely deforested prior to World
War II, and the species does not appear to have been present when Allied invasion forces landed on
Okinawa in April, 1945 (Baker 1948). According to McCune (1975):
A wide variety of tropical deciduous and evergreen trees grew on the Ryukyus in the past, but unfortunately
almost all the forests have been cut over at least once, and replanting has been sporadic.
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The Bonin Islands were visited in 1827 by Captain Beechey and the naturalists of the HMS Blossom (Vigors
1839). They recorded the first Bonin Islands Pigeon on Peel Island (now Chichijima), but due to a delay in
completing the section on Mollusca, the results were not published until 1839, when Vigors named the bird
Columba metallica. In the meantime, Kittlitz (1832 [1832–33]) found it on Peel Island a year later, when he
called there on his round-the-world voyage. Therefore his name, C. versicolor, takes priority.
There was no further zoological investigation of the islands for 60 years, and the last specimen was
collected on September 15, 1889, on Nakōdo-jima, by P. A. Holst (Seebohm 1890a). The cause of extinction is
unknown, but like the Ryukyu Pigeon, it may have been due to destruction of the forests. A number of other
pigeons of the Izu Islands, the Ryukyu Islands and Okinawa have become very rare for the same reason
(Gibbs et al. 2001).
Subfossil remains have revealed that a small species of pigeon referable to the genus Columba once existed
on Mauritius (Hume 2011). The Mauritius Wood Pigeon was briefly mentioned in some early accounts,
which generally only describe the ease with which it could be caught. It had a short, robust tarsometatarsus,
indicative of an arboreal species, so it probably fed on seeds and fruit in the forest canopy. It probably disap-
peared around 1730 as a result of over-hunting, predation from Black Rats Rattus rattus and severe
deforestation. Nothing else is known about this bird.
The discovery of subfossil remains of a large pigeon related to the Mauritius Pink Pigeon Nesoenas mayeri
confirmed the status of the Réunion Pink Pigeon (Mourer-Chauviré et al. 1999), which hitherto was known
only from Dubois’s rather vague account (Dubois 1674). When staying on Réunion in 1671–72, he made the
following remark:
. . . others [pigeons] of a russet red colour, a little larger than European pigeons, with the beak larger, red at base
near the head, the eyes surrounded by a fiery colour, as in the pheasants. At a certain season they are so fat that
one does not see the uropigium (croupion); they taste very good.
Mourer-Chauviré et al. (1999) undertook a morphometric analysis of the subfossil remains, and showed
that the Réunion species was larger than the Mauritian form, but how else it might have differed is now
impossible to say. Pigeon and doves were mentioned by various visitors to Réunion (Cheke & Hume 2008),
but most leave very few diagnostic details. The last reports stem from 1704 (Feuilley 1705), so the Réunion
Pink Pigeon had disappeared within 30 years, despite being considered common in the late 1600s. The
arrival of the Black Rat Rattus rattus in the early 1670s, which had reached plague proportions by 1676,
followed by cats (Hume 2011a), appears to have been the major factor in their rapid disappearance, as defor-
estation was negligible at this time.
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Subfossil remains of a small pigeon, with comparatively reduced wing elements but long and robust legs,
were discovered in caves on Rodrigues along with the Solitaire Pezophaps solitaria. They were described by
Milne-Edwards (1874) as belonging to a small pigeon with a sternum that is unique amongst the
Columbidae.
Leguat (1708) made the only detailed description of the bird in life:
The pigeons here are somewhat less [i.e. in size] than ours, and all of a Slate colour, fat and good [to eat]. They
perch and build their Nests upon Trees; they are easily taken, being so Tame, that we have had fifty about our
Table to Pick up the Melon-Seeds which we threw them, and they lik’d mightily. We took them when we pleas’d,
and ty’d little Rags to their thighs of several Colours, that we might know them again if we let them loose. They
never miss’d attending us at our Meals, and we call’d them our Chickens. They never built their Nests in the
Isle, but in the little Islets that are near it. We suppos’d ’twas to avoid the persecution of the Rats, of which there
are vast Numbers in this Island.
Originally, Milne-Edwards (1874) had found what he believed to be two separate species, the second refer-
able to the Madagascar Turtle Dove Nesoenas picturata. A few years later, a second series of bones were
found, and Günther & Newton (1879) commented on these as follows:
A metatarsus seemed to [Milne-Edwards] to afford evidence of a second species, which he believed to be Turtur
[Nesoenas] picturatus of Madagascar, Réunion and Mauritius. We have before us two metatarsi, not differing
from that described by M. Milne-Edwards, and do not see the reason why they should be separated from C.
rodericana, to which, evidently, all the other bones of our series belong.
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Shelley (1883) referred these subfossil remains to Alectroenas (as have many authors), but this is certainly not
the case, as other subfossil remains have now been found confirming the presence of Alectroenas on
Rodrigues (see p. 184).
Leguat (1708) and Tafforet (1726) both noted the presence of rats, and it appears they exterminated the
Rodrigues Turtle Dove sometime between 1726 and 1761. Nothing further is known about this bird.
The Madagascar Turtle Dove Nesoenas picturata picturata was thought to have been a native species on
Mauritius, as the type description was based on a Mauritius-caught bird. However, it had been introduced
from Madagascar from at least 1770 (Cheke & Hume 2008). Subfossil remains have shown that Mauritius
once harboured a distinct species (Hume 2011a), which had disappeared by around 1730 as a result of
over-hunting, predation from introduced mammals and severe deforestation. The Mauritius Turtle Dove
differed from the Madagascar species in having more robust legs and smaller wings, and was probably
more terrestrial.
The Madagascar Turtle Dove Nesoenas picturata picturata is an adaptable species that is distributed
throughout Madagascar. There are also races on Anjouan in the Comoros (N. p. comoroensis); Glorioso Island,
Aldabra and, formerly, Cosmeledo, Astove and Assumption (N. p. copperingi); on the Amirante Islands (N.
p. aldabrana); and on the Seychelles (N. p. rostrata) (Gibbs et al. 2001).
The nominate race, either naturally or by human agency, has become established on many archipelagos,
often hybridising with local populations (Feare & Gill 1995; Skerrett et al. 2001). These introductions have
resulted in the extinction of some phenotypically distinct subspecies by genetic swamping, most notably on
the Seychelles and possibly the Amirante Islands; pure N. p. rostrata and N. p. aldabrana may no longer occur
(Feare & Gill 1995; Baptista et al. 1997; Skerrett et al. 2001). On the Seychelles, hybrid populations now
occupy Praslin, Mahé, Cousin and Cousine, and pure-bred populations are supposed to survive only on
Bird and Aride Islands, but there is evidence that even these populations show signs of hybridisation. The
Seychelles race is now probably extinct.
The Amirante Islands are a group of small atolls and coral islands situated south-west of the Seychelles in
the southern Indian Ocean. The Amirante subspecies of Madagascar Turtle Dove, aldabrana, once occurred
on Alphonse, Poivre, Desroches and probably on other islands within the island group (Skerrett et al. 2001).
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The race was originally described from captive-bred birds held by Edward Newton while he was colo-
nial secretary on Mauritius (Benson 1999). The parent Amirante birds were sent by the Seychelles Chief
Commissioner Swinburne Ward to Newton in 1868, but what eventually happened to Newton’s breeding
stock cannot now be established. It was last reported in 1950s on Alphonse, where the birds were deliber-
ately exterminated as a pest on drying copra (Skerrett et al. 2001). Skerrett et al. consider it unlikely that the
Amirante birds disappeared due to hybridisation with the nominate race N. p. picturata, introduced from
Madagascar (contra Hume 2011a; Hume & Walters 2012).
It is likely that other subspecies once occurred on islands in the southwestern Indian Ocean region, as
reports of Nesoenas turtle doves are known from Farquhar, Providence and St Pierre (Lionet 1985; Skerrett et
al. 2001).
The Macropygia cuckoo doves are a poorly known genus of pigeons that occur in South-east Asia, Indonesia,
eastern Australia and the Philippines east to the Solomon Islands and Vanuatu (Gibbs et al. 2001). The
discovery of subfossil remains in the Marquesas and the Society Islands has now shown that the genus
formerly occurred in Polynesia, with the closest living species being Mackinlay’s C
uckoo Dove Macropygia
mackinlayi in Vanuatu (Steadman 1992). Steadman suggests that many more species are likely to have once
occurred on the hundreds of islands between these two island groups.
The Marquesan Cuckoo Dove was a large species with long slender legs, and the proportions of the wing
compared to leg suggest that the species may have been more terrestrial than other members of the genus.
The subfossil remains were found in archaeological deposits, so the birds were certainly hunted by the
Polynesians and probably disappeared shortly after colonisation of the islands, long before the arrival of
Europeans. The Polynesians brought pigs and dogs, and inadvertently introduced the Pacific Rat Rattus
exulans, which would have been a dangerous predator. Steadman suggests that the inherent tameness of
Brown Cuckoo Dove M. phasianella in Australia would have been exaggerated in island forms, and perhaps
nesting close to the ground would have made the birds extremely vulnerable to extinction.
The Society Islands Cuckoo Dove is known only from the island of Huahine, but probably had a much
greater distribution in eastern Polynesia in the past (Steadman 1992). It is the largest of any Macropygia
species, either living or extinct, and had long, slender legs, which suggests that it may have been a largely
terrestrial bird. It is known only from the tibiotarsus, so until more material is discovered the flying capa-
bility of this species must remain unknown. Like its congener in the Marquesas (see above), the arrival of
Polynesians and their commensal mammals exterminated the population soon after human settlement.
Description 39–41cm (15.5–16in). Large dove with long, broad wings and a long graduated tail. Male: head bluish-grey; display
area at back of neck iridescent bronze, green or purple depending to the light; back tinged s late-grey with olive-brown; lower
back and rump greyish blue becoming greyish-brown on uppertail-coverts; two central tail feathers brownish-grey, rest white;
wing-coverts brownish-grey with irregular subterminal spots; primaries and secondaries darker g reyish-brown; secondaries
edged with white; throat and breast rich pinkish rufous, becoming paler on lower breast to white on abdomen and under tail
coverts; bill black; iris carmine red, naked orbital ring purplish red; legs and feet red. Female similar to male but overall duller,
greyish-brown head neck and back; reduced display iridescent markings; underparts b uff-brown, less rufous and tail shorter;
legs and feed paler red; iris orange red, naked orbital ring greyish blue. Immature similar to adult female, with scapulars,
wing-coverts, feathers of foreneck and breast tipped with white; lacks display iridescent markings; legs and feet dull red; iris
brownish with narrow ring of carmine.
The loss of the Passenger Pigeon, perhaps once the most numerous bird on Earth, is the most damning of all
human-caused extinctions; it defies belief that a species so abundant could be exterminated in such a short
period of time. Before the settlement of the continent by Europeans, the range of the Passenger Pigeon
extended through the deciduous forests of North America, from the Great Plains east to the Atlantic, north
to extreme southern Canada, and south to Virginia and northern Mississippi. In winter the birds migrated
south to the south-eastern United States and the Caribbean slope of Central Mexico (though occasionally
further north, in areas such as Indiana, Pennsylvania and Massachusetts). Stragglers occurred in British
Columbia, north-eastern Baffin Bay, and south to Bermuda, Cuba and the Valley of Mexico (Schorger 1955;
Gibbs et al. 2001). They were vagrants to Florida, and there are records from the British Isles, though there
are doubts about their authenticity. Audubon himself imported 350 live birds to England in March 1830,
and distributed them amongst several noblemen (Saunders in Yarrell 1884), and many other imports took
place. It is likely that most if not all British records involved escaped birds (Alexander & Fitter 1955).
The bird was highly migratory, and bred in immense colonies through much of its breeding range;
however, these vast colonies appeared to be irregular, and the birds were not to be found in them every
year, or in the same place (Schorger, in Scott 1947). Nests were often found in twos and threes, and small
colonies were scattered through the birds’ range. But the numbers in flocks could be staggering; Wilson
(1808–14) estimated 2,230,272,000 birds in a flock, and Schorger in 1871 calculated 136,000,000 in a Wisconsin
nesting area that covered 2200km2. Greenway (1967) considered that in making allowance for isolated pairs
breeding on the periphery, about half as many again should be added to reach a conservative total. This
infers that Wilson’s estimate is not implausible; but Audubon (1831), who probably never saw a great
nesting, certainly did not believe Wilson’s figure, and divided it by two! There can be no doubt that reports
of vast flocks that darkened the sky, and that the weight of their numbers in trees caused branches to
collapse to the forest floor, are based on factual observations (see Kalm, in Gronberger 1911; Wilson 1808–14;
Forbush 1912; Bent 1932).
Decline
The Passenger Pigeon was in decline from the early 1800s, with this accelerating during the latter half of the
19th century. This coincided with an increase in commercial exploitation, particularly following the devel-
opment of new railroads and telegraph systems after the American Civil War. By this time the birds were
continually hunted during migration or nesting periods, for meat, fertiliser, and even sport (Hume 2015a).
An unpublished letter written on June 11, 1851 by Thomas Cawley from Cleveland, Ohio to his sister in
New York gives a flavour of the killing sprees associated with Passenger Pigeon migrations:
Pidgeons are very plenty & for the week past every day from day light till dark there has been continual firing at
the poor birds. Even while I write there are hundreds of men & boys in & around the city popping away. They
kill hundreds in the streets & as long as this warfare continues, I don’t have many hands [hired hunters] to kill
it, seems to me that every body has bid farewell to mutton chop & live on pidgeon pie.
By 1870, birds had ceased breeding in large concentrated groups except in the northern states around the
Great Lakes. Furthermore, as the price of their corpses decreased due to market saturation, the live-trap-
ping of birds for shooting tournaments became fashionable (Hume 2015a). For example, from over a million
birds taken in 1878 at one of the last great nestings at Petosky, Michigan and elsewhere, around a quarter
were sent to the shooting market (Mitchell 1935). This practice only ceased when there were no more
pigeons available. The last large nesting took place in Ohio in 1885 (Trautman 1940).
The last wild specimen recorded was taken at Babcock, Wisconsin, between 9 and 15 September 1899
(Schorger 1938), or March 24 1900 in Pike County, Ohio (Fuller 2000). It is possible that a few survived
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later, as there were a number of sight records between 1900 and 1907, but those cases that were investi-
gated proved to be in error, with most referring to the Mourning Dove Zenaida macroura. However, the
Passenger Pigeon continued to survive, but not thrive, in captivity.
Martha
The original flock held at the Cincinnati Zoological Gardens came from Michigan in 1875, and consisted of
26 birds, with about half males and half females (S. A. Stephan, in Deane 1907). Around five or six escaped
shortly after. The pigeons raised 23 young while in captivity, but only ever laid one egg at a time. By 1908,
only three birds were still alive; two males and a single female, that were all well beyond breeding age. Of
these, the female, Martha, was born at the Cincinatti Zoo, but later joined a flock kept by Professor Charles
O. Whitman of the University of Chicago. She was returned to the dwindling Cincinatti flock in 1902
(Shufeldt 1915). Whitman photographed a number of the pigeons, before the last of his birds died in the
winter of 1907 (Fuller 2015). On 1 September 1914, just before one o’clock in the afternoon, the last indi-
vidual, Martha, then aged 29 and having outlived the rest of her kind by four years, perished, thus adding
the Passenger Pigeon to the ever-increasing list of extinct species (Shufeldt 1915; Greenburg 2014; Fuller
2015).
Habits
The Passenger Pigeon diet comprised beech mast, acorns and chestnuts, though a wide range of other foods
was also taken, including invertebrates. Mast is produced abundantly every two to five years, but it is very
rarely produced in abundance in consecutive years. High levels of mast availability were essential for
successful breeding, and as mast is produced in autumn, and the pigeons bred in spring, there needed to be
a considerable residue for the spring and summer while the young were being reared. This occurred only in
colder areas where there was enough snow cover to prevent the mast being exploited by resident species
over the winter.
The Passenger Pigeon exhibited rather subdued p re-nesting behaviour (in captivity at least), without the
bowing, strutting or b ill-contact seen in other pigeons (Craig 1911). Breeding occurred from March until
September, most intensely between April and May. The nest was a flimsy construction made from fine
twigs, and usually one but occasionally two white eggs were laid. Incubation lasted about 12–13 days, and
the squabs fledged within 12–14 days (Deane 1896). Vocalisations included crowing, chattering, twittering
and croaking, while a soft, toneless clucking was made during mating (Blockstein 2002).
The St Helena Pigeon was a large, distinctive pigeon with disproportionately short wings and robust legs,
all of which are indicative of flightlessness. Only the Dodo Raphus cucullatus, Solitaire Pezophaps solitaria and
Viti Levu Giant Pigeon Natunaornis gigoura had evolved a similar condition. Olson (1975b) considered these
characters sufficient to warrant generic status. Rowland et al. (1999) and Ashmole & Ashmole (2000)
suggested (on grounds of skeletal morphology) that the St Helena Pigeon was closest to the Streptopelia
turtle doves, which are adept colonisers of remote oceanic islands and archipelagos. Once established on
the island, the St Helena Pigeon evolved characters associated with reduced flying capabilities, most
notably a general increase in size, reduction of the pectoral girdle (bones of the shoulder) and increased
robustness of the tarsometatarsus, with widely spaced trochleae (knuckle bones). The bird was almost
certainly terrestrial and would have
nested on the ground. Nothing else is
known about it. St Helena Pigeon
It is difficult to determine whether Dysmoropelia dekarchiskos
this species disappeared in relatively
recent times. According to Olson (1975b),
the ‘small kind of blue dove’ observed by
Forster on St Helena in the 1770s
(Ashmole 1963b) was probably the intro-
duced Zebra Dove Geopelia striata and
not an endemic form, and that introduc-
tions had been made very early in the
island’s history. Linschoten in 1589 (in
Tiele 1885) found ‘hennes, partridges
and doves’, and in 1634 Peter Mundy
found ‘little speckled ginny Hennes,
partridges and Pigeons’ (Temple 1914);
the doves and pigeons may have referred
to the endemic D. dekarchiskos. However,
being a large and distinctive terrestrial
species, the St Helena Pigeon would
surely have been mentioned in more
detail by the astute observer Mundy, so it
169
had probably become extinct at least before his visit in 1634. The reason for its demise is unclear, but being
flightless and confined to the ground it would have fallen easy prey to humans and their commensals.
Quail doves are terrestrial birds typically associated with dense, shady undergrowth (Gibbs et al. 2001).
Wetmore (1920) believed the extinct Puerto Rican species to be closely related to the G rey-headed Quail
Dove Geotrygon caniceps of Cuba (G. c. caniceps and Dominican Republic (G. c. leucometopius).
Compared with the widespread Ruddy Quail Dove G. montana, the Puerto Rican bird was more slender
in the tarsometatarsus. Owing to the apparent abundance of subfossil remains, the Puerto Rican Quail Dove
is thought to have once been common on the island, but it disappeared due to deforestation of the island
after human settlement.
The genus Caloenas comprises one surviving species, the widespread Nicobar Pigeon C. nicobarica, which is
divided into two subspecies. The Nicobar Pigeon is a distinctive terrestrial bird, being large and heavily
built with long wings and a short tail. It regularly migrates large distances between oceanic islands, and its
range extends from the Andaman and Nicobar Islands in the Bay of Bengal to Palau and the Solomon
Islands in the South Pacific (Gibbs et al. 2001). The genus also includes two recently extinct species, the enig-
matic C. maculata (see below) and the Kanaka Pigeon C. canacorum.
The Kanaka Pigeon was first described from subfossil remains found on New Caledonia, but recent work
(Steadman 1989a, 2006b) has shown that it also occurred on Tonga and probably on Vanuatu and Fiji as
well. The Kanaka Pigeon was 25% larger than the extant Nicobar Pigeon, being approximately 38cm in total
length, and wing proportions show that it was quite capable of flight (Balouet & Olson 1989). A large terres-
trial pigeon such as this would have no doubt made excellent eating and, as a result, probably became
extinct throughout its range due to over-hunting.
The relationships of the Spotted Green Pigeon have long been shrouded in mystery. It was thought to bear
no close relationship with any extant genera save a rather superficial resemblance to the Nicobar Pigeon
Caloenas nicobarica. Wagler (1827) considered that it might have been the young of this species, of which
Rothschild & Hartert (1901) commented:
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The most peculiar Caloenas maculata – correctly identified as a Caloenas by Wagler – is certainly not the young
of C. nicobarica, as the young are almost quite like the adults, and not spotted. It is extraordinary that the home
of this bird is not yet discovered, and we suggest the possibility – although there were two specimens – that it is
an abnormality. The bird is well figured in the first volume of the Bulletin of the Liverpool Museums, where the
type is preserved.
Of the two specimens originally collected, one is now lost. One of us (JPH) wrote about the species to the
late Derek Goodwin, an expert on pigeons and doves, who replied that he was unaware of it, but that it
appeared to be a distinct species, perhaps related to Caloenas (D. Goodwin pers. comm.). Gibbs et al. (2001)
thought that due to its distinct morphology, the bird may be better regarded as a distant relative of Ptilinopus,
Ducula or Gymnophaps, and warranted generic status.
Recent mtDNA analysis has shown that the Spotted Green Pigeon is a distinct taxon, closely related to
Caloenas nicobarica, and basal to the Dodo Raphus cucullatus (see p. 155) and Rodrigues Solitaire Pezophaps
solitaria (see p. 158) (Heupink et al. 2014). The short, rounded wings reported in Hume & Walters (2012)
appear to be due to the loss of the main primaries (H. van Grouw, pers. comm.), so it probably had long
wings, with a long tail and short and stout tarsi. This indicates that it was probably an arboreal species, not
a terrestrial one, with the ability to perhaps migrate between islands (Heupink et al. 2014; contra Hume &
Walters 2012). The bill was typical of a frugivorous pigeon.
Gibbs et al. (2001) and Hume & Walters (2012) discuss a bird known to the Tahitians as a titi; they
described it as a speckled green-and-white bird that had a loud call, titi (Henry 1928). This was perhaps a
reference to the Spotted Green Pigeon. However, the name titi is an onomatopoetic name throughout East
Polynesia for petrels and shearwaters (Steadman 2001); the provenance of the Spotted Green Pigeon
remains unresolved.
The Norfolk Island Ground Dove was illustrated by John Hunter (1793), Vice-Admiral of HMS Sirius of the
First Fleet and later second governor of New South Wales (van Grouw & Hume 2016). The illustration was
reproduced by Hindwood (1965) and again in Forshaw (2016), and represents the only known image of the
bird. No skin specimens exist. Hindwood (1965) also quoted several letters and other papers proving that
an extinct dove formerly occurred on Norfolk Island. Hindwood describes it is as follows:
The bill is black with a slight indication of a cere; the legs and toes are reddish-flesh; the crown, throat and neck
white; the back of the neck, mantle, back, breast, belly and tail are a deep warm brown, the wings being the same
colour with deep purple coverts or shoulder-patches.
Further confirmatory evidence that a dove lived on Norfolk Island is contained in contemporary published
material. In a letter written from the island on October 28, 1788 (printed in the Kentish Gazette on June 6,
1789), it is stated that:
The wild fowl are pigeons, quails, doves, plovers, corlieus, parrots, paroquets and a black bird with a web foot
that burrows in the ground, with a great variety of other smaller birds; the pigeons we used to take as many of
them by hand as we chose, but they are now a little shy.
In March, 1788, soon after the settlement of the island, Hunter (1793) wrote:
We saw pigeons, parrots, parroquets, doves, and a variety of other birds, in great numbers, and so very tame,
that they might be knocked down with a stick.
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The following extract, from a letter written by a non-commissioned officer on Norfolk Island in August 1790
also mentions doves (Historical Records of New South Wales 1892):
There is some little variety of Birds. A Parrot, Paroquet, Pidgeon, Dove, Hawk, and Owl, with many small birds.
The Parrot is very curios; the Paroquet common with no great share of beauty; the Pidgeon beautiful; the Dove
delicate; the Owl and Hawk quite common. Amongst the small Birds there is but one worthy of note, which is
not unlike a Robin, having a rich vermilion breast.
It is certain that a dove, as distinct from the large pigeon (Norfolk Island Pigeon Hemiphaga novaeseelandiae;
see p. 187), once occurred on Norfolk Island, and as subfossil material has now been found (Meredith 1985;
Holdaway & Anderson 2001), it confirms that the Norfolk Island Ground Dove inhabited the island prior to
European settlement. Being a predominantly terrestrial species and tame and confiding, it was doubtless
soon exterminated by over-hunting and predation from feral cats and rats. Nothing is known about its
habits, but it was probably an inhabitant of coastal forests (Higgins & Davies 1996).
Taxonomy
Latham (1801b) described under Columba norfolciensis three quite distinct birds, of which two are now iden-
tified as the Common Emerald Dove Chalcophaps indica and the White-headed Pigeon Columba leucomela; the
third is the Norfolk Island Ground Dove (Forshaw 2016). Because it is uncertain that the bird illustrated by
Hunter is the one described by Latham (Goodwin 1983; Forshaw 2016), Latham’s name was formally
suppressed (ICZN 2010), leaving the bird illustrated by Hunter without a name; hence the new generic
assignation and specific epithet (see Forshaw 2016).
The Polynesian Ground Dove Alopecoenas erythropterus has a wide distribution in the South Pacific, and
once occurred in two widely separated island archipelagos, the Tuamotu Islands and the Society Islands. It
now survives only on the Tuamotus, where two subspecies have been described. Nominate A. e. erythrop-
terus is considered critically endangered, whereas the White-headed Polynesian Ground Dove A. e. albicollis
(see below) is probably extinct.
The Society Islands Ground Dove was described and illustrated from specimens collected during all
three voyages of Captain James Cook to the South Pacific, between 1768 and 1779. Latham (1783, 1790)
called them ‘Garnet-winged Pigeons’ based on the coloration of the mantle and wing-coverts. Most of the
skin specimens are now lost, but one specimen from Tahiti was purchased by the collector William Bullock,
and is still in Leiden. There is some variation in the descriptions and illustrations, and as they were obtained
on both Moorea and Tahiti, these two island populations may have represented distinct subspecies.
The birds were once abundant (Murphy 1924), but no Gallicolumba has been seen in the Society Islands
since the original specimens were collected in the 18th century (Holyoak 1974b). No doubt o ver-hunting,
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habitat destruction, and introduced mammalian predators were responsible for the Society Islands Ground
Dove’s extinction. Nothing is known about the bird’s habits. It was called the ‘ oo-oo widou’ by the Mooreans
(Sharp 1906a).
Two forms of Polynesian Ground Dove A. erythropterus are currently recognised (Gibbs et al. 1998), although
a third as-yet undescribed race (or perhaps species) formerly occurred on the Society Islands (see above).
Subfossil remains have also been found on Mangaia in the Cook Islands, which may represent a fourth
taxon (Steadman 1989b).
The nominate, now considered Critically Endangered (Collar et al. 1994), originally occurred on a number
of islands in the Tuamotu Archipelago. The w hite-headed race from the islands of Hao, Hiti and perhaps
Tahanea was a medium-sized, short-tailed terrestrial bird. It was originally abundant, but it rapidly disap-
peared, primarily due to predation by introduced cats and rats everywhere that humans settled. It seems to
have been last seen in the 1950s, but the rediscovery of populations of the nominate race on small islands
provide hope for the continued survival of A. e. albicollis.
Habits
The White-headed Polynesian Ground Dove occurred in dense forest with a mature understorey comprising
tall shrubs, ferns and grasses, in low scrub, and in Pandanus forest with open ground vegetation (Gibbs et al.
2001). It was a terrestrial species, feeding on fallen seeds, buds of low-growing vegetation, and a variety of
invertebrates.
The Tanna Ground Dove has not been seen since the Forsters illustrated and described a specimen (now
lost) on Cook’s second voyage, obtained on Tanna in southern Vanuatu. Greenway (1967) doubted that the
bird was confined to Tanna, a small island with relatively few extant birds. There seems, however, to be no
question of an error of locality, for Georg Forster wrote: ‘Tanna. Female 17 August 1774’ on the margin of the
drawing, and his father specifically recorded a new species of dove as having been shot on the island.
Stresemann (1950) pointed out that the German edition of Forster’s travels contains a note which is not in
the Italian or English editions (translation by JPH):
173
17.8.1774: Behind these fields we came to a forest, where there was however no other kind of trees than those
which we, at the beach, had already found. On the other hand a pigeon of a new kind was shot.
Latham (1823) commented further:
This seems to be between the two former [the two races of Polynesian Ground Dove; see p. 172] for it has
the white streak over the eye, as in the first described, and the fore parts to the breast white as in Var A. The fore-
head, sides, and fore part of the neck, are also white; legs dull crimson; belly and back reddish black; tail rounded.
This last was brought from the Island of Tanna; and in the collection of Sir Joseph Banks.
The above description by Latham (1823) and that of Forster (1844) differ slightly, and Salvadori (1893)
suggested that Latham’s description was of a male, and Forster’s of a female. Stresemann (1950), however,
believed that Forster’s specimen was a male.
The Tanna Ground Dove probably disappeared due to predation by introduced rats, as its probable close
relation on Santa Cruz, A. sanctaecrucis, has disappeared for the same reason over much of its range (Gibbs
et al. 2001). Nothing more is known about it.
Doves of the genus Alopecoenas are small to medium, rather plump pigeons, with comparatively small
wings and long, unfeathered tarsi (Gibbs et al. 2001). They are generally terrestrial species, which makes
them extremely vulnerable to anthropogenic changes, so many species have either become extinct or are
critically endangered. The Henderson Island Ground Dove was similar in some proportions to the
wide-ranging Shy Ground Dove A. stairi, except that it was larger, the wings were shorter and the tarsi were
more robust (Worthy & Wragg 2003). This suggests that it was a poor flier at best, and like the other extinct
Henderson Island terrestrial pigeon species, it probably died out due to o ver-hunting soon after humans
arrived on the island.
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This ground dove, based on the tarsometatarsus, was the largest Alopecoenas species known (Steadman
1992) and also the most widely distributed species in eastern Polynesia, occurring sympatrically with
smaller species on the Cook Islands, Society Islands, Tuamotus and Marquesas (Steadman 2006a). Its
remains have been found in midden archaeological sites, so it was certainly hunted by the Polynesians.
Other Alopecoenas forage and nest on or near the ground, and require primary forest in order to survive, so
the destruction of habitat and predation from introduced mammals would have contributed to its
extinction.
This was one of the largest Alopecoenas doves, with long but slender tarsi (Balouet & Olson 1989). It is only
known from very few skeletal elements, so whether it had reduced flying ability is now difficult to estab-
lish. Like most other extinct New Caledonian bird species, the New Caledonian Ground Dove probably
disappeared rapidly after the arrival of humans and their commensal animals.
This dove was a large species with a heavy bill. It is known from just two specimens, one taken in San
Cristobal (now Makira) in 1882 and the other by the Whitney South Seas Expedition on Ramos Island on 5
September 1927. Very few subsequent searches have been made, and no birds have been seen since the
Whitney Expedition (Gibbs et al. 2001). It appears to have succumbed to habitat destruction and predation
by introduced rats, cats, pigs and dogs, and also o ver-hunting. On Makira most of the lowland forest has
now been destroyed, whereas Ramos remains comparatively forested, but hunters on adjacent islands make
regular trips to collect game (Buckingham et al. 1995).
Habits
Very little was recorded about this ground dove. It appears to have been a bird of the lowlands as both
specimens were collected below 300m, but congeneric species occur in forest at all altitudes (Gibbs et al.
2001).
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This species is known from just two specimens, collected in 1891 by the Menage Scientific Expedition; even
then it was described as extremely rare (Bourns & Worcester 1894). However, reports by locals indicate that
it was common before the 1970s, and there are unconfirmed reports from 1995, but other b leeding-heart
pigeon species have been introduced (Gibbs et al. 2001) and it might well be these introductions that have
been reported. Surveys in the 1990s failed to locate it (Collar et al. 2001). T awi-tawi had almost been
completely logged by 1994, and hunting increased in the 1970s. The bird is rumoured to survive on small
off-shore islands, but these may not be insufficiently large to maintain viable populations. If the Sulu
Bleeding-heart does survive, the future for it is very bleak indeed.
Habits
Almost nothing recorded. Like other Gallicolumba pigeons, it presumably inhabited the forest floor, prefer-
ring primary forest with dense undergrowth, and fed on fruit, seeds and invertebrates.
The relationships of this pigeon are obscure. It was the largest Henderson Island pigeon, once occurring
sympatrically with the extinct imperial pigeon Ducula harrisoni and ground dove Alopecoenas leonpascoi, and
the only surviving Henderson pigeon species, the Henderson Island Fruit Dove Ptilinopus insularis. Skeletal
characteristics suggest that Bountyphaps obsoleta was more closely related to the T ooth-
billed Pigeon
Didunculus strigirostris and the Nicobar Pigeon Caloenas nicobarica than to other genera (Worthy & Wragg
2008).
It is remarkable that Henderson Island, with an area of just 37km2 and a maximum height of 33m, could
have supported four species of pigeon, but similar scenarios are known on tiny Moheli in the Comoros, and
Réunion and Mauritius in the Mascarene Islands (Hume 2011a). The differing ecologies of these species
presumably provided full utilisation of limited terrestrial and arboreal niches on Henderson Island. The
proportions of the available skeletal remains suggest that B. obsoleta had comparatively reduced wings
compared with body size, and had weak powers of flight but was not flightless. The subfossil remains were
discovered in Polynesian middens, so these birds probably disappeared due to o ver-hunting shortly after
the human occupation of the island.
purplish-red. It is still not established whether the crest was held erect as in the Goura crested ground pigeons; all of the skin
specimens of Microgoura meeki have the crest prostrate.
All known specimens of the Choiseul Ground Pigeon Microgoura meeki were collected by Albert Stewart
Meek in January 1904, and the bird has not been seen with certainty since. The American Museum of
Natural History’s Whitney South Seas Expedition searched for the bird in 1927 and 1929 without success
(Tennent 2009). They were told by local people that the bird had not been seen for a long time. The expedi-
tion searched mainly in the mountains, though it seems unlikely that Meek managed to get far inland. It
seems more likely that the bird inhabited the coastal swamps, and it was there that Parker (1967a) searched
for it in the 1960s, but failed to find it. Parker (1967a, b; 1972a) and Tennent (2009) discuss and summarise
most of what is known about the bird. Both examined the collector Hamilton Hamlin’s notes of the Whitney
Expedition, which in 1927 collected for six days opposite Moli Island, towards the north-west end of
Choiseul. Hamlin wrote on 21 November:
Mr. Beck has found that the natives savvy [the pigeon] but they give the impression that it is rare.
Four days were spent subsequently at the southern end of Choiseul Bay, but there was no sight of M. meeki
during this period. In 1929 Hamlin returned to Choiseul for a more thorough search. No mention of the
pigeon occurs in journal entries made at northern Choiseul Bay, Nio, or Senga, opposite Ruiana Island
(probably modern Lengatura), two camps on the north-eastern coast, nor from inland areas reached from
these camps. However, at Sasamunga on the southern coast Hamlin wrote on 7 October:
Enquiries about the Microgoura resulted in some new information: the birds they know here from my descrip-
tion is called ‘kukuru-ni-lova’, lit. ‘pigeon-belong-ground’, and is recalled only by the older men, who say that
cats, introduced since the advent of the Mission, have destroyed so many that they cannot remember when one
was last seen in the bush. And for the past ten years practically all the bush people have moved down to the salt
water. The big river basin [Kolombangara River] we traversed yesterday is said to have been a good place for
them. The birds were easily caught by the boys in their hands after they had found a low-branched tree in which
the pigeons roosted in twos and threes and fours by noting the manure on the ground underneath; they would
simply wait their opportunity and seize them while sleeping.
This area was searched for
several days but no Microgoura
was found. Hamlin also called
briefly at several south-coast Choiseul Ground Pigeon
villages east of Sasamunga. At Microgoura meeki
Toi-Toi near Katurasele, villagers
confirmed that they had not
recently seen the pigeon, and
that feral cats had been known
to prey on it. At Sambi (probably
modern Zongga) and Rorwai
(Ruravai), older inhabitants
recognised the description of the
bird and remarked on its beauty,
imitating its call by a low trilling
sound. Three natives of Taora in
eastern Choiseul told Hamlin
that they had seen Microgoura
that year. Five days’ search in
this locality revealed no trace of
the bird. Inhabitants of Kumbura
village on the northern coast of
eastern Choiseul stated that they
too had seen the bird that year.
Eighteen days’ search revealed
nothing. Parker (1967a)
commented that:
177
It is significant that most of the localities at which there was native recognition of Microgoura are in the vicinity
of riverine flats and swamps . . . It is highly probable that this pigeon, like its New Guinea relatives in the genus
Goura, inhabits riverine and flooded forest and other swampy areas. There is no mention among the Whitney
fieldnotes for Choiseul of any collecting having been done in swamps, which may help to explain the expedition’s
failure to obtain this pigeon.
Parker (1972a) visited the area again in 1968 in an attempt to discover the bird. He wrote:
Only the older men of Choiseul to whom I spoke recalled Microgoura, usually from areas along the sheltered
south coast. All who knew the bird said that they had not seen it for many years and that cats, introduced to keep
down rats, had finished the species. Dogs used on pig-hunts would also catch them. The most recent record I
obtained was from the inhabitants of Sasamungga on the Kolambangara River (apparently a former stronghold
of Microgoura) of a small roost seen during World War II, i.e. in the early 1940s. This would confirm the pres-
ence of Microgoura in the Kolambangara basin at the time of the Whitney Expedition’s search, when in spite of
native advice and help the expedition failed to find it. From other parts of the southern coast it seems to have
disappeared earlier, during the 1920s and 1930s . . . Its absence from the area of Choiseul Bay by 1927–9, when
Beck and Hamlin of the Whitney Expedition called there, may have been because of the development of the
coconut plantations and the almost complete removal of the original vegetation. My chief hope had been to find
Microgoura on the swampy, almost undisturbed, islands of Rob Roy and Wagina, the former uninhabited, the
latter inhabited only since the Gilbertese settlement of 1964. I failed to find any trace of the bird there. The
Gilbertese, who did not recognise my picture of Microgoura, remarked that they had found feral cats common
on Wagina when they arrived.
Parker goes on:
I spent from 26 January to 7 February 1968 on the western coast of Malaita making enquiries among the natives.
I found that no one knew of Microgoura at all, though people sometimes at first confused my coloured picture of
that species with such crested birds as Aviceda subcristata (kito or ito, the Crested Hawk) and Reinwardtoena
crassirostris (gwalotê, the [Crested Cuckoo Dove]). Although one cannot say even now that Microgoura
meeki is extinct, the likelihood of its survival is small. The feral cat, its reported predator, has been widespread
in the Solomon Islands for several decades.
Goodwin (1983) reported that David Roe of the National Museum, Honiara, Solomon Islands, informed
him of the bird’s continued existence based on reliable local sources, and that it is rare and confined to a
coastal area with limestone cliffs and sparse vegetation. This seems highly unlikely, however, and the
species is now considered extinct (BirdLife International 2011).
Taxonomy
The relationships of this bird are obscure. Gibbs et al. (2001) suggest that it was closely related to the mono-
typic Thick-billed Ground Pigeon Trugon terrestris, and Goodwin (1983) thought that it was a link between
Trugon and the Goura crested pigeons, although it differs markedly from the latter genus.
Habits
Little is known about the Choiseul Ground Pigeon. Meek (1913) commented that the bird made no nest, but
laid a single dark creamy-white egg on the ground, small in proportion to the size of the pigeon. He was
satisfied that it did not occur on Bougainville, but was told by local boys that it was found on both Ysabel
and Malaita. Gibbs et al. (2001) recall a village chief’s description of a pigeon that had disappeared when his
father was a boy, and which had a ‘beautiful rising and falling whistling call given from the roost every
evening’. Parker (1972a) writes:
Two informants, one from Sasamungga and one from Kiala, told me that stones were often found in the gizzard
of Microgoura. According to Chalmers and Gill (1885) stones were also found in the gizzard of the related New
Guinea crowned pigeons, Goura . . . One of my Choiseul informants described the gizzard stone of Microgoura
as being of gold, thereby signifying perhaps that such stones were regarded as valuable.
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The Tooth-billed Pigeon Didunculus strigirostris is a distinctive, peculiar pigeon with a robust hooked bill
with tooth-like notches on the mandible, which occurs only on Savai’i, Upolu and Nu’utele in Samoa (Gibbs
et al. 2001). It is considered vulnerable due to hunting and forest fragmentation. A second Didunculus
species, the Tongan T ooth-billed Pigeon, was described in 2006 from cave deposits on Eua, Tonga, and from
archaeological sites on Tongatapu, Lifuka, Ha’ano, ‘Uiha and Ha’afeva (Steadman 2006b). It is a much larger
species, and as in other Pacific genera once considered monotypic, such as Caloenas, it demonstrates that the
genus was more widespread in the past. The Tongan Tooth-billed Pigeon disappeared some time after the
colonisation of the islands by Polynesians, probably as a result of over-hunting, habitat loss and predation
by introduced mammals.
This species is known from one specimen, a female, one of a pair shot in 1953 in Mount Canloan Natural
Park, Negros (Ripley & Rabor 1955). The other, a presumed male, was lost in the undergrowth after the
shooting. There have been no further sightings despite recent surveys (Brooks et al. 1992), and high hunting
pressure coupled with almost complete deforestation of Negros provide little hope for its continued
survival.
The type was collected at approximately 1,100m in a tall fruiting tree, but the forest had been cleared up
to 1,000m, so if this was a lowland species and forced to the edges of its normal range, little or no suitable
habitat now remains (Collar et al. 2001, Gibbs et al. 2001).
Ebon Island Fruit Dove Ptilinopus porphyraceus marshallianus Peters & Griscom
Ptilinopus marshallianus Peters & Griscom 1928, pp. 104–06 (Ebon Island, Marshall Islands)
Specimens The type specimen is in Cambridge, Massachusetts.
Status Known only from the type. The population is extinct.
Range Ebon Island, Marshall Islands, South Pacific.
Description 23cm (9in). Forehead, lores and crown deep p inkish-purple, bordered in front of eye and rear of crown with
yellow; back g reyish-green; wings dark glossy green; reduced emarginated tip to first primary; rump and uppertail-coverts
dark olive; uppertail dark green; chin and throat creamy white shading to greenish-grey on ear-coverts, neck and sides of breast;
upper belly greenish-grey shading to dark olive; variable indigo patch on central upper belly; vent and undertail-coverts
orange; broad and bright yellow apical t ail-band; bill dull green; iris yellow-buff; legs and feet purplish-red.
This bird’s status was based on a single specimen preserved in spirit, discovered in the Museum of
Comparative Zoology, Harvard. Aware of the possible effects of immersion in alcohol on the colour of
feathers, Peters & Griscom (1928) carefully compared the bird with a specimen of the closely related Grey-
green Fruit Dove Ptilinopus purpuratus. The comparison specimen, of the race P. p. chrysogaster, had also
been preserved in alcohol. Their careful diagnosis of the bird was as follows:
the bright gamboge [yellow] colour on the centre of the abdomen and undertail-coverts [of the specimen of
chrysogaster] has faded out to a pale dirty buff, and the yellow wash on the rest of the underparts is only faintly
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indicated in a pale buffy tinge. In the type of P. marshallianus [Ebon Island Fruit Dove] this pale buffy tinge
is entirely wanting. The underparts, therefore, cannot have had the yellow wash so obvious as in P. chrys-
ogaster, P. hernsheimi or even P. coralensis [westen Society Islands race of G rey-green Fruit Dove,
Kosrae race of P urple-capped Fruit Dove and Atoll Fruit Dove, respectively]. The centre of the abdomen,
vent and undertail-coverts of the type of marshallianus are, however, a much clearer and brighter buff, and
noticeably deeper on the u ndertail-coverts with a salmon tinge. It is highly likely, therefore, that these parts
might actually have been orange in life.
Peters & Griscom further point out that the bird had the ‘centre of the abdomen, vent, undertail-coverts and
a broad terminal band on the tail bright gamboge yellow’.
Ripley & Birckhead (1942) synonymised the bird with P. porphyraceus hernsheimi, the race of Purple-capped
Fruit Dove endemic to Kosrae Island in the Caroline Islands, and this assertion has often been followed.
However, Ripley & Birkhead’s arguments are not convincing:
It is identical in pattern with hernsheimi, except that it lacks the yellowish coloring throughout. As it is known
to have been in alcohol for many years before being made into a skin, and as the supposed locality, Ebon Island,
Marshall Islands, was never very positive, it seems likely that it is simply a specimen of hernsheimi with a
wrong locality and with the plumage changed by the action of alcohol.
These assertions are baldly contradicted by Peters & Griscom (1928). They were well aware of the effects of
immersion in alcohol on feather pigments, and there is good evidence that the specimen did in fact come
from Ebon. The island is one of the southernmost of the Marshall Group, and has the richest vegetation of
any Marshall Island atoll (Amerson 1969). It was the hub of the Micronesian Mission, and it was the resident
Rev. B. G. Snow who collected the type and presented it to the Museum of Comparative Zoology, along
with a large collection of fish and marine invertebrates obtained on Ebon.
Furthermore, an overlooked report by Rev. E. T. Doane in 1861 (see Olson 1992a) made the following
account about Ebon’s avifauna:
This atoll is the home for a few varieties of birds. But in this feature of the island, the contrast is as wide between
the ‘low coral’ island and the ‘high volcanic’ one – as between their natural features. The high islands of
Micronesia are largely supplied with the feathery tribe, but this atoll can claim only a few birds – and with two
or three exceptions these are all water fowl. There are a few Columbidae, Carpophaga [Ducula] oceanica,
which manage to elude the keen search of the natives. These birds are occasionally heard cooing away in the tops
of some quite isolated b read-fruit tree. A Cuculus [doubtless L ong-tailed Koel Urodynamis taitensis] gives
forth occasionally its sharp whistle – and these, with the addition of another land bird whose species I have not
been able to learn – are all the feathery songsters this atoll can claim.
Olson (1992a) believed that the third unknown bird was P. p. marshallianus and, if so, the subspecies almost
certainly represents a population, now extinct, endemic to Ebon Island.
Extinction
The Marshall Islands were visited very briefly by Europeans as early as 1529, and again in 1767, but Ebon
was not discovered until May 25, 1824 by the American ship Nantucket under Captain George Ray, who
named it Boston Island (Findlay 1886). The islands were known to whaling fleets, so it might be that escaped
rats were responsible for the population’s demise. Nothing more is known about the bird.
Nuka Hiva R
ed-moustached Fruit Dove Ptilinopus mercierii mercierii
(Des Murs & Prévost)
Kurukuru Mercieri Des Murs & Prévost 1849, p. 266 (Valley of Mohana, Nuka Hiva Island, Marquesas group)
Specimens The type specimen is in Paris.
Status Extinct. Not recorded since 1849.
Range Nuka Hiva, Marquesas Islands, South Pacific.
Description 22cm (8.5in). A brightly coloured, relatively small, compact dove: forehead and crown bright purplish crimson;
throat yellowish; nape, neck and breast silver-grey tinged with green; underparts bright golden yellow; back and wings rich
green with golden tinge; tail dark green with broad, whitish terminal band.
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The Nuka Hiva Red-moustached Fruit Dove is known from a single specimen, collected in 1849 in the
Valley of Mohana, Nuka Hiva, an island in the Marquesas. It was probably already rare at this time. Holyoak
& Thibault (1984) point out that the Whitney Expedition did not find it in 1921–23, and nor did they them-
selves in 1972 or 1975. Nothing is known about this bird in life or why it became extinct, but the introduction
of cats and rats on the island were no doubt largely responsible for its extinction.
Like the nominate R ed-moustached Fruit Dove on Nuka Hiva, little is known about this bird. The type was
collected by Garrett (Salvadori 1892) and a few other specimens were taken during the course of the nine-
teenth century; the Whitney South Seas Expedition collected 14 specimens (Ripley & Birckhead 1942),
despite it being considered rare in 1921–22, and it has not been seen since. Predation by the introduced
Great Horned Owl Bubo virginianus has been cited as the cause of its extinction, but rats and cats, which
were introduced long before the owl, are more likely candidates (Gibbs et al. 2001).
Habits
The habits of this subspecies are better documented than those of the Nuka Hiva race. It was described as
arboreal, and more rapid and agile in flight than the White-capped Fruit Dove Ptilinopus dupetithouarsii,
which is still extant on the Marquesas Islands; the doves were sometimes seen foraging together in the tops
of tall trees (Holyoak & Thibault 1984). The diet comprised fruits, vines and seeds. The R ed-moustached
Fruit Doves occurred in mountain forests, generally higher up than the W hite-capped, often at the head of
canyons. An immature bird was taken in November (Ripley & Birckhead 1942), so breeding probably
occurred in September and October.
Olson (1986c) discovered a very important and hitherto overlooked manuscript by Andrew Bloxam, made
during the voyage of HMS Blonde in 1825. It contained the only notes made about the birds collected on
Mauke in the Cook Islands, some of which disappeared soon after. Included in his notes is an excellent
description of a dove, of which the type is now lost, that may be an extinct subspecies of Cook Islands Fruit
Dove Ptilinopus rarotongensis. Bloxam’s account (see Olson 1986c) reads:
Length 8¾ Inch. Bill 5/8 inch. Legs red, covered with feathers nearly to the toes. Bill short, brownish. Tongue
entire, sharp pointed. Forehead & top of head a beautiful deep lilac approaching to pink. Hind head & neck all
round & upper part of breast a powdered grey. Upper part of wings, tail, & back, green of various and beautiful
shades & tints. The last ¾ inch of the tail a dusky pale white bar, slightly tinged with green. Wing and tail
beneath pale ash. Lower part of belly and vent yellow. Upper part of belly yellowish, with a slight tinge of pink or
dark lilac in the middle. Red berries were found in its maw.
The Cook Islands Fruit Dove is known from Raratonga (P. r. rarotongensis) and ‘Atiu (P. r. goodwini), of which
the latter differs mainly in the reduced or absent red b
elly-patch (Gibbs et al. 2001), being replaced by a few
orange feather tips (Holyoak 1974a). Olson (1986c) noted that Bloxam’s description of the Mauke bird as
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having a belly ‘with a slight tinge of pink or dark lilac in the middle’ accords better with P. r. goodwini than
with the nominate, as would be expected on biogeographical grounds. There is a possibilty that the Mauke
population may have been subspecifically distinct, and would therefore be cited as Ptilinopus rarotongensis
byronensis.
The genus Alectroenas, the blue pigeons, occurred widely over the islands of the western Indian Ocean, and
according to DNA analysis, is closely related to the New Caledonian C loven-feathered Dove Drepanoptila
holosericea, with the Ptilinopus fruit doves representing the ancestral group (Shapiro et al. 2002; Pereira et al.
2007). Blue pigeons occur in Madagascar, the Seychelles, and Aldabra and the Comoros respectively. They
are closely related, and best regarded as a superspecies (Greenway 1967; Goodwin 1983). In the Mascarenes
there was one species on Mauritius, probably one on Réunion (see below) (Mourer-Chauviré et al. 1999),
and another on Rodrigues (Hume 2011a).
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Specimens
Sonnerat (1782) originally named the bird Pigeon Hollandais – Dutch Pigeon – probably on account of its
blue, white and red plumage, the colours of the Dutch flag (Newton 1879), but he used only French names,
allowing Scopoli (1786) to describe it formally. Two S
onnerat-derived specimens originally came to Paris,
where Temminck (1813) saw them, much the worse for wear, having been exposed to the fumes of sulphuric
acid, which dissolved some of the feathers. By the time Milne-Edwards & Oustalet (1893) made the same
comment in the 1890s, only one Sonnerat specimen remained. A third specimen arrived in Paris in 1800
collected by one Colonel M. Mathieu, which became part of the Dufresne Collection (Cheke & Hume 2008).
Dufresne was a dealer in skins, who had for some time been Conservator of the Cabinet of National History
of the Empress Josephine. The University of Edinburgh eventually acquired the specimen about 1816,
which later became part of the Royal Museum of Scotland. The fourth skin, the last taken and one of the
three surviving specimens, was collected by E. Geoffroy in the Savanne Forest in 1826, and given to Julien
Desjardins (Oustalet 1897), principal founder of the Mauritius Natural History Society; it is still in the
Mauritius Institute.
The Mauritius Blue Pigeon survived alongside humans and introduced predators for over two centuries,
so it was almost certainly deforestation that caused its extinction – the main threat to blue pigeons today
(Baptista et al. 1997). The Mauritius Blue Pigeon was extremely rare by the mid-1820s, but it may have
survived in remote places until 1837 (Hume 2011a). Desjardins (1832) made the last comment about this
species in 1832:
. . . one still finds [the Mauritius Blue Pigeon] towards the centre of the island in the middle of those fine
forests which, by their remoteness, have escaped the devastating axe.
Habits
Milbert (1812) in 1801 was the only observer to make comments about the Mauritius Blue Pigeon in life. He
stated that it lived near river banks and ate fruit and freshwater molluscs. Later authors have doubted this
statement, but Goodwin (1983) remarked that in view of the number of predominantly frugivorous pigeon
genera that contain species known to eat molluscs and other invertebrates (e.g. Ptilinopus, Gallicolumba), he
saw no reason to doubt Milbert’s statement.
Johannes Pretorius in 1666–69 attempted to keep the Mauritius Blue Pigeon alive in captivity, but all
attempts failed (Hume & Winters 2015). He also confirmed that this species exhibited warty carunculations
on its face, as do other Alectroenas species:
The pigeons are beautiful in colour with crests on the head and warts on the face. I have tried to raise juveniles
and to tame adults, but they always die under my hands.
One individual was brought alive in the 1790s to the Netherlands and lived for three months (Tuijn 1969).
Arnout Vosmaer (translated in Tuijn 1969) described it:
These dark-blue [pigeons] with lead coloured head-feathers, which they can turn upwards just as a collar, were
sent to me from the Cape, but originated from an Isle Mauritius. Presented to the court by the Baron I. N. E. van
Lynden 1790, they were called Pavillons Hollandais. One only I have received alive but died later after a few
months from dropsy. In the evening 11 till 12 o’clock and many times during the night it made nice sounds 10 to
12 times quickly after another like Baf Baf [pronounced barf barf], and during the day a kind of cooing sound.
As in other blue pigeons, it probably occurred in pairs or small flocks, moving locally or seasonally in
response to available food sources, inhabited humid, montane evergreen forests, and foraged predomi-
nantly on fruit, never alighting on the ground to feed.
Evidence for the existence of an Alectroenas blue pigeon on Réunion rests entirely on a few rather vague 17th
century accounts (Hume 2011a). The first was a report by the Dutch navigator, Willem Bontekoe (Bontekoe
1646; translated by JPH), who in 1619 reported:
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We found large numbers of doves of the kind which has blue wings. They let themselves be taken by hand, or we
knocked them down with sticks and canes, without their making any effort to fly away. In one day we killed a
good 200, brought them to the fire and boiled and roasted them for the sick as well as for those in health.
Alectroenas is the only b
lue-winged pigeon genus, and as two n
ow-extinct blue pigeon species once occurred
on the neighbouring islands of Mauritius and Rodrigues, it is likely that Réunion once harboured one as
well (see Mourer-Chauviré et al. 1999; Hume et al. 2006). Dubois (Dubois 1674) in 1671–72 may have also
observed them, and this is the last time that they were reported. The Réunion Blue Pigeon had probably
disappeared by around 1700 due to o ver-hunting and predation by introduced cats, as in 1704 the compe-
tent observer Feuilley (1705) failed to mention them.
How the Réunion species differed from other Alectroenas species is not known. Because the distinctive
white patterning seen in insular blue pigeons from the Seychelles, the Comoros and the Mascarenes was not
mentioned, it may have been more similar to the Madagascar species, A. madagascariensis, which is gener-
ally all blue (Hume et al. 2006; Cheke & Hume 2008).
A single tarsometatarsus and a possible femur referable to the genus Alectroenas has shown that a species of
blue pigeon once occurred on Rodrigues (Hume 2011a). It was larger than any of the extant Alectroenas
species, but smaller than its nearest relative, the Mauritius Blue Pigeon A. nitidissima. It was not mentioned
by the competent observers Leguat in 1691–93 (Leguat 1708) or Tafforet (1725–26) in 1725–26. The island
was infrequently used as a refurbishment station by various shipping nations from at least 1601 (Cheke &
Hume 2008), so it is possible that the species became extinct before Leguat arrived in the 1690s. Leguat
noted the presence of large numbers of rats when he arrived on the island, and these would have been
formidable predators.
Being restricted to Rodrigues, which measures just 18km long by 6.5km wide, the Rodrigues Blue Pigeon
would have been extremely susceptible to any anthropogenic changes to the island.
While working in the India Office Records administered by the British Library in London, D. R. Stoddart (in
Stoddart & Benson 1970) came across the following passage in an unpublished manuscript by a Moresby,
written in 1821–22:
Jean de Nova i.e. Farquar and Providence . . . like the Amirantes, Coetivy and Alphonse are the resort of Millions
of Birds of which, the Frigate Bird, the Fou, a beautiful small white gull, a variety of various coloured Gannet,
and the Tropic Bird are the principle: in S. Pierre and Providence a species of small blue pigeon are in great abun-
dance, and so seldom disturbed that they do not fly at Man’s approach, but are knock’d down with Sticks, we
found them excessively good eating, these birds build and nest on the Mapou tree and other Dwarf trees which
cover the surface of the islands . . .
Stoddart & Benson add:
The ‘small blue pigeon’ must have been a species of Blue Pigeon Alectroenas. We know of no other reference to
its occurrence on Farquhar, Providence or St. Pierre. Stoddart spent 19 September 1968 on Farquhar, traversing
most of the atoll land . . . but saw no blue pigeons.
Unless subfossil remains are discovered, it is impossible to say whether the pigeon of these islands was
distinct, or that it was referable to an extant Alectroenas species.
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Status Extinct. Known from Holocene subfossil remains. Date of extinction unknown.
Range Viti Levu, Fiji.
The discovery of a giant flightless pigeon, almost rivalling the Dodo Raphus cucullatus and Solitaire
Pezophaps solitaria of the Mascarene Islands in size, is one of the more exciting discoveries of recent years. It
is the largest known pigeon from the Pacific region, and it shows that flightless pigeons were not confined
to the Indian Ocean (Worthy 2001).
The Viti Levu Giant Pigeon was closely related to the Goura crowned pigeons, which are terrestrial but
have retained the ability to fly; however, it dwarfed them in size, except in the dimensions of the wing.
These were disproportionately small, thus the pigeon was totally flightless, and would have nested and
foraged on the forest floor. The bill was large and deep, suggesting that it was quite capable of taking large
fruit, seeds and invertebrate species. Upon the arrival of people on the island, the Viti Levu Giant Pigeon
would have been extremely vulnerable to o ver-hunting, and as for the Dodo and Solitaire, introduced
mammalian predators would have been serious predators of eggs and chicks.
The imperial pigeons Ducula spp. are large to very large arboreal forest birds. Quite capable of covering
large distances over water, they feed almost exclusively on fruit (Gibbs et al. 2001). They have suffered from
over-hunting and deforestation over much of their range, and island species are particularly vulnerable.
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The Lakeba Imperial Pigeon was larger than the largest extant imperial pigeons, Marquesas Imperial
Pigeon D. galeata and New Caledonian Imperial Pigeon D. goliath. The proportions of the limbs also differed,
being comparatively longer in D. lakeba (Worthy 2001), and not the typical short, stout tarsi seen in arboreal
pigeons. Long tarsometatarsi in pigeons are often characteristic of more terrestrial groups e.g. Gallicolumba,
Microgoura but it is impossible to say with any confidence that this was the case with this species.
Holyoak & Thibault (1978a) proposed that the genera Ducula and Ptilinopus are allopatric throughout the
Pacific, yet the discovery of the Lakeba and ‘Uvea species (see below), which were sympatric with Pacific
Imperial Pigeon D. pacifica, suggests this is an artefact of negative human influence (Balouet & Olson 1987).
The Wallis Islands are situated 385km west of Samoa and 385km n orth-east of Fiji, which gives an idea of
the ability of the imperial pigeons to colonise remote islands. The ‘Uvea Imperial Pigeon was sympatric
with the Pacific Imperial Pigeon Ducula pacifica, but it differed greatly in its larger size, being just as large as
the Lakeba Imperial Pigeon D. lakeba (Balouet & Olson 1987; Worthy 2001). Unlike this species, however, the
tarsometatarsus was not long, but short and robust, as in D. pacifica. The presence of the ‘Uvea Imperial
Pigeon in archaeological middens indicates that these birds were regularly eaten, and it is likely that
over-hunting was the sole cause of extinction.
This was larger than any extant imperial pigeon, and equal in size to the extinct ‘Uvea Ducula david and
Lakeba D. lakeba Imperial Pigeons. The discovery of the Henderson Island Imperial Pigeon has resolved the
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disjunct range of large Ducula pigeons within the Pacific, with the largest species occurring in New
Caledonia and the Marquesas Islands, and none in between (Worthy & Wragg 2006). The former presence of
Ducula species on ‘Uvea Island and on Lakeba, Fiji, plus a number of a s-yet undescribed species, indicates
that their present distribution is an artefact of human impact.
Unusually for a Ducula pigeon, it seems the Henderson Island Imperial Pigeon had reduced wing
elements and long legs, suggesting that it was a terrestrial species, with poor flying ability. This would have
made it extremely vulnerable to human hunting pressure and predation by introduced mammalian
predators.
The Spotted Imperial Pigeon Ducula carola is divided into three well-differentiated races endemic to the
Philippines, where the species is rare and declining due to severe deforestation and over-hunting (Gibbs et
al. 2001). The Negros subspecies D. c. nigrorum occurred on Negros and Siquijor. According to Ripley &
Rabor (1956) it was considered common on Negros in both the 1890s and 1950s, but was not recorded in
1991 (Collar et al. 1999). On Siquijor it is known from a single specimen taken in 1952, and has never been
seen since. Both populations may now be extinct.
Habits
The Spotted Imperial Pigeon is gregarious and nomadic, occurring in flocks of up to ten birds, but also
singly and in pairs. It travels long distances in response to food availability. On Negros it preferred lowland,
closed-canopy primary forest from near sea level in September to November up to an altitude of 2,000m in
April to May (Gibbs et al. 2001), but it also inhabited selectively logged forest. It fed on a variety of fruit, and
large numbers of birds would gather on fruiting trees. The breeding season probably occurred from
February to July, but a young bird was taken in December on Siquijor.
The New Zealand Pigeon or Kererū was originally divided into three subspecies, but these have now been
given full species status; Hemiphaga novaeseelandiae on North and South Islands, and smaller offshore islands
on New Zealand, H. chathamensis on the Chatham Islands, and H. spadicea on Norfolk Island (Holdaway et
al. 2001). There may once have been a population on Lord Howe Island (see p. 378), and another on Raoul
Island (see p. 189).
Norfolk Island was first discovered by Captain Cook in October 1774, after which Phillip Gidley King
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arrived from Sydney on 6 March 1788 to set up a penal settlement. Convicts were imported periodically
until June 1856, when descendants of the Bounty mutineers were relocated from Pitcairn to Norfolk and
Philip Islands. Cattle had already been introduced, and these settlers brought sheep, horses, pigs, rabbits
and poultry. Black Rats Rattus rattus also became established and appear to have exterminated the Pacific
Rat R. exulans, indicating that the island had previously been visited, but not settled, by Polynesians.
The pigeon
Early settlers called the Norfolk Island Pigeon the ‘Wood Quest’ (Moore 1985), but few observations seem to
have been made of the bird. Hunter (1793) wrote:
On our first landing, we found a great number of pigeons that were so tame, that we knocked them down with
sticks; but latterly they quitted the low boughs, and generally harboured about the tops of the pines: when
plucked and drawn, they weighed from three-quarters to one pound each.
Hunter’s account infers that even as early as the 1790s, the pigeons had learnt to avoid humans, ultimately
to no avail.
In 1838, Ensign Best (in Moore 1985) mentioned that he and one of his companions killed at least 72
pigeons during 17 hunting expeditions, with the largest bag of 25 taken by two men on September 18, 1838.
Moore (1985), followed by Hume & Walters (2012) thought this was a reference to the Norfolk Island Pigeon,
but it appears not to be the case. Forshaw (2016) suggests that these pigeons were probably feral pigeons or
Rock Doves Columba livia, which had been introduced as a food source. Best (in Forshaw 2016) further
commented:
Nov. 1, 1838. . . . A man we met offered to show me a pair of fine Wood Quests. We went with him, found them,
and I killed one, the other flew away. Giving up the chase of the truant we descended the cliffs to a small bay of
which there was an island abounding with sea birds . . . Ascending the cliffs . . . in another attempt I killed the
Wood Quest.
March 21, 1839. I made one of a party who went in search of Wood Quests. We left the settlement with Storey for
our guide about eleven and went first to Cascade, from hence I went along the road to Mount Pitt (Lugard and
Storey keeping in the gullies to the right) and reached the summit without seeing a bird in the Wood Quest shape
. . . On our road homeward the order of march was changed, Tyssen and I going with Storey while the others kept
the road. We had not gone far when up a tree which the other two had passed I heard a Wood Quest’s cry and it
was killed. This was the only one we found.
Best’s accounts show that the Norfolk Island Pigeon had become extremely scarce by the late 1830s (see
Forshaw 2016). The rapid disappearance of the pigeon was thought to have been corroborated by Peale
(1848), who failed to find it when he visited the island in 1839 with the United States Exploring Expedition,
but it appears that Peale never actually visited Norfolk Island at this time. Therefore, the last unequivocal
mention of the bird was by Ensign Best on March 21, 1839 (Tennyson & Martinson 2007; Forshaw 2016).
Over-hunting, deforestation and predation by introduced mammals, especially cats, are the likely causes
for this bird’s extinction (Forshaw 2016).
Captive birds
A number of Norfolk Island pigeons arrived alive in England. A report of living birds was made by Vigors
(1833) in 1833, when he received a pair at the Zoological Society, London. What happened to these birds is
not known. Furthermore, Forshaw (2016) mentions two captive individuals that were held in the aviaries of
the Earl of Derby, a female that died on December 9, 1835, and a male on December 28, 1836. The latter was
the last record of the bird in captivity.
Habits
Backhouse (1843) reported on the Norfolk Island pigeon’s diet, and the ease at which they could be caught.
When describing the island on March 14, 1835, he stated:
In woody gullies, the Norfolk Island Cabbage-tree Areca sapida [Norfolk Island Palm Rhopalostylis baueri]
abounds . . . The seeds furnish food for the Wood-quest, a large species of pigeon, which has a bronzed head and
breast, and is white underneath, and principally slate-coloured on the back and wings. This bird is so
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unconscious of danger as to sit till taken by a noose at the end of a stick; when one is shot, another will sometimes
remain on the same bough, till itself is also fired at.
Forshaw (2016) points out that Best’s description of hearing a ‘Wood Quest’s cry’ is unusual, as Hemiphaga
pigeons are usually silent.
The New Zealand Pigeon or Kererū Hemiphaga novaeseelandiae is known to have colonised remote Norfolk
Island and the Chatham Islands, with each population given subspecific status. A population on Lord Howe
Island (see below) was also probably subspecifically distinct, although no specimen exists. A single humerus
referable to this genus was discovered on Raoul, in the Kermadec Islands (Worthy & Brassey 2000), which
may also represent an endemic subspecies. It was extinct by 1887 when the curator of Auckland’s Museum,
T. F. Cheeseman, visited the island (Cheeseman 1891), but he obtained some details about the bird from the
islanders:
The earlier settlers on Sunday Island found a large fruit pigeon very abundant on their first arrival; but its
numbers were gradually thinned, and it was finally exterminated, partly by settlers themselves and partly by the
wild cats introduced by them. A Mr. Johnson, who resided on the island about fifteen years ago, states that it
exactly resembled the New Zealand species in size and colour, and he has no hesitation in considering it to be the
same.
It was exterminated due to over-hunting and predation by cats, but nothing else is known about it.
There is some confusion about the distribution and taxonomy of the Norfolk Island Kaka. All of the speci-
mens with provenance data came from Philip Island, but two species were described, N. productus and N.
norfolcensis, based on variations in the plumage, and in the case of N. norfolcensis on a specimen with an
abnormal deformity of the bill (Pelzeln 1860a; see p. 429). This led some ornithologists, most notably
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Rothschild (1907a), to speculate that N. norfolcensis might have been an inhabitant of Lord Howe Island. All
of these wild assumptions have fortunately now been quelled, and as the plumage of the Norfolk Island
Kaka is extremely variable and bill deformities were apparently not uncommon in these parrots, N. norfol-
censis has been rightfully synonymised under N. productus.
The Norfolk Island Kaka was first mentioned by J. R. Forster on Cook’s second voyage to the South
Pacific, after the discovery of Norfolk Island in October, 1774 (Forster 1844). A penal colony was established
from 1788 to 1814, and again from 1825 to 1854 (Hazzard 1984), and it appears that the Norfolk Island Kaka
declined rapidly due to over-hunting. It was sometime during the second period that the last wild birds
disappeared, as a comprehensive ornithological survey by Abel Best in 1838–39 (Moore 1985) failed to
mention it. The bird may have survived longer on Philip Island (Gould 1865), but the habitat was destroyed
by the introduction of rabbits, pigs and goats (Coyne 2010). The ease with which the Norfolk Island Parrot
could be killed, and the fact that it was probably en route to becoming behaviourally flightless, is reported
by Gurney (1854):
I have seen the man who exterminated the Nestor productus from Philip Island, he having shot the last of that
species left on the island; he informs me that they rarely made use of their wings, except when closely pressed:
their mode of progression was by the upper mandible; and whenever he used to go to the island to shoot, he would
invariably find them on the ground, except one, which used to be sentry on one of the lower branches of the
Araucaria excelsa [Norfolk Island Pine Araucaria heterophylla], and the instant any person landed, they would
run to those trees and haul themselves up by the bill, and, as a matter of course, they would remain till they were
shot, or the intruder had left the island. He likewise informed me that there was a large species of hawk [Brown
Goshawk Accipiter fasciatus] that used to commit great havoc amongst them, but what species it was he could
not tell me.
A breeding population of Brown Goshawk was extirpated on Norfolk Island during the late 18th century,
and sometime prior to the 1850s on Philip Island (Tennyson & Martinson 2006).
The last specimen of this species, a captive bird, died in a cage in London sometime after 1851 (Greenway
1967). Gould (1865) spoke of its plight sometime in the 1850s:
I regret to state that the native haunts of this fine bird have been so intruded upon, and such a war of extermina-
tion been carried on against it, that it is now entirely extirpated, and a few stuffed specimens are the only
mementos of its having existed. Until lately it still lived on Philip Island (an islet lying off Norfolk Island), but
in this small domain it is no longer to be found.
Habits
Almost nothing is known about the biology of this parrot. Gould (1865) provided the only details:
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During my stay at Sydney [around 1838] I had an opportunity of seeing a living example in the possession of
Major Anderson, and was much interested with many of its actions, which were so different from those of every
other member of the family, that I felt convinced they were equally different and curious in a state of nature. This
bird was not confined to a cage, but permitted to range over the house, along the floors of which it passed, not
with the awkward waddling gait of a parrot, but in a succession of leaps, precisely after the manner of the
Corvidae. Mrs. Anderson, to whom I am indebted for the little I could learn respecting it, informed me that it
was found among the rocks and upon the loftiest trees of Philip Island, that it was so tame as to be readily taken
alive with a noose, and that it fed upon the blossoms of the w
hite-wood tree, or white Hibiscus, sucking the honey
of the flowers: the mention of this latter circumstance induced me to examine the tongue of the bird, which
presented a very peculiar structure, not like that of the true h oney-feeding Parrakeets . . . with a b rush-like
termination, but with a narrow horny scoop on the under side, which, together with the extremity of the tongue,
resembled the end of a finger with the nail beneath instead of above: this peculiarity in the structure of the organ
is doubtless indicative of a corresponding peculiarity in the nature of the food upon which the bird subsists. Mrs.
Anderson told me that it lays four eggs in the hollow part of a tree, but beyond this I was unable to ascertain
anything respecting its nidification.
The call was described as a ‘hoarse, quacking, inharmonious noise, sometimes resembling the barking of
a dog (Tennyson & Martinson 2007).
The Kea Nestor notabilis was supposedly identified from the vast number of subfossil bones collected on the
Chatham Islands by Forbes in the 1890s (Forbes 1892b). However, further study by Dawson (1959)
concluded that they actually represented the Kaka N. meridionalis, indicating that this species was able to
colonise remote island groups, including Norfolk Island, Philip Island, and doubtfully Lord Howe Island
(see p. 430). Millener (1999) re-examined the material, and showed that the Chatham Islands Kaka was
indeed an endemic, undescribed species, and suggested that it had reduced flying capabilities, hitherto
known among parrots only in the Norfolk Island Kaka Nestor productus, Kakapo Strigops habroptilus and the
Broad-billed Parrot Lophopsittacus mauritianus of Mauritius (Hume 2007).
Using molecular and morphological evidence, Wood et al. (2015) confirmed that the Chatham Islands
Kaka was a distinct taxon, and sister to the New Zealand mainland Kaka N. meridionalis. It was about the
same size as the latter, but had larger leg elements and a larger pelvis. Wood et al. did not find that the
Chatham Islands Kaka had reduced flying capabilities (contra Millener 1999), but that the increase in size of
the pelvic and leg elements was probably a result of more time spent feeding on the ground. Isotopic anal-
ysis indicated that the Chatham Islands Kaka was mainly herbivorous, but it may have also seasonally
preyed on the eggs and chicks of other Chatham Island birds. It was widespread in the islands and presum-
ably mainly an inhabitant of forests, which were extensive prior to Polynesian occupation (Tennyson 2015),
but being a generalist it probably was capable of taking advantage of other environments.
The terrestrial habits of the Chatham Islands Kaka and its probably tame and confiding nature would
have made it vulnerable to introduced predators, and it appears to have succumbed to Polynesian over-
hunting, deforestation and predation of its eggs and chicks by introduced Pacific Rat Rattus exulans at some
point between the 13th and 16th centuries (Wood et al. 2014).
Conuropsis carolinensis (p. 186) occurred as far north as Virginia, and the Macquarie Island Parakeet
Cyanoramphus novaezelandiae erythrotis (p. 170) inhabited the sub-Antarctic island of Macquarie. This family,
along with the Columbidae and Rallidae, has suffered more losses than any other bird group. Demand for
the pet trade, hunting for food and habitat destruction have brought many species close to extinction.
Parrots and macaws endemic to islands have been particularly vulnerable, where a disputed number of
species have become extinct.
The Cebu Hanging Parrot disappeared due to the almost complete deforestation of Cebu in the 19th century.
In 1873, it was still common in woods near Toledo, but even then the forest was rapidly disappearing (Rabor
1959), and only small patches of forest survived. Bourns & Worcester (in McGregor 1909a), who searched
for it in 1888, found it very rare, and after several days’ work in the great coastal groves near Carmen found
only a single specimen.
In 1906, when McGregor and his assistants were collecting (McGregor 1909a), they had difficulty in
finding any forest at all. Although some r e-forestation has taken place, Rabor (1959) considered that by 1947
no primary forest remained. As no serious ornithological study was conducted on Cebu between 1907 and
1947, the date of extinction of all the Cebu endemics has been arbitrarily set as 1906. However, subsequent
studies (e.g. Brooks et al. 1995; Collar 1998) have shown that not all the birds thought to have disappeared in
1906 have in fact done so, and that Rabor was unduly pessimistic.
Rand (1949) reported the existence of a number of specimens of Cebu Hanging Parrot in captivity, which
had been caught in 1929, and said that some of these survived until 1943 in London Zoo. Unconfirmed
reports from 2004 are in all likelihood referable to other subspecies introduced to the island, as hanging
parrots are regularly captured for the bird trade and transported between the islands (Forshaw 1989). The
endemic Cebu race is almost certainly extinct.
Habits
The hanging parrot inhabited primary forest of the lowlands and hills, though it also occurred in coconut
groves and in secondary growth. Like other races of the species, it probably fed in the upper storeys of the
forest on blossoms, nectar fruits and seeds.
Little was recorded about this bird. Hachisuka (1934) remarked that most of the original forest had been
cleared from the island, and that the hanging parrot subsequently appeared in coconut groves, fruit trees
and bamboo clumps. It seems to have disappeared due to deforestation, and perhaps due to collection for
the pet bird trade. Rumours that it might still survive have not been substantiated; a supposed specimen
collected in 1954 proved to be an escaped cage bird (Rand & Rabor 1960), and recent observations are prob-
ably of escaped subspecies from elsewhere.
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Since its discovery, the little-known Sangihe Red-and-blue Lory appears to have been rare, and was still
considered so in the late 1970s (Forshaw 1989). Sangihe’s lowland areas have been cleared for coconut plan-
tations, and the lories were forced into mountain forests (Meyer & Wiglesworth 1898). Heavy trapping of
birds around 1950 then reduced the remaining population to very low numbers (Collar et al. 2001). Searches
between the late 1970s and 1986 failed to locate any birds (Forshaw 1989), but small populations were
discovered in 1995 (Juniper & Parr 1998). However, these appear to be hybrids between Sangihe birds and
escapees of the Talaud subspecies E. h. talautensis. It is probable that no pure-bred Sangihe Red-and-blue
Lories survive today.
Habits
These were social birds that sometimes roosted in large flocks. They occurred in forest and cultivated lands.
Their diet consisted of pollen, nectar, fruit and insects, and they nested in hollows high in tall trees (Juniper
& Parr 1998).
Sinoto’s Lorikeet was larger than any known Vini species. There appears to be size variation between speci-
mens from different islands, with birds being larger on Hiva Oa and Tahuata than on Ua Huka (Steadman &
Zarriello 1987).
Sinoto’s Lorikeet was sympatric with the Conquered Lorikeet V. vidivici (see below) and the extant (but
endangered) Ultramarine Lorikeet V. ultramarina on Tahuata and Ua Huka at least. Their presence in a
number of archaeological sites suggests that the two species became extinct soon after the arrival of the
Polynesians.
The Conquered Lorikeet was larger than any extant Vini species, but smaller than the sympatric Sinoto’s
Lorikeet V. sinotoi. The introduction of the Pacific Rat Rattus exulans by Polynesians may have impacted on
the populations of both these lorikeets. The extant Ultramarine Lorikeet V. ultramarina is also threatened by
predation from introduced Black Rats Rattus rattus (Juniper & Parr 1998).
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The type and other specimen of this lorikeet were collected before 1860, and there have been no subsequent
confirmed sightings. Sarasin (1913) said that, at the time of writing, the bird was believed to survive in the
northern forests near Oubatche. According to Greenway (1967), MacMillan failed to find it when collecting
for the American Museum of Natural History, but no date is given for this expedition. A report by a local of
two birds having been seen west of Mount Panié was made to Tony Stokes in June 1976 (in Juniper & Parr
1998), but no further information exists and the record remains unconfirmed.
The New Caledonian Lorikeet inhabited the upper levels of dense forest; otherwise nothing is known of
its habits. The reason for the species’ disappearance is unknown.
The Red-fronted Parakeet Cyanoramphus novaezelandiae is a polytypic species which ranges over New
Caledonia, New Zealand and the outlying Kermadec, Chatham and Antipodes Islands, and Auckland
Island (Juniper & Parr 1998). The Macquarie Island Parakeet inhabited the rather inhospitable, treeless,
sub-Antarctic island of Macquarie. It was discovered early in the 19th century by sealers, who killed the
birds in large numbers for food (Taylor 1979).
Remarkably, the parakeets survived this onslaught, as well as predation from introduced cats and dogs,
and were still considered common in 1820. Some were taken alive to Sydney, Australia, by sealers (Porter
1934). A Russian expedition under Bellinghausen visited the island in 1820, when 20 specimens were
collected by naturalists for museums, including one live bird (Taylor 1979). They were said to still be abun-
dant in 1877 and 1880, but disappeared very suddenly during the following decade. It is likely that the bird
became extinct for reasons similar to those of the Macquarie Island Banded Rail Hypotaenidia philippensis
macquariensis (see p. 102) – the introductions of the Weka Gallirallus australis and domestic rabbits (Taylor
1979). The last definite sighting of the Macquarie Island Parakeet was in 1890.
Habits
There are no trees on Macquarie Island, so the birds nested under bunches of tussock grass on the ground or
in burrows (Cumpston 1968). They were particularly common on the shore, feeding on invertebrates living
in heaps of seaweed (Taylor 1979) and also on various seeds. Little else seems to have been recorded of their
habits.
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Salvadori (1891b) described this bird from a skin specimen, brought back from the voyage of the Herald, one
of a number of ships sent out to locate the ill-fated HMS Erebus and HMS Terror, the two ships disastrously
led by Sir John Franklin while searching for the North-West Passage.
Hindwood (1940) claimed that the extinction of this parakeet was brought about by the islanders, who
shot and trapped the birds because of the reputed damage they did to the gardens and crops. The islanders
were entirely reliant on crop-growing for subsistence and trade with passing ships, and the parakeets were
extremely destructive. Robert Etheridge (Etheridge 1889), in the report of the Australian Museum expedi-
tion to Lord Howe in 1887, remarked:
the island was infested in former years by a parakeet . . . said to have existed in very large numbers, doing
considerable damage to crops, and having gradually disappeared about ten years ago.
Hill (1870) observed a solitary pair in 1869, which proved to be the last record of their occurrence, and
stated:
The paraquet . . . was a nuisance to the cultivators, once appearing in flocks; now I saw but a solitary pair in
their rapid flight through the foliage, and recognised them only by their peculiar noise.
A similar species, C. cookii, still survives on Norfolk Island; both were once considered conspecific with
the New Zealand Kakariki C. novaezelandiae, but they are now generally accorded specific status (McAllen &
Bruce 1988; Christidis & Boles 2008).
The Lord Howe Parakeet occurred in forested habitats and areas of cultivation (Higgins 1999), and prob-
ably fed on seeds, fruits, buds, and leaves of native trees (Garrett et al. 2010).
The ability of Cyanoramphus parakeets to reach isolated islands and island archipelagos is remarkable for
such a small bird, but they are extremely vulnerable to human disturbance, especially from the introduction
of predatory mammals. Holdaway et al. (2010) discovered a single coracoid referable to a Cyanoramphus
parakeet on the New Zealand subantarctic Campbell Island, which strongly suggests that a resident popu-
lation may once have occurred there. No skins were collected, nor observations made of the bird, but as the
Brown Rat Rattus norvegicus was possibly introduced in 1810, when the island was first discovered, it would
not have taken this vermin long to exterminate a population of parakeets. When Robert McCormick,
surgeon on the ill-fated Erebus and Terror expedition (see Lord Howe Parakeet above), visited Campbell
Island in 1840, he reported that no land birds were present (Miskelly 2000). This suggests that the parakeet
had disappeared in the intervening years. It is not known how the Campbell Island Parakeet differed from
other species; further subfossil remains are needed to confirm its taxonomic status.
Nothing is known of this parrot’s habits or why it became extinct. Both surviving skins of this parrot are
believed to have been collected by Georg Forster in 1773 or 1774 during Captain Cook’s second voyage
(Stresemann 1950). However, Medway (2009), based on Joseph Banks’s travel diary, thought that they were
collected in November 1777 during Cook’s third voyage. It seems likely that no collecting took place on
Raiatea in 1768 on Cook’s first voyage, as the native people were very hostile, so Forster probably obtained
both specimens on the second. There is some confusion as to the locality, however, but Forster (1844) refers
to a ‘Psittacus pacificus’ seen in Otaheite (Tahiti) and Oriadea (Raiatea), from which Greenway (1967)
presumed that he found and collected Cyanoramphus zealandicus on Tahiti (see below) and C. ulietanus on
Raiatea. Uncharacteristically, Forster did not regard the two species as different.
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Locality
That the Black-fronted Parrot once occurred on Tahiti is beyond doubt; Greenway (1967) states:
Neither Cook’s nor Banks’ published diaries mention parrots on Tahiti itself . . . yet there can be no question that
C. zealandicus existed there. Indeed Forster’s remark in his ‘Observations’ . . . to the effect that the Tahitians so
prized the crimson parrot feathers brought from the Friendly Islands (Tonga) that they gave their treasured pigs
for them might lead one to believe that there was no green and red parrot on Tahiti; the remark has no such
significance, nor apparently has his silence on the subject of parrots in the Society Islands.
It has long been extinct; there is no tradition of a green and red parrot on the island (Greenway 1967). As
the locals apparently greatly prized the feathers, over-hunting may have been at least partly responsible for
its disappearance.
Habits
According to Des Murs (1845, 1849), Marolles shot three birds on Tahiti in 1844, at Port Phaeton on the
isthmus of Taravao. The bird was rare at that time, and only to be found on the isthmus and in the moun-
tains of Tahiti-iti. Marolles saw only about four or five individuals in total, and the locals told him that the
parrots lived in big trees on inaccessible scarps and deep valleys. Nothing else is known about it.
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The Paradise Parrot was a beautiful grass parakeet, which appears to have been restricted and declining
even before European settlement (Forshaw 1989). It was susceptible to droughts, suffering particularly from
a severe one in 1902, which may have affected food availability, and by 1915 it was thought to have disap-
peared. However, after careful searches, a few more birds were located in 1918 (Chisholm 1922), and in
1922, Henry Jerrard (see Olsen 2007) took a series of famous photographs of a nesting pair.
The bird was last recorded with certainty in November 1927. Although a number of sightings have been
recorded, including five birds in 1990 (Kiernan 1993), the Paradise Parrot is generally considered extinct.
Various reasons have been suggested for its extinction, which include habitat alteration (drought,
over-grazing, the increase of fire, the spread of introduced prickly pears, and clearance of Eucalyptus trees),
and also disease, trapping for the pet trade, and predation by introduced species.
Their habit of commonly inhabiting roads and tracks made them easily accessible to trappers (Chisholm
1924), and large numbers were exported overseas (Higgins 1999). Alex Chisholm, who started a publicity
campaign to preserve the bird, initiated what was to become Australia’s first survey of an endangered
species in 1916 (Crome & Shields 1992). In 1927, plans to start a captive breeding and release programme
was discussed, but by this time the parakeet was effectively extinct (Anon 1927).
Habits
The Paradise Parrot inhabited open savanna woodland and shrubby grassland, and was particularly associ-
ated with river valleys (Higgins 1999). It lived in pairs or small family groups, with nests made in
hollowed-out termite mounds, or occasionally in a river bank or tree hollow, and often at or near ground
level.
Chisolm (1924) observed a pair feeding:
The method of feeding was to seize a grass bent near its base and run it through the bill until the seed was
reached. In odd cases the grass-stem was thick; then the bird would climb on and bend it down with weight. They
kept fairly close together all the time, and did not speak. Eventually, there was a noise in the distance, whereupon
the male gave a sharp little call and the two flew off in company, disappearing among trees one hundred yards
away.
Four to five eggs were laid on a mat of crumbled earth, and only the female incubated the clutch. Diet
consisted almost entirely of the seeds of grasses and herbaceous plants. The flight of this parrot was consid-
ered rapid and undulating, but it spent almost all of its time on the ground.
The Pacific Islands Eclectus Parrot was larger and more robust in the cranium and leg elements but had
comparatively smaller wings than the extant Eclectus Parrot Eclectus roratus (Steadman 2006c). The latter
occurs on the Solomon Islands and Papua New Guinea west to the Moluccas (Juniper & Parr 1998), so E.
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infectus establishes the genus east of the Solomon Islands. It may have survived until the 18th century, as a
bird described as ‘all green’, which is indicative of a male, was drawn in 1793 in a report on Vava’u, Tonga,
by the Malaspina expedition (Olson 2006). It was never seen again, so presumably died out due to
over-hunting and predation from introduced mammals.
Thirioux’s Grey Parrot was described from subfossil remains collected by Etienne Thirioux on Mauritius in
the early part of the 20th century (Holyoak 1973). Initially it was placed in the genus Lophopsittacus, and
considered to be a small parrot related to the Broad-billed Parrot L. mauritianus (see p. 205). Re-examination
of the material, however, has indicated that it was in fact a grey, long-tailed P sittacula-derivative (Hume
2007), sympatric with but larger than the extant Echo Parakeet P. echo.
They were particularly abundant, easy to catch, and most sought after as game:
. . . some of the people went bird hunting. They could grab as many birds as they wished and could catch them by
hand. It was an entertaining sight to see. The grey parrots are especially tame and if one is caught and made to
cry out, soon hundreds of the birds fly around ones’ ears, which were then hit to the ground with little sticks.
Despite this persecution, grey parrots remained reasonably common until the 1750s, but the population
must have crashed shortly afterwards, as they were last mentioned in 1759 (Cossigny 1764). It was during
the 1730s that the French instigated large-scale slash and burn forest clearance and this no doubt had a
serious effect on tree-cavity nesting species, including parrots.
A grey parrot, probably conspecific with Thirioux’s Grey Parrot, was mentioned on Réunion by a number
of observers, but as yet no fossil remains have been discovered to determine its taxonomic status. Like its
Mauritian counterpart, it was common and regularly hunted for food, but also appears to have been also
persecuted for damage to crops. Dubois (1674) said of it in 1671–72:
. . . grey parrots, as good [to eat] as the pigeons & sparrows [fodies, Foudia delloni], grey parrots, pigeons and
other birds, bats [Pteropus fruit bats], do plenty of damage, some to cereals some to fruit.
It became extinct sometime after 1732 (Cossigny 1732–1755) and it appears that no live or dead bird was
ever taken out of Réunion.
The Seychelles Parakeet was an insular representative of the Alexandrine Parakeet Psittacula eupatria, which
occurs throughout the Indian subcontinent and South-east Asia and with which it has often been consid-
ered conspecific. However, osteological and molecular studies of the Seychelles bird confirm its
distinctiveness (Hume 2007; Kundu et al. 2011). Jackson et al. (2015) further showed that it was sister-species
to the Alexandrine Parakeet. The Seychelles Parakeet was reported on three of the main Seychelles islands,
Mahé, Praslin and Silhouette (Skerrett et al. 2001), and it was considered to be abundant in 1811 (Prior 1820).
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Newton (1867b) was told in 1866 that the bird had been practically exterminated because of its taste for
ripe maize, and remarked:
The cocoa-nuts are now planted more than halfway up the mountain, and it is probable that in ten years none of
the native forests will remain . . . and here we saw the ‘Cateau vert’ [P. wardi] at the edge of the forest, in a place
some 600 or 700 feet high, where was a patch of maize; but they had been so often fired at that they would not
come within shot.
Newton never saw this green parakeet on Mahé, but whilst there he was informed that:
The ‘Cateau vert’, from the constant persecution against it brought on by its unfortunate partiality for ripe
maize, was said to be nearly exterminated.
He also suggested forest destruction as a partial reason for its demise, but Greenway (1967) pointed out that
Mahé rises almost perpendicularly from the sea to a height of 610m, and that much forest must have
remained on the slopes. But Mahé is a small island, only 20km2 in area. Birds were still present in 1870, at
which time some skins were sent to Cambridge University. In June 1881, H. M. Warry collected skins and
Marianne North (North 1894; see below) described and illustrated two birds from Silhouette in 1883,
including a juvenile (Cheke 2013c). These were the last known from that island, and the last record was of a
bird shot by Abbott on Mahé in March 1893 (Ridgway 1895; Skerrett et al. 2001). Nicoll (1908) made no
mention of seeing them in 1906, so presumably they had disappeared by that time.
Vesey-Fitzgerald (1940, 1941) spent some time on the Seychelles between 1931 and 1936, but did not find
the species. Regardless, it was still considered to be possibly extant, as Peters (1937 [1932–48]) wrote:
‘Formerly abundant in the Seychelles; now confined to the island of Silhouette. Possibly extinct’. However,
it is highly unlikely that the bird survived after 1906.
Habits
The Seychelles Parakeet was a bird of native forest, but adapted to open, cultivated areas once forests were
cleared. It occurred in flocks, and its diet included fruits (Skerrett et al. 2001). Prior (1820) gave a brief
account:
Several birds of handsome plumage are found in the woods. Among others a considerable number of green
parrots, not remarkable for their imitative powers . . .
Around the end of 1883, the botanical illustrator Marianne North painted and described a captive pair of
Seychelles Parakeets when visiting Mahé, the largest island in the Seychelles (North 1894). The birds had
been brought from neighbouring Silhouette, and North’s rather scathing remarks about their stupidity is
more a reflection of many island birds’ inherent tameness. This account is little-known, yet it is the only one
mentioning these parakeets in captivity:
I went one day to their house [a Doctor B and his wife], and painted their parrots, which came originally from
Silhouette: queer, misshapen birds, with enormous beaks and patches of red and yellow badly put on, one of them
having a black ring round its neck [male]. Both were quite helplessly bullied by common pigeons, which came
and ate their food, while they jabbered in a melancholy way, and submitted. They had absolutely no tops to their
heads, which perhaps accounted for their stupidity. They had a stand on the back verandah, where they slept and
fed. They were not tied up, but went and stole their own fruit off the neighbouring trees.
The Rodrigues Ring-necked Parakeet was first mentioned by Leguat (1708) in 1691–93, who stated:
There are an abundance of green and blew Parrots, they are of a middling and equal bigness; when they are
young, their Flesh is as good as that of young Pigeons.
In reference to his house on Rodrigues, Leguat continued:
You see’ twas between two Parterres [garden constructions], and upheld by a great Tree, which also cover’d it
on the side of the Sea. This tree bore a Fruit something like an Olive [= Bois d’Olive Cassine orientale]; and the
Parrots lov’d the Nuts of it mightily.
The parrots were so tame and easy to catch that Leguat refrained from hunting them, even taking one to
Mauritius, when he departed Rodrigues in 1693:
Hunting and Fishing were so easie to us, that it took away from the Pleasure. We often delighted ourselves in
teaching the Parrots to speak, there being vast numbers of them. We carried one to Maurice Isle, which talk’d
French and Flemish.
Tafforet (1725–26) stated that there were three kinds of parrot on the island. The first was the Rodrigues
Parrot Necropsittacus rodericanus; the second was Psittacula exsul; ‘The second species is slightly smaller and
much more beautiful, because they have their plumage green like the preceeding, a little more blue, and
above the wings a little red as well as their beak’; the third is referable to a female P. exsul. Interestingly, the
two surviving skins exhibit blue coloration, and not the green described by Tafforet. This blue morph is the
result of the total absence of the pigment psittacin, which in parrots means blue feathers are produced
rather than green ones; parrot breeders use the term ‘true blue’ for this condition, and it is the most common
colour aberration in Psittacula and other parrots (Martin 2002). Sometimes psittacin levels are diluted,
producing a colour somewhere between green and blue (this is termed ‘parblue’). Newton & Newton (1876)
described P. exsul as being ‘greyish glaucous’, with green or blue reflections depending on the angle of light.
This suggests that the colour aberration in the two skins of the Rodrigues Ring-necked Parakeet was due to
a ‘parblue’ mutation rather than a ‘true blue’ one (Hume & van Grouw 2014).
Extinction
The first specimen of P. exsul was received by Alfred Newton in 1871 (Newton 1872), but by this time the
parakeets had become pitifully scarce. One was seen on September 30, 1874 in forests on the s outh-western
side of the island (Slater 1879a), and a few were seen in 1875 (Caldwell 1875), when the last specimen was
collected (Newton & Newton 1876). A devastating series of cyclones struck the following year
(North-
Coombes 1971), which perhaps wiped out the last few survivors (Cheke 1987). Contrary to
Greenway’s (1958) suggestion that P. exsul might survive on offshore islets, the Rodrigues islets are too
small to support viable populations of birds.
Subfossil remains were discovered in caves on Rodrigues, and show that this parakeet, compared with
the Ring-necked Parakeet P. krameri, which is considered a close relative, was larger, had a robust head and
jaws, large leg elements, but comparatively short wings. Osteologically, it appears to represent a diminutive
Alexandrine Parakeet P. eupatria rather than a large P. krameri (Hume 2007), and Tafforet’s (1725–26) account
(above) describing a little red above the wings (shoulder-patch) is characteristic of P. eupatria, and not P.
krameri.
However, a molecular study has confirmed that Psittacula exsul is not only closely related to the Mauritius
Echo Parakeet P. echo and Réunion Ring-necked Parakeet P. eques, and part of the same clade, but that the
Rodrigues bird was ancestral to them as well (Jackson et al. 2015).
Status Extinct. Known only from contemporary accounts and illustrations. A supposed skin is in Edinburgh. Last recorded in
1732.
Range Réunion, Mascarene Islands.
Description 40cm (16in). Generally green, darker on back, yellowish on underside; two collar rings, one pink, one b lackish-blue;
underside of tail brownish-grey edged yellow; upper bill red, lower bill blackish-brown; iris yellow; feet blackish-grey.
There has been much confusion regarding this bird, as it has never been clear how it differed from the rare
but extant Echo Parakeet Psittacula echo of Mauritius, to which it was evidently closely related. However,
there seems little doubt that such a bird did occur on Réunion. Dubois (1674) specifically referred to
‘Perroquets Verts . . . ayant un collier noir’ (green parrots with a black collar) on the island. Rothschild (1907a)
eloquently explained the position:
There has been considerable confusion with regard to this parrot. It was first asserted that it occurred on both
Bourbon [Réunion] and Mauritius. Then Professor Newton separated the Mauritius bird as Pal. echo.
Salvadori, however . . . reunited the Bourbon and Mauritius birds, while quite unaccountably stating only
Mauritius as the habitat. The Abbé Dubois describes this bird as follows: ‘Green parrots as large as pigeons
having a double collar’. Now the species of Palaeornis [= Psittacula] from Rodriguez, the Seychelles, and the
mainland of Africa are all distinct, and the other land birds of Mauritius are and were different from those of
Bourbon. I therefore feel quite certain that Professor Newton is right, and that his Palaeornis echo is distinct
from P. eques, though, unfortunately, we do not know in which way the two forms differed.
The Réunion Ring-necked Parakeet is known from descriptions and paintings only, but at least three
specimens may have reached Paris during the latter half of the 18th century (Cheke 1987). The illustration
by Martinet in Buffon (1779) is poor, but a far superior illustration was executed by Barraband (in Levaillant
1805a; see Hume 2007). From this illustration, there are few, if any, distinct plumage differences between the
Réunion and Mauritius populations, which may well have been conspecific. A recent mtDNA analysis of a
skin held in the Royal Museum, Edinburgh, has shown that it may be a unique Réunion Ring-necked
Parakeet (Jackson et al. 2015). It differs genetically from P. echo (and whatever it is diverged from P. echo
around 0.6 mya). However, circumstantial evidence suggests that the skin was collected on Mauritius and
not Réunion (Cheke & Hume 2008; Cheke & Jansen 2016); thus the small genetic difference may be a result
of inter-island migration and a continued mixing of genetic material. A number of other Mascarene birds,
including Psittacula parakeets (Mourer-Chauvire et al. 1999; Cheke & Hume 2008; Hume 2007, Hume 2014)
appeared to migrate between Mauritius and Réunion, and each island population was conspecific. There is
also some suggestion that the Edinburgh specimen might have been the one used for Martinet’s plate, but
there is no unequivocal confirming evidence that this is the case.
Early extinction
Considering the available habitat that still remains on Réunion and the fact that P. echo survives on
Mauritius, the Réunion Ring-necked Parakeet died out very early in Réunion’s post-human history. The
birds must have still been fairly common during the early years of the 18th century as Borghesi (Lougnon
1970), in 1703, stated that:
Still in this Isle, I have observed numerous sorts of perroquets, of large or of small size: some green, the others
grey, and others mottled of different colours.
It must have died out extremely rapidly as the last mention of it was in 1732 (Cossigny 1732–1755).
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Almost nothing appears to be known about this parrot in life, or the reasons for its extinction. A few captive
birds arrived in France during the late 18th century, by which time it was probably already extinct in the
wild. It was first mentioned by Dubois (1674) in 1671–2:
Parrots a little bigger than pigeons, with plumage the colour of squirrel fur, a black hood on the head, the beak
very large and the colour of fire.
All Réunion parrots were hunted regularly and they appear to have fat cycles associated with vertical
migration. This was clearly evident to Feuilley (1705) in 1704, who reported:
There are several sorts of parrot, of different sizes and colours. Some are the size of a hen, grey, the beak red
[Mascarene Parrot]; others the same colour the size of a pigeon [Thirioux’s Grey Parrot; see p. 199], and yet
others, smaller, are green [Réunion Ring-necked Parakeet; see p. 201]. There are great quantities, especially
in the Sainte-Suzanne area and on the mountainsides. They are very good to eat, especially when they are fat,
which is from the month of June until the month of September, because at that time the trees produce a certain
wild seed that these birds eat.
The Mascarene Parrot was mentioned for the last time on Réunion at the beginning of the 18th century
(Hume 2007; Cheke & Hume 2008), and appears to have survived longer than any other Réunion parrot. At
least three or four captive birds were alive in Europe around the latter half of the 18th century, of which two
were preserved as skins. Mauduyt (1784) mentions live birds in Paris:
The Mascarin is found at Ile Bourbon [Réunion]; I have seen several alive in Paris, they were rather gentle
birds; they had in their favour only that the red beak contrasted agreeably with the dark background of their
plumage; they had not learnt to talk.
By the 1790s, Levaillant (1805a) said it was very rare, and that he had seen three specimens, one in
Mauduyt’s possession, one in the cabinet of Abbé Octave Aubry, and the third in the Paris Museum. Of the
surviving skins, one, the type, is in Paris, and a second is held at Vienna. The latter has been stated to be
‘partially albinistic’ but this is certainly not the case. Partial albinism does not exist in nature, and the pres-
ence of partial white feathering is a result of other mutations (see van Grouw 2013). Interestingly, the white
feathers exhibited in the Vienna skin are a result of poor diet, which was due to this individual’s probable
long period in captivity (Hume & van Grouw 2014). During the 18th century, the dietary requirements of
parrots was poorly known, so this individual of Mascarinus was probably unable to extract sufficient quan-
tities of the amino acid tyrosine from its food, which is necessary for normal melanin synthesis; hence white
feathers in the plumage. The o ften-quoted last living specimen was depicted by Hahn (1834) in the menag-
erie of the King of Bavaria, but the specimen was not preserved and there is no evidence for the species
surviving after 1800 (Hume 2007). Hahn may not have even seen the bird himself, but made the following
comment (from Hume 2007):
Mascarene Parrot
Mascarinus mascarin
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bill red; base of bill black; head and throat b luish-grey; remainder of body brownish red, lower parts paler; tail
white at base, remaining half brownish red; feet blackish. ORIGIN: Madagascar [in error]. WHERE FOUND:
Unknown. FOOD: Fruits. BREEDING: Unknown. CHARACTERS: It is not very lively and its call is harsh.
There is a living example in the menagerie of King Maximilian of Bavaria at Munich on which the illustration is
based.
The Mascarene Parrot was almost certainly extinct by 1800. Peter Mundy (Temple 1914) spoke of ‘russet
parrots’ on Mauritius, which may refer to this species, but no confirmatory fossil remains have been found.
Taxonomy
The relationships of this aberrant parrot have been difficult to determine, and it has been duly placed in a
number of genera, most recently (and surprisingly) nested within the subspecies of the Lesser Vasa Parrot
Coracopsis nigra of Madagascar, the Comoros and the Seychelles, based on molecular evidence (Kundu et al.
2011). This extraordinary placement was questioned by a number of authorities (e.g. Hume & Walters 2012;
Joseph et al. 2012; Safford & Hawkins 2013), as the Mascarene Parrot differs drastically in morphology.
Because doubts were expressed about the results, Podsiadlowski et al. (2017) undertook a more stringent
DNA analysis and concluded that Kundu et al. 2011 were in error; the samples used were contaminated.
Furthermore, Podsiadlowski et al.’s analyses have shown that Mascarinus was in fact a part of the Psittacula
radiation, forming a clade along with Psittacula and Tanygnathus. The authors further recommended that
Tanygnathus, Mascarinus and Psittacula should be merged into a single genus, Psittacula. This confirms the
previous work of other authors who suggested a close relationship between these groups (Forbes 1879;
Garrod 1874; Oustalet 1897; Forshaw 1989; Hume 2007).
This was the largest of the parrots found on Rodrigues. The all-green Rodrigues Parrot was first mentioned
by ship’s mate Julien Tafforet in 1725–26:
The parrots are of three kinds, and in numbers. The largest are larger than a pigeon, and have a very long tail, the
head large as well as the beak. They mostly come on the islet [Île aux Mat = Île Gombrani] which is to the south
of the island, where they eat a small black seed, which produces a shrub whose leaves have the smell of the
orange-tree, and come to the mainland to drink water. The second species is slightly smaller and more beautiful,
because they have green plumage like the preceding [Rodrigues Parrot], a little more blue, and above the wings
a little red as well as their beak [male Rodrigues Parakeet Psittacula exsul]. The third species is small and
altogether green, and the beak black [female Rodrigues Parakeet].
When Pingré (1763) stayed on Rodrigues in 1761 to observe the Transit of Venus, the Rodrigues Parrot had
become rare:
The perruche [Rodrigues Parakeet] seemed to me much more delicate [in flavour]. I would not have missed
any game from France if this one had been commoner in Rodrigues; but it begins to become rare. There are even
fewer perroquets [Rodrigues Parrot], although there were once a big enough quantity according to François
Leguat; indeed a little islet south of Rodrigues still retains the name Isle of Parrots.
This was the last time it was mentioned, so must have died out shortly after. It presumably disappeared due
to forest clearance, over-hunting, and probable rat predation of eggs and chicks. Cats would have also been
dangerous predators.
Milne-Edwards (1867a) described this species from a portion of maxilla (upper bill). Later, Günther &
Newton (1879) described a n ear-complete set of skull and postcranial bones. These subfossil remains were
assumed to refer to the parrot described by Tafforet (1725–26). It was considered to be a large cockatoo-sized
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Rodrigues Parrot
Necropsittacus rodericanus
bird, with a disproportionately large head and jaws and long tail (see Hume 2007). Hachisuka (1953), based
on Tafforet’s account, correctly asserted that the colour of the bird was a uniform green.
Réunion Parrot
Necropsittacus? borbonicus Rothschild 1907a, p. 62, pl. 8
Status Extinct. Known only from Dubois’s description.
Range Réunion, Mascarene Islands.
Dubois (1674) in 1671–72, who appears to have been the only eye-witness of this bird, describes it briefly:
Body the size of a large pigeon, green; head, tail and upper parts of wings the colour of fire.
Rothschild (1907a) muddled the description of this bird with that of the Rodrigues Parrot Necropsittacus
rodericanus, and accordingly placed it into the same genus. There is no evidence to support this. He further
confused the issue, and described another species, Necropsittacus francicus (see p. 444), based on the account
of Dubois, but gave the provenance as Mauritius, not Réunion.
What Dubois actually saw on Réunion is now difficult to interpret. His descriptions of other Réunion
birds can be trusted, so perhaps this species was an endemic parrot that disappeared rapidly after being
first observed in 1671–72, or the possibility exists that it was an escaped cage bird (Greenway 1967). Fuller
(1987) warns of the potential hazards when trying to determine the original parrot diversity on islands:
One problem with early reports of parrots on islands is that there can be no certainty whether the birds referred
to were actually endemic or whether they were individuals belonging to a widespread species that had been intro-
duced by human agency.
The Broad-billed Parrot was formally described from a single mandible collected along with the first Dodo
remains in the Mare aux Songes marsh in 1865 (Owen 1866b). Owen concluded that a c ockatoo-sized parrot
had once been a contemporary of the Dodo, but it was now extinct.
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Subsequently, Newton (1875b) reported the discovery of an illustration of a large parrot in the unpub-
lished journal of Wolphart Harmanszoon, who voyaged to Mauritius in 1601–02 in the ship Gelderland.
There are other vague descriptions and illustrations, and there is no doubt that they all refer to the
Broad-billed Parrot. It was first mentioned soon after the discovery of Mauritius in 1598, and was referred to
as the Indian Crow, which may have been due to a deep raucous call (Hume 2007). It was a large species, as
recorded in the Het Tweede Boeck (1601):
. . . Is a bird which we called the Indian Crow, more than twice as big as the parroquets, of two or three colours.
Joris Laerle gave a detailed sketch of Lophopsittacus from life in 1601 (Hume 2003), but left no written
description. The bird had a distinct non-erectile frontal crest, comparatively reduced wings and a long,
bifurcated tail. Despite suggestions to the contrary (e.g. Hachisuka 1953), there is no evidence that this
species was flightless anatomically (Hume 2007), although it would not have been a strong flier. It also
exhibited the greatest size sexual dimorphism known in any parrot. The coloration is debatable. Johannes
Pretorius in 1666–69 (in Hume & Winters 2015) attempted to keep Broad-billed Parrots in captivity on
Mauritius, but they never survived. His mention of them being beautifully coloured is also interesting:
The Indian ravens are very beautifully coloured. They cannot fly and are not often found. This kind is a very bad
tempered bird. When captive it refuses to eat. It would prefer to die rather than to live in captivity.
Hoffman in 1673–75 (1680, in Cassel 1931; Hume 2007) also left a description:
There are also geese, flamingos, three species of pigeon of varied colours, mottled and green perroquets [para-
keets], red ravens with hooked-beaks and with blue heads, that cannot fly well and are known by the Dutch as
Indian ravens.
It is remarkable that a large, poorly volant but aggressive parrot survived alongside potential predators
such as introduced macaque monkeys and rats for decades, but it appears to have finally succumbed to
human disturbance a short time after Hoffman’s account.
Habits
The Broad-billed Parrot had developed huge jaws for cracking palm and other forest tree nuts, comparable
in size with those of the world’s largest parrot, the Hyacinth Macaw Anodorhynchus hyacinthinus (Hume
2007), and was not w eak-jawed or frugivorous as postulated by Holyoak (1971). Hachisuka’s (1953) state-
ment that the bird was probably nocturnal was based solely on the fact that that he believed the flightless
Kakapo Strigops habroptilus and the Australian Night Parrot Geopsittacus occidentalis occupied a similar
ecological niche. This is no evidence to support this at all.
Broad-billed Parrot
Lophopsittacus mauritianus
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The relationships of the Broad-billed Parrot are unclear. A number of authorities have suggested a close
relationship between Lophopsittacus, Necropsittacus and Psittacula (Milne- Edwards 1874; Berlioz 1946;
Newton and Gadow 1893; Holyoak 1973), and Berlioz (1940) thought that this species was closely related to
Necropsittacus of Rodrigues. Morphological evidence (Hume 2007) suggests that all of the endemic
Mascarene genera – Lophopsittacus, Necropsittacus and Mascarinus – are clearly nested within the Psittaculini.
The Glaucous Macaw is a little-known species, and very few skin specimens survive. It formerly inhabited
Paraguay, the provinces of Misiones and Corrientes in northern Argentina, south-western Brazil and
possibly also the Artigas area of north-western Uruguay (see Collar et al. 1992). It was reported as common
only in Uruguay during the second half of the 18th century; otherwise it appears to have been rare over
much of its range. It was formerly in demand as a cage bird, but King (1981) reported that bird dealers in
Asunción, Paraguay, had been unable to obtain specimens for at least three decades. One was seen in
Uruguay in March 1951, when a single bird was observed perched on a fence-post in northwest Artigas
(Nores & Yzurieta 1983; Collar et al. 1992). No macaws have been seen there since, despite several surveys;
the area has now been destroyed by eucalyptus plantations
(Collar et al. 1992). Another observation of a macaw was made
in the early 1960s in Paraná, Brazil, which may have been refer-
able to the Glaucous Macaw, and reports of nesting pairs were
made by a local who did not want to disclose the area for fear of
hunters and trappers collecting the birds. Very few were kept in
zoological gardens, with a handful of captive birds reported
from the 1860s to the early 1920s. But even some of these may
have been confused with the similar Lear’s Macaw
Anodorhynchus leari (Forshaw 1989). The reasons for its disap-
pearance are not known, but trapping and deforestation are
likely to have been contributing factors.
Habits
The Glaucous Macaw inhabited subtropical gallery forest, little
of which survives in south-eastern Brazil, but considerable
tracts remain in other parts of the species’ former range, e.g. in
south-eastern Paraguay and north-eastern Argentina. It
appears to have been inextricably linked to riverine habitats
and fringing forests along major rivers, and may have
wandered into lightly wooded savannas, as do other
Anodorhynchus macaws (Forshaw 1989; Collar et al. 1992).
Almost nothing was known of its habits. In 1805, Azara (1802–
1805) and d’Orbigny (1835) reported that it nested in holes in
trees and enormous holes that they dug out of cliffs, and laid
two eggs. Apparently no further observations on its breeding
were ever made. It was probably dependant on the Yatay Palm
Glaucous Macaw
Butia yatay, the only palm in its range with the appropriate size Anodorhynchus glaucus
of food nut (Yamashita & Valle 1993).
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The confusion surrounding the original diversity of the Antillean macaws has been extraordinary, and no
other group of extinct birds has aroused such overzealous misinterpretation. The evidence on which the
majority of species have been described is, at best, inadequate, and based entirely on the vaguest of accounts.
Three species, the Cuban Macaw Ara tricolor, Lesser Antillian Macaw Ara guadeloupensis (see p. 209) and
Puerto Rican Macaw Ara autochthones (see p. 211) are known from skins and subfossil bones (Wetmore
1937a; Olson & Maíz López 2008). There is illustrative and comparatively well-documented evidence for a
species of macaw on Jamaica; otherwise the presence of endemic macaws on other West Indian Islands is
extremely dubious.
The primary reason for the confusion is that macaws were regularly transported from the South
American mainland to the West Indies, and often between islands, both by the Carib Native Americans of
the Antilles and by early European settlers (Olson & Maíz López 2008). South American macaws were taken
to Jamaica before being transported to Europe (Clark 1905d), so it is hardly surprising that if an imported
mainland species escaped, it could subsequently be construed as being an island endemic. Forshaw (1989)
provided a cautionary summary, the facts of which were not considered with enough care by a variety of
earlier taxonomists:
It is clear from the writings of early travellers that macaws, in addition to Ara tricolor, formerly occurred in the
West Indies, but there is supporting evidence only from the Virgin Islands. Other species have been named, but
it cannot be established with certainty that they were members of this genus or were, in fact, different from
extant species found in Central and South America.
It was Clark (1905a, d) who first examined the early documentation concerning the macaws of the Greater
and Lesser Antilles. Compared with some authors (e.g. Rothschild 1905b, 1907a), Clark was conservative,
describing only two new species, Ara guadeloupensis (see p. 209) and Amazona martinicana (see p. 214), both
of which he justified as being valid. All other supposed macaw species known only from accounts are open
to alternative explanation.
Wiley & Kirwan (2013) published an exhaustive review of the macaws of the West Indies, and
provided strong evidence to suggest that only the Cuban Macaw Ara tricolor, Puerto Rican Macaw Ara
autochtones and perhaps Montserrat Macaw Ara sp., which is known from a single coracoid (Williams &
Steadman 2001), formerly occurred in the West Indies. The last-named may be referable to the Lesser
Antillean Macaw Ara guadeloupensis (see p. 209). Wiley & Kirwan further support, but with reservation,
the existence of the Jamaican Yellow-headed Macaw Ara gossei (see p. 380), yet Gosse (1847) was unsure
if it was conspecific with A. tricolor. It might conceivably have been an imported Cuban Macaw (Olson &
Maíz López 2008).
Province to the west, but apparently the birds there disappeared after a hurricane in 1844 (Olson & Suárez
2008). It was still easily encountered in 1849, but Gundlach recorded it as very rare shortly after.
The last known pair of macaws were supposedly shot (but not collected) at San Francisco de la Vega, Isla
de Pinos (and not Zapata swamp on the south coast; see Wiley & Kirwan 2013) in 1864 (Zappey, in Bangs &
Zappey 1905), but this hearsay information was given to Zappey by the plantation owner where the birds
were shot (Olson & Suárez 2008). The Cuban Macaw may have survived until 1885, for Gundlach (in Cory
1886a) believed it was still extant in the southern Cuban swamps. Wiley & Kirwan (2013) provide strong
evidence that specimens were still being shot in 1889 in central Cuba. Although Gundlach found the flesh
evil-smelling and foul-tasting, over-hunting for food by the locals, collecting for the aviculture trade and
perhaps habitat destruction were probably the reasons for its extinction.
Habits
Little is known of the Cuban Macaw’s habits. Local people reported to Gundlach (1876) that it nested in
holes in palm trees, and lived in pairs and family groups. He noted that it had a loud vocalisation similar to
the macaws of Central America (Gundlach 1893). It fed on fruits, palm fruit, seeds of the naturalised
Chinaberry tree Melia azedarach, tender shoots and buds (Wiley & Kirwan 2013). Small populations appar-
ently bred in scattered localities. Olson & Suárez (2008) suggest that the dorso-ventrally compressed skull,
which in other macaws is an adaptation for powerful muscle attachment, is an indication that this species
fed on the extremely hard-shelled nuts of palms, and the bird’s distribution may have been linked closely to
availability of this food source. In the Zapata swamp, the palm flora is dominated by Cuban Royal Palm
Roystonea regia and Bull Thatch Palm Sabal maritima (Wiley & Kirwan 2013), which produce large fruits
favoured by parrots.
Until recently, the Lesser Antillian Macaw was known only from written accounts and illustrations.
Williams & Steadman (2001) tentatively assigned an ulna found in an archaeological site on Marie-Galante
to this species. It is slightly smaller than mainland macaws. However, Olson & Maíz López (2001) doubted
this assignation, and thought the specimen was more likely referable to an Amazona parrot. The recent
discovery of an ungual or claw on Marie-Galante Island confirms that an endemic macaw once occurred on
Guadeloupe (Gala & Lenoble 2015). The claw is Pleistocene in age and radiocarbon dated at 10,690 years,
predating the human colonisation of Guadeloupe by around 5,300 years ago. Gala & Lenoble also show that
the macaw was not transported to Guadeloupe by native Americans as a trade or food item. It is very likely
that the same species also occurred on nearby islands and not confined to one, so the Guadeloupe Macaw is
better referred to as the Lesser Antillian Macaw. It formerly occurred on Guadeloupe, on Marie-Galante to
the southeast and possibly other larger islands to the south. Rothschild (1905, 1907) described a second
species of macaw from Guadeloupe, Anodorhynchus purpurascens, but this was based erroneously on a
second-hand source describing the Guadeloupe Amazon Amazona violocea (Lenoble 2015).
Several authors describe the Lesser Antillian Macaw in considerable detail, and all are agreed on its
appearance (Wiley & Kirwan 2013; Lenoble 2015). The earliest account was given by Valdes (1535, in
Greenway 1967), who reported that Ferdinand Columbus recorded red parrots ‘as big as chickens’ on
Guadeloupe in 1496.
Du Tertre (1654, in Clark 1905a) said:
We have in Guadeloupe three of the parrot kind, viz. Macaws, Parrots and Parrakeets, each different from those
which inhabit the neighbouring islands, for each has its parrots different from those of the others in size, voice
and color. The Macaw is the largest of all the parrot tribe; for although the parrots of Guadeloupe are larger than
all other parrots, both of the islands and of the mainland, the Macaws are a third larger than they. The head,
neck, underparts, and back are flame color. The wings are a mixture of yellow, azure, and scarlet. The tail is
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wholly red, and a foot and a half long. The natives hold the feathers of the tail in great esteem; they stick them in
their hair, and pass them through the lobe of the ear and the septurn of the nose to serve as mustaches, and
consider themselves then much more genteel and worthy of the admiration of Europeans.
This bird lives on berries, and on the fruit of certain trees, but principally on the apples of the manchioneel,
which is a powerful and caustic poison to other animals. It is the prettiest sight in the world to see ten or a dozen
Macaws in a green tree. Their voice is loud and piercing, and they always cry when flying. If one imitates their
cry, they stop short. They have a grave and dignified demeanor, and so far from being alarmed by many shots
fired under a tree where they are perched, they gaze at their companions who fall dead to the ground without
being disturbed at all, so that one may fire five or six times into the same tree without their appearing to be
frightened.
The natives make use of a stratagem to take them alive; they watch for a chance to find them on the ground,
eating the fruit which has fallen from the trees, when they approach quietly under cover of the trees, then all at
once run forward, clapping their hands and filling the air with cries capable not only of astounding the birds, but
of terrifying the boldest. Then the poor birds, surprised and distracted, as if struck with a thunderbolt, lose the
use of their wings, and, making a virtue of necessity, throw themselves on their backs and assume the defensive
with the weapons nature has given them – their beaks and claws – with which they defend themselves so bravely
that not one of the natives dares to put his hand on them. One of the natives brings a big stick which he lays
across the belly of the bird, who seizes it with beak and claws; but while he is occupied in biting it, the native ties
him so adroitly to the stick that he can then do with him anything he wishes. The flesh of this bird is very tough,
and considered by many unwholesome, and even poisonous. I never had any ill effects from it, although we
inhabitants often eat it.
Du Tertre also illustrated the bird (rather crudely), along with the Guadeloupe Amazon (see p. 213) and
Guadeloupe Parakeet (see p. 212). In 1664, Du Tertre said the birds ate machioneel apples, but he later (1667)
corrected himself and said that they were eaten only in times of necessity. He explained that the male and
female were ‘inseparable companions’ and rarely seen apart; that they bred once or twice a year in a hole in
a stump, and laid two eggs the size of a partridge. The colour is not stated. The supposed twice-yearly
breeding is more likely explained as staggered breeding, not unreasonable for t ropical-island birds.
Labat
Labat (1742, quoted by Clark 1905a) gave a remarkably similar description:
The feathers of the head, neck, back and underparts are flame colour, the wings are of a mixture of blue, yellow
and red, the tail, which is from fifteen to twenty inches in length is wholly red . . . it talks very well, if it is taught
when young; it is amiable and kind, and allows itself to be caressed . . . There are Macaws, Parrots and Parrakeets
in each of our islands, and it is easy to tell from their plumage from which island they have come. Those from
Guadeloupe are generally larger than the others, but the parrakeets are smaller.
Labat’s important comment that each island’s parrot population was recognisably different eliminates the
possibility that the extinct macaws observed in the West Indies were all escapees of South American species.
Furthermore, the amiable and docile nature referred to by Labat is quite consistent with the observations of
Du Tertre, and is not characteristic of South American macaws.
Subfossil remains
Williams & Steadman (2001) tentatively assigned an ulna found in an archaeological site on M
arie-Galante
to this species. It is slightly smaller than mainland macaws. However, Olson & Maiz López (2008) doubted
this assignation, and thought the specimen was more likely referable to an Amazona parrot. Olson & Maíz
also gave good reasons to doubt the existence of any of the macaws known only from accounts. After
confirming the validity of the Puerto Rican Macaw A. autochthones (see p. 211) they state:
Although human-caused extinction on islands has in many cases been rampant, we very much doubt that ‘each
Greater Antillean and Lesser Antillean island once sustained one or two indigenous if not endemic species of
Ara’ (Williams & Steadman 2001). There is no credible evidence in support of such a belief and the only rela-
tively concrete evidence now available – the archeological record – suggests just the opposite – that there may
have been only one other macaw in the Antilles apart from the Cuban Macaw Ara tricolor.
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Paintings
Daubenton’s (1779 [1765–1780]) plate (no. 12) shows a bird that closely resembles the descriptions of Labat
and Du Tertre, suggesting that at least one specimen of this species must have found its way to Europe. Two
copies of Daubenton’s plate in the Rothschild Library at Tring differ somewhat in intensity of colour but not
of pattern. That the bird in Daubenton’s plate was not assigned a name by Boddaert (1783) is probably due
to the fact that Linnaeus (1766) quotes it in his description of the Scarlet Macaw Ara macao, a bird that A.
guadeloupensis seems to have resembled closely, except for a completely red tail and far more red on the
scapulars and tertials. However, Linnaeus’s description is clearly of A. macao and not of Daubenton’s bird.
The Guadeloupe Macaw was evidently a close relative of the Scarlet Macaw, and was possibly a derivative
of it.
There is also a painting dating from 1626 by Roelandt Savery, now in the Natural History Museum in
London. This is of a group of birds, most notably a Dodo Raphus cucullatus, but also including a macaw that
agrees with the description of A. guadeloupensis. Parrots were imported into Europe from all parts of the
world around this time. However, caution must be taken when interpreting illustrative evidence such as
this. The macaw depiction probably included some artistic licence, and that it may have been painted from
memory, plagiarised, or simply altered to suit the composition of the painting.
This macaw was described from a single tibiotarus of a juvenile bird from the island of St Croix (Wetmore
1937a), but it is not even certain if it occurred naturally on the island. Nothing more was known about it
until the discovery of p re-Columbian archaeological material on Puerto Rico (Olson & Maíz Lopez 2008).
The bird was decidedly larger than the Cuban Macaw Ara tricolor, but the coloration, unfortunately, remains
unknown.
All parrots were particularly sought after, both as food, but more importantly, as highly prized objects
for trading, so it is likely that the Puerto Rico population was exterminated for these reasons.
The Puerto Rican Parakeet has long been considered a race of Hispaniolan Parakeet Psittacara chloroptera, or
sometimes conspecific with it, and doubts were expressed that it ever occurred on Puerto Rico. Olson (2015)
provided an exhaustive study based on subfossil remains, skins and historical information, which showed
that not only was P. maugei a distinct species, differing in many aspects from P. chloroptera, but that it origi-
nally occurred on Puerto Rico and also Vieques, an island off the east coast. On Puerto Rico, it appears to
have disappeared by 1860 (Greenway 1967), despite being considered common in the 1790s (Olson 2015). In
1873, Gundlach (1878a, b) knew of it only by legends and that it occurred in forests on the eastern part of the
island, whereas Stahl (1887) stated in 1883 that:
[Psittacara maugei] is long extinct. Existing old men remember hearing from their parents that this little parrot
existed in ancient times and in places flocks occurred even in the plantations, wreaking damage, but through
untiring pursuit were soon eradicated.
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It was evidently abundant on Mona Island around 1875 (Olson 2015), but the last specimen, a female,
was taken in late February or early March 1892 by the collector W. W. Brown (Cory 1892b). It most probably
became extinct within a decade of Brown’s visit, as Bowdish (1902–1903) failed to mention it from his 1901
visit. The extinction of the Puerto Rican Parakeet is believed to have been due to a combination of forest
destruction, persecutin for crop damage, and perhaps disease (Olson 2015).
Habits
On Mona Island, Brown noted that the island had very few trees and no fresh water (Cory 1892b), so the
Puerto Rican Parakeet may have seasonally migrated between Mona and mainland Puerto Rico. The
Hispaniolan Parakeet feeds on a wide range of fruits, seeds, nuts, flowers and the flesh of fruits (Juniper &
Parr 1998); as P. maugei was larger with distinct bill morphology, most notably in the tomial notch on the
rostrum being shallower and the hooked tip less ventrally directed, suggesting that its bill was more weakly
constructed, it may have fed on softer food than the Hispaniolan bird (Olson 2015).
The Guadeloupe Parakeet is known from the descriptions by Du Tertre (1667) and Labat (1742), and a
woodcut depicting this bird, along with the Guadeloupe Amazon (see p. 213) and Lesser Antillean Macaw
(see p. 209). Greenway (1967) quoted Du Tertre, who mentions three species of parrot on Guadeloupe of
differing sizes: ‘Those which we call Perriques are the small Perroquets, all green and as big as magpies’.
Labat’s quote is equally brief:
Those of Guadeloupe are about the size of a blackbird, entirely green, except a few small red feathers, which they
have on their head. Their bill is white. They are very gentle, loving, and learn to speak easily.
Bond (1956) suggested that the birds seen by Labat were captive Cuban Parakeets Aratinga euops, with the
red of the underwing-coverts concealed, but the Cuban Parakeet had not been described at this time.
Marien & Koopman (1955) considered the Guadeloupe Parakeet to be related to the Green Parakeet A.
holochlora, which occurs in Mexico and Central America, and believed Labat to be a trustworthy observer.
Marien & Koopman were convinced that a small parakeet formerly existed on Guadeloupe; in view of
the distance between the ranges of A. holochlora and A. labati, it was best to consider them as separate
species.
There is little reason to doubt that a parakeet formerly occurred on Guadeloupe, but the differences may
or may not have been specifically distinct. L’Herminier (in Bond 1956) failed to find the birds in the early
19th century, so the Guadeloupe Parakeet presumably disappeared some time between Labat’s visit in the
1740s and then. The reasons for its extinction are unclear.
The Painted Conure is a widespread, generally common species occurring in distjunct populations in
Panama and northern South America (Juniper & Parr 1998). The Sinú Painted Conure was restricted to the
Sinú River valley and was last reliably recorded in 1949. Almost nothing was recorded about its habits. It
was not found during searches in 2004 and 2006 (ProAves Colombia 2008). It may well be extinct, but if it
still survives this bird is under threat due to habitat loss and hunting associated with armed conflict in the
region; the future of the Sinú Painted Conure appears bleak indeed.
DNA analysis has shown that the Sinú population is probably specifically distinct (Joseph & Stockwell
2002), but has at present been retained as a subspecies of Pyrrhura picta pending further study (SACC 2007).
The surviving West Indian amazons are found on Cuba, Jamaica, Hispaniola and Puerto Rico, and three
islands in the Lesser Antilles – Dominica, St Vincent and St Lucia (Juniper & Parr 1998). They are all consid-
ered endangered, primarily due to trapping for the pet trade, over-hunting for food and habitat destruction
(BirdLife International 2011).
The Guadeloupe Amazon was first described by Du Tertre in 1664 (translated in Clark 1905c):
The Parrot of Guadeloupe is almost as large as a fowl. The beak and the eye are bordered with carnation. All the
feathers of the head, neck, and underparts are of a violet color, mixed with a little green and black, and changeable
like the throat of a pigeon. All the upper part of the back is brownish green. The long quills are black, the others
yellow, green and red, and it has on the wing‑coverts two rosettes of a rose color. When it erects the feathers of its
neck, it makes a beautiful ruff about its head, which it seems to admire, as a peacock its tail. It has a strong voice,
talks very distinctly, and learns quickly if taken young. It lives on the wild fruits which grow in the forests,
except that it does not eat the manchioneel. Cotton seed intoxicates it, and affects it as wine does a man, and for
that reason they eat it with great eagerness. The flavour of its flesh is excellent, but changeable, according to the
kind of food. If it eats cashew nuts, the flesh has an agreeable flavour of garlic; if “bois des inde” it has a flavour of
cloves and cinnamon; if on bitter fruits, it becomes bitter like gall. If it feeds on genips, the flesh becomes wholly
black, but that does not prevent it from having a very fine flavour. When it feeds on guavas it is at its best, and
then the French commit great havoc among them.
Buffon (1770–1783, translated in Clark 1905c) was informed in 1779 that the parrot was very rare, and
summed up the probable reason for its extinction:
We have never seen this parrot, and it is not found in Cayenne. It is even very rare in Guadeloupe to-day, for
none of the inhabitants of that island have given us any information concerning it; but that is not extraordinary,
for since the islands have been inhabited, the number of parrots has greatly diminished, and Du Tertre remarks
in particular of this one that the French colonists wage a terrible war on in season, when it is especially fat and
succulent.
The Guadeloupe Amazon appears to have been related to the Imperial Amazon Amazona imperialis, which
survives in the steep mountain forests of Dominica. Guadeloupe is less mountainous than Dominica, and
harbours a larger human population, so pressures on the birds would have been greater. Although common
in the first half of the 18th century, the Guadeloupe Amazon declined rapidly and was last recorded in 1779
(Clark 1905c; del Hoyo & Collar 2014); it probably became extinct shortly after.
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Habits
Du Tertre (1667, in Clark 1905c) gave some brief breeding details:
We had two which built their nest a hundred paces from our house in a large tree. The male and the female sat
alternately, and came one after the other to feed at the house, where they brought their young when they were
large enough to leave the nest.
The amazon of Martinique is far more poorly attested than the amazon of Guadeloupe. Clark (1905c)
pointed out that all the early writers on Martinique mentioned parrots, but none of them described them.
The only useful account is that of Labat (1742):
The Parrot is too common a bird for me to stop to give a descripton of it . . . [The parrot] of Dominica has some
red feathers in its wings, in its tail and under the throat. That of Martinique resembles it, except that the top of
the head is slate colour with a little red.
Rothschild (1905b) identified the Dominican bird as the Red-necked Amazon Amazona arausiaca. This bird
survives on the island in mountain forest, although at lower altitudes than those inhabited by the Imperial
Amazon A. imperialis (Juniper & Parr 1998).
The Martinique Amazon, like its extinct relative on Guadeloupe, was considered common in the first half
of the 18th century, but over-hunting rapidly exterminated the birds (Clark 1905c; del Hoyo & Collar 2014);
the last record was in 1779 and it presumably died out shortly after that date.
Only three specimens, two males and a female, of this race of Puerto Rico Amazon Amazona vittata were
ever collected. These were all taken in 1899 by W. W. Brown; the bird has never been seen since, although
Wetmore searched for it in 1912 (Wetmore 1916, 1927). He wrote (1916) that both the Culebra Island and
nominate Puerto Rico Amazons had been common fifteen years previously, but by the time of writing had
disappeared from all localities including Culebra Island, save for a few places on the north-east coast of
Puerto Rico. The Puerto Rico Amazon is considered critically endangered today, while the Culebra Island
Amazon was never seen again. The Culebra birds were persecuted for crop damage, and probably suffered
high mortality rates from hurricanes because of deforestation (Juniper & Parr 1998).
An extinct population of Puerto Rico Amazon also occurred on the island of Vieques, but no specimens
were collected. As the Culebra birds were given subspecific status, it is possible that the Vieques population
was also subspecifically distinct. Wetmore (Wetmore 1916, 1927) also searched unsuccessfully for this bird
in 1912.
Description 36cm (14in). Generally green, abdomen washed bluish; primaries deep blue; speculum bright scarlet; feathers of
hind neck edged black; throat feathers mixed with yellow; forehead, chin and upper throat greyish, bordered with dull pale
yellow on crown; sides of head dull yellow; thighs grey; tail green, base yellow marked with red, with light greenish-yellow
terminal band; bill whitish horn, tip dusky; legs and feet dark grey.
The type specimen of this race of Yellow-shouldered Amazon Amazona barbadensis was brought alive from
Aruba in 1882, and described by Lawrence (1883) from the living bird. Although he requested the skin to be
preserved when the bird died, this was not done. However, he was assured that the bird’s plumage had not
altered in any way during its lifetime. The population on Aruba subsequently died out, but no further
specimens were ever collected.
No mention of the race is made by either Peters (1937 [1931–48]) or Greenway (1967). Voous (1957) stated
that the Aruba population probably disappeared between 1944 and 1947, the last individuals being either
poisoned because of their destructive habits in fruit gardens, or sold for high prices. But Voous subsequently
discovered that two had been reported by a Mr. B. Hartwell, so a few may have survived after this date.
Voous doubted whether Lawrence’s type actually came from Aruba, and there is considerable individual
and geographic variation in the species, so it is uncertain whether or not A. b. canifrons actually represents a
valid taxon.
The Carolina Parakeet was originally distributed across the deciduous forests of North America, from
Florida and southern Virginia west to eastern Texas, Oklahoma, Kansas and Nebraska. Individuals were
occasionally found as far north as Maryland, Pennsylvania, New York and the Great Lakes, Wisconsin and
Colorado. The species has been divided into two subspecies, separated by the Appalachian Mountains.
As early as 1831, Audubon (1831) noted that the Carolina Parakeet was rapidly diminishing in numbers,
and from that point onwards the range gradually declined from east to west, the birds disappearing as
European settlers moved westwards. According to Ridgway (1916) they were last seen in 1856 in Ohio, in
1851 in Indiana and in 1878 in Kentucky, the latter being the latest sighting east of the Mississippi River,
except for a handful remaining in Florida.
The last skin specimen was collected on Paget Creek, Brevard County, on the east coast of Florida by Dr
E. A. Mearns on April 18, 1901 (Greenway 1967); in April 1904, Chapman saw two flocks, consisting of a
total of 13 birds, at Taylor’s Creek on the north-east side of Lake Okeechobee in Florida (McKinley 1985).
This was the last definite sighting in the wild. Unconfirmed sightings of single birds were reported in
Missouri in 1905 and Kansas as late as 1912, and there were vague reports from Louisiana in 1910 (Widmann
1907; Oberholser 1938), but to all extent and purposes, the Carolina Parakeet was by now extinct in the wild.
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Unconfirmed sightings
In 1920 a Florida resident, Henry Redding, reported a flock of about 30 near Fort Drum Creek, and there
were persistent rumours and sightings in Florida as late as 1938, but careful investigation by specialists
failed to discover them (Vogt et al. 1939, Allen 1940). These late sightings have to be treated with caution, as
by that time several exotic parrots had established feral populations. Sprunt & Chamberlain (1949) main-
tained that birds still inhabited the Santee Swamp near Charleston, South Carolina, in 1938, though it has
been suggested that these may have been a feral population of escaped Australian parrots. The matter can
probably never now be satisfactorily settled, as this area was subsequently destroyed by the construction of
a power plant. Despite the uncertainties, Snyder & Russell (2002) provide convincing evidence that the
Carolina Parakeet probably survived until at least 1937–38 in Florida, and around the same time in South
Carolina and Georgia.
Captive birds
Carolina Parakeets were often kept in captivity where they bred regularly, but mortality of adults and juve-
niles was high. As late as 1898, two flocks of parakeets numbering 76 birds were captured for the avicultural
trade, by placing netting over two roosting sites in hollow cypresses (Snyder & Russell 2002). Demand for
the birds dramatically increased as they became rarer; this may have accelerated their decline to extinction
(Forshaw 1989). Even the noted American ornithologist Robert Ridgway kept breeding birds, which
produced young in 1902 (Amadon 1966).
Florida was the main parakeet-collecting area, and many live specimens reached Europe. During the
latter half of the 19th century, a free-flying population of parakeets was maintained for some time in
Germany (Kolar 1972). Ultimately, all captive birds either failed to breed, succumbed to disease, or perished
due to inadequate knowledge of their housing requirements. The last known bird died in Cincinnati Zoo on
February 21, 1918, a male called ‘Inca’, who outlived his mate, ‘Lady Jane’, by a year (Fuller 2000).
Relationships
Evidence recently obtained from DNA has shown that the Carolina Parakeet was related to a clade of
parrots that included the Sun Parakeet Aratinga solstitialis, Golden-capped Parakeet A. auricapillus and
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Nanday or Black-hooded Parakeet Nandayus nenday (Kirchman et al. 2012), of which the Sun and Golden-
capped Parakeets have similar yellow-and-orange head plumage and blue in the wings. The Carolina
Parakeet was also thought to have been closely related to the Monk Parakeet Myiopsitta monachus (Snyder
2004), but this is not supported by molecular evidence.
Habits
The Carolina Parakeet occurred in dense flocks, which often occurred together with the extinct Passenger
Pigeon Ectopistes migratorius (Blockstein 2002; see p. 166). The movement of the birds was described by
Charles Bendire (Bendire 1895):
Their flight, which is more or less undulating, resembles that of the Passenger Pigeon and again that of the
Falcons; it is extremely swift and graceful, enabling them, even when flying in rather compact flocks, to dart in
and out of the densest timber with perfect ease.
In Ohio, Kentucky and Mississippi, the birds were found only in well-wooded river valleys (Wilson 1808 in
1808–14), while in other areas, Wilson was assured by residents that birds did not occur in the surrounding
neighbourhood; he traversed considerable tracts of country without seeing them. Abert (in McKinley 1960)
describes the habitat of the parakeets between the Canadian and Arkansas Rivers in October 1845:
After a long march through misty low lands, where sycamore trees seem to arch to the heavens, and gaudy paro-
quets were circling round with rapid flight and screaming loudly among their lofty branches, we forced our way
through the tangled undergrowth of spice-wood and smilax, and reached the banks of the Canadian as the last
rays of the sun were disappearing.
It appears that the most typical habitat for the Carolina Parakeet was heavily forested ‘bottomlands’
bordering large rivers, and cypress swamps, which were then common in the southern states (Bendire
1895). Their food was largely seeds, and they had a fondness for thistles Cirsium lecontei and Cocklebur
Xanthium strumarium, which were abundant. They also ate various blossoms and seeds of cypress, maple
and elm, but were unfortunately fond of raiding orchards in an apparently capricious manner. Wilson (1810
in 1808–14) reported:
I have known a flock of these birds to alight on an apple tree, and have myself seen them twist off the fruit, one by
one, strewing it in every direction around the tree, without observing that any of the depredators descended to
pick them up. To a Parakeet which I wounded and kept for some considerable time, I very often offered apples,
which it uniformly rejected, but burs, or beech nuts never.
Second-hand accounts of breeding indicate that two to five white eggs were laid in a hole in a large tree
some distance from the ground. Both sexes fed the young in captivity, but they were careless parents, and
often failed to rear the young (Childs 1905, 1906). Reports that they roosted in hollow trees, and in large
numbers, are probably true since they were observed to do so in captivity. Bent (1940) reported that they
used their bills for holding on to the inside of the tree. Some authors reported hibernation (presumably in
the hollow trees), but the birds were apparently constantly seen during the winter (Greenway 1967), so this
seems doubtful. Furthermore, Kirchman et al. (2012) suggest that the feathered cere in Conuropsis was an
adaptation to survive the cold. Vocalisations comprised loud screams that could be heard over long
distances, and quieter chattering (Snyder & Russell 2002).
While flying, Edward McIlhenny (in Bendire 1895) remarked:
While on the wing they chatter and cry continually; this cry sounds like ‘qui’ with the rising inflection on the i;
this is repeated several times, the last one being drawn out like ‘qui-i-i-i’.
Cuculidae (Cuckoos)
The Old World cuckoos are cosmopolitan in distribution, with the highest diversity in the tropics. The
family has been divided into up to 28 genera. They are generally arboreal birds, although some, like the
Greater Roadrunner Geococcyx californianus, are terrestrial. A minority of species are brood-parasites, laying
their eggs in the nests of other birds, but most make their own nests. In general, the cuckoos are not threat-
ened, but some island subspecies have declined due to deforestation.
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The St Helena Dwarf Cuckoo was a diminutive species, considerably smaller than the African Chrysococcyx
emerald cuckoos, a genus from which it may have been derived (Olson 1975b). However, characters of the
humerus are distinct, so the placement of this bird in its own genus is tentative, pending the discovery of
further diagnostic material.
Cuckoos can migrate over open water (Long-tailed Koel Urodynamis taitensis, for example, regularly
winters on remote Polynesian islands), so it is not surprising that a cuckoo successfully colonised St Helena.
It was probably a forest bird, so would have suffered considerably after the arrival of humans at the begin-
ning of the 16th century, when most of the forests were destroyed. The diminutive size of the St Helena
Dwarf Cuckoo suggests that the foster parents parasitised by the cuckoo were also small, perhaps a diminu-
tive forest passerine that is also now extinct (Rowland et al. 1998).
Delalande’s Coua was known principally from Île S ainte-Marie, off the east coast of Madagascar, and was
considered ‘not very rare’ in 1831–32 (Sganzin 1840), but it is also stated to have occurred on the adjacent
mainland, especially on the Point-à-Larré. No birds have been collected since 1834, however, when Bernier
obtained one for Paris Museum (Greenway 1967).
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Habits
The bird seems to have fed mainly on molluscs, and Ackerman (1841) gave a good account of Delalande’s
Coua’s feeding technique in the wild (translation by JPH):
Often in movement it was like that of the magpie, and this bird gives an air of promptness, I will say some impa-
tience, that characterises its walk, as it jerks and often jumps both at the same time; thus it goes from branch to
branch, rock to rock, to seek the Agathines [snails] which forms its principal food. When it finds one of them,
whatever size, it carries it up to a large stone, and while holding the shell by the opening, it strikes it on the stone
turning and raising the head sometimes to the right, sometimes to the left; when it recognizes that the shell is
broken by the sound of the crack, it turns it over, body up, and, with its beak, withdraws the mollusc which he
swallows at once. If the opening is not large enough for the entire body of the snail to pass through, the coua
strikes it again until the shell is sufficiently broken.
Ackerman (1841) further describes a captive specimen:
I had a coua for a few months, in a rather large birdcage where it lived in good harmony with the other birds, and
had become almost familiar. It distinguished my voice rather well and came when I called it; so through the
netting I showed him a snail, and it flew in all directions and sang as it does in the forest. This song limited itself
to a ‘crouou’ in decrescendo that it repeated especially often when happy or impatient. After having made known
his wish for the snail, I gave it to him, and then he walked away while holding it with its beak, uttered his
‘crouou’ several times in continuation, after which he broke the shell . . . When eating the mollusc it still sang a
Delalande’s Coua
Coua delelandei
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little, and when it did not have anything left, it came to see me for more. I forgot to say that the bird wiped its
beak each time, as it was soiled by the sticky matter from the snail.
Among a collection of subfossil remains of birds from Madagascar, M ilne-Edwards & Grandidier (1895)
discovered a tarsometatarsus of a very large coua, larger than both the Giant Coua Coua gigas and the extinct
Delalande’s Coua C. delalandei. Nothing else is known about it, but a bird of this size would probably have
been regularly hunted for food and for feathers by the Malagasy. The date of its extinction is unknown.
This is the largest member of the genus Coua, living or extinct. Characteristics of the available skeletal mate-
rial show that Bertha’s Coua was a large, s wift-running species and perhaps one of the heaviest (Goodman
& Ravoavay 1993). Although no pectoral elements have been recovered, it is likely that this species was
incapable of strong flight, and may have led a totally terrestrial existence. Couas are regularly hunted on
Madagascar for food and feathers, so over-hunting may have been the main reason for its extinction.
The island of Assumption in the Seychelles once had a population of Madagascar Coucal Centropus toulou.
Its taxonomic status is unresolved (some authorities consider it undifferentiated from the mainland
Madagascar nominate).
The Aldabra race of this coucal is still extant and not considered threatened, but the Assumption bird
appears to have disappeared in the 1920s, and is now considered extinct (Benson & Penny 1971). Assumption
Island has been almost completely destroyed by guano extraction, so any likelihood of this race surviving
today is extremely remote. Apparently birds from Aldabra birds repopulated Assumption after the extinc-
tion of the C. t. assumptionis, and a population of several hundred birds is now established.
Habits
Nicoll (1906b) gave the only details:
A common though rather retiring bird, found breeding at the time of our visit [around March 1906]. One nest
contained two eggs and was a large domed structure made of dry grass, placed high up in a bush. The bird sits
with its long tail over its head, inside the nest. I only saw the male sitting; in fact I did not see a female at all on
Assumption. This species is very tame, and I almost caught one individual alive with a noose on the end of a
stick. It utters a long bubbling note which is audible for some distance.
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Tytonidae
The Puerto Rican Barn Owl was described by Alexander Wetmore (1920) when working on a considerable
collection of cave bird bones collected by H. E. Anthony on Puerto Rico. This bird was found only in a cave
near Utuado on the island. It was a small barn owl of the group that includes Ashy-faced Owl Tyto glaucops
of Hispaniola, T. (alba) insularis of St Vincent, Bequia, Union, Carriacou and Grenada, T. (a). bargei of Curaçao,
and T. (a). nigriscens of Dominica.
Wetmore described it as being most like the Ashy-faced Owl, but with the internal head of the talon
larger and much longer. These West Indian barn owls appear to represent a natural group, of which the
Puerto Rico bird represents a now-extinct species between the Hispaniola and Lesser Antilles forms. The
reasons for its disappearance are unknown, but it was almost certainly due to habitat modification after
humans arrived on the island. However, the Puerto Rican Barn Owl may have survived into the historic
period. There are reports of c ave-roosting owls from as recently as 1912, which may have been references to
this bird.
This owl, although clearly a member of Tytonidae, is very distinct, and pending the discovery of more mate-
rial, might belong to a separate genus altogether (Balouet & Olson 1989). Very few bones have been
preserved; the material we do have represents a juvenile and probably one adult.
The bones were found in Gilles Cave, near Bouloupari on the west coast, along with various reptile and
bird remains, including the extinct New Caledonian Giant Scrubfowl Sylviornis neocaledoniae (see p. 40), the
New Caledonian Wood Rail Gallirallus lafresnayanus (see p. 109) and the New Caledonia Owlet-nightjar
Aegotheles savesi (see p. 395). Barn owls regularly nest in caves, so it is likely that the owls’ presence accounts
for the accumulation of the remains of the smaller species, which were regurgitated as owl pellets. The
reasons for the extinction of the New Caledonian Barn Owl are unknown.
The West Indian barn owls were the largest of the genus Tyto, but smaller than some extant strigid owls, and
contrary to popular belief, none were flightless or even poorly volant (Suárez & Olson 2015). Their
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distribution throughout the West Indies was closely linked to the availability of their favoured mammalian
prey, hutias (Capromyidae) and giant rice rats Oryzomys sp. (Cricetidae), and their extinction coincided
with the human-caused disappearance of these large rodents.
The Hispaniolan Giant Barn Owl was an extremely large barn owl, with the tarsus as robust as that of a
Snowy Owl Bubo scandiacus, but much longer (Wetmore 1922a). The tarsometatarsus was larger than T.
noeli, approximately the same size as T. pollens, but less robust than either of these species, with a shorter
carpometacarpus (Suárez & Olson 2015; see below). It was discovered in cave deposits along with a number
of now-extinct West Indian mammals, so it was probably a major predator of these animals. It had compara-
tively long toes compared with Tyto noeli (see below), which may have enabled it to prey on arboreal
primates and thick-skinned edentates (Steadman & Hilgartner 1999), but more probably it was a predator
of large rodents (Suárez & Olson 2015), as were the other giant barn owls. The Hispaniolan Giant Barn Owl
probably occurred throughout Hispaniola in pre-human times (Suárez & Olson 2015), and may have
persisted until the late 18th century. On February 1, 1788, while climbing to the summit of La Selle, Haiti,
three French residents at night heard ‘hollow cries imitating the human voice that they attributed to some nocturnal
bird, as they had seen feathers resembling those of a swan at the edge of sort of a den or cavern’ (Wetmore & Swales
1931; Suárez & Olson 2015), and another sighting was reported in the 1970s (Suárez & Olson 2015). However,
it is now almost certainly extinct and its disappearance can be blamed on deforestation and the loss of prey
species.
Noel’s Giant Barn owl (Barbudan Barn Owl) Tyto noeli Arredondo
Tyto noeli Arredondo, 1972a: 416 (part). (Type locality: “Cueva del Túnel”, Término Municipal de La Salud, La Habana [now
Mayabeque] Province, Cuba); Suárez & Olson 2015, p. 540–541
Tyto noelli Arredondo, 1972a: 428, table 5 (lapsus)
Tyto neddi Steadman & Hilgartner 1999, p. 76 (Pocket, Gun Shop Cliff, Two Foot Bay, Barbuda); Hume & Walters 2012, p. 193
Specimens Subfossil remains are in Cuba and Washington, D. C.
Status Extinct. Known only from subfossil remains.
Range Western Cuba, south-central Jamaica and Barbuda, West Indies.
Noel’s Giant Barn Owl T. noeli was the smallest and least robust of the West Indian Tyto species, and exhib-
ited pronounced sexual dimorphism (Suárez & Olson 2015). It is the commonest Tyto species found in West
Indian fossil deposits, and also had the largest range, occurring in Cuba, Jamaica and Barbuda. It was the
first of the West Indian giant barn owls to be described from the Lesser Antilles, and was sympatric with T.
pollens and T. cravesae on Cuba. As Barbuda was once connected to Antigua, it is likely that this owl also
occurred there (Steadman & Hilgartner 1999), and it may have also occurred on other islands in the Lesser
Antilles that held populations of hutias and giant rice rats, its principal food source. Barbuda once
harboured three species of owl – the Burrowing Owl Athene cunicularia, American Barn Owl T. furcata and T.
noeli, all differing in size and ecology, which enabled them to coexist and utilise the island’s food resources.
The small, terrestrial Burrowing Owl feeds on invertebrates, reptiles and amphibians; the American Barn
Owl takes reptiles and small–medium mammals, while T. noeli ate large rodents and insectivores.
The extensive anthropogenic changes that have taken place on Barbuda resulted in the extinction of all of
the larger endemic rodents, which resulted in the loss of Noel’s Giant Barn Owl’s preferred large prey. It
was probably a combination of deforestation, introduced invasive animals and loss of prey that caused this
bird’s extinction.
This was the largest and most robust of all the West Bahaman Giant Barn Owl
Indian giant barn owls, but the rarest in the fossil Tyto pollens with Bahaman Hutia
record, known from just 11 mainly fragmentary Geocapromys ingrahami
specimens (Suárez & Olson 2015). Its distinctive-
ness from T. ostologa of Hispaniola and T. noeli of
Cuba, Jamaica and Barbuda was first recognized
by Olson & Hilgartner (1982), and although new
material has shown that there is an overlap in size
between it and T. ostologa, the former can be distin-
guished by the much more robust elements and
other characters (Suárez & Olson 2015). The tarso-
metatarsus of the Bahaman Giant Barn Owl was
almost twice the width of that of the extant
American Barn Owl Tyto furcata, but it was
comparatively short, being only about 15% longer
(Wetmore 1937b; Brodkorb 1959). The carpometa-
carpus was especially large and strong, which
suggests that the wing was proportionately larger
than Tyto ostologa (Suárez & Olson 2015). As with
all of the extinct West Indian barn owls, its remains
were discovered in cave deposits along with the
remains of large, endemic rodents, particularly the
hutia Geocapromys ingrahami, which comprised its
principal food source.
There is a possibility that this giant barn owl
survived into historic times. It was said to occur in
old-growth Caribbean Pine Pinus caribaea and nest
in burrows, at least on Andros Island, and was
contemporary with human occupation of the
island. It may have given rise to a native legend of
an entity called the ‘chickcharney’, a mischievous leprechaun-like nocturnal imp, which had three toes and
the ability to rotate its head 360° (Popov & Popov 1997). It was said that, if disturbed, chickcharneys would
unleash terrible misfortune.
This species of giant barn owl was only recently described from fossil localities in western Cuba (Suárez &
Olson 2015). It was larger than Noel’s Barn Owl Tyto noeli, but smaller and less robust, with the tarsometa-
tarsus more elongate than in Bahaman Barn Owl T. pollens (both of which it was sympatric with on Cuba). A
single cave deposit formed by Craves’s Giant Barn Owl contain juveniles of the extant Desmarest’s Hutia
Capromys pilorides and extinct Cuban Coney Geocapromys columbianus, which were favoured prey items.
Considering that this owl shared Cuba with two other giant barn owls plus an incredible diversity of other
avian raptors (Suárez & Olson 2015), how they shared potential food resources remains a mystery.
Like all giant West Indian barn owls, its disappearance can be blamed entirely on anthropological
changes to the island.
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Scops owls of the genus Otus have colonised all of the main island groups in the Indian Ocean, and have
evolved into a number of insular species and subspecies (Fuchs et al. 2008). It is from this group that the
endemic Mascarene genus Mascarenotus was derived (Mourer-Chauviré et al. 1994). The Mascarene owls
were characterised by large size, at least 42cm in total length in the Mauritius and Réunion species (Hume et
al. 2006), though smaller in the Rodrigues birds, whereas owls of the genus Otus attain a maximum total
length of 28cm. The Mascarenotus owls also had long legs, presumably adaptations for catching reptiles and
possibly small birds. The skeletal proportions of the Mascarene species exhibited remarkable parallel evolu-
tion with some extant species, such as the Puerto Rican Screech Owl Gymnoglaux lawrencii, Cuban Screech
Owl O. nudipes, and the extinct endemic Hawaiian genus Grallistrix (Mourer-Chauviré et al. 1999), presum-
ably as a result of these islands lacking terrestrial mammalian predators.
The Mauritian Lizard Owl disappeared some time after 1837, probably due to an increase in deforesta-
tion during the 19th century, as it had survived alongside introduced predators for centuries. Clark (1859)
made the final comment on the bird, and stated that it was extinct by then, after being formerly plentiful:
In September 1837 several inhabitants of the Savanne area told me they had seen owls in their forests; Dr
Dobson, of the 99th Regiment, assured me he had killed one in the woods of Curepipe. It could well happen that
in a few years, the species, if it is one, will have disappeared completely because of the destruction of our forests,
and the large number of poachers who roam the woods that remain.
It was a large, w
hite-faced, eared owl, with bare tarsi; a drawing by the 18th century artist Jossigny (see
Cheke & Hume 2008) is the only known illustration of the genus.
It appears that no owls were reported historically on Réunion, but the discovery of subfossil remains
confirms that a species, akin to the Mauritius Mascarenotus sauzieri and Rodrigues M. murivorus lizard owls,
once occurred there (Mourer-Chauviré et al. 1994). The Réunion Lizard Owl Mascarenotus grucheti was
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probably a secretive inhabitant of dense and remote forests. It was approximately the same size as M. sauz-
ieri but had slightly more reduced wing elements (Mourer-Chauviré et al. 2006). The short wings and long
legs may have been an adaptation for hunting in the dense forested areas once so prevalent on the island,
and analogous to forest-adapted Accipiter hawks, which have similar wing and leg proportions.
It is very likely that the Réunion Lizard Owl disappeared some time in the 1700s as a result of severe
deforestation, which accelerated during the 18th century on Réunion (Cheke & Hume 2008).
The Rodrigues Lizard Owl was briefly mentioned by Leguat (1708) in 1691–93, and described by Tafforet
(1726) in 1725–26, who stated:
A bird is seen which is very like the brown owl, and which eats the little birds and small lizards. They live almost
always in the trees; and when they think the weather fine, they utter at night always the same cry. On the other
hand, when they find the weather bad they are not heard.
This owl was originally placed in Athene, and has long been called a ‘little owl’ because of it, but the generic
placement is in error. Günther & Newton (1879) stated it would have been larger than the Southern Boobook
Ninox boobook (averaging 30cm in total length), but smaller than the Long-eared Owl Asio otus (averaging
36cm in total length), and that ‘the power of flight of this bird cannot have been much impaired. On the
other hand, the size and strength of its legs, as compared to the size of the body, and especially with the
length of the wing, was much more than in any of the species of owls, the skeletons of which we have been
able to compare’. These are now known to be specialised adaptations for forest life within this genus
(Mourer-Chauviré et al. 1994).
Leguat (1708) called them ‘Hiboux’, the French name for an eared owl, so presumably this species, like
the Mauritius species Mascarenotus sauzieri, had ear tufts.
The astronomer Pingré (1763), who was on Rodrigues to observe the Transit of Venus, did not mention
owls, so they presumably disappeared some time between 1726 (when they were reported by Tafforet) and
the time of Pingré’s visit in 1761. The rapid disappearance of the owl is something of a mystery. All of the
Mascarene lizard-owls probably nested in tree-holes, so deforestation would have been particularly devas-
tating (Hume 2013, 2014a). However, it is more likely that introduced cats and perhaps rats may have
finished off the birds.
The avian fossil record of Madeira is poorly known, despite the collection of substantial subfossil remains
(Pieper 1985). Rando et al. (2012) described a new species of owl from Madeira, which was related to the
Eurasian Scops Owl Otus scops. The Madeira Scops Owl O. mauli was similar in size and general morphology
to O. scops, but differed substantially in the relative proportions of the hindlimbs. The total leg length (femur
+ tibiotarsus + tarsometatarsus) was 20.5% greater, but the bones were more slender. Rando et al. suggest
that the Madeira Scops Owl was a ground-dwelling inhabitant of the laurel forests, feeding on the abundant
invertebrate life, and it may have taken reptiles and birds as well; it probably also nested close to or on the
ground. When Portuguese settlers arrived in the 15th century, the laurel forests were burnt off to make way
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for farming, and rats and mice were introduced. The loss of habitat, predation of its eggs and young, and
competition with rodents for limited invertebrate food supplies almost certainly caused the extinction of
the Madeira Scops Owl during the 16th or 17th centuries.
A second species of scops owl may have once occurred in Madeira, as poorly preserved subfossil remains
of an Otus were discovered on Porto Santo, an island to the northeast of Madeira (Rando et al. 2012). The leg
bones were variable, being longer, similar in size or slightly smaller than O. mauli. The bones either repre-
sent Otus mauli or a close relative, but until better material becomes available, Rando et al. describe the Porto
Santo owl as Otus cf. mauli.
Another now-extinct owl, the São Miguel Scops Owl Otus frutuosoi, once occurred on São Miguel in the
Azores (Rando et al. 2013). It was smaller than the Madeira Scops Owl, and had weak flying capabilities, so
it was probably a ground inhabitant in the laurel forests. It disappeared shortly after the colonisation of the
island by humans around 2,000 years ago.
The discovery of an endemic Hawaiian genus of strigid owls, all now extinct, exemplifies the importance of
the fossil record in ascertaining the original faunal diversity on oceanic islands. Based on characters of the
skull, the stilt owls appear closest to the genus Strix, but differ considerably in the proportions of the
post-cranial bones (Olson & James 1991).
The Kaua’i Stilt Owl was the size of the Tawny Owl Strix aluco at 37–43cm (14.5–17in) in total length, but
the wings were relatively more reduced, whereas the tibiotarsus and tarsometatarsus were much more
elongate, giving the bird a ‘stilt-like’ appearance. The toes were very robust, even more than in the large
Great Grey Owl Strix nebulosa. This suggests that the feet were incredibly strong, and that the owls probably
dispatched their prey by crushing it. Overall, Grallistrix, like Mascarenotus (see p. 224) and some other owl
genera, parallel the skeletal proportions of f orest-dwelling hawks of the genus Accipiter, which are specialist
bird-predators; in the absence of this genus the owls have occupied the vacant niche (Olson & James 1991).
These specialised owls no doubt suffered serious declines after human colonists arrived on the Hawaiian
Islands. The Polynesians cleared the lowland forests and deprived the owls of food, and if the owls were
roosting or nesting on the ground, the introduction of the Pacific Rat Rattus exulans may have been a major
predator of eggs and chicks.
This species was small, comparable in size with the Oahu bird (see below), but the femur was shorter,
whereas the tarsometatarsus was longer and more gracile. Subfossil remains of this owl were found at alti-
tudes ranging from 305m to 1,000m on the southern side of the Mount Haleakala crater. It is possible,
therefore, that this species occupied varied habits and altitudes on pre-human Maui, much the same as the
Laughing Owl Ninox albifacies did on New Zealand (Worthy & Holdaway 2002).
This species was smaller than the species on Kaua’i and Molokai, equal in size with Grallistrix erdmani of
Maui, but it had more robust leg elements and a shorter tarsometatarsus. The fossil localities where this
species have been found are in lowland coastal sites, which suggests that roosting occurred at times on open
sand dunes or in low dune vegetation (Olson & James 1991). This would have made the owls particularly
vulnerable to predation by introduced mammals.
The Molokai Stilt Owl was a large species, greater in size than the species on Oahu and Maui. There is a
good series of tarsometatarsi available, which show that size sexual dimorphism was present; in owls the
female is generally larger than the male. Like Grallistrix orion, subfossil remains were found in coastal
deposits, so they probably roosted or even nested in sand dunes or low dune vegetation.
The recently discovered Pernambuco Pygmy Owl is yet another species restricted to remnant forest patches
in the Atlantic forest of Brazil. It was first discovered in 1980 in Reserva Biológica de Saltinho in Pernambuco,
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a tiny protected area of 4.8 km2 composed primarily of old secondary forests; an individual was also tape-
recorded there in November 1990, and a single individual was observed on November 19, 2001 eating a
large cicada high in the canopy near the edge of a forest belt at Usina Trapiche, Sirinhaém (Silva & Coelho
2002). This is the last known recording. Silva & Coelho strongly suggested that a viable population could
not survive in the remnant forest in which it was observed, and since then continued deforestation has
reduced the habitat of the owl even further. Despite intensive searches using call playback surveys in both
in Pernambuco and Alagaos states since 2004, there have been no further authenticated records. It is likely
that the owl is now extinct (Pereira et al. 2014; Lees & Pimm 2015).
Habits
Little was recorded about its habits. It occurred in forest up to 150m in altitude, and was observed in the
canopy of secondary forest, suggesting that it might be able to utilise this modified forest type (Silva &
Coelho 2002). It fed on insects, but probably took small mammals, birds and reptiles as well. The primary
call comprised a short series of 5–7 notes uttered at around 4.2 notes per second.
The tiny Elf Owl Micrathene whitneyi is the world’s smallest owl; the species is divided into four subspecies,
the nominate in mainland south-west United States, one in Texas and Mexico, one at the the southern end of
Baja California, and M. w. graysoni on Socorro Island in the Revillagigedo Islands. The Socorro Owl was
considered common in 1932 (Jehl & Parkes 1982), and a large population apparently still existed in 1958, as
this was reported to Villa (1960), although they were not observed by him. However, the bird’s numbers
appear to have crashed shortly after, with the last record occurring around 1970.
A search for the subspecies in 1978 and 1981 was unsuccessful (Jehl & Parkes 1982). Sheep were intro-
duced to Socorro in 1869 with devastating effects on the vegetation (Brattstrom & Howell 1956), and the
establishment of military personnel, followed by feral cats in 1953 (Jehl & Parkes 1982), must have all
contributed to the Socorro Elf Owl’s demise.
Habits
Little was recorded about the bird. Like the other Elf Owl subspecies, they were insectivorous, foraging at
night, dawn and dusk. Their call was a high-pitched whinny or chuckle.
Burrowing Owls Athene cunicularia occur over the drier parts of the Americas from southern Canada to
Tierra del Fuego, including a number of islands. Their terrestrial habits have made them vulnerable to the
effects of human activities, and their nests at the end of long tunnels makes them susceptible to predation
by introduced mammals. Little was recorded about the Marie-Galante subspecies. It appears to have never
been common, and became extinct shortly after mongooses were introduced to the island in the late 19th
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century; this predatory mammal is thought to have been solely responsible for the bird’s extinction (Bond
1950).
Only six specimens of A. c. guadeloupensis exist, all collected by L’Herminier early in the 19th century
(Greenway 1967). Bond (1950) pointed out that there is no evidence that the bird ever occurred on
Guadeloupe, as had originally been thought (Ridgway 1874), but it was in fact confined to M arie-Galante.
Habits
All observers agreed that the bird was nocturnal, and lived in old drains, holes in cliffs, and ruined walls. It
was locally called C
oo-coo from its hoot at night. Captive birds frequently uttered a chattering cry, but they
did not hoot. Ober (in Lawrence 1878b) was told the owls fed on lizards and mice.
The Bermuda Saw-whet Owl was probably derived from the Northern Saw-whet Owl Aegolius acadicus, but
differed in the proportions of the skeleton. The mandible was much smaller, the tarsometatarsus was of
equal length but more robust, and the wings were slightly reduced (Olson 2012b). Olson suggests that the
small mandible was indicative of the Bermuda Saw-whet Owl being diurnal and having a small head; it
would not have needed the high degree of development of the auditory apparatus, which gives a nocturnal
owl the characteristic ‘big-headedness.’ It probably fed on the smaller endemic birds, lizards and large
insects, which Bermuda once had in abundance. Northern Saw-whet Owls are known to roost in conifer
trees, and nest in the abandoned holes of flickers (a group of small woodpecker). The Bermuda Saw-whet
Owl probably utilised the Bermuda Cedar Juniperus bermudiana (Olson 2012b), and the extinct Bermuda
Flicker Colaptes oceanicus (see p. 243) would have provided plenty of nest sites, especially in the soft-trunked
Bermuda Palm Sabal bermudana (Olson 2012b, 2013).
The Bermuda Saw-whet Owl was almost certainly still extant when Europeans first arrived on the island.
Olson (2012b) reports the account of William Strachey, who was exiled on Bermuda in 1609–1610, and stated
that an owl was present. A Captain John Smith (Lefroy 1981; Olson 2012b) provided more information, and
wrote in 1623 that:
. . .there were a kinde of small Owles in great abundance, but they are now all slaine or fled.
The Bermuda Saw-whet Owl had been present on Bermuda for at least 80,000 years and remained rela-
tively morphologically unchanged from the mainland form (Olson 2012b). Human destruction of habitat,
which resulted in the decline of suitable roosting and nesting trees, plus the introduction of predatory
mammals, resulted in the extinction of the owl in just over a decade.
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The Lord Howe Boobook was first mentioned by MacDonald (1853), who stated that ‘a small species of owl
prowls about at night’. Little else was recorded about the bird, though Etheridge (1889) stated:
N. boobook frequents thickets of scrub on the hill sides during the day and may be found around the settlements
at night, probably on the lookout for mice, otherwise its food is a mystery, unless it be nocturnal insects.
The southern Australian subspecies of Southern Boobook Ninox boobook was released on the island in the
1920s, as were Australian Barn Owls Tyto alba delicatula and Masked Owls Tyto novaehollandiae, in an unsuc-
cessful attempt to exterminate the Black Rat Rattus rattus, which had escaped from the steamship SS
Makambo when it grounded in June 1918.
Hindwood (1940) implies that the native boobook was extant at the time of writing. Calls believed to be
of the endemic race were reported up to the 1950s, but may in fact have represented the calls of any of the
owl introductions. The Lord Howe Boobook probably disappeared as a result of rat predation of eggs and
chicks and predation of adults by introduced owls, and perhaps competition with the introduced owls for
food. Being a tree-cavity nester, deforestation must also have impacted on the species.
The Norfolk Island Boobook was first mentioned by John Hunter, Captain on the Sirius transporting
convicts to Norfolk, who later became governor of New South Wales (Percival 1949). He stated:
The owls, which have very handsome plumage, make a noise like one man calling to another, and they pronounce
the word “yaho” very distinctly.
The decline of the Norfolk Island Boobook began with the anthropogenic changes to its forest habitat.
Boobooks are tree nesters, so deforestation would have been particularly detrimental to the population. As
the forests became fragmented, there was also an increase in competition for nest-hollows with introduced
honeybees and Crimson Rosellas Platycercus elegans (Garnett & Crowley 2000).
By 1986, the subspecies had been reduced to a single bird, a female called Miamiti (Anon 2004). The bird
was now effectively extinct, but in order to conserve at least some of the genes, two males of the nominate
New Zealand race N. n. novaeseelandiae were introduced to pair with the last female (Olsen et al. 1989), as
DNA sampling showed that this subspecies was genetically closest to the Norfolk population (Norman et al.
1998). One male disappeared almost immediately, but the second male successfully mated with the Norfolk
female and young were fledged in 1989 and again in 1990. Miamiti, the last p ure-bred Norfolk Island
Boobook, died some time in 1996, but a small hybrid population of 12–13 birds survives on the island
(Double & Olsen 1997), which are slowly increasing (Higgin 1999; Garrett et al. 2010). This tragic scenario
epitomises the finality of extinction, when a l ast-ditch effort to save a species can only result in hybrid birds.
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Habits
The owl appeared to favour the island’s subtropical rainforest, which was almost completely cleared in the
19th century following human settlement, and by 1909 was confined to the gullies surrounding Mount Pitt
(Olsen et al. 1989). It nested in nest-hollows in large mature trees, and first bred when three to four years old
(Olsen 1996); eggs were laid from September to October.
Balouet & Olson (1989) described the subfossil remains of a Ninox owl, which was much smaller than the
Australian N. boobook, and more similar to the New Zealand Boobook N. novaeseelandiae. The owl is known
from a left femur and two tibiotarsi from Pindai Cave, on the west coast of the mainland, and a partial ulna
from Gilles Cave, which is situated about 7km inland and also on the west coast. In terms of biogeography,
it is no surprise that a member of this genus once occurred on New Caledonia, as populations also occurred
on remote Lord Howe and Norfolk Islands.
Because nocturnal birds are so poorly known on New Caledonia, Balouet & Olson suggest that the New
Caledonian Boobook Owl might still exist, but no mention was ever made of the bird and the reasons for its
disappearance (or extreme rarity) are unclear.
Both races of Laughing Owl disappeared rapidly when the human population of New Zealand increased,
at first from the vicinity of settlements and then throughout their range; it is believed that introduced
predators were primarily responsible for the decline (Greenway 1967). There seems to have been a
marked decrease before 1880, with the last official specimen record in July 1914, when a dead individual
was found. The Laughing Owl was not yet extinct, however. Photographic evidence has shown that the
birds were still breeding in 1916 near Albury, south Canterbury, and remained common throughout the
1920s (Worthy & Holdaway 1993). One individual, an immature, was also photographed alive between
1909 and 1912 at its nest by Cuthbert and Oliver Parr (see Fuller 2000). Oliver Parr records also seeing
owls around 1922–24, but this remains unconfirmed, and later reports at Lake Thompson in 1930 and
Mount Maud in 1939 (Williams & Harrison 1972), and Fiordland in 1956 (Hall-Jones 1960) are also
unconfirmed. A remarkable record concerns egg fragments, found in Waianakarua in 1960 (Williams &
Harrison 1972).
Their staple diet was supposed to be the Pacific Rat Rattus exulans, and Buller (1906) suggested that the
scarcity of this species, apparently killed off by introduced rats, may have been responsible for the owl’s
demise, a view that is not generally accepted. Gurney (1896) however, found that a Laughing Owl captive
in England much preferred small birds to rats or mice as food, and birds provided the staple diet before
humans colonised New Zealand, though they also took a range of other foods as well (Worthy & Holdaway
2002). As the owls roosted and nested on the ground (Williams & Harrison 1972; Worthy & Holdaway 2002),
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Myers (1923) believed that stoats and weasels preyed on them, and they were also likely to have been
vulnerable to predation by cats and rats.
It was thought that laughing owls formerly also inhabited the Chatham Islands (Forbes 1892b), but
Milliner (1999) has shown that this was based on erroneous identification of fossil material.
Relationships
A recent DNA study has shown that the Laughing Owl is nested within the boobook genus Ninox and basal
to the extant Morepork N. novaeseelandiae, thus not warranting a separate generic placement (Wood et al.
2016). The authors conclude that the Laughing Owl was probably the first in a double invasion of New
Zealand by the same Australian owl lineage, but because the niche of a small nocturnal predator was
already occupied by the New Zealand Owlet-nightjar Aegotheles novaezealandiae (see p. 235), the laughing
owl evolved into a larger nocturnal predator to utilise different food resources.
Habits
Laughing Owls were generally found in and around rocky limestone areas, either in open country or along
forest margins and scrub, but Worthy & Holdaway (2002) have shown that they occupied a variety of habits,
including forest. They hunted for food over open country, and probably spent much time on the ground,
when roosting or nesting in rock fissures (Williams & Harrison 1972). Smith (1884) lamented that the species
would be extinct soon after the date of writing, and that it was only in the last couple of years that anything
had become known of its habits. He believed that the birds nested in high rock crevices, not on the ground.
The breeding season began in September, with both parents incubating the eggs for a period of 25 days, the
female more frequently. They nested in crevices or small caves in limestone cliffs. Usually three eggs were
laid, but sometimes there was a second clutch. The young appear to have been fed on invertebrates, particu-
larly worms.
The call was reported to be a loud coo-e-e, while captive birds variously uttered a low chuckle like a
turkey, a cat-like mew, a puppy-like yelp, and a tuneless whistle (Gurney 1896; Greenway 1967). The owl
had very elongated legs, and unlike most other strigid owls the tarsometatarsus was longer than the femur.
This suggests that the Laughing Owl was a terrestrial species that had evolved into a large nocturnal pred-
ator feeding on a predominantly ground-dwelling fauna (Wood et al. 2016). Their diet varied depending on
geographical location, feeding predominantly on petrels and parakeets in areas where they were abundant,
and frogs and lizards in others (Worthy & Holdaway 2002). They also preyed on rails, pigeons, small birds,
lizards (including Tuatara Sphenodon punctatus), the endemic terrestrial mystacinid bats and large inverte-
brates, including the flightless Giant Weta Deinacrida sp. (Wood et al. 2016) and even the Morepork and
extinct New Zealand Owlet-nightjar were taken (see p. 235).
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Laughing Owls began to disappear from North Island in the first half of the 19th century, or even earlier,
and became extinct soon after 1889, in which year there was a sight record at Waikohu, Te Karaka (Buller
1905). Another was observed in 1890 (Oliver 1955), and a bird was recorded nesting in a hut before 1892
near Porirua harbour (Buller 1892). One was heard at Easter 1927 at Wairaumoana, the southwestern leg of
Lake Waikaremoana (Blackburn 1927), (and not 1925, contra St Paul & McKenzie 1977), and a very doubtful
sighting was made near Purangi, Waitara Range around 1930 (Williams & Harrison 1972).
Oliver (1930) stated that only two specimens were ever collected in North Island, the first by a taxider-
mist called Martin on Mount Egmont, Taranaki, in about 1856, and the type from Wairarapa District, about
50 miles from Wellington in either 1868 or 1869. Both are now lost (Worthy 1997).
The bird was apparently w ell-known to the Maoris, who said it was once found in several districts.
Subfossil and midden bones are known from various sites in North Island. These show that it was larger
than the South Island form (Gill 1996), although Buller (1906) claimed that the North Island form was
smaller. Worthy & Holdaway (2002) doubt the validity of this subspecies, and note that size differences
appear to be a clinal increase from north to south.
Rothschild (1907a), after examining the type, found it to be a s ubadult, and, like Buller, concluded that the
tail had been tampered with; Rothschild confirmed that the tail had been replaced with that of an Australian
Ninox.
Caprimulgidae (Nightjars)
The nightjars are a family of nocturnal or crepuscular birds with short bills but extremely large mouths,
with which they catch insects on the wing. The plumage is cryptic, providing effective camouflage when
sitting motionless during nesting and periods of inactivity. They have a cosmopolitan distribution and the
family is divided into up to 14 genera, depending on the authority. Nightjars are also endemic to islands
and island groups, where they are threatened by habitat destruction.
All four specimens of this nightjar were taken in the western part of the island, in Westmoreland and
Freeman’s Hall, in Trelawny. Cleere (1999) gave the following details about the four specimens: the first, a
male, was taken in about 1844; the second, also a male, at Savanna la Mar in the Bluefields area of
Westmoreland in August 1858; the third was a possible juvenile female taken in September 1859 at Freeman’s
Hall near Albert Town in Trelawny; and the fourth was a male taken near Linstead around November 1860.
This species was probably always rare, for neither Gosse (1847) nor any of his correspondents in the 1830s
mentions it, although the last known specimen was taken in 1860.
The extinction of the bird is believed to be due to predation by introduced mammals. The mongoose was
not introduced to Jamaica until 1872 (Bond 1950), so presumably the bird was exterminated by rats before
this date. Greenway (1967) suggested that a few might still survive, though Stuart Panton (in Bond 1950)
considered them to be definitely extinct. However, there are unconfirmed reports from the Milk River and
Hellshire Hills (Cleere 1999), but if it does still exist it must be extremely rare.
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Aegothelidae
Habits
Nothing was recorded about the habits of the Jamaican Pauraque. It was sedentary, crepuscular and
nocturnal, presumably nesting on the ground and feeding on insects (Cleere 1999, 2010). All specimens
were taken in the lowlands on the drier south side of the island, in dry limestone forest, s emi-arid wood-
land or in more open country.
This nightjar was intermediate in size between the Least Poorwill S. brewsteri (17–21cm) and Jamaican
Poorwill S. americana (23–25cm), with the tarsometatarsus comparatively slender as in S. brewsteri (Olson
1985b). Its remains were discovered in cave deposits alongside the extinct endemic insectivorous mammals
of the genus Nesophontes and some extinct microchiropteran bats, all deposited by the Barn Owl Tyto alba. It
is likely that the Cuban Pauraque was preyed on by the owls, which still occupy the caves. Olson (1985b)
suggests that a small, cryptic and nocturnal nightjar that occupied the little-explored arid regions east of the
type locality of Daiquirí may still exist.
Aegothelidae (Owlet-nightjars)
The Aegothelidae, containing a single genus, Aegotheles, are nocturnal, insectivorous birds, related to the
nightjars and frogmouths. New Guinea is the family’s diversity h ot-spot, but they also occur in Australia,
the Moluccas, on New Caledonia and formerly in New Zealand. Like nightjars they are cryptic birds,
remaining motionless on branches or on the ground during the daytime.
The New Zealand Owlet-nightjar was the largest known species in the family, living or extinct, and weighed
an estimated 150–200 g, making it also the heaviest in the genus. The wings were comparatively reduced,
but the keel of the sternum was strong, indicating that flight was possible, albeit perhaps laboured (Rich &
Scarlett 1977). It had long legs and robust feet, which strongly suggest that it was terrestrial, probably
spending all or most of the time on the ground. It appears to have been widely distributed before human
settlement on both North and South Islands, and its remains are often found in the pellets of the extinct
Laughing Owl Ninox albifacies, which may have been its chief predator.
It was thought to have disappeared before European settlement (but see below) and has not so far been
found in association with Maori middens, probably because of its small size and nocturnal habits. Holdaway
et al. (2002) obtained a series of carbon i sotope-derived ages from o
wlet-nightjar bones, which indicate that
a population decline occurred prior to Polynesian settlement. This decline was probably consistent with the
arrival of the Pacific Rat Rattus exulans, a terrestrial-bird predator that colonised Pacific islands as a
stow-away on Polynesian boats. It appears that the rats may have colonised New Zealand prior to human
settlement on unmanned drifting boats, in which the Polynesian occupants had perished during the voyage.
A poorly flighted, ground-inhabiting owlet-nightjar would have been easy prey for the rats, and it seems to
have disappeared soon after their arrival.
Live observations
Despite its probable extinction before European colonisation, Potts (1871; see also Tennyson & Martinson
2007) mentioned some tantalising 19th-century accounts, which suggest that the owlet-nightjar may have
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New Zealand Owlet-nightjar survived into the historic period. The bird in ques-
Aegotheles novaezealandiae tion was described by Ellman (1861) as a ‘Little
Owl Strix parvissima’, but based on observations
and accounts only. Ellman described it as very
scarce, not larger than a starling, the head very
large, seen among the forests and exceedingly shy,
and Potts (1871) reported it as ‘about the size of a
kingfisher, being exceedingly gentle when
captured’, to name just a few. Unfortunately no
specimens were saved to confirm their identity.
The New Zealand Owlet-nightjar is now certainly
extinct.
Apodidae (Swifts)
The swifts are charecterised by their long, narrow,
scythe-shaped wings, are are among the fastest
flying birds. They spend most of their lives on the
wing. Although superficially they appear similar
to swallows, they are in fact more closely related to
the hummingbirds.
A number of swifts use saliva to glue their nests
to vertical surfaces, the main ingredient for
bird’s-
nest soup, which has resulted in their
decline in many areas. Island species are also
affected by habitat destruction.
There are about 26 species of Aerodramus swiflets, widely distributed across southern Asia, Oceania,
north-eastern Australia and the Indian Ocean (Chantler & Driessens 2000). They are cave-nesting species,
known for the supposed delicacy of their nests, a principal component of soup; the birds have widely
reduced in numbers due to over-harvesting. The Mangaia Swiftlet is the first species of Aerodramus to be
described from subfossil remains; while similar to the extant ’Atiu Swiftlet A. sawtelli, an endemic on ’Atiu,
the nearest island to Mangaia, it was larger and differed in characteristics and proportions of the skeleton
(Steadman 2002a). Along with all extant Aerodramus, the Mangaia Swiftlet was probably able to echolocate
in order to navigate in the dark of the caves.
Mangaia has numerous caves but swiftlet remains are rare, which may have been a direct result of
Polynesian use of the caves as shelters and burial grounds. It is unlikely that the birds were hunted as food,
but continued human disturbance and perhaps introduced avian predators may have caused their disap-
pearance. The Cook Islands may well harbour more extinct species of Aerodramus swiftlets, and have now
become the first island group east of Vanuatu known to have had multiple swift species (Steadman 2002a).
Trochilidae (Hummingbirds)
The hummingbirds are restricted to the Americas, though they occur from Alaska in the north to Tierra del
Fuego in the south, and throughout the Caribbean. Their taxonomy is complex, and the family has been
divided into more than 100 genera. They exhibit a diverse range of bill shapes and sizes, and one species,
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Trochilidae
Mangaia Swiftlet
Aerodramus manuoi
the Bee Hummingbird Mellisuga helenae, is the smallest known bird at just 5cm (2in) long. Many species
have been described from single trade specimens, which have subsequently proved to be hybrid individ-
uals. Gary Graves of the United States National Museum in particular has done much to resolve these
taxonomic problems (see Appendix 1). Some species with restricted ranges are in extreme danger of extinc-
tion due to habitat destruction, the island endemics particularly so.
This species had long been regarded as a doubtful member of the genus Amazilia. Weller (1999b) examined
the type (formerly in the Gould Collection, now at Tring), comparing it with species of Chlorostilbon and
Amazilia, and found that none of these genera occurred together geographically, nor could they have
produced a hybrid resembling C. elegans, thus confirming the species’ validity.
Nothing is known about its habits or why it disappeared. We are not even sure of its provenance. If it was
an inhabitant of a West Indian island, it is likely that human modification of habitat and introduced preda-
tors caused its extinction.
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Alfaro’s Hummingbird is a close relative of the Indigo-capped Hummingbird Amazilia cyanifrons, and it is
sometimes considered a race of it. It is known only from the Miravalles Volcano in Costa Rica; there are no
recent records. Weller (2001) considers that A. alfaroana may represent an extinct species, pointing out that it
has unique plumage features not found in any of its relatives. Nothing was recorded about its habits, and
the reasons for its disappearance are unclear.
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Brachypteraciidae
Description Male 12cm (4.75in); female 10cm (4in). Exhibited a high degree of sexual dimorphism, perhaps the greatest found
among hummingbirds; the male and female were originally thought to be of different species. Male bright rufous-orange, flight
feathers dusky; crown reddish yellow, with dusky lores; immature male spotted rufous on crown. Female dark blue-green
above, with bluish tinge on crown; underparts whitish, throat densely spotted with dark green, extending onto flanks.
This race was originally described as a full species (Gould 1870), but it was subsequently relegated to
subspecific rank. The differences between it and the nominate S. f. fernandensis of Isla Róbinson Crusoe are
slight and fall within the range of individual variation. There is also doubt as to the true provenance, as
Gould obtained his specimen from the botanist Friedrich Leybold, who may not have even been on the
island (Hahn et al. 2009). Furthermore, the supposed last record was in 1908, but during a thorough survey
undertaken in the late 19th century, not a single specimen was observed (Johow 2003).
The nominate population is considered critically endangered, due to deforestation, predation by intro-
duced rats and cats, invasions of exotic plants which reduce the numbers of its key nesting tree, Luma
apiculata, and severe erosion caused by goats and rabbits. The Alejandro Selkirk Island Firecrown probably
disappeared for the same reasons.
Habits
Little was recorded about its habits. On Isla Róbinson Crusoe the birds inhabit forests, thickets, and gardens,
but the remnant native forests appear to be crucial for successful breeding. They feed predominantly on
nectar, but insects are also taken. Two white eggs are
Bogotá Sun-angel
laid in a small c up-shaped nest typically 3–4m above Heliangelus zusii
ground, and almost exclusively in Luma apiculata.
Brachypteraciidae (Ground-rollers)
The ground- rollers are endemic to Madagascar. The
family is divided into three genera. They are secretive,
terrestrial birds that live in rainforest as well as sub-arid
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thorny scrub. They all have restricted ranges, and have declined due to the severe deforestation that has
taken place on Madagascar.
The Ampoza G round-roller is known from a single humerus collected in Ampoza, in south-west Madagascar
(Goodman 2000). Measurements of the humerus indicate that it was more robust than any surviving ground
roller. The two extant species of Brachypteracias, the Scaly G round-roller B. squamiger and Short-legged
Ground-roller B. leptosomus, are humid, mesic forest inhabitants. As the type locality of B. langrandi is now
situated in dry deciduous forest, Goodman hypothesised that was a possible victim of natural desiccation,
in which the original, more humid mesic forest gradually disappeared. Nothing else is known about this
species.
Halcyonidae (Kingfishers)
The kingfishers are generally small to
medium-sized fish- and insect-eating Mangareva Kingfisher
Halcyon gambieri gambieri
birds, which inhabit wetlands and
forests in Africa, Asia, and Australasia,
and have colonised remote islands
and island groups in the Indian and
Pacific Oceans. The family is presently
divided into 12 genera. Kingfishers
confined to islands have suffered from
habitat destruction and introduced
predators, and some are in danger of
extinction.
Mangereva Kingfisher
Halcyon gambieri gambieri
Oustalet
Halcyon Gambieri Oustalet 1895, p. 182
(Mangereva, Gambier Archipelago,
Tuamotus)
Specimens The type specimen is in Paris.
Status Known only from the type, collected
in 1841.
Range Mangereva in the Gambier Islands,
Tuamotu Archipelago, South Pacific.
Description 20cm (8in). A green and white
kingfisher, white on the neck and underparts,
and bluish green on back, wings and tail:
head and neck russet-buff with some
greenish-blue feathers on crown; broad green
eye-band; chin and underparts white; mantle,
back, rump, wings and tail blue; bill black;
legs and feet grey; iris brown. Differed from
H. g. niauensis by being darker on head,
upperparts greenish rather than blue, and
lacking tinge of buff on neck band and chest.
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Upupidae
The Gambier Islands form a single encircling reef enclosing five large and several smaller islands. All the
islands of the group are steep, rugged and forested. The main island is Mangereva, about 6.4km long, and
the highest peak, Mount Duff, is 380m high. Mangereva’s endemic kingfisher was the nominate race of the
Tuamotu Kingfisher; the other race, H. g. niauensis on Niau, is critically endangered (Gouni & Zysman 2007).
The type of the Mangereva bird was collected in 1841 by Dumont d’Urville on the voyage of the Astrolabe
(Oustalet 1895). None was found by the Whitney South Seas Expedition in 1922 (Murphy 1924), nor by any
subsequent visitors. There can be no doubt that it is extinct. Ernest Quayle, one of the Whitney Expedition
members (quoted by Holyoak & Thibault 1977a), noted considerable deforestation, and pests such as cats,
rats, goats and dogs present on the island, which are the main factors effecting the Niau population today
(Seitre & Seitre 1992).
Habits
On Niau, the kingfisher forages in various habitats such as limestone forests, wetlands and open coasts, but
also in areas modified by humans. They excavate nest holes in dead or decaying palm trunks, and feed on
insects and small lizards (Gouni & Zysman 2007).
This race of the Sulawesi Dwarf Kingfisher C. fallax was last seen in 1997, and a thorough survey in 1998
failed to locate any birds (Riley 2002). It was reported to have occurred on the Talaud Islands, but this is in
error. Habitat destruction appears to be the primary cause of its disappearance, and intensive logging and
subsequent conversion of land to agriculture has severely depleted its habitat.
Almost nothing was recorded about this bird. The nominate race on Sulawesi occurs in dry lowland
primary forest up to an altitude of 1,000m, but generally at approximately 600m. This habitat has been
hardest hit by human disturbance.
Upupidae (Hoopoes)
The Common Hoopoe Upupa epops is widely distributed in Africa and Eurasia, and is the only surviving
member of the family. It is a distinctive bird with an erectile crest and a long, decurved bill. It feeds exclu-
sively on the ground and has a strong odour, which acts as a defence against predators.
This was a very large hoopoe, with longer and more robust cranial and wing elements than the extant
Common Hoopoe Upupa epops, from which it is probably derived (Olson 1975b). The distal wing elements
are reduced, indicating that it had restricted powers of flight. The bill is also larger and heavier, and notice-
ably more decurved than in Common Hoopoe.
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Ashmole (1963b) discovered the first subfossil remains of the St Helena Giant Hoopoe, but assumed they
belonged to the Common Hoopoe, a bird associated with grasslands and open forests. Olson (1975b)
suggested that a native hoopoe would have had a similar ecology, negating the assumption that St Helena
was formerly covered entirely by heavy forest in p re-European times (see Ashmole & Ashmole 2000). This
is further supported by St Helena’s only surviving endemic bird, the Wirebird Charadrius sanctaehelenae,
which is an open grassland inhabitant. The St Helena Giant Hoopoe probably occurred in areas of open
ground with reduced vegetation cover within the forested areas (Rowland et al. 1998).
Fossil remains of the St Helena Giant Hoopoe have been found on the saline semi-desert of Prosperous
Bay, which was originally bordered by a Scrubwood scrub region and a Dry Gumwood region, these once
providing suitable habitat for the birds (Lewis 2008). Their reduced flying capabilities probably restricted
their movements around the island, so they were unlikely to have moved far from the feeding areas (Lewis
2008).
The robust limbs coupled with a large, long bill are adaptations for strong probing, no doubt utilised
when the bird was foraging for insects (such as the recently extinct Labidura herculeana, the world’s largest
earwig) and other invertebrates. The arrival of Europeans on St Helena in 1502 sealed the fate of the bird. It
was probably exterminated by introduced rats and cats, severe deforestation, and perhaps loss of inverte-
brate prey items. The St Helena Giant Hoopoe may have survived until around 1640 (Lewis 2008).
Bucerotidae (Hornbills)
The hornbills are distinctive birds with large, decurved bills, often brightly coloured and sometimes with a
casque on the mandible. The family is divided into 14 genera, distributed widely in Africa, Asia, and
Melanesia. Some species are in critical danger of extinction, primarily due to habitat destruction.
Status Probably extinct. Last confirmed report in 1971. The status of this subspecies is uncertain, and considered by some
authorities to be a colour morph.
Range Ticao, Philippines.
Description 65cm (25.5in). The Ticao bird was very much larger than nominate P. p. panini (Kennedy et al. 2000). The adults
exhibit sexual dimorphism. Male head and neck cream-white; ear-coverts and throat black; upperparts black, glossed green;
upper breast white, lower breast and u
ppertail-coverts rufous brown; tail whitish buff with a broad black tip; bill dark reddish
black with yellow ridges; casque blackish red; bare ocular skin white tinted bluish; iris red; legs and feet dark brown. Female
similar except head and entire underparts black; bare orbital skin blue; bill duller.
The Ticao Tarictic Hornbill was a race of Penelopides panini, which occurs on the islands of Panay, Negros,
Masbate and Guimaras in the Philippines. It is declining or has been extirpated from a number of islands,
mainly due to forest destruction and o ver-hunting. In 1902 the Ticao race was considered abundant
(McGregor 1909a), and a large series of skins and some eggs were collected. Since then, almost all the forest
on the island has been replaced by plantations and human settlements. The last observation of the hornbill
was made in 1971 (BirdLife International 2011). This bird is now very likely to be extinct.
Habits
The Ticao Tarictic Hornbill was an inhabitant of primary evergreen forest, and nested in tall trees; the female
was typically sealed into the nesting cavity and fed by the male (McGregor 1909a). The diet was predomi-
nantly fruit, but small-
vertebrate and invertebrate prey was also consumed, particularly during the
breeding season.
Steadman (2006a) confirmed the presence of a hornbill on Lifou in the Loyalty Islands. This bird was smaller
than Blyth’s Hornbill A. plicatus, the hornbill found in New Guinea and Indonesia. This is the first evidence
for a hornbill species east of the Solomon Islands main chain, and confirms that hornbills are capable of
long-distance, cross-water dispersal (Diamond 2002).
The subfossil remains were discovered in archaeological middens, and as hornbills are considered good
eating, it is probable that the Lifou birds were hunted to extinction by Polynesians. They would probably
have also been vulnerable to deforestation and the Polynesian introduction of the Pacific Rat Rattus exulans.
Picidae (Woodpeckers)
The Picidae have a wide distribution including some island archipelagos, but are absent from Australia,
Madagascar, New Zealand and the majority of the polar regions. Woodpeckers are divided into up to 30
genera (depending on authority). They are generally forest birds, but some species inhabit treeless zones
such as deserts and rocky scree hillsides. Many are threatened due to habitat destruction, and the largest
members of the family all required large areas of undisturbed forest – the Imperial Woopecker Campephilus
imperialis, Ivory-billed Woodpecker C. principalis and Cuban I vory-billed Woodpecker C. bairdi; all are now
almost certainly extinct.
The Northern or Yellow-shafted Flicker Colaptes auratus has a wide distribution, occurring over most of
North America, Mexico, Cuba, and the Cayman Islands (Short 1982). It has been divided into up to nine
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subspecies, with some authorities considering the Red-shafted Flicker Colaptes cafer to be a subspecies of it
(Winkler & Christie 2002). A probable derivative of the Northern Flicker once occurred on Bermuda.
The Bermuda Flicker Colaptes oceanicus was smaller than all mainland and Cuban subspecies of Northern
Flicker, but it was larger than the extant Grand Cayman bird C. a. gundlachi (Olson 2013). The morphology
of the sternum and carpometacarpus indicate that the Bermuda Flicker had reduced flying abilities, and the
discovery of juvenile bones shows that the population was once a resident, endemic species (Olson 2013).
Olson also suggests that the more open forested habitat and recently burned forests preferred by the
Northern Flicker (Moore 1995; Winkler & Christie 2002) may have been maintained on Bermuda by light-
ning-caused fires. The Bermuda Flicker may have excavated its roosting and nest holes in the endemic
Bermuda Palm Sabal bermudana, which was once one of the dominant trees on the island (Olson 2012, 2013).
Other species may have utilised abandoned flicker holes, including the extinct Bermuda Saw-whet Owl
Aegolius gradyi (Olson 2012; see p. 229). Flickers are generally terrestrial foragers feeding on invertebrates,
especially ants, fruit and seeds (Moore 1995; Winkler & Christie 2002), so presumably the Bermuda birds
would have had similar feeding habits. It is likely that the Bermuda Flicker survived until at least the early
1600s, as Olson (2013) quotes the account of a Captain John Smith, written in 1623:
Neither hath the Aire for her part been wanting with due supplies of many sorts of Fowles . . . numbers of small
birds like Sparrowes and Robins, which have lately beene destroyed by the wilde Cats, Wood-pickars, very many
Crowes. . .
There is a possibility that Smith’s ‘Wood-pickars’ were the migratory Yellow-bellied Sapsucker (Sphyrapicus
varius), which also occurs in small numbers on Bermuda, but more importantly the account shows that feral
cats were present. These ferocious predators contributed significantly to the extinction of the Guadalupe
Flicker Colaptes cafer rufipileus (see below), so no doubt on Bermuda, a naive, poorly volant species that
habitually foraged on the ground would have been quickly eliminated.
The Red-shafted Flicker C. a. cafer occurs across North America from Alaska to Nicaragua, and is divided
into a number of geographical races (Short 1982). The Guadalupe race of this species was discovered by Dr
Edward Palmer in 1875 (Ridgway 1876b). Even at the time of its discovery it was comparatively rare, and
apparently confined to the pine woods at the north end of the island (Bent 1939). W. E. Bryant (1887) found
it in a restricted grove of cypress on top of the ridge on Mount Augusta, the highest point on the island, and
also occasionally in other places where there were stands of pine or palm. However, Henry Kaeding (1905)
found the bird ‘rarely seen’ when he visited the island in 1897.
The last twelve skins and six clutches of eggs of the Guadalupe Flicker were collected in May and June
1906 by W. W. Brown (Thayer & Bangs 1908), who estimated that there was a population of 40 birds, but it is
unclear whether this was before or after Brown had collected the twelve. Incredibly, this idiotic act of
thoughtless greed gave no consideration to the fact that it could be the coup de grâce for the population, a
case similar to that of the Guadalupe Caracara Polyborus lutosus (when nine were shot from the probable last
flock of 11; see p. 93). Between 1906 and 1922 there is no information, but after 1922, when naturalists of the
California Academy of Sciences visited Guadalupe, they failed to find any flickers (Greenway 1967). None
has been seen since, though a number of people have searched for them.
The main reason for extinction is thought to have been destruction of the habitat by goats; present in 1886
(Finlay 1886), these numbered 6,000–8,000in 1906 (Thayer & Bangs 1908). The island was also overrun by
cats, and Brown in 1906 reported that the birds were very tame and fell easy prey to them. Thayer & Bangs
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Picidae
(1908) remarked that the flicker bird was probably ‘doomed to speedy extinction, and will be the next of the
Guadaloupe [=Guadalupe] birds to go’. Unfortunately, their prediction came true.
The White-bellied Woodpecker D. javensis has a wide distribution in Southeast Asia, from western India
through to Japan, and has been divided into 14 subspecies (Winkler & Christie 2002). It inhabits a range of
forest types, including evergreen and deciduous, but can also tolerate degraded, secondary forest. Some
subspecies are in decline and the endangered Korean race, D. j. richardsi, is now extirpated on the Japanese
island of Tsushima. The Cebu White-bellied Woodpecker, D. j. cebuensis, was first collected by Bourns &
Worcester (1894) on 22 June, 1892 during the Manage Scientific Expedition to the Philippines. Only two
more specimens were ever collected. By the time the subspecies was recognised as distinct (Kennedy 1987)
it was long gone – the last record appears to have been in the 1940s (Delacour & Mayr 1946) or 1950s
(Paguntalan & Jakosalem 2008). This large woodpecker disappeared due to the severe deforestation that
has taken place on Cebu.
At a meeting of the Zoological Society of London, held on August 14, 1832, specimens were exhibited of a previ-
ously undescribed Woodpecker, remarkable for its extraordinary size. These specimens, the male of which
measured two feet in length, were said to have been obtained by Mr. Gould from ‘that little explored district of
California which borders the territory of Mexico’ a statement which serves as a good illustration of the vague
ideas of American geography that prevailed among naturalists of those days. Mr. Gould made a felicitous choice
of name when he called this bird Picus imperialis, for it is by far the largest and most striking member of the
Woodpecker family in the world.
So wrote E. W. Nelson (1898), after an expedition into the mountains to find this bird. On shooting at a pair,
he further commented:
The male was only winged and as we approached threw himself over on his tail, with outspread wings, presenting
a warlike front of threatening beak and talons. It was impossible not to admire the courage and defiance shown by
the fierce glow of his golden-yellow eyes and upraised flaming crest . . . The birds were surprisingly easy to stalk,
even after being hunted and shot at for several days, but were difficult to secure because they are powerful, hard-
muscled creatures possessed of remarkable vitality. They showed considerable attachment to one another and
245
when one was shot the other members of the flock remained scattered about on the trees for a short time calling
each other at intervals. Wounded birds fought with savage courage.
One cannot help wondering what survival mechanism prompted the evolution of such behaviour. One also
feels frustrated that with the bird’s extinction, we shall never know the answer.
The Imperial was the largest woodpecker in the world. Widespread and almost total destruction of its
specialised habitat in open pine forest, particularly the removal of large old and dead trees in which it
nested, is thought to be the principal reason for its disappearance. The last confirmed record was from
Durango between April-May 1956, when William L. Rhein filmed a foraging and flying female Imperial
Woodpecker (see Lammertink et al. 2011). This remarkable silent footage was only recently discovered.
There were unconfirmed sightings through the 1970s (Collar et al. 1992), and Tanner (1964) made a dedi-
cated search for it in 1962, but was unsuccessful. Lammertink et al. (1996), reviewed all post-1956 reports,
and concluded that the species had survived into the 1990s in the central part of its range, but also that its
continued survival was extremely unlikely.
Historically, it was not rare but contraction of its range had been occurring for some time – it had not
been seen in Jalisco since the 19th century, but was reported as being rare there in 1897 (Nelson 1898). Early
disappearance from some localities was probably due to hunting. Locals regarded the young such a deli-
cacy that they cut down trees in order to get the nests (Lumholtz 1903). The birds were also thought to have
medicinal properties; in Durango they were shot to obtain the bill, which was supposed to ‘draw’ sickness
from the body (Tanner 1964). Although a total of about 120 scientific specimens have been collected, this is
not likely to have had a significant effect on the population. After the discovery of Rhein’s film and using
photographs taken on the 1956 exhibition, Lammertink et al. (2011) pinpointed the exact location and inter-
viewed surviving local residents. They were informed by one that the birds were considered a pest by
logging companies, and the locals were encouraged to kill them. This included application of an agricul-
tural poison to the trunks of foraged
trees. No doubt this persecution, along Imperial Woodpecker Campephilus imperialis
(female left, male right)
with severe deforestation, hastened
the end of the woodpeckers. The site
where Rhein obtained the unique film
of the bird and all surrounding areas
has now been completely logged.
Habits
Although it has been claimed that the
bird was found in mountains down to
1,525m, nearly all records are from
between 1,920m and 3,050m. Its
habitat consisted of open forest of
large pines with many dead trees
intermixed, broken by patches of
grassy parkland (in this its habits were
similar to Cuban Ivory-billed
Woodpecker C. bairdi but differed from
the American version C. principalis,
which favoured swampy forests). The
birds foraged almost exclusively by
scaling and knocking chunks of bark
from dead pine trees to obtain large
insect larvae.
Nelson (1898) recorded them
feeding:
They fly from tree to tree with rather
slow, heavy wing strokes similar to
those of a Crow, and when about to
alight, by an added impulse, glide
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upward along the trunk in a graceful curve and firmly grasp the bark or smooth wood. After a short pause and a
glance around, they ascend the trunk in little runs of from one to three feet, with alternating pauses, usually
keeping along the main stem of the tree, but when searching for food sometimes travelling out on the larger
branches. At such times they were often seen clinging, back down, to the lower side of the branch, chiseling away
with powerful blows.
A breeding pair required a very large area of pristine mature forest, approximately 26km2 (Tanner 1964),
and outside the breeding season the birds apparently formed small groups of 6–12 individuals, which
foraged over a wider area, apparently in response to food availability (Lammertink et al. 1996). Nest holes
were excavated in dead pine trees, with holes situated up to 20m off the ground on the main trunk, and
apparently always in Pinus montezumae. Breeding seems to have occurred during February to June, eggs
having been found in February, young from March to as late as June, though fledged young had occurred as
early as April. Only one set of eggs (with two in the clutch) seems ever to have been found, and these were
accidentally broken soon after collection (Nelson 1898). The Rhein film footage (Lammertink et al. 2011)
revealed that when scaling a tree, the climbing strides were rather slow – no surprise considering the size of
the bird – but in flight the wing beats were remarkably rapid. The habitat comprised mixed coniferous
forest of Pinus, Psuedotsuga and Juniperus, with a few oaks Quercus sp. The filmed female perched and
foraged on the largest dead or recently dead Durango Pines Pinus durangensis. Rhein also reported that the
bird made cackling notes and ‘toy trumpet’ sounds.
So much has been written on this bird that this account makes no attempt to be comprehensive (see Tanner
1942 and Collar et al. 1992; for more recent summaries see Gallagher 2005 and Steinberg 2008).
The numbers and distribution of the Ivory-billed Woodpecker began to decline noticeably towards the
end of the 19th century, though most early ornithologists indicate that it was never common. By 1855 it had
disappeared from North Carolina and from nearly the whole of the country west of the Mississippi delta.
By 1915, it had gone from Texas, Arkansas and Alabama, and from most of Florida, Georgia and South
Carolina, surviving only in the larger swamps. By 1930, it was known only from the Santee swamp in South
Carolina and a few scattered localities in Florida and north-eastern Louisiana. It is generally believed that
the species was dependent on virgin forests, and had retreated when these were felled. Unfortunately, the
history of logging in the United States is insufficiently documented for a satisfactory correlation to be made;
nevertheless most authorities do not doubt this conclusion. However, Snyder (2007) blamed direct hunting,
both for food and for specimens, as the cause of extinction. He noted that it was the encroachment of hunters
and collectors into hitherto remote forested areas that seriously impacted woodpecker numbers, as popula-
tions had previously disappeared from forests due to hunting that had not been logged.
The last known population disappeared by at least 1948, with the clearance of about 311km2 of virgin
forest by the Singer Sewing Machine Company Tract in Louisiana for soya bean cultivation (Collar et al.
1992). Short & Horne (1990) suggest that the last observation occurred in the 1950s, when a pair was seen
feeding in beetle-killed pines in Southern Georgia, whereas Dennis (1979) believed that reported sightings
in the 1970s in modified forests were also genuine, although King (1981) wrote:
Unconfirmed sightings or tape recordings of this woodpecker’s characteristic call, every few years, reinforce
hopes of its continued existence, but the specific locations of such occurrences are unknown or are not being
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publicised and few specific programs are aimed directly at this woodpecker’s conservation. The exact date of its
extinction, whether already passed or in the future, is unlikely to be ascertainable.
Birds were recorded using a tree cavity in central Florida in 1968–69 (Agey & Heinzmann 1971), but no
evidence was ever produced to confirm this. An unconfirmed sighting in 1999 of a pair along the Pearl River
in south-east Louisiana (Gallagher 2005) prompted the US Fish and Wildlife Service to improve potential
Ivory-billed Woodpecker habitat by purposely selecting trees for killing in order to encourage beetle larvae.
By 2002, 11 sites from 3 to 40 acres in size had been managed, but Hurricane Katrina devastated the area; no
confirmed sightings of woodpeckers were ever made.
Recent sightings
A recent supposed filmed sighting (Fitzpatrick et al. 2005) made worldwide news, but the evidence is far
from convincing and most ornithologists dismiss its authenticity. It appears to represent the Pileated
Woodpecker Dryocopus pileatus (Collinson 2007), the largest woodpecker alive in the United States today.
Short (1982) considered that competition with Pileated Woodpeckers played a considerable part in the
decline of the Ivory-billed Woodpecker once the latter’s habitat began to disappear, and thought it unlikely
that the Ivory-bill could be saved in the presence of D. pileatus.
Despite the number of reports, supposed photographic evidence and sound recordings, not one undis-
puted observation of a living Ivory-billed Woodpecker has been made since 1944.
Gotelli et al. (2012) devised a new statistical method for estimating the probability of the survival of the
Ivory-billed Woodpecker into recent times, using dated museum specimens from geo-referenced localities.
Based on specimens collected between 1853 and 1932, a noticeable decline occurred after a peak of collecting
between 1885 and 1894. The authors hypothesise that if a population of <100 occurred between 1929–1932
– Tanner (1942) estimated that c.22 birds were alive in the southeastern United States during this period –
the probable extinction date was no later than 1980, but more likely nearer to 1960. Gotelli et al. conclude
that:
. . . these results suggest that there is virtually no chance the Ivory-billed Woodpecker is currently extant within
its historical range in the southeastern United States. The results also suggest conservation resources devoted to
its rediscovery and recovery could be better allocated to other species.
Recent mitochondrial DNA analysis (Fleischer et al. 2006) has shown that the I vory-billed Woodpecker of
the mainland United States is specifically distinct from the Cuban Ivory-billed Woodpecker Campephilus
bairdii, and along with the Imperial Woodpecker Campephilus imperialis form a monophyletic group. Each
should be regarded as distinct species, and this taxonomic arrangement is followed here.
Habits
The Ivory-billed Woodpecker inhabited swampy forests and river floodplains, usually, but not exclusively,
where oak and sweet gum trees predominated. In certain areas, such as Florida, drier areas were also used,
and the type of country in which the bird was found varied considerably with locality. It was very seden-
tary, the birds apparently spending their entire lives within a few miles of the place where they were hatched
(Bent 1939).
Ivory-billed Woodpeckers fed mainly on wood-boring beetle larvae. They scaled the bark from dead
trees to search for the insects that live between the bark and the sapwood. The birds also dug into the inner
heartwood to obtain deeper-living borers; however, this food source was less frequently used by the Ivory-
billled Woodpecker than by the Pileated.
The Ivory-billed Woodpecker is also recorded as feeding on berries and acorns (Bent 1939). The species
varied greatly in the timing of the breeding season, which extended from January to April. The nest was in a
hole excavated in a living or dead tree, and the tree species involved varied with locality. In Florida it was
usually in a cypress, in northern Louisiana more usually in red maple, sweet gum or one of several species
of oak. The eggs were china-white (like those of all woodpeckers), with between one and five recorded in a
clutch. However, more than three was unusual, and rarely was more than one young successfully fledged.
According to Bent (1939), no description was ever published of the chick or juvenile plumage.
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This woodpecker is thought to have occurred historically over much of Cuba, but it was only ever definitely
observed in the eastern and western extremities of the island, including the Pinar del Rio Mountains, Sierra
de Moa, Guantánamo, and along the Hanábana River (Collar et al. 1992). In later years it was confined to a
small area of pine forests in the n orth-east, mainly in the Sierra de Moa, Sierra del Cristal and Sierra de
Nipe. Shull’s (1985) claim that it once occurred on the Isle of Pines is, in the opinion of Collar et al. (1992),
unfounded.
A steady decline
The bird suffered a steady decline after it was discovered in the east of the island in 1863 (Collar et al. 1992),
and Gundlach (1874a) noted the species’ increasing rarity from year to year; by the end of the 19th century it
was already very rare and hard to find, having disappeared from several of its former localities. Barbour
(1943) considered it to be virtually extinct. In the Sierra del Cristal, the last sighting was in 1920. The Sierra
de Moa was the bird’s last stronghold, when Dennis (1948) observed a trio in April 1948. In 1956 a popula-
tion of only 12 or 13 was estimated to remain in Oriente Province (Lamb 1957). In 1977 it was estimated that
no more than eight pairs, and probably fewer than six pairs, survived.
The last recorded sightings were in 1986, when a few were seen in the hilly pine forest of Ojito de Agua,
and the last record was made on March 16, 1987, when a single female was seen in the same area (Lammertink
1995); searches in 1990 and 1991 proved unsuccessful. The forests of the Sierra de Moa are now thought to
be too degraded to support any birds, the only hope being a narrow corridor between the Cupeyal Reserve
and Ojito de Agua and the upper reaches of the Rio Jaguani, but even if a few birds survive there, which
seems unlikely, there is little hope that the species can be saved.
John Dennis (1948), the only person to photograph the bird alive and observe a breeding pair, gave a
poignant account as to the fate of the bird:
Whatever the future holds for the Cuban Ivory-bill, one thing seems to be certain, and this is that on the long
path of evolution from the origin of a species to its total oblivion, the I vory-bill in Cuba has a slight edge on the
American bird as the two go down to extinction – this, in face of the fact that Cuba was settled a hundred years
earlier than our country and has suffered more from deforestation as the country was cleared for grazing and the
sugar industry. The bird has not survived in Cuba, however, due to any innate gentleness on the part of Cubans
toward birds. Just as in our country, the Ivory-bill was killed for every conceivable reason – sport, food, trophies,
curiosity. Not as many were killed by professional collectors, simply because the birds vanished so quickly from
all accessible parts of the country. The fact that a few roadless wilderness areas have remained up until the
present time as well as the apparent ability of the birds to meet changing conditions seem to have allowed a few to
survive. If present trends continue these few birds will quickly disappear.
Habits
The Cuban Ivory-billed Woodpecker occurred in mixed pine and hardwood forests, mainly in the moun-
tains, though it formerly also occurred in the lowlands (Lamb 1957). Roosting and breeding took place
almost exclusively in cavities in mature pines, though the woodpecker also fed in mixed forests. This
required large tracts of virgin forest with a plentiful supply of dead and dying trees, from which they
obtained their main source of food, beetle larvae, particularly longhorn beetles (Dennis 1948). The breeding
season occurred from March through to June. The Cuban Ivory-billed Woodpecker was reported to form
groups of eight or more individuals at certain times; otherwise they generally occurred in pairs.
249
The North Island Bush Wren is poorly known and represented by just three skins.The type, an immature,
was obtained in 1850; a second specimen, an adult, was collected soon after, both from the Rimutaka Range,
and a third was, according to the registration in Liverpool, a male collected by J. von Haast at Taupo, North
Island, in 1866 (Fisher 1981). Subfossil remains have also been discovered, which show that the species was
formerly widespread (Worthy & Holdaway 2002).
There have been a number of valid reports of the North Island subspecies, from the southern Rimutaka
Range in 1918 and the Te Urewera region, up to at least 1955. Unconfirmed sightings occurred on from
Kapiti Island in 1911, in the Huiarau Range in 1925, near Lake Waikareiti in 1949, and in the Aniwaniwa
Valley in 1955 (Edgar 1949; St. Paul & McKenzie 1977; Miskelly 2003). There have been no sightings since.
Its extinction can be attributed almost solely to introduced predators such as cats, rats, the Stoat Mustela
erminea and Weasel M. nivalis, all of which have been especially devastating to g round-nesting birds.
Range Formerly occurred on Stewart Island, Solomon Island, Big South Cape Island, and South Cape Island, New Zealand.
Description 8.75cm (3. 35in). Smaller than South Island race X. l. longipes, back less suffused with green; brown of head
extending further down neck; superciliary streak less distinct; and chin less white. The pure white egg was also much larger,
being 21.5mm x 15.5mm, compared with 18mm x 13.5mm in X. l. longipes (Stead 1936).
The islands south-west of Stewart Island, New Zealand, are remote, and in 1913 remained almost undis-
turbed. In that year, two of them, Big South Cape Island and Solomon Island, were visited by Guthrie-Smith
(Guthrie-Smith 1925), who found a bush wren there. He did not distinguish it from the bush wren of South
Island, so Stead (1936), after visiting some of the islands, including Solomon Island, collected some speci-
mens which he described as a new race. It was recorded from Stewart Island, South Cape Island, Big South
Cape Island and Solomon Island, and probably occurred on two others. It was observed on Stewart Island
in 1951 (Dawson 1951), but feral cats appear to have exterminated them shortly after this date. It was still
reasonably common on Solomon Island until the early 1960s, but had disappeared by 1965.
The last stronghold was Big South Cape Island, but the Black Rat Rattus rattus invaded the island in 1965
and wiped out the population. Blackburn (1965) remarked that in September 1964 six specimens were trans-
ferred from Big South Cape Island to Kaimohu Island by the New Zealand Wildlife Service, and four were
seen there on February 25, 1965 but there were no young. The last of these birds died in 1972.
Habits
On Solomon Island the birds were largely terrestrial, and they often nested in holes in the ground, or some-
times in fallen logs or clumps of fern. Guthrie-Smith (1925), who photographed a nest and breeding birds in
1911 (Guthrie-Smith 1925), noted that any nest tending to get very wet was meticulously kept dry after
every shower, with the birds carrying wet feathers out from the nest and taking in dry ones. Two dispropor-
tionately large eggs were laid, and both parents shared in incubation. The young were fed on moths, flies
and other insects. The call was described as a faint rasping sound and a loud ‘cheep’ when alarmed or
excited. Guthrie-Smith further commented on its habits:
The activities of the bird are altogether restricted to movements two or three feet from the ground. They never
stray more than a few yards from the tangle where safety lies. Even when close to his own nest, I have watched
the male wait and hesitate, unable to harden his heart sufficiently to dare to make a forward movement.
They are at least as active, however, on their legs as on their wings. The hop of the bush wren is a remarkable
performance. During the first salutary movement the bush wren carries himself parallel to the earth; at the
termination, however, of each leap he telescopes upwards on his toes, momentarily erecting himself in the oddest
way to his full height. When the two movements are blended in rapid action, what with his whitish feet, short
toes and long thin legs, and tightly folded body plumage, he resembles in no small degree a barefooted bairn
running on sands with tucked-up garments firmly fastened around the waist. He passes through the darkling
underscrub like a forest gnome, like a woodland brownie.
The South Island Bush Wren appears to have been widespread in South Island, but had become absent or
rare from most of its range by 1955 (Oliver 1955). The last authenticated reports were from Arthur’s Pass in
1966 and Nelson Lakes National Park in 1968 (Cresswell 1968). A few reports have subsequently been made
from Fiordland and Nelson Lakes, but they remain unsubstantiated. Its disappearance was due to preda-
tion by feral cats, rats, weasels and stoats.
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Habits
Little is known about its behaviour. Oliver (1955) stated that the bird was found ‘sparingly’ in damp forests
of the Southern Alps, at a high altitude and up to the limits of the forest. In the Fiordland area, however, it
came down almost to sea level. The bush wrens were seen in parties of two or four, flying from tree to tree.
They fed on the ground as well as on trees, and had a peculiar habit of bobbing their heads.
Few nests were ever found, but one was recorded by Potts in 1870 near the headwaters of the Havelock
River. It was beneath the moss-covered roots of a ribbonwood tree, and looked like a thick tuft of moss. The
pouch-like nest was concealed beneath this covering, composed almost entirely of fern rootlets; it was
extraordinarily well hidden. Although ready for use, this nest contained no eggs.
The loss of the Stephens Island Wren represents yet another senseless extinction of an insular bird species,
perhaps the definitive exemplar of what can go wrong when humans interfere with an ecosystem. Edward
Lukins, an ardent natural history collector, made a prophetic account about the island’s bird fauna in
October, 1894, just prior to the completion of the lighthouse on the island (see Medway 2004a):
It was with considerable interest I found so many of the more rare of the New Zealand birds on Stephens Island
and I hope they may long continue to thrive on this inaccessible island where they will certainly be unmolested
by stoats and weasels.
But it was cats, not stoats and weasels, that would ultimately be the culprits.
In 1894 the cat belonging to Mr. D. Lyall, the lighthouse keeper on Stephens Island, brought in 11 speci-
mens of a little bird, which were given to H. H. Travers. Ten of these were forwarded to Rothschild, who
named the species jointly after Lyall and Travers (Rothschild 1894b). Much has been made about Lyall’s cat,
Tibbles, exterminating the entire population of the wren – a good story, but probably untrue. Galbreath &
Brown (2004) have shown that although Lyall’s cat had caught a number of birds, cats were feral on the
island by 1894 and would have quickly eliminated the population. Buller (1905) was told by Lyall in around
1895 (see Galbreath 1989) that cats had gone wild and were making ‘sad havoc among all the birds’. The cats
were extremely numerous by 1897, with the island ‘swarming with them’ by 1898 (Travers in Medway
2004a). Robert Cathcart, a replacement lighthouse keeper, shot over a hundred in eight months in 1899, but
noted that the cats still seemed as plentiful as ever (Medway 2004a). It was not until 1925, when most of the
birds on Stephens Island had been exterminated, that the island was finally cleared of cats. Over-collecting
by the lighthouse keepers and a rush to obtain specimens by professional collectors may have also helped to
finish off the birds (Worthy & Holdaway 2002).
A feud developed between Walter Rothschild and Walter Buller over the naming of the Stephens Island
Wren (Buller 1895, 1906; Rothschild 1895a; Watola 2008). Buller (1895) had described the bird from a single
individual, and was ready to publish, but delays in the artwork and communication from New Zealand
resulted in Rothschild forestalling Buller in a most ungentlemanly way, and describing the species first
(Rothschild 1894b). Buller (1895) made the following comment about his discovery of the Stephens Island
Wren:
I have lately received a single specimen of species of Xenicus entirely distinct from the two forms (X. longipes
[South Island Bush Wren] and X. gilviventris [Rock Wren]) inhabiting the mainland . . . My correspondent
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on the island informs me that the bird is semi-nocturnal in its habits, and that he has seen three examples, all of
which were brought in at different times by the cat.
Habits
Almost nothing was recorded about the habits of this bird. Dieffenbach (1843) noted that Stephens Island
was covered with dense forest from the coastal shores to the summit of the mountain, and Lukins (in
Medway 2004a) recorded that ‘the gnarled cedars’, Kohekohe Dysoxylum spectabile, ‘and other dwarfed
trees indicate that high winds are customary there’. Rothschild (1907a) gave the only direct information
about the bird, quoting a letter from Travers:
The bird itself has only been seen on two occasions alive, when disturbed from holes in the rocks, and was not
obtained. It is nocturnal, runs like a mouse and very fast, and did not fly at all.
An account of the Stephens Island and its once-abundant bird fauna was made by F. W. Ingram (in Medway
2004a), who spent a total of 12 months between April 1892 and January 1894 assisting with the construction
of the lighthouse. The situation for most of the birds was about to irrevocably change:
The saddle-back, native thrush, native crow, the robin, two kinds of coo [cuckoos], one long tailed and the other
with a striped breast, also kaka, pigeons, moreporks, two kinds of wrens [Stephens Island Wren and South Island
Rifleman Acanthisitta chloris chloris] (very small birds), and I also got one land-rail. There were hundreds of
parakeets and tuis (or parson birds), also the moke moke (or bell bird). . .
The population of wrens on Stephens Island is now known to be relictual, as remains of the bird are fairly
common in owl deposits from both North and South Islands (Worthy & Holdaway 2002); this bird once had
a wide range across New Zealand. Millener (1989) demonstrates that the osteological evidence clearly
proves that the bird was flightless, and the peculiarly flattened cranium, flat, wide bill and distinctive leg
bones suggest a specialised but undetermined ecology (Worthy & Holdaway 2002). Long retained in
Xenicus, this species is now placed in its own genus, Traversia (Millener 1989, Turbott 1990).
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The flightless South Island Stout-legged Wren was the heaviest of the New Zealand wrens, with reduced
wings, a robust pelvis and stout legs. Millener (1988) found subfossil remains in association with those of
four other species of the family, representing four genera. Reconstruction of the early Holocene flora from
the sites where the bones were found indicate that the wren inhabited mixed podocarp broadleaf forest,
which formed the dominant vegetation over much of New Zealand before the arrival of the Polynesians; it
also ranged into subalpine scrub. This wren’s remains are known only from South Island, and it appears
that the closely related Pachyplichas jagmi (see below) replaced it in North Island. However, Worthy &
Holdaway (2002) suggest that they were the same species, only differing in size.
All of the extinct New Zealand wrens, being predominantly terrestrial and with poor or no flying capa-
bilities, were vulnerable to introduced predators, and the extinction of both stout-legged wrens was
probably due to the introduction of the Pacific Rat Rattus exulans after the Polynesian colonisation of New
Zealand. No doubt the Stout-legged Wren fed on invertebrates, using its powerful legs and claws to scratch
in leaf litter and in rotten wood (Tennyson & Martinson 2007).
The North Island Stout-legged Wren was smaller than its South Island counterpart P. yaldwyni, but other-
wise the birds were similar. Numerous subfossil remains of this species and P. yaldwyni have been found in
Laughing Owl Ninox albifacies cave deposits (Worthy & Holdaway 2002), indicating that the owl was prob-
ably the main natural predator of these birds. Subfossil remains have also been found in cave pitfall traps;
being flightless, once they had fallen in, they would have had no way of getting out.
This species is known from four localities and from fewer than six individuals (Worthy & Holdaway 2002).
It was a diminutive bird with stout legs, tiny wings and a reduced sternal keel, suggesting that it was prob-
ably flightless (Milliner & Worthy 1991). It was perhaps heavier than either surviving New Zealand wrens
but weighed less than the extinct stout-legged wrens. The bill was different from that of any other acan-
thisittidid, being both long and decurved. This was presumably an adaptation for foraging, with the
Long-billed Wren probing in crevices, mats of dense vegetation or leaf litter for invertebrate prey.
Reconstructions of the palaeoenvironment around the fossil localities suggest that the bird was an inhab-
itant of h
igh-altitude scrubland and montane beech forest (Milliner & Worthy 1991; Worthy & Holdaway
2002), similar to the extant but endangered Rock Wren Xenicus gilviventris. It appears that the L ong-billed
Wren became extinct before the arrival of Europeans, and may have been an early casualty of the Polynesian-
introduced Pacific Rat Rattus exulans.
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Pittidae (Pittas)
The pittas are medium-sized, generally terrestrial forest inhabitants, divided into three genera. They are
stocky birds, with a stout bill, short tail, and long legs and feet. The centre of distribution is Asia and
Australasia, but some species occur in Africa. Being forest birds, a number are in danger of extinction due to
loss of habitat.
The Black-faced Pitta P. anerythra occurs on Bougainville Island, on Papua New Guinea, and on Choiseul
and Santa Isabel in the Solomon Islands. It is only known to survive with certainty on Santa Isabel, where it
is considered common. The Bougainville population was also common when it was first collected at the
turn of the 20th century, but it has not been recorded since 1938 (Erritzoe & Erritzoe 1998), despite subse-
quent searches for it. The reason for its disappearance is unclear, as the species can tolerate a range of
introduced predators.
Habits
Little was recorded about the Bougainville race of this pitta. An egg collected in mid-May was creamy-
white, spotted with brownish-red and underlying greenish-mauve (Erritzoe 2003); a female was also taken
in breeding condition in late July. The nominate P. a. anerythra from Santa Isabel is a wary bird and occurs in
primary forest remnants and secondary forest, as well as gardens and small plots. The call of the nominate
is distinctive, and is described as a double rasping kree-kreeee-o generally issued at dawn or dusk (Lambert
& Woodcock 1996).
The Choiseul Black-faced Pitta has not been seen in recent years, and may be extinct. Unconfirmed reports
from the neighbouring islands of Kolombangara and Vangunu are unlikely, as these islands are in a different
biogeographic subregion (BirdLife International 2011).
The Choiseul subspecies was collected in forested mountain valleys, and coastal and alluvial plains
(Mayr 1935, 1945), which have now suffered from severe deforestation. The continued logging in the
lowlands of Choiseul means this bird is now probably extinct. Nothing was recorded about its habits.
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Furnariidae (Ovenbirds)
The ovenbirds are a large, complex family of birds which have been divided into three subfamilies and a
number of genera. They are so-called because some species build elaborate, oven-like clay nests, but most
build nests out of sticks and grasses. They are distributed in Central and South America, where a number of
species are endangered due to deforestation.
The Scaly-throated Earthcreeper U. dumetaria occurs in Argentina, Bolivia, Chile, Peru and Uruguay
(Ridgely & Tudor 2009). It is a wide-ranging species and generally not threatened throughout most of its
range. However, the Peruvian race U. d. peruana is only known from two specimens, taken in the early 1950s
at Puno, Peru, and the bird has never been found since. The Puno region is vast, so there is still a chance that
it might exist, but continuing habitat destruction threatens any prospect of l ong-term survival.
Habits
Little is known about this bird, except that it was confined to the high montane open shrubland. It was a
terrestrial bird, nesting in burrows.
The taxonomy of the Cranioleuca spinetails is complex and unresolved. The Stripe-crowned Spinetail C.
pyrrhophia is a widespread inhabitant of marshes, river islands and várzea forest (Ridgely & Tudor 2009)
and is divided into three subspecies. The northern population, from the northern Bolivian Andes, is known
from a few specimens and has not been recorded since the 1950s. It appears to have suffered from habitat
destruction and may now be extinct.
undertail-coverts brown; tail pale orange-rufous, with the central rectrices darker dorsally; bill, upper mandible black, lower
mandible paler; iris brown; legs and feet greyish-olive. Female exhibited reversed sexual dimorphism and had a longer bill than
the male.
The Cryptic Treehunter, along with the Alagoas Foliage-gleaner Philydor novaesi (p. XXXX), Pernambuco
Pygmy Owl Glaucidium mooreorum (p. XXXX) and Alagoas Curassow Mitu mitu (extinct in the wild) all
occurred in remnant Atlantic forest, a once vast and diverse forested biodiversity hotspot that covered 150
million hectares along the eastern seaboard of Brazil. Due to relentless deforestation for sugar cane produc-
tion and cattle ranching, less than 8% survives, and most of what is left is badly fragmented, with over 83%
of fragments less than 50 hectares in size (Ribeiro et al. 2009). These isolated fragments are still under threat
and thousands more hectares are disappearing every year.
The Cryptic Treehunter remained unknown to science until 2014, when specimens collected in 1986 at
Murici, Alagoas,and presumed to have been the similar looking Alagoas Foliage-gleaner, subsequently
turned out to belong to a new taxon (Lees & Pimm 2014; Mazar-Barnett & Buzzetti 2014). The Cryptic
Treehunter occurred in two separate patches of remnant forest, in Murici, and in Frei Caneca in Pernambuco,
and in 2004 numbered less than 20 birds, perhaps even fewer (Barnett & Buzzetti 2014). Continued defor-
estation reduced suitable habitat further, and extensive searches have failed to rediscover the bird. It was
last seen in February 2005 in Frei Caneca, and in April 2007 in Murici (Pereira et al. 2014), and is now consid-
ered probably extinct. The extinction of the Cryptic Treehunter can be blamed entirely on deforestation; the
loss of the bromeliads on which it foraged would have drastically reduced its food supply.
Habits
The Cryptic Treehunter preferred dense, humid forest, with a profusion of epiphytes, especially bromeliads
(Mazar-Barnett & Buzzetti 2014). They were found singly, in pairs or in association with large mixed-species
flocks, and moved between the mid-levels to the subcanopy searching almost exclusively in bromeliads for
invertebrate food. The song was described as a dry rattle followed by louder rasping notes.
The Alagoas Foliage-gleaner occurred in the same two remnant patches of Atlantic forest in Alagoas and
Pernambuco as the Cryptic Treehunter Cichlocolaptes mazarbarnetti (see p. 256). It was first discovered in
1979 in Murici, Alagoas, where it was considered conspicuous and easy to locate (Teixeira & Gonzaga 1983),
and a new population was discovered in April 2003 in Frei Caneca, Pernambuco (Lees et al. 2014; Pereira et
al. 2014). Subsequent searches witnessed a rapidly declining population due to uncontrolled deforestation.
The last bird at Murici was observed in 2007 and the last confirmed sighting was at Frei Caneca on
September 13, 2011, when an individual was filmed (Lees et al. 2014); this appeared to be a lone individual
that was seen between 2007 and 2010. Despite further intensive investigation, there have been no more
observations, and the Alagoas Foliage-gleaner is considered probably extinct (Lees et al. 2014; Pereira et al.
2014; Lees & Pimm 2015). Deforestation and the subsequent loss of bromeliads from which it mainly
obtained its invertebrate food, are the reasons for its disappearance (Lees et al. 2014).
Habits
The Alagoas Foliage-gleaner occurred in lowland evergreen forest and humid, hilly secondary forest up to
550m (Remson 2003). Birds were seen singly, in pairs and small groups, and also with mixed-species flocks.
They gleaned insects, including beetles, grasshoppers and ants, from dead leaves, live foliage, debris and
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bark, and pulled beetle larvae from dead wood (Remson 2003; Lee et al. 2014); like the Cryptic Treehunter,
they had a preference for foraging in bromeliads. An immature was collected in March, suggesting that
breeding occurred in January or February, and moulting birds were observed in February (Ridgely & Tudor
1994). The vocals comprised a descending series of whistles uoo-uoo-uoo and a thoorr alarm call.
This cryptic species has not been relocated since the type material was collected between 1955 and 1956
(Wetmore & Phelps 1956), despite extensive surveys in 1990 and 1996 (Boesman 1998). The entire type
locality has now been cleared for coffee plantations, and forest clearance for agriculture continues unabated.
There are apparently some still-forested areas adjacent to the type locality where a small population may
persist, but this antpitta’s long-term status seems bleak, if it survives at all.
Habits
Very little is known about the species’ habits. It was collected in dense cloud-forest at elevations of 1,800–
2,100m, and being cryptic in coloration would be difficult to locate.
The discovery of the unique type of Antioquia Brown-banded Antpitta, around 130 years after it was
collected, epitomises the need to evaluate all early collections of birds. It is closely related to the extant but
endangered nominate Brown-banded Antpitta G. m. milleri, which occurs 160km to the south (Salaman et al.
2009) and was itself only rediscovered in 1994, after a lapse of 52 years (Donald et al. 2010; see p. 500). The
type locality of the Antioquia bird has undergone extreme ecological change – almost complete deforesta-
tion due to coffee-growing and other types of agriculture. If this race still survives, it must be critically
endangered.
Virtually nothing is known about this bird. It was collected at an altitude between 1,800–2,800m, and the
stomach contained insects.
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Maluridae
Maluridae (Grasswrens)
The grasswrens and fairywrens occur only in Australia and New Guinea, and have been divided into six
genera and 28 species. They are small, terrestrial birds, feeding on the ground or in low vegetation. Some
species have declined due to habitat disturbance and degradation, and droughts. Two or perhaps four
subspecies of the Thick-billed-billed Grasswren Amytornis textilis are considered probably extinct.
Several subspecies of the Thick-billed Grasswren are poorly known, and the now extinct Namoi Valley
subspecies is no exception. Bennett (in North 1904) described the bird’s habitat and the probable reason for
its extinction:
For the past two years, 1885–6, I have been continually travelling over the country in which they were formerly
abundant, and during all that time I have only met with a pair of these birds. This disappearance is, I think, due
to their weak powers of flight, and to the occupation and stocking of the country and the burning off of the large
areas of dense porcupine grass amongst which they could always be found. In former years I have often found
their nests; they were generally placed in a tussock of porcupine grass, but sometimes I have discovered them in
brush fences running through the mallee.
The last confirmed record of the Namoi Valley Thick-billed Grasswren was of two specimens taken on
Pimpara Lake Station, near Milparinka, in the Upper Western Region of western New South Wales in
October 1912, and it is now probably extinct (Higgins et al. 2001).
Habits
Little is known about the Namoi Valley birds. Gould (in Parker 1972) made the following account:
259
The only place in which I observed the Textile wren was the plains bordering the lower Namoi . . . On the lower
Namoi, where it is very abundant, it is found in all those parts of the plains that are studded with scrubs and
clumps of a low shrub-like tree, resembling the Barilla [Salsola kali] of the coast, through and among which it
creeps with astonishing rapidity . . .
The Southwestern Thick-billed Grasswren disappeared due to overgrazing by livestock and introduced
mammals in the 1880s, followed by a series of devastating droughts in the 1890s (McAllen 1987). It was last
collected in September 1910 (Carter 1910; Black 2011a).
Another two populations of grasswrens have been described, but they have only recently been consid-
ered subspecifically distinct (Black 2011a; Black et al. 2011). A. t. carteri once occurred on Dirk Hartog Island
off Western Australia, and inhabited dense acacia shrublands. Birds have not been recorded there since
October 1918, and disappeared due to rat predation. The northern inland population A. t. giganturus
occurred in arid-zone salt-tolerant shrublands, and has not been seen since the last specimen was collected
in 1908.
Habits
Little was recorded, but the southwestern population of grasswrens inhabited dense thickets within a
variety of eucalyptus tree communities (Black 2011a).
Euler’s Flycatcher Lathrotriccus euleri is fairly common and quite widely distributed over South America
and the islands of Trinidad and, formerly, Grenada, with five races currently recognised (French et al. 1991).
The Grenadan race has not been recorded since the early 1950s, and is probably extinct.
The reasons for the disappearance of this taxon are uncertain, but are likely to be habitat destruction and
possibly introduced predators. Bond (1956) claimed that only two specimens were ever collected.
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Tyrannidae
Habits
Little was recorded about the Grenada subspecies of Euler’s Flycatcher. The nominate race is generally a
middle and lower forest-canopy inhabitant, remaining inconspicuous in the undergrowth, only to dart
from a perch to catch insects. The call is a loud hoarse chee–chi–wi–wi–wi.
The poorly known Bearded Tachuri P. pectoralis has a disjunct distribution, with populations occurring on
either side of the Amazon Basin (Collar & Wege 1995). The Bogotá subspecies P. p, bogotensis is known from
261
a few specimens taken in the early and mid-20th century (Ridgely & Tudor 2009). It was restricted to the
Suba marshes north of Bogotá city, and disappeared due to the extensive conversion and drainage of its
habitat. There were possible s ight-records in the 1990s in remnant marshes in the Bogotá area (Collar &
Wege 1995), but the bird has not been recorded with certainty since the 1950s.
Habits
Little was recorded about the Bogotá birds, but the species forages and breeds in tall reeds and grasses, and
always in the proximity of water. It is insectivorous, and birds are usually found in pairs. The nest is
constructed from fine rootlets interwoven with grass.
Turnagridae (Piopios)
The piopios are an ancient group of birds, and their relationships are somewhat obscure. They have been
placed in a number of different families over the years, but their true affinities may lie closest to the bower-
birds (Ptilonorhynchidae) (Olson et al. 1983; Christidis et al. 1996). However, recent work (Worthy &
Holdaway 2002) suggests a relationship closer to the whistlers (Pachycephalidae), whereas Johansson et al.
(2011) suggest that the piopios are orioles. Higgins et al. (2006) consider the Turnagridae of uncertain affini-
ties, and it is probably best to maintain this position until their affinities can be resolved. The North and
South Island Piopios were traditionally regarded as races of a single species, but they are now considered
specifically distinct (Holdaway 1999; Worthy & Holdaway 2006).
The South Island Piopio Turnagra capensis was discovered in 1769 on Cook’s first voyage of discovery to the
Pacific. It was described by the naturalist and artist Anders Sparrman (Sparrman 1787), who joined the
expedition at the Cape of Good Hope. Sparrman chose the specific name capensis, which was based on a
blunder, as he had muddled the labels of his specimens, and mistakenly believed that the specimen origi-
nated from the Cape (Worthy & Holdaway 2002).
to be made. Unconfirmed reports of sightings were made in forest inland from Patea in 1923 and from
between Gisborne and Wairoa on May 7, 1947 (Allison et al. 1949), from Southland District 17 in December
1947 (Dunckley & Todd 1949), and in the Nelson district in January 1948 (Moore 1949). The last supposed
sighting was on May 24, 1963 (Knox & Walters 1994), based on an account by Child (1965), but this is consid-
ered unreliable; the South Island Piopio had almost certainly disappeared some 60 years before.
The extinction of the South Island Piopio can be attributed to habitat destruction and the increase in
dogs, cats, stoats, weasels and rats during the late 19th century, which predated the adult birds as well as
taking eggs and chicks.
Habits
Settlers named the bird the ‘thrush’ on account of its size and brown plumage, which recalled the Song
Thrush Turdus philomelos of their homelands, although it lacked anything like the thrush’s song, having
merely a long-drawn plaintive whistle, and other calls (not unlike those of the New Zealand Pipit Anthus
novaeseelandiae). Potts (1872) gave a good description of the Piopio’s habits:
It seems to delight in those openings which are found in river-beds, between long belts of tutu and other scrub;
there it may be observed either hopping along the ground or fluttering about the lower sprays of shrubs, flying
out to the spits of sand, or drifted trees, that lie stranded in the river-bed. On some of the longer formed spits,
that are becoming clothed with vegetation, it searches amongst the burry Acaena, snips off the fruit stalks of
moss, picking the seed of some trailing Veronica. Its progress on the ground is usually deliberate; it hops with
both feet together, a slight flutter of the wings, and a flirt of the tail accompanying each motion; when approached
too closely, it leaves its perch, always descending at first, as though safer when on or near the ground; if it would
rise on the wing, a momentum is gained by a succession of hops.
It inhabited forest and scrub, mainly in valleys, and was strongly territorial, driving off other birds as well
as those of its own species. It was not shy, and one of its endearing traits was to enter the tents of bushmen,
looking for scraps of food. It is likely that its tame nature, especially in the presence of dogs, cats and stoats,
was one of the reasons for its decline. Its food consisted of insects, spiders, seeds and fruit. Potts (1872)
recorded the nest as being between four and twelve feet (1.2 and 3.6m) from the ground, a small compact
structure of moss and twigs, lined with fine grasses, or sometimes tree-fern down. Potts also said that the
birds double-brooded, with no more than two eggs observed in any nest, but four eggs were found in the
ovary of a female killed at Christmas.
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The Stephens Island population of piopio has generally been relegated to the synonymy of Turnagra capensis,
despite having a shorter wing and tail, but recently its subspecific status has been restored (Medway 2004b).
Twelve specimens are known; the subspecific name was thought to be based on juvenile specimens as they
were smaller than the mainland Turnagra species. Medway provided good evidence for the race’s validity,
further supported by the New Zealand Checklist Committee (Gill et al. 2010). Stephens Island covers an
area of just 2.6km², and is situated just 3.2km from the mainland, so the evolution of a distinct subspecies is
quite remarkable. However, piopios were poor over-water dispersers; the founding population must have
been isolated for some time.
The Stephens Island Piopio numbered in the hundreds in 1894. Lukins (in Medway 2004a) recorded that:
. . . the native thrush, now almost extinct on the mainland, is so numerous that there was scarcely a bush in
which at least one could not be seen . . .
But the cats introduced to the island during the establishment of its lighthouse had multiplied to hundreds
by that time, and this appears to have spelt the end for the resident piopios. The last specimen was taken on
January 7, 1897; they had certainly gone by 1898.
Less is known of the North Island Piopio than its South Island congener. This may be because the North
Island Piopio was not described until 1865 (Schlegel 1865), and the specimen was from an unknown locality.
Buller (1869) also described the bird (in ignorance of Schlegel’s name), and further stated that he had shot a
specimen in Kaipara district in 1852 (Buller 1887–88).
The North Island Piopio was considered common at the time of European arrival in New Zealand, but it
soon began to decline, and by 1892 had nearly disappeared; the last specimens were taken in 1887 (Medway
1968). However, Sopp (1957) reported possible sightings at Hopuruahine, Lake Waikaremoana in 1938, on
the shore of Waikare-iti in May 1946, on the Wanganui River on 11 November 1952, near Lake Waikare-iti on
9 April 1953, and the last observation, also at Lake Waikare-iti, on 9 April 1955. Mead (1950) observed birds
from 14–17 March 1950 on the Wanganui River, and Bell & Singleton (1974) reported a parent bird feeding
young in March 1970, but this remains unconfirmed. Regardless of these late observations, the North Island
Piopio was probably extinct by 1902 (Higgins et al. 2006). It disappeared for the same reasons as the South
Island species, a combination of habitat loss and predation by introduced cats, dogs, rats and mustelids.
Habits
Much of what we know about the North Island Piopio is based on the observations of Buller (1887–88), who
noted that it lived in forest, chiefly undergrowth, and rarely flew far. It fed on insects, worms and berries. Its
call was a short whistling cry, quickly repeated, p io-pio – hence the bird’s name. The nest and eggs do not
seem to have been described, but probably did not differ from those of the South Island species.
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Meliphagidae
Mead (1950), if his account can be believed, gave the last account of this bird. His four-day trip in March
1950 took him up the Wanganui River, with a survey party including a Maori captain, Andrew Anderson,
who had spent forty years on the river. The latter described the bird as
. . . inquisitive and tame, yet in a way shy. It would often appear when a wild pig was killed, but on noticing that
it had been observed, would fly away, frequently returning and flying past for one more look before going away
altogether.
In the course of the journey the party saw several individuals, and heard others singing in the trees. Mead
ended with a recommendation that, as so little was known about the bird, someone with more time than he
had ought to undertake a study of the bird and its habits ‘as soon as possible’. Evidently nobody managed
to do so in time.
Meliphagidae (Honeyeaters)
The honeyeaters are widely distributed, with the centre of distribution being Australia and New Guinea.
They also occur on remote islands, including New Zealand, Samoa and Tonga. The family has been
divided into 42 genera with approximately 182 species. Some island species are at risk due to habitat
destruction, but only one species, the Chatham Island Bellbird Anthornis melanocephala, is known to have
become extinct.
The Chatham Island Bellbird was discovered by Dieffenbach (Gray 1843) when he visited the Chatham
Islands in 1839. Buller (1872–73) obtained a specimen there in 1855, and it was still common in 1871, when
Travers found it on Chatham, Pitt, Mangere and Little Mangere Islands (Hutton 1872), but principally on
Mangere Island. Twenty-five years later it was reported to be disappearing fast by a collector who shot all
he could find for financial gain (Oliver 1955).
The Chatham Island Bellbird became extinct in the early years of the 20th century, possibly as a result of
the destruction of the native trees in which the birds lived. Cats and rats may also have contributed to their
extinction, and possibly o ver-collecting for the museum trade. It probably disappeared from Chatham and
Mangere Islands first, but survived longer on Little Mangere Island, where a few pairs were reported in
1906 (Fleming 1937). Greenway (1967) was at a loss to explain their disappearance, since Cockayne (1904)
had found in 1900 that there was substantial forest remaining, and the actual disturbance must have been
relatively minimal at the time the birds vanished. The species was almost certainly extinct by 1924, when
Archey & Lindsay (in Oliver 1955) failed to find any. It is likely that an introduced disease rapidly extir-
pated the birds, since there was a sudden population decline before the onset of specimen collecting and the
introduction of cats and rats; the Hawaiian honeycreepers declined in a similar fashion.
Habits
The Chatham Island Bellbird bred in August, September and October; the nest was in a tree or bush, or
under the head of a tree-fern, at about 3.5m from the ground. The only nest ever preserved is composed of
fine sticks, grass and roots, lined with grass. The eggs were pink with large reddish-brown blotches at the
large end, and there were three to a clutch. Travers (Hutton 1872) noted that the song was richer and fuller
than that of the New Zealand Bellbird A. melanura.
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Dasyornithidae (Bristlebirds)
The bristlebirds were once placed in the family Acanthizidae, but have now been given family status
(Christidis & Boles 2008). There are three species, all d
ull-coloured, terrestrial birds, endemic to Australia.
As they prefer coastal environments they are extremely vulnerable to anthropogenic disturbance.
The Rufous Bristlebird occurs in coastal and sub-coastal areas of southern Australia, with two extant
subspecies, nominate D. b. broadbenti in South Australia and Portland, Victoria, and D. b. caryochrous in
Victoria (Seymour et al. 2003). A western population, D. b. litoralis, was endemic to the s outh-western coast
of Western Australia, between Cape Mentelle and Cape Naturaliste (Johnstone & Storr 2004). It was consid-
ered moderately common in the late 19th century (Storr 1991), but numbers drastically declined. It was last
recorded in 1908, when a specimen was collected (Johnstone & Storr 2004). A systematic search for it in
February 1922 was unsuccessful (Whitley 1971), and despite rumours of its continued existence, none have
been recorded for over a century. It must now be extinct.
It disappeared due to the repeated burning of its shrubland habitat for conversion to pasture (Carter
1924), and cats may have been serious predators, especially as the birds predominantly foraged on the
ground.
Habits
It was a skulking bird, difficult to see and flush from the undergrowth, and was generally solitary, only
occasionally seen in pairs (Higgins & Peter 2002). The preferred habitat was stunted dense coastal shrub-
lands on cliff tops and sand dunes (Milligan 1902 ; Carter 1924), where it hunted for snails (Milligan 1902)
and probably insect larvae, fruit and seeds. It probably foraged in leaf litter and terrestrial vegetation.
Milligan gave a detailed account of the bird:
The bird has two distinct calls – alarm notes and song notes. The former it utters when closely pursued and
pressed, and resembles the words ‘pink, pink, pink.’ The latter is a series of clear, liquid, thrush-like notes . . . The
bird was most difficult to flush or even see, and it was only in the afternoon of the second day’s pursuit that I
obtained a momentary glimpse of it as it rushed across a kwagga [wallaby] track in the scrub with tail elevated.
My next sight of it was on the fourth day of pursuit, just prior to shooting it . . . Its motion on this occasion
(perhaps due to its being severely pressed) was distinct from that observed of the former. Appearing, as it did,
running at top speed across one of those beautifully rounded sand hills (which abound on the coasts), with its tail
depressed below the plane of the body, and its dwarf rounded wings used as an aid to its running, its toes just
touching the ground, and its neck stretched to the utmost, the bird reminded me very much of the Lyre Bird in
similar circumstances.
When first encountered, the Lord Howe Island Gerygone was considered common (Sharland 1929), but by
1924 it was almost extinct (Hindwood 1940). Like so many of Lord Howe Island’s birds, this species disap-
peared after Black Rats Rattus rattus came ashore from the wreck of the Mokambo, on Ned’s Beach in 1918,
but the birds survived for more than a decade after their arrival (Garnett et al. 2010). Owls were imported to
kill the rats, but may have fed on the endemic birds as well. McAllan et al. (2004) suggest that disease intro-
duced by exotic birds may also have been a contributing cause, as the closely related Norfolk Island
Gerygone G. modesta has survived alongside these predators for many years.
Habits
Hindwood (1940) commented that it was ‘known locally as “Rain-bird” because it was very active after
rain, and also “Pop goes the Weasel” on account of the supposed resemblance of its call’. It was insectivo-
rous, actively hunting insects among foliage of trees (Hull 1909), and the birds occasionally entered houses
to glean insects and spiders from the walls and ceilings (Hutton 1991). The d ome-shaped nest comprised
dry bark, fibres, leaves, grass, moss and wool, bound together with spiderwebs and suspended from a twig.
The clutch consisted of three pink-tinged, brown-speckled eggs.
Callaeidae (Wattlebirds)
The wattlebirds of New Zealand are an ancient group whose relationships are unclear. The family comprises
three distinct monotypic genera. They are characterised by the presence of large wattles behind the bill, and
are or were long-legged, poorly volant birds that habitually foraged on the ground. Only the South Island
Kokako Callaeas c. cinerea and Saddleback Philesturnus carunculatus still survive. Habitat destruction and the
introduction of predatory mammals were the primary reasons for their decline and disappearance.
The Huia was the largest member of the New Zealand endemic family, the Callaeidae or wattlebirds. Its
relationships are obscure (Shepherd & Lambert 2007; Lambert et al. 2009); it was confined to North Island,
and its origins are as ancient and obscure as those of the moas and kiwis.
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The bird was remarkable in the fact that it exhibited extreme sexual dimorphism in the bill, the male’s
being short and straight and the female’s long and curved. When the first specimens reached the ornitholo-
gist John Gould, he believed that the sexes were different species, calling the female H. acutirostris and the
male H. crassirostris (Gould 1837a).
The Huia was sacred to the Maoris, whose chiefs had the tail feathers in their headdresses; elaborately
carved wooden boxes were made in which these precious feathers were kept. In Europe, feathers and bills
were also made into gold-mounted brooches and other ornaments. The fashion for feathers played a crucial
part in the bird’s downfall. Its fate may have been sealed by the visit of the Duke and Duchess of York (the
future King George V and Queen Mary) at the turn of the 20th century, when a Maori woman took a feather
from her hair and placed it in the band of the Duke’s hat, thus creating a fashionable demand for Huia
feathers (Higgins et al. 2006). These soon commanded a price of up to £5 each, a hefty sum at that time.
Decline
The Huia once occurred throughout much of North Island, confirmed by the discovery of subfossil remains
(Worthy & Holdaway 2002), but Maori habitat destruction and perhaps o ver-hunting had reduced its range,
and it was limited to southern North Island during the latter half of the 19th century. European wholesale
destruction of the lowlands for agriculture occurred at this time (Buller 1905), and it appears that this
specialised bird required undisturbed, old-growth forest, and could not survive in regenerating secondary
growth (Morris & Smith 1995); it is probable that forest destruction was the most important reason for the
Huia’s decline. Buller (1905) quoted the Maoris as saying ‘You have permitted the killing of the Huia under
heavy penalty, and yet you allow the forests, whence it gets its subsistence, to be destroyed’.
As the bird became rarer, the demand for specimens increased, and thousands were collected and
exported overseas. Greenway (1967), who recorded that there were 65 surviving skins in the world’s
museums, suggested that collecting would not have had an adverse effect on Huia numbers as Maoris had
been hunting the birds for generations. Greenway was wrong; Maori traditional protocols permitted
hunting only during the season when the feathers were in peak condition, not y ear-round. Fuller (2000) also
disagreed with Greenway’s suggestion, and noted that
he had not taken into account the numbers taken just for Huia Heteralocha acutirostris
their tail feathers, and specimens lost during collection; (female left, male right)
indeed, there are 119 skins in New Zealand museums
alone.
The decline of the Huia in the latter half of the 19th
century is poorly known, and most of the surviving
written accounts are from 1890 to 1907, since it was at
this time that ornithologists in New Zealand were first
becoming concerned about the bird’s fate. However, as
early as 1870, Buller (1887–88) described the bird as one
of the country’s rarest and most valuable species, adding
that soon it would exist only in museums and collec-
tions. By the early 1900s, the Huia was effectively
extinct. Myers (1923), with the help of a founding
member of the Ornithological Society of New Zealand,
Robert Stidolph, spent six or seven years collecting
information on possible sight records, and these
amounted to just seven between 1907 and 1922; the last
confirmed sighting was by the ornithologist W. W. Smith
on December 28, 1907.
However, although considered extinct since 1907, a
recently unearthed second-hand report strongly
suggests that the Huia survived until at least 1924.
Stidolph was told by a Captain Val Sanderson that in
1924, a shepherd reported a sighting of a Huia near the
boundary of Whareroa Farm on the Kapiti Coast, north
of Wellington (Wood 2016). Stidolph recorded in his
diary:
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Callaeidae
Capt. Sanderson said a shepherd at the edge of the bush on Mt. Wainui near Paekakariki was whistling for his
dogs when a male huia came to his call. This was only six or eight weeks ago.
Stidolph and Sanderson went on several expeditions in 1924 to locate the bird, but unfortunately all of their
efforts were in vain.
Habits
Buller (1887–88) gave the best accounts of the bird and recorded that:
The Huia never leaves the shade of the forest. It moves along the ground, or from tree to tree, with surprising
celerity by a series of bounds or jumps. In its flight it never rises, like other birds, above the tree tops, except in
the depth of the woods, when it happens to fly from one high tree to another . . . They are generally met with in
pairs, but sometimes a party of four or more are found consorting together . . . this species builds its nest in
hollow trees, forming it of dry grass, leaves and the withered stems of herbaceous plants, carefully twined
together in a circular form, and lined with softer material of a similar kind.
The Huia occurred mainly in montane and lowland hardwood-podicarp forests with dense understorey,
and occasionally in Nothofagus forest; the birds also made some seasonal altitudinal movement (Higgins et
al. 2006). The breeding season began in early summer, and the nest was a large, saucer-shaped structure
measuring around 35cm in diameter, with a small depression lined with finer material (Gill & Martinson
1991). Two to four eggs were laid; they had a greyish background, with brown and purplish markings.
Huias fed chiefly on insects, particularly the grub of the Huhu Beetle Prionoplus reticularis, and also preyed
on various species of wetas, other insect larvae, and fruit. Much of their foraging was done on rotting wood,
decaying bark, mosses and lichens.
The Huia had well-developed musculature for gaping, allowing the jaws to be prised open in order to
pull apart the substrate with considerable force (Burton 1974). This method of feeding was communal; the
male attacked the tree with his strong chisel-like bill, while the female inserted her long, slender bill into
crevices. However, the supposed cooperation in feeding, where the male opened up larval burrows in order
for the female to extract the grub, is unfounded. The sexually dimorphic bill was an adaptation to reduce
competition between the sexes, and to exploit a wide range of food in varied microhabitats (Moorehouse
269
1996). Lambert et al. (2009) showed that the long bill of the female only developed as the bird aged, and was
in fact indistinguishable from that of the male in juveniles.
Apparently the birds were largely silent, but calls have been described as an array of peculiar whistles,
and a soft, melodious, flute-like song (Higgins et al. 2006). The call was penetrating and could be heard in
dense forest from up to 400m away.
The Kokako Callaeas cinerea, the Saddleback Notiomystis cincta and extinct Huia Heteralocha acutirostris are
members of the ancient endemic New Zealand wattlebird family, the Callaeaidae, whose relationships are
still poorly known (Ewen et al. 2006). The Kokako formerly occurred widely on North and South Islands,
New Zealand, but it declined significantly during the mid-20th century, being last officially recorded on
South Island in 1967 (Clout & Hay 1981). There have been a number of unconfirmed reports since then but
no direct evidence for its continued survival (Bain 2009); it was officially declared extinct by the New
Zealand Department of Conservation on January 16, 2007. The North Island population is considered
endangered, with around 1,500 birds remaining (BirdLife International 2011), and there is a large conserva-
tion effort underway to save it from extinction.
The Kokako is a poor-flying terrestrial bird that has suffered particularly from deforestation and intro-
duced predators, especially cats, Stoat Mustela erminea and Black Rat Rattus rattus, and from competition
and nest predation by the B rush-tailed Possum Trichosurus vulpecula. Introduced deer and goats also destroy
suitable foraging habitat for the birds, by over-grazing choice food plants (Innes et al. 1999). The South
Island Kokako disappeared more rapidly than its North Island congener as that it spent more time on the
forest floor, making it more vulnerable to introduced predators (Scofield 2009).
Habits
The South Island Kokako favoured mature podocarp and hardwood forests, where it glided from tree to
tree, preferring to move around by leaping and hopping from branch to branch. It would scramble up tall
trees to gain height in order to glide to a new location, where it would almost crash-land before continuing
in leaps and bounds. It was predominantly herbivorous; foraging on the forest floor or on low branches for
various leaves, flowers, fruit and nectar, but it also took fern fronds and invertebrates (Heather & Robertson
1997).
The bird had a resonating clear, organ-like song that could be heard for miles, with breeding pairs often
singing together in the morning in a b ell-like duet. Buller (1872–73) gave a very good account of the bird
when it was still common, but he also prophesied its impending doom:
The Glaucopis cinerea, or Crow [Callaeas cinerea], of the Middle Island, is rarely found below an altitude of
two or three thousand feet, and indeed, is found in greatest numbers at and above the higher of these altitudes, in
the glens of the Fagus forest. I am inclined to think these birds pair for life, as they are almost invariably found in
couples at all seasons of the year. They are extremely active, hopping with long strides along the ground, and
from branch to branch, in their search for insects. Their chief food, however, consists of sow-thistle and other
succulent herbs, and it is remarkable that, in eating such substances, they hold them with the fist precisely as a
parrot holds its food, tearing off and swallowing large fragments. The note of this bird is wonderfully sweet and
plaintive, and, during the breeding season, its song is one of the most varied and beautiful of all New Zealand’s
birds. It appears, however, always to be pitched in a minor key. The male birds are very pugnacious, fighting,
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whenever they meet, with the greatest determination. They are still numerous in the forests adjoining the station,
but I fear wild cats are likely to clear them out within a few years.
Notiomystidae (Stitchbird)
The New Zealand Stitchbird Notiomystis cincta was until recently placed in the Meliphagidae (honeyeaters),
but mitochondrial DNA has shown that the species is monophyletic and warrants placement in its own
family, the Notiomystidae (Driskell et al. 2007), it being more closely related to the New Zealand
wattlebirds.
The New Zealand Stitchbird N. cincta was confined to North Island and adjacent islands, but declined as the
forests were cleared. It also suffered from predation by Black Rats Rattus rattus, and possibly introduced
avian diseases (Angehr 1984). The bird has not been recorded on the mainland since 1880.
The last natural population of the species survives on rat-free Little Barrier Island, from which a translo-
cation programme has met with limited success. Birds on the island are sometimes considered a separate
race, N. c. hautara (Mathews 1935a); they differ in having a shorter wing and more intense yellow on the
breast of the male. Splitting this population as a race means that the mainland nominate is therefore consid-
ered extinct (Angehr 2011).
Habits
The nominate race once occurred in a range of forest types, but being an exclusive tree-cavity nester
(Guthrie-Smith 1925) it appears to have required large expanses of mature forest in order to survive. It was
known to forage in the lower forest storey, feeding predominantly on nectar, but it also took fruit and inver-
tebrates (Angehr 1984; Castro et al. 1994b). Buller (1872–73) wrote of the nominate:
They can hover like great bees or humming birds in front of blossoms. With the sunlight falling full on the
splendid gold of the outspread wings, or the deep blacks and pure whites of head and neck, the male then appears
not a bird but a huge brilliant tropical butterfly – a magnificent creature indeed.
The species has a peculiar and unique breeding ecology, being the only bird species known to copulate
while facing each other (Anderson 1993; Castro et al. 1996). The call of the male is a loud piercing whistle
see–si–ip, with a varied low warbling song; the alarm call, rendered stitch stitch, is the origin of the European
name for the bird.
Cinclosomatidae (Quail-thrushes)
The quail-thrushes are restricted to Australia and New Guinea, where five species are recognised. They are
terrestrial, weak-flying birds that take to the wing only when pressed. They are generally insectivorous, and
forage on forest floors, and also nest on or near the ground. This has made them vulnerable to introduced
mammalian predators.
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The Spotted Quail-thrush is divided into three subspecies, the nominate C. p. punctatum in south-east
Australia, C. p. dovei in eastern Tasmania, and C. p. anachoreta, which was endemic to the Mount Lofty
Ranges in South Australia.
The Mount Lofty Spotted Quail-thrush appears to have been restricted in range, but it was considered
reasonably common in the early 20th century. However, it gradually disappeared, becoming rare by the
1930s (Ashby 1932), and extremely rare by the 1950s (Terrill & Rix 1950). Since then there have been only a
handful of sightings. There were a few in the 1960s, one in 1971, one in 1977 (Higgins & Peter 2002), and one
in 1981, when a single breeding observation was made (Garnett & Crowley 2000). One was seen at Mount
Bold in 1982 (Garnett et al. 2010), and there was a final record in 1984 (Blakers et al. 1984). Subsequent general
surveys of the Mount Lofty region have failed to locate the species (e.g. Barrett et al. 2003).
The probable extinction of the bird can be blamed almost entirely on severe deforestation, fragmentation
of habitat, and general habitat degradation (Close 1972; Garnett & Crowley 2000). The effects of large
numbers of feral cats, dogs and foxes are worsened by the opening up of primary forest, and by the Spotted
Quail-thrush’s habit of foraging on the ground. Little of the original habitat remains, and there is little likeli-
hood of this race being rediscovered.
Habits
Little is known about the habits of this bird. It was shy and wary, difficult to approach, and recorded singly,
in pairs, and in small family groups (Bransbury 1984), preferring sclerophyllous forest dominated by
Eucalyptus trees, heathland on stony slopes, and ridges in valleys and gorges (Schodde 1954). Eggs were laid
from August to September (Ford 1983), with the birds nesting in a well-disguised cup built on the ground
(Higgins & Peter 2002). The birds’ diet comprised a variety of insects (especially beetles and grasshoppers),
and seeds of various plants were also consumed (Lea & Gray 1935).
The extant subspecies forage on the ground, turning over leaf litter, bark and small sticks, or gleaning
insects from low vegetation (Green 1995); the Mount Lofty subspecies almost certainly foraged in the same
manner.
The Pied Currawong Strepera graculina is divided into five subspecies, of which only the Lord Howe Island
subspecies, S. g. crissalis, is considered endangered (McAllan et al. 2004). The western race was isolated and
occurred on the grassy plains in southwest Victoria, extending west to the South Australian border (Garnett
et al. 2010). It was common in the Grampians in 1925 (Ashby 1927). However, the southeastern subspecies,
S. g. nebulosa, appears to have hybridised the Western Pied Currawong out of existence (Garnett et al. 2010).
Native Australian burning regimes formerly maintained the habitat of the Western birds, but after the aban-
donment of such practices since the late 18th century an increase in tree density occurred; this was before
systematic tree-clearing was universally practiced (Barr & Cary 1992; Garnett et al. 2010). This favoured S. g.
nebulosa, and encouraged their invasion. Garnett et al. suggest that the extinction of pure-bred birds of the
western races occurred before 1900, before the species was even described (in 1913).
Campephagidae (Cuckoo-shrikes)
The cuckoo-shrikes are forest birds that are widely distributed in tropical Africa, Asia and Australasia. The
family has been divided into at least 8 genera, with approximately 85 species. Males are predominantly
grey or grey and white, whereas the some females superficially resemble cuckoos, hence the name. Being
forest inhabitants, some island species and subspecies are endangered or extinct due to deforestation.
The Blackish Cuckoo-shrike is divided into three subspecies, two of which are now possibly extinct; only
the nominate C. c. coerulescens of Luzon and Catanduanes is known for certain to survive.
A bird from the population on Marinduque was first collected in 1971 (DuPont 1972), but the race has not
been recorded since, and might be extinct. The Philippines have suffered severe deforestation, especially in
the lowlands, a habitat favoured by the cuckoo-shrikes. If it still survives, this race must be critically endan-
gered. Nothing is known about its habits.
The Cebu Blackish Cuckoo-shrike has not been seen since McGregor (1909b) collected it in 1906. Its extinc-
tion can be attributed to the almost complete deforestation of the island. Although some Cebu birds once
thought to be extinct were rediscovered during the 1990s, Brooks et al. (1995) considered this race to be
extinct. However, an unconfirmed report was made in 2000 (Pedregosa in Paguntalan & Jakosalem 2008). If
it still exists in what little forest remains on Cebu, it must be exceedingly rare.
Very little was recorded about the habits of this cuckoo-shrike. It occurred in primary forest in the hills,
and McGregor (1909) found it fairly common, occurring in flocks in high trees in forests.
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The Bar-bellied Cuckoo-shrike C. striata occurs in India, South-east Asia and the Philippines. The Cebu race
has not been seen since McGregor (1909b) collected it in 1906. Almost the entire island has been deforested.
Brooks et al. (1995) considered the race extinct, but like the Cebu Blackish Cuckoo-shrike C. coerulescens
altera, there was an unconfirmed report of its continued existence in 2000 (Pedregosa in Paguntalan &
Jakosalem 2008). Little of its habits was recorded. It frequented primary forests of lowlands, hills and moun-
tains, but also occurred in secondary growth (Rabor 1959).
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black above with soft fluffy plumage; rump fawn-buff; uppertail-coverts dark grey; tail black, tipped with white; lesser and
median wing-
coverts white; primaries and greater w ing-
coverts black, edged with white; cheeks and ear-coverts
yellowish-white; under surface light ochraceous buff; throat white. Female browner than male with blackish centres to feathers
of head; uppertail-coverts with sandy margins; and w ing-coverts washed sandy.
The Long-tailed Triller L. leucopyga is common, occurring in New Caledonia, the Solomon Islands, and
Vanuatu. The Norfolk Island race was common when Hull (1909) observed it in 1908–09, but it died out
within the next 30 years or so, and was last recorded in 1942. Its extinction was due to Black Rat Rattus rattus
predation and severe deforestation (Higgins & Cowlin 2006). Predation by rats increased in 1951 after the
island’s cat population was hit by distemper (Tennyson & Martinson 2007). The construction of an airport
in the centre of the island, which resulted in the clearance of a large area of remnant forest, probably finished
the bird off (Garnett & Cowley 2000).
Habits
The triller was an inhabitant of Norfolk Island’s subtropical rainforest, which has now been virtually
cleared. The breeding season began in September, and usually two, sometimes three eggs were laid in
November and December, but if conditions were suitable the birds would also breed from January to March
(Hull 1909). The nest was a shallow cup, made from lichen, moss and fibrous roots, and lined with finer
material (Higgins & Cowlin 2006). Hull (1909) gave an account of its habits:
This is one species peculiar to Norfolk Island, where it is found in considerable numbers. A bright and lively
little bird, in general appearance it closely resembles Lalage tricolor [White-winged Triller], but is smaller
. . . It perches on stumps or fences, from which it pounces on g round-insects, or flying upwards, takes others on
the wing. Owing to its habit of collecting an early breakfast amongst d ew-drenched grass, its otherwise
snow-white breast becomes discoloured with the reddish dust which settled on the g rass-blades overnight. Its
chief note is a single harsh chirp, like the first or last staccato of a cicada.
Oriolidae (Orioles)
The orioles have a wide distribution in Europe, Africa, Asia and Australia. The family has been divided into
two genera, but further divisions are likely. They are medium-sized birds, with the males tending to be
brightly coloured. In general the orioles are not threatened, and only one island subspecies is known to
have become extinct.
The Philippine Oriole O. steerii occurs widely throughout the Philippines, with six subspecies currently
recognised. In March 1877, A. H. Everett visited the island of Cebu (Tweeddale 1877). Everett remained on
the island for several weeks and made some important discoveries, including the endemic race of this
species.
Most of Everett’s specimens were collected in March and were obtained near coal mines, about 19km
north of the port of Cebu City and some 9km inland. In 1892, Bourns & Worcester (1894) found it ‘exceed-
ingly common in the small amount of forest left in Cebu’. The Cebu Oriole has not been seen since McGregor
(1909b) collected it in 1906, and Paguntalan & Jakosalem (2008) failed to find any after extensive surveys.
Deforestation was the primary factor in its extinction.
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The Lord Howe Fantail disappeared from the island between 1924 and 1928, probably as a result of the
escape of rats from the wreck of the Mokambo in 1918. It was very nearly extinct in 1924, when a local resi-
dent, Harry Payten, stated that it had been ‘practically wiped out’ (Hindwood 1940), and by 1928 it appears
to have completely disappeared (Sharland 1929).
Habits
The Lord Howe Fantail occurred in open forest, where it was formerly common (Hutton 1991). Hull (1909)
published some interesting observations on these birds:
On my arrival at Lord Howe Island, 3rd October, 1907, Mr. J. B. Waterhouse informed that he had found a
Fantail’s nest near his house. On the following day I visited the spot, and was shown a nest which the birds had
built a few weeks previously, but abandoned on completion. It was placed on a horizontal twig of a small tree,
about fifteen feet from the ground. A fresh n esting-place had been chosen in a small prickly shrub, overgrown
with clematis, the nest being almost hidden amongst the v ine-leaves, securely fastened in the fork of a small
branch of the shrub. The female was sitting, and reluctantly slipped off at our approach. The nest contained two
eggs, and, thinking that the full complement had not been laid, I left it until the 8th October. The female was still
sitting on that date, but no more eggs had been added to the original two, which were found to be slightly
advanced in incubation. This Fantail is very tame, and fond of frequenting the vicinity of dwellings, where it
will often enter the kitchen and capture flies from the walls.
The nest was a cup of decayed wood fibres and fine grass, bound together with cobwebs and lined with
grass. Three eggs (sometimes only two) were laid; these were creamy white, spotted and streaked with pale
brown and slate markings, forming a cap at the larger end. Eggs were laid in October and November (Hull
1909).
The Rufous Fantail Rhipidura rufifrons inhabits the eastern coastal ranges of Sulawesi through Australia and
New Guinea to the Solomon Islands. The Guam population was once common and widely distributed on
the island (Seale 1901). However, the Brown Tree Snake Boiga irregularis was introduced to the island in the
1940s; the fantail population had disappeared from most of southern Guam by the late 1960s, but could still
be found in the southern coastal regions in 1977 (Drahos 2006). They must have disappeared shortly after as
they survived only in the forests of the n
orth-eastern cliffs a few years later (Jenkins 1983). The birds were
still considered common there in 1978–1979, but they rapidly disappeared, and were last recorded in 1984
or 1985. The disappearance of this bird can be attributed almost entirely to predation by the Brown Tree
Snake, but Drahos (2006) considered its final rapid decline to extinction was caused by predation by rats
and monitor lizards as well as the snake, perhaps assisted by residual insecticides.
Habits
The Guam Rufous Fantail occurred originally in all habitats on Guam, with a preference for forest under-
storey and scrub, especially Leucaena leucocephala forests, but it did not occur in the southern savannas
(Baker 1951; Jenkins 1983; Drahos 2006). They were active birds, continually spreading their fanned tails,
foraging in pairs or small groups for insects in the understorey. Prey was mostly taken on the wing. They
often associated with the now extinct Guam Flycatcher Myiagra freycineti (see p. 280), with the birds sharing
similar habitat preferences (Drahos 2006). The nest was a neat and compact construction, made from fine
vegetable fibre glued together with m
ucous-like saliva, with a projection of fibres from the base (Seale 1901),
and situated 1–2m above the ground. The breeding season was from January to April, and two dull white
eggs were laid, each with a ring of brownish spots diffused around the centre or nearer the large end
(Jenkins 1983). The song was a melodious array of notes.
Monarchidae (Monarchs)
The monarchs (or monarch-flycatchers) are birds with glossy black males and b rown-and-white females,
occurring in the Marquesas, the Society Islands and the Cook Islands. Although five species are listed by
Mayr et al. (1986), the number of valid species still remains to be resolved. They are so similar morphologi-
cally that many earlier writers (e.g. Sharpe 1879) failed to distinguish between them.
Maupiti is a small, high island that was w ell-wooded in the 19th century, but the ravages of human occupa-
tion have left little of the original forest, and introduced animals including rats are in abundance. At some
time in the latter half of the 19th century, the Maupiti Monarch disappeared. Nothing is known about its
habits.
Habits
The monarchs occurred in high-elevation forested valleys, but also utilised secondary and degraded forest,
though this is probably an artefact of the complete destruction of its lowland habitat, with the last popula-
tion surviving outside what would have been its normal range. The birds foraged for insects in the dense
understorey.
This species was restricted to forested valleys at the western end of Ua Pou in 1922 (Murphy & Mathews
1928), when it was discovered by the Whitney South Seas Expedition. By 1975 it was only found in areas
above 450m, and the population was estimated at 150–200 pairs; 5–7 pairs were also recorded in native
forest in the upper Hohoi valley (Coates et al. 2006). However, the population quickly collapsed, and the
observation of two immatures in March 1985 in the Hakahetau valley is the last authentic record (Thibault
& Meyer 2001); searches for it in 1989, 1990, 1998 and 1999 were unsuccessful. It may now be extinct. The
introduction of the tree-climbing Black Rat Rattus rattus, the rapid spread of the invasive legume, Leucaena
leucocephala, which destroys suitable habitat, plus the destruction of lowland forest appear to have caused
its extinction.
Habits
Little was recorded about its habits. It occurred in native forest dominated by Hibiscus tiliaceus, Fagraea
berterorana and Pandanus tectorius (Coates et al. 2006).
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The only surviving monarch in the Marquesas, P. motanensis, occurs on Mohotani, where the population is
considered to be stable (Holyoak & Thibault 1984). Although the island has suffered from deforestation and
the introduction of feral cats and the Pacific Rat Rattus exulans, the key factor in its survival is that the island
is free of Black Rats Rattus rattus. Birds on the island of Nuku Hiva were not so fortunate. They were rare in
the 1930s, and had disappeared by 1975, when Holyoak & Thibault (1977) failed to locate the species. Two
supposed observations by hunters in 1975 are extremely doubtful. Nothing is known of its habits. It disap-
peared for the same reasons as the Ua Pou Monarch.
The Eiao Monarch was originally considered a subspecies of Iphis Monarch Pomarea iphis, but it has now
been given full specific status (Cibois et al. 2004). It was restricted to the small, uninhabited island of Eiao in
the Marquesas Islands, where it occurred in low-elevation dry forest, foraging for insect prey in dense
underbrush.
It was numerous in 1922 when discovered by the Whitney South Seas Expedition (Murphy & Mathews
1928), but despite a number of thorough ornithological surveys it has not been seen since 1977 (Coates et al.
2006). The species is now almost certainly extinct (Cibois et al. 2004). It occupied lowland dry forest, which
was heavily degraded after the introduction of sheep in the early 1880s (Thibault & Meyer 2001), and feral
cats and rats no doubt contributed to its decline. Thibault (1989) suggested that the release of the
Chestnut-breasted Munia Lonchura castaneothorax coincided with the disappearance of Eiao Monarch, and
that an introduced disease may possibly have been involved in its demise.
The Celestial Monarch is divided into two subspecies, with the nominate occurring on a number of islands
in the Philippines. The race H. c. rabori was confined to Negros, with one record (a specimen) from Sibuyan,
taken on October 10, 1892 from an unspecified locality (Bourns & Worcester 1894). It occurred in the canopy
or the middle storey of subtropical or tropical moist lowland forests up to 750m, where it foraged for insects.
The bird was considered common in the early years of the 20th century, but was rare by 1959. The Negros
population continued to decline due to the severe deforestation of the lowlands, and has not been recorded
since the 1990s (Collar et al. 1999). It may be extinct.
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Habits
This bird occurred in all habitats on Guam except the southern savannas, skulking and foraging for insects
in thick vegetation. It was often seen in association with the Guam Rufous Fantail Rhipidura rufifrons uraniae
(see p. 276), but it tended to forage higher in the canopy (Drahos 2006). The birds apparently bred
year-round, but the main breeding season was from January to June, with less activity at other times of the
year, especially in November and December (Jenkins 1983; Drahos 2006). The nest was well hidden, and
was constructed from bark, lichen and fine grasses bonded with mucous and spider silk (Drahos 2006). Two
dark cream eggs spotted with r ufous-brown dots and splotches were laid, though sometimes only a single
egg was laid (Jenkins 1983). It aggressively defended its territory, sometimes driving away the much larger
Micronesian Starling Aplonis opaca, and it could be located by its loud contact call, which comprised a series
of one to six whistles.
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Corvidae
Corvidae (Crows)
The crows have a worldwide distribution, except for the extreme polar regions. The family is diverse and
includes crows, jays, treepies and choughs. At least 21 genera with more than 120 species are recognised.
Despite persecution, most species remain unthreatened, but forest species and species restricted to islands
and island groups are vulnerable to habitat destruction and predation by introduced mammals.
The discovery of the New Zealand ravens and Hawaiian crows from subfossil remains has shown that the
family was once distributed widely across the Pacific, and that island populations were extremely vulner-
able to anthropogenic changes. The Chatham Island Raven averaged larger than the mainland New Zealand
Raven C. m. antipodum (see below), and the bill was long, broad and slightly curved. The bird reached a size
of 72cm in total length, and weighed between 833g and 955g or even up to 1kg (Holdaway 1989), making it
one of the largest passerines known (Scofield et al. 2016). Its remains are commonly found in coastal sites on
the Chatham Islands (Gill 2003), and, although not conclusive, this suggests that the Chatham Island Raven
may have been a coastal bird, perhaps seasonally scavenging amongst seal colonies, and foraging at other
times in the lowland coastal forests for fruits and invertebrates (Worthy & Holdaway 2002). Holdaway
(1989) further suggests that it made a large stick nest in trees or on cliffs.
Forbes (1892c) placed the Chatham Island Raven in its own genus, Paleocorax, based on cranial differ-
ences, but Worthy & Holdaway (2002) consider this taxonomic placement unwarranted. A recent molecular,
osteological and fossil distributional study has further shown that the New Zealand ravens form a sister-
taxon relationship with a clade of Australian birds, Australian Raven Corvus coronoides, Little Raven C.
mellori, and Forest Raven C. tasmanicus, and that the endemic New Zealand ravens diverged from the
Australian clade in the Pleistocene, around 1.73 mya (Scofield et al. 2016); the diversity of New Zealand
ravens also stemmed from a single invasion. This event coincided with the start of a period of glaciation
that saw a marked decrease in global temperature worldwide, which may have opened up niches for birds
such as the New Zealand ravens to utilise (Atkinson & Millener 1991). There is also some evidence to
suggest that the New Zealand ravens were more widespread during the last glacial maximum (Tennyson &
Martinson 2007).
As the divergence of the Chatham Island Raven and New Zealand Raven (see below) occurred only 0.15
mya, and osteological differences are not distinct with much overlap, Scofield et al. (2016) consider these to
be races of a single species, with the South Island population (formerly race pycrafti) a junior synonym and
not a valid subspecies. The tarsometatarsus of the New Zealand ravens was elongate compared to
Australian Corvus, which suggests that they were more terrestrial, but unlike previous studies that
suggested they were poor fliers (e.g. Pycraft 1911), they exhibited no reduction in wing length, suggesting
that they flew strongly (Scofield et al. 2016). Furthermore, the New Zealand ravens exhibited an ossified
palate as well as other adaptations in the cranium (Scofield et al. 2016), which further suggest that they had
a specialised feeding strategy (see New Zealand Raven below).
The arrival of the Morimori followed by the Maori, both of whom cleared the forests and introduced
potential predators, and the loss of food supply, particularly carrion (Holdaway 1989), led to the raven’s
extinction.
Despite being smaller on average than the nominate Chatham Island Raven C. m. moriorum, the New
Zealand Raven was still a large bird, with a long but rather broader and straighter bill than its Chatham
Islands congener (Gill 2003). Both were morphologically typical corvids, but the long tarsometatarsus
suggests that these birds may have occupied a more terrestrial niche than other ravens. Like the Chatham
Island birds, remains on New Zealand are numerous in coastal fossil deposits, and the birds would have
been extremely vulnerable to human-induced changes of lowland habitat.
All of the New Zealand ravens were perfectly adept fliers, and a single bone (attributed at the time to the
South Island race, which has now been synonymised with the North Island one) has been found on the
remote Auckland Islands (Worthy & Holdaway 2002). However, it is not known whether this bone repre-
sented a resident population or was that of a vagrant to the islands.
Habits
The coastal distribution of New Zealand Raven fossil remains is congruent with the former distribution of
an extinct clade of New Zealand Sea-lion Phocarctos hookeri and haul-out areas of Southern Elephant Seal
Mirounga leonina, which suggests that the New Zealand Raven was seasonally reliant on pinniped breeding
colonies for food (Scofield et al. 2016). The bill was larger and heavier than Australian ravens, and Scofield et
al. suggest that the specialised ossified palate of the New Zealand Raven was evidence that they were facul-
tative scavengers of carrion associated with these vast sea-lion colonies, which once occurred on the beaches
of North Island, eastern and southern South Island and Stewart Island. They may have also taken advan-
tage of Haast’s Eagle Aquila moorei (see p. 88) kills (Holdaway 1989). Placostylus ambagiosus land snails in the
far north of New Zealand could have provided an alternative food source when sea-lion carrion was
unavailable (Scofield et al. 2016).
The Puerto Rican Crow is known from a single right ulna discovered in a cave deposit on Puerto Rico, and
from other remains on St Croix in the Virgin Islands (Wetmore 1937a). It was a small species, decidedly
smaller than the White-necked Crow Corvus leucognaphalus, a species it was once sympatric with on Puerto
Rico (Wetmore 1920) and which has now itself been extirpated on the island (Madge & Burn 1993). The
endemic Puerto Rican Crow may have occupied a different niche, perhaps having a different-shaped bill
and favouring lowland habitats. It must have disappeared soon after the island was first colonised by
humans.
One of the many interesting aspects about Hawaiian palaeornithology is the potential for discovery of
subfossil remains that completely undermine present understanding of supposed avian biogeography in
the region. The discovery in the 1970s and 1980s of new species of corvids in the Hawaiian Islands illus-
trates this. The Hawaiian crows were previously known only from a single species on the island of Hawaii,
but the fossil record has now shown that this group of birds radiated into a number of forms, and that the
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Petroicidae
The fossil record shows that a second large species of crow formerly occurred on Oahu, where it was sympa-
tric with the Deep-billed Crow C. impluviatus, and also on Molokai (James & Olson 1991). The Robust Crow
was larger than the extant Hawaiian Crow C. hawaiiensis, and had a longer, straighter and shallower bill,
and differed from C. impluviatus in the shape of the bill and characters of the p
ost-cranial bones.
The fossil localities on Oahu and Molokai where C. viriosus was found are both situated in lowland,
coastal sites, so it may have been a predominantly lowland species. The bill shape suggests that it may have
foraged in different habitats or on different food items than those preferred by the Deep-billed Crow. The
lowlands of all the main Hawaiian Islands have been altered drastically by human occupation; the Robust
Crow is just one of the many species that disappeared before the arrival of Europeans in the late 18th
century.
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Although this subspecies is poorly differentiated from the nominate Hooded Robin M. c. cucullata, it is
regarded as valid in recent work (Schodde & Mason 1999). It occurred on Melville Island and Bathurst
Islands, where it was considered common in the early 20th century (Mathews 1914), but has only been
observed on a few occasions in recent years. Changes in the firing régime of the vegetation by Native
Australians, which is now infrequent, appear to have led to a decrease in foraging area for the robins
(Higgins & Peter 2002). There is no other known reason for the bird’s disappearance. It was last recorded in
December 1991 on Bathurst Island, and January 1992 on Melville Island (Fensham & Woinarski 1992).
Surveys to locate the subspecies in 1996 and 2000 were unsuccessful.
Little was recorded about the bird’s habits. It was conspicuous and bold, favouring tall, dense, grassy
understorey in Acacia scrublands, where it foraged for insects.
A large series of birds were obtained by the professional collector Hyojiri Orii in August 1922 on the Daitō
Islands, which were subsequently deposited in the Yamashina Institute for Ornithology. From the collec-
tion, the Japanese ornithologist Nagamichi Kuroda described a new subspecies of Varied Tit Sittiparus varius
orii, endemic to Kita Daitō-jima and Minami Daitō-jima, and named it in honour of the collector (Kuroda
1923). These are the only specimens known. The bird appeared to be numerous when first collected, but the
subsequent destruction of almost all of the original forests for sugar cane production and military construc-
tion of airfields after human settlement in the 1930s resulted in its rapid extinction (Takagi & Akatani 2011);
the Daitō Varied Tit was last reliably recorded in 1938 (Harrap & Quinn 1996). Searches were made in 1984,
1986 and subsequently, but without success (Brazil 1990; Gosler & Clement 2007).
Habits
Nothing was recorded about the habits of the Daitō Varied Tit, but other subspecies generally inhabit broad-
leaved evergreen forests, feeding on invertebrates, especially lepidopteran larvae (Yamaguchi 2005). They
nest in tree cavities, lay between three and eight eggs, and the young fledge after 16 days.
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Mohoidae
Mohoidae (Mohos)
One of the most extraordinary discoveries of recent years was the resolution of the true relationships of the
Hawaiian ’o’os (pronounced o h-ohs), always previously considered to be members of the honeyeater family,
the Meliphagidae. Phylogenetic analysis from mitochondrial DNA sampling places them among a clade of
unusual passerine families – the waxwings (Bombycillidae), silky flycatchers (Ptiligonatidae), and the Palm
Chat Dulus dominicus of Hispaniola, which is placed in its own family, Dulidae (Fleischer et al. 2008). Their
resemblance to meliphagids was a result of convergent evolution, despite their morphological, behavioural
and ecological similarities. The Kioea Chaetoptila angustipluma (see p. 289), which differed considerably in
coloration from the other ‘o’os, was especially reminiscent of the Meliphaga honeyeaters of Australia and
New Guinea. The familial status given to the Hawaiian ‘o’os is not without consequence; it means that this
is the only family of songbirds in the last two centuries to have suffered complete extinction.
The Kaua’i ’O’o survived longer than any other member of the Mohoidae, but it seems that it, too, is now
extinct. It was common in the 1890s, but there was a drastic decline in the early 1900s (Munro 1944), and it
was feared extinct until its rediscovery in the Alaka’i Swamp on Mount Wai’ale’ale in the 1960s (Richardson
& Bowles 1964). Confined to the Alaka’i (Pratt 1994), its numbers continued to decline; Conant et al. (1998)
made a brief study in July 1975, but by 1981, only a single pair remained. In 1982, Hurricane Iwa devastated
parts of Kaua’i, and the female disappeared. The male, possibly the last of the Mohoidae, continued to call
from an old nest site but to no avail, and was last seen in June 1985 (Sykes et al. 2000). A possible vocal
record was made on April 28–29, 1987, which may have been the same bird, but there have been no recent
sightings despite extensive surveys (Conant et al. 1998).
The extinction of this species was caused by a number of factors (see Sykes et al. 2000). Habitat destruc-
tion and predation by Black Rats Rattus rattus were probably significant, but the introduction of pigs, which
are widely distributed and have penetrated remote mountain areas, may have been the main culprits. Pigs
open up forests by forcing paths through the understorey, and by their actions of wallowing provide
breeding pools for mosquitoes, which hitherto were not found in high montane forests. Nearly all Hawaiian
birds are susceptible to avian malaria Plasmodium relictum and avian pox Poxvirus avium; it seems that the
mosquito was responsible for finishing off the Kaua’i ’O’o.
Habits
The Kaua’i ’O’o was common in all forests, from s ea-level to the highest elevations (Munro 1944). It was
more insectivorous than the other species, and its short, stiff, pointed tail was a woodpecker-like adaptation
for bracing itself against the trunk of trees while foraging (Perkins 1903). The diet comprised native cock-
roaches, spiders, prognathogryllid crickets, beetles and their larvae, and they would search on and under
the bark to secure prey items. Nectar was regularly taken, particularly from lobelias, Ohia Metrosideros poly-
morpha, and from introduced bananas. Their flimsy and untidy nests were made in the topmost branches of
trees, using dead twigs and dry fern fronds, but J. L. Sincock (in Sykes et al. 2000) recorded three cavity nest
sites, all in natural hollows in Ohia trees. Vocalisations were varied and complex, and included a loud
two-note phrase, oh-oh, that could be heard up to 100m away (Sykes et al. 2000).
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The Oahu ’O’o disappeared very early in the ornithological history of the Hawaiian Islands, and virtually
nothing is known about it. However, Andrew Bloxam, naturalist on HMS Blonde and collecter of the first
Oahu ’O’o in 1825, kept a diary and made notes, which provides some information (Olson 1996a). Bloxam
stayed on Oahu for a month, from May 6 until June 7, but he did not encounter the ’O’o himself and had to
pay a high price for the specimens. Like the Hawaii ’O’o (see below), the Oahu bird could easily be kept in
captivity, and Bloxam had the chance to keep some during his one-month sojourn (from Olson 1996a):
Their note is a harsh chirp of two or three different tones. They feed principally on the nectar of the Bowers of the
Eugenia Malaccensis. I kept some alive which the natives brought me, tho’ almost entirely destitute of feathers,
for three of four weeks, feeding them principally upon sugar & water. They took flies, however, which came into
their cage, with great quickness & adroitness. They are apparently a strong & hardy bird, but the cold would
probably . . . hinder them from being taken to England. They are now very scarce in all the islands. I did not see
even one in the different excursions I made, & the natives asked a high price for the very few they brought to me
& and almost the whole of these were destitute of feathers. I preserved only one tolerable specimen the whole time
I was upon the islands – & even from that some of the yellow feathers had been plucked out.
It is generally believed that the last three known specimens were collected by J. K. Townsend and Herr
Deppe in the hills beyond Honolulu (in the Nu’uanu Valley) in January 1837 (Olson & James 1994a), twelve
years after Bloxam’s visit. However, a female collected by Deppe has September 7, 1840 written on the
specimen label (Sykes et al. 2000).
The bird was searched for in the 1890s by various naturalists, including the competent Perkins (1903),
but no trace was found. It is certainly extinct, and probably was for most of the 19th century. The Oahu ’O’o
disappeared for similar reasons as the Kaua’i birds, and also due to o ver-hunting for its valuable yellow
plumes.
The Hawaii ’O’o was first discovered on Cook’s third voyage, and was the largest member of the genus.
Perkins (1893) reported that it was still a common bird when he observed it, but it frequented the tallest
lehua trees and was thus difficult to observe at close hand. Munro (1944) found it common above Kaawaloa
in 1891 and 1892, but it had disappeared by 1894, apparently due to much of the native forest being cleared
for coffee plantations. Perkins (1903) explained that it was fast disappearing as a result of the forest being
opened up by browsing cattle, making it unsuitable for the birds. This coincided with an increase in hunting
for the prized yellow plumes and demand for museum skins – a staggering 1,000 were taken in 1898 alone
(Henshaw 1902). Henshaw collected around 23 specimens between 1900 and 1902, the last individual being
taken on May 13, 1902 at Ka’au Crater by an M. L. Walton (Sykes et al. 2000). Perhaps o ver-hunting was the
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Mohoidae
Habits
It was reported by all observers to be an extremely shy
and timid bird, inhabiting the tops of the tallest trees,
though occasionally, where it was not persecuted, it
foraged in the undergrowth. It was predominantly
nectarivorous, favouring arborescent lobelias and Ohia
Metrosideros polymorpha, and had a long tubular tongue
for the purpose, but it was also fond of the red fruit of
the ’ie’ie vine Freycinetia arborea, and various insects and
caterpillars.
Perkins (1903) noted that the Hawaii ’O’o was easily
kept in captivity on a diet of sugar-cane juice, and it
would become tame; old native bird-catchers would
keep a tame one as a decoy, to attract others to sticky
bird-lime traps. Some were released on other islands
around 1892, and those released on Kaua’i were still
alive a year or two later. In their dipping flight they
were reminiscent of a Magpie Pica pica (Wilson & Evans
1890–99); while foraging the wings were flapped at
great speed, producing a unique, continuous buzzing
sound (Munro 1944). The call could be heard up to half
a mile away (Perkins 1903), was usually disyllabic, and
varied according to the season. The nest was never
discovered, but Perkins (1893) was told that birds were
seen entering tree hollows.
Robert Perkins, who spent weeks camping in the Hawaii ’O’o
mountains of the Hawaiian Islands, b are-foot and on Moho nobilis
horseback, provided wonderful descriptions of the
habits of the Hawaii ’O’o and other Hawaiian birds
before their extinction; ornithology has much to thank him for. His notes on the Hawaii ’O’o’ are typical of
his observations (Perkins 1903):
The Oo is very agile in its movements when feeding amongst the Ohia flowers, gliding and darting from bloom
to bloom and from limb to limb with great rapidity, at one moment one side, the next far away on the other side of
some giant tree. It is very intolerant of the scarlet Iiwi (Vestiaria), and will at intervals suspend its feeding to
chase away any of these that have ventured into its own tree, or it will even leave this, however profuse to be in
blossoms, to drive from some distant tree one that it has chanced to spy there. This aggressiveness appears so
wanton and unnecessary, and so frequently interrupts its own feeding, that one suspects it must be an ancient
habit, which has survived from a time when either nectar-producing flowers were scarcer, or the birds which feed
upon them were much more numerous. In attacking the other n ectar-eating birds it makes a savage rush on the
one that excites its animosity, and in making this attack it frequently erects its tail with the long twisted feathers,
and also raises its wings, so that the yellow axillary feathers and u ndertail-coverts of the same colour can be
distinctly seen . . . it presents a very fine appearance.
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The Molokai ’O’o was last observed by Munro (1944) in 1904, when about six birds were seen. He was told
in 1915 that birds were still to be found, but later searches found none, in spite of the fact that, as late as 1949,
tracts of apparently suitable forest survived right up to the top of Mount Olokui (Collar et al. 1994). However,
Black Rats Rattus rattus were seen in the trees in daytime (Richardson 1949) and were evidently a great
threat to the birds. The species is now certainly extinct.
Habits
Although less persecuted than its Hawaii Island congener, this was a restless and timid bird, but it was
inquisitive and could be beckoned by imitating its call. Perkins (1903), who again made excellent field notes,
wrote that the bird was:
. . . seen numerously in the Ohia trees, when these are in flower, but this is clearly because its favourite lobelia
nectar is unattainable. Thus on one of my visits to Molokai of about a score of Oo that I watched while they were
feeding, at a time when the Ohias and lobelias were both in flower, one only was seen to visit the former, though
these were attractive enough to other birds. This Oo also visits the flowers of the banana in some of the deep
gulches and valleys, and is said to also eat the fruit. To insect food, when the nectar of the lobelias is abundant, it
pays little attention. Thus of ten shot at the lobelias and dissected not one contained a trace of insect food,
although these were secured on different days and at a very different times of day. On the other hand, the one
mentioned above as visiting the Ohia flowers at this season contained a considerable amount of insect food.
Its call was loud and could be heard at an altitudinal distance of more than 900m. It was described as a
five-syllable owów–owów–ów, the last being the loudest and almost a shriek.
Maui is the least ornithologically collected island of the Hawaiian Islands; as recently as 1973 a new genus
and species of Hawaiian finch, the Po’ouli Melamprosops phaeosoma, was discovered there (Casey & Jacobi
1974) (see p. 352). In 1828 Judd (1880) was accompanied over the mountains of Maui by natives who had
been trained to catch ’o’os, and evidently knew them well. On June 9, 1901, Henshaw (1902) heard and saw
a male ’o’o n orth-east of Olinda on Maui, at an elevation of 900m. No further observations were made until
the 1970s, when several sightings of ’o’os were reported by untrained observers in the forests of eastern
Maui.
In 1973, J. D. Jacobi (pers. comm. to S. R. Sabo) heard calls like those of an ’o’o, but did not see the bird. In
May 1980, Sabo and J. M. Scott saw what they thought was an ’o’o, and on May 10, 1981, Sabo got good
views of a bird which he positively identified as Bishop’s ’O’o (i.e. the same species as occurred on Molokai)
in the Ko’olan Forest Reserve (Sabo 1982).
There is a little scepticism about the recent reported sightings, but there is no doubt that an ’o’o occurred
on Maui until recent times; subfossil remains have been discovered that confirm its existence (James &
Olson 1991). Whether it was indeed conspecific with the Molokai bird, or a different species, will never be
known, as extensive field surveys undertaken in recent years have failed to find any evidence of the birds.
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Mohoidae
This large, attractive member of the Mohoidae is known from just four skins, the last taken by the collector
Mills in 1859 (Munro 1944). It has never been recorded since. It is not certain where Mills obtained his
Kioeas, but many of his other Hawaiian bird specimens came from Olaa, so it is likely that his specimens
came from the same locality. Perkins (1903) doubted this provenance and believed the Kioea to be confined
to the high plateau between the mountains and the upper level of forest, where the type was collected, and
where Peale and Pickering of the United States Exploring Expedition saw it (Peale 1848).
The Kioea was unusual in that it differed markedly from birds in the genus Moho in tail shape and colora-
tion. The suite of anthropological habitat changes and faunal introductions to the Hawaiian Islands were
the reasons for its extinction. Why this species disappeared so rapidly, while the Hawaii ’O’o Moho nobilis
remained common until the end of the 19th century, is a mystery.
Habits
Virtually nothing is known about the habits of this bird. Peale (1848) made a brief comment:
This rare species was obtained on the Island of Hawaii. It is very active, and graceful in its motions, frequents
the woody districts, and is disposed to be musical, having most of the habits of a Meliphaga [Honeyeater]; they
are generally found about those trees which are in flower.
Kioea Chaetoptila angustipluma
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Perkins (1903) remarked that it frequented the flowers of the Ohia, and had much the same habits as the
’O’o, the call being a loud ‘chuck’.
The genus Chaetoptila was long thought to be monotypic, but the discovery of subfossil remains on Oahu
and Maui have shown that this genus was originally widely distributed throughout the archipelago. James
& Olson (1991) not only collected subfossil remains of Kioea C. angustipluma on Hawaii, but further speci-
mens tentatively assigned to this species were discovered on Oahu and Maui. The former at least probably
represents a distinct species (Olson & James 1982). Most interesting was the discovery of another Chaetoptila
species, which had a much narrower bill, and occurred sympatrically with C. angustipluma on Maui (James
& Olson 1991). Like the ducks and geese, ibises and owls, this is yet another example of a Hawaiian bird
radiation event which hitherto would never have been recognised.
The nominate race of the Golden Swallow T. euchrysea, endemic to Jamaica, was considered common in the
19th century (Gosse 1847), but it gradually declined over the next century. It was rare and localised in the
1890s, and was observed only in Cockpit Country east across the central highlands to the Blue Mountains in
the latter half of the 20th century (Raffaele et al. 1998). The last major roosting site was destroyed in 1987,
with the last confirmed sighting made in 1989. These seems little hope for this bird’s continued survival.
The subspecies T. e. sclateri of Hispaniola is still extant, and although numbers have declined (King 1981)
it remains locally common (BirdLife International 2011). The reason for the disappearance of the Jamaica
population is not clear, but deforestation due to shifting agriculture is the prime reason for the decline of the
Hispaniola race (Stattersfield et al. 1998).
Habits
The swallow occurred in humid montane pine forests and at s ea-level, where it bred in a variety of sites
including old woodpecker holes, natural holes in dead pines and caves. It utilised human dwellings;
Hispaniola birds nest in bauxite mines (Townsend et al. 2008). It tolerated secondary forest, and sometimes
fed over c ane-fields and open, deforested country (Stattersfield et al. 1998). It was insectivorous, and foraged
alone or in small groups, flying close to the ground. The call was described as a soft, disyllabic tchee weet.
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Phylloscopidae
Macrosphenidae (Crombecs)
The crombecs are restricted to Africa. The family is divided into six genera, with 18 species. They are forest
birds that are susceptible to habitat disturbance, although in general they are not threatened. One subspe-
cies may have become extinct in recent times.
Chapin’s Crombec is considered by some authorities to warrant specific status (Sibley & Monroe 1993),
though Fishpool & Collar (2006) retain it as a race of White-browed Crombec S. leucophrys. It is quite distinct,
however, and further analysis is required to substantiate its status. The bird was restricted to the Lendu
Plateau in DR Congo, where three specimens were collected by J. M. Vrydagh in 1942 (Schouteden 1947); it
has not been recorded since. The Lendu Plateau has now been completely deforested (Louette 1989), so the
likelihood of finding an extant population is remote (Fishpool & Collar 2006).
Habits
Little was recorded on the habits of this bird. Vrydagh (1949) recorded them as not uncommon in 1942, and
occurring in gallery forest in dense undergrowth, where they foraged for insects (see Fishpool & Collar
2006).
The Canary Islands Chiffchaff P. canariensis was formerly treated as a subspecies of the Common Chiffchaff
P. collybita, but it is now elevated to the rank of full species on the basis of vocalisations, wing morphology
and mitochondrial DNA analysis (Helbig et al. 1996). The Canary Islands Chiffchaff has been divided into
two subspecies, with the nominate occurring in the western Canary Islands, and an eastern race on
Lanzarote and, doubtfully, Fuerteventura; no specimens were ever taken on Fuerteventura, and its presence
there is based on hearsay evidence only. The Lanzarote Chiffchaff appears to have been rare since its
discovery, and was confined to the Haría Valley (Simms 1985). There it occupied broom thickets in higher
areas with fresh growth; it was insectivorous, foraging in the undergrowth or occasionally hovering in
classic Phylloscopus fashion.
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The exact reasons for its disappearance are not known, but much of its habitat was cleared for agricul-
ture, and this may have been the primary reason for its decline. It was supposedly recorded in 1986 (Clement
1995), but it is more likely it had disappeared before then. There have been no further records.
The Aldabra Brush Warbler was restricted to a tiny part of Gionnet, at the west end of Malabar (Middle)
Island, Aldabra (Prŷs-Jones 1979; see Safford & Hawkins 2013 for a comprehensive review). Prŷs-Jones esti-
mated that its greatest distribution was never more than a 50m wide strip extending for 9km along the coast
– perhaps one of the smallest ranges of any known bird species. However, Safford & Hawkins (2013) suggest
that this was actually an over-estimate, and that it was probably restricted to an area of 4.5 km, making the
distribution even smaller.
An adult female with its nest and three eggs were collected on 11 December 1967; a male was collected a
few days later in the same place (Benson & Penny, 1968). Another specimen was mist-netted and ringed in
April 1974, but otherwise the species was not seen again until Prŷs-Jones’s survey of 1974–1976, when only
five individuals were found – three males and two females (Prŷs–Jones 1979). The females paired with a
male each the following season, but no young were ever found, and after November 1975, only five unmated
males were seen (Hambler et al. 1985). In February 1977, just two males were definitely known to survive,
and by October 1983 this had fallen to one (Collar and Stuart 1985). This was the last record. The species is
now extinct (Safford & Hawkins 2013).
Its disappearance remains something of a mystery, but cats and Black Rats Rattus rattus are found on
Middle Island. Rats are common and undertake considerable nest predation among passerines (Frith 1976),
and it seems likely that they were mainly responsible for the extinction of the Aldabra Brush Warbler. The
last population was restricted to areas where Pandanus palm was abundant, which may have provided
some protection against rats (Safford & Hawkins 2013). A secondary factor may have been the abundance of
the Aladabra Giant Tortoise Aldabrachelys gigantea. Giant tortoises did not tend to occur in the dense vegeta-
tion favoured by the warbler, hence the abundant leaf-litter, but opening up of the scrub by introduced
goats may have drastically altered the situation (Prŷs-Jones 1979; Safford & Hawkins 2013). Goats and giant
tortoises are now absent from potentially suitable remaining habitat on West Malabar (Kennerley & Pearson
2010).
Habits
The warbler occurred in areas of rugged pavé and champignon limestone rock, covered by dense mixed
scrub including stands of Pandanus tectorius, Dracaena reflexa and Pemphis acidula (Prŷs-Jones 1979; Safford &
Hawkins 2013). It was a skulking bird, rarely flying long distances, and entirely insectivorous, feeding on
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Acrocephalidae
spiders, ants, caterpillars, moths and other invertebrates (Benson & Penny 1968), which it obtained by
picking, leap-snatching, and probing in leaf litter (Prŷs-Jones 1979). Breeding coincided with the wet season,
from October to January. A nest with three eggs was found in December 1967, about 0.6m above ground in
the base of a young Pandanus, but other nests were found in Pemphis acidula at 1.6m and in Mystroxylon aethi-
opicum at 3.2m (Benson & Penny 1968). The eggs were pale purplish, spotted with dark brown. The birds
were usually silent. Two types of call were recorded, a short chak or scolding chatter, and a harsh, nasal chirr,
most often used by the female, but always given after one or two chak calls (Benson & Penny 1968; Skerrett
et al. 2001). Males sometimes sang from an open perch.
The Guam Reed Warbler was endemic to Guam, and like the Guam Rufous Fantail Rhipidura rufifrons
uraniae (see p. 276), Guam Flycatcher Myiagra freycineti (see p. 280) and Guam Bridled White-eye Zosterops
conspicillatus conspicillatus (see p. 302), the population was exterminated by the introduced Brown Tree
Snake Boiga irregularis (Reichel et al. 1992; Kennerley & Pearson 2010); it appears to have been restricted in
range, and generally found only in wetland environments (Baker 1951; Reichel et al. 1992; Pratt et al. 1979).
The last confirmed observation was in 1969 in the Agana Swamp, but it was considered fairly common
there in 1967 or 1968 (Reichel et al. 1992); the population must have disappeared extremely rapidly. Searches
in the 1970s and 1980s failed to find any birds (Kennerley & Pearson 2010; Saitoh et al. 2012). Habitat loss
including fires and drainage of the wetlands, pesticide use and additional introduced animals such as rats
and cats all helped contribute to its extinction (Reichel et al. 1992; Kennerley & Pearson 2010).
Habits
The Guam Reed Warbler occurred in wetland and wetland-edge vegetation comprising Tangantangan
Leucaena leucocephala thickets, limestone forest edges, marshes and freshwater wetlands, especially in tall
reeds. It skulked in dense thickets, where the birds were more often heard than seen (Baker 1951; Reichel et
al. 1992). The call was a loud and distinctive chuck or tchack, with the male most often singing from an
exposed perch (Kennerley & Pearson 2010). The song was maintained for long periods and it was loud,
varied and complex.
The two known specimens of the Astrolabe Warbler remained undescribed until Holyoak & Thibault
(1978b) recognised them as distinct. The specimens were collected during the expedition commanded by
Captain Dumont d‘Urville aboard the two ships Astrolabe and Zélée, which visited the South Pacific in 1838–
39. The type is stated to have come from Mangereva, Gambier Islands, and the other specimen from
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Nouhera (= Nuku Hiva in the Marquesas Islands), but Holyoak & Thibault believed this to be an error. They
considered that the specimens resembled warblers from Micronesia, and were different in structure from
Polynesian species. It is also unlikely that two identical specimens would have been taken on two such
widely separate islands. They therefore decided that both localities were erroneous and that the specimens
were obtained in Micronesia; the Astrolabe visited Losap, Truk, Guam, Yap and Pelelin in this region.
Acrocephalus species of very different appearance are known from Guam and Truk. Holyoak & Thibault
suggested that:
. . . the high volcanic island of Yap, from which no warbler is known, could be the source of the Astrolabe speci-
mens. The first real ornithological exploration of Yap took place many years later when Hartlaub and Finsch
(1872) described some of the endemic land birds and a warbler could have become extinct before then.
They described the bird as a race of the similar Nightingale Warbler Acrocephalus hiwae, which is the only
surviving Acrocephalus in the Marianas, but it continues to decline and is considered Critically Endangered
(Camp et al. 2009; del Hoyo & Collar 2017).
However, mitochondrial DNA and morphological analysis shows that the Astrolobe Warbler may have
come from the Gambier Islands after all (Cibois et al. 2011), with the bird being only rather distantly related
to Nightingale Warbler (and being much closer to Northern Marquesas Warbler A. percernis). Cibois et al.
discuss a number of travellers’ descriptions and lost specimens of reed warblers from the Gambier Islands,
and how the local name komako, referable to reed warblers in the Marquesas, was still used in the Gambiers
in 1934 for a bird that was no longer present.
This warbler probably disappeared from the Gambier Islands some time in the mid-19th century, almost
certainly due to deforestation and the introduction of predators.
Pagan Island once harboured a distinct population of Acrocephalus warbler, but no birds have been recorded
since the 1970s, and it may now be extinct. The birds were almost certainly gone by 1981, when an extensive
survey took place (Reichel et al. 1992). They had been reduced to a very small population during the 1960s,
and appear to have been restricted to the area around the island’s two lakes.
Severe habitat modification was the primary threat to the birds as wetlands were drained for agriculture
and feral livestock destroyed the understorey, essential for their breeding success. Predation from rats and
cats plus an increase in land use by the military all contributed to the bird’s decline. The lake shore vegeta-
tion was almost gone by 1979, and volcanic eruptions in 1981 destroyed much of the remaining woody
vegetation. There seems little chance that this bird survives.
The Aguijan Reed Warbler was first collected on 29 February 1940, when it was considered common
(Yamashina 1940). By the time of the next intensive bird survey organised by the U.S. Fish and Wildlife
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Service in 1982, the population was estimated at between four and 15 birds (Engbring et al. 1986), and by the
mid-1990s it was 1–2 (Marshall et al. 2011). Extensive surveys to locate the birds in 2000, 2001, 2008 and 2009
were unsuccessful (Esselstyn et al. 2003; Camp et al. 2009); the Aguijan Reed Warbler has not been seen since
the mid-1990s and is now probably extinct.
Its disappearance can be attributed to habitat destruction during the Japanese Administration of the
island from 1914–1945, when intensive farming resulted in large-scale vegetation clearance (Reichel et al.
1992). Goats were introduced in the 1950s, which destroyed the understorey, and this may have been a
major factor in the warbler’s decline.
Habits
There are no wetlands on Aguijan, so the warbler was restricted to native forest, where they were particu-
larly associated with dense canopy cover (Engbring et al. 1986), and more recently disturbed areas vegetated
by groves of trees and thickets (Marshall et al. 2011).
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The Tahiti Warbler Acrocephalus caffer is a large, long-billed warbler, of which a number of races occur in the
Society and Marquesas Islands. Holyoak & Thibault (1978b) were the first to realise that the bird from
Huahine was subspecifically distinct from A. c. caffer on adjacent islands. The five specimens of Huahine
Warbler are presumed to have been collected on that island by Andrew Garrett, probably in 1869; his locality
data are considered to be reliable (Gräffe 1873; Holyoak & Thibault 1978b).
The date of the bird’s disappearance is unknown. No warbler was found on Huahine by the Whitney
South Seas Expedition in 1921, nor has one been seen subsequently (Kennerley & Pearson 2010), but rats are
present in great numbers, so it is likely that they would have been serious predators of eggs and chicks.
Habits
Like many of the extinct or endangered Pacific Acrocephalus warblers, little is known about its habits. The
nominate on Tahiti breeds almost exclusively in bamboo thickets and secondary forest in river valleys and
hillsides up to 1,700m (Pratt et al. 1987). Its diet is varied, mainly insects, but it will take crayfish, snails and
nectar (Thibault 1988).
There can be no doubt that an Acrocephalus warbler once occurred on Raiatea. It was first collected in 1773
by the Forsters on Cook’s second voyage, when the father, J. R. Forster, wrote a description apparently
meant to refer to a warbler painted by his son, Georg. There is some confusion as to the provenance,
however, as the bird depicted by Georg was said to have been shot on Tahiti, whereas J. R. Forster’s descrip-
tion was of a bird collected on Raiatea. There are also slight differences between the description and the
plate. In an attempt to clarify the confusion, Lysaght (1959) stated:
Georg Forster’s bird agrees to some extent with C. caffra caffra (Sparrman) 1786 from Tahiti, but is larger, the
exposed culmen in the drawing being 34mm, whereas in the birds measured by Murphy and Mathews it is only
25–28mm. J. R. Forster states that the bird was eight and a half inches long and the size of a common starling.
His measurements were clearly not taken from this drawing of a bird nine and a quarter inches in length;
furthermore he states that his bird came from Ulietea [Raiatea], an island in the Society group some way from
Tahiti. Murphy and Mathews do not refer to any race from there and it is possible that Forster’s description
concerns a bird which has since become extinct; it was apparently meant to refer to the species painted by Georg
Forster since the description begins with a note on the drawing.
Stresemann (1950) considered that the type of the Tahiti Reed Warbler A. c. caffer was probably collected on
Raiatea, not Tahiti, but Holyoak & Thibault (1984) remark:
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Stresemann (1950) suggested that Sparrman’s type of A. c. caffer . . . was more probably shot at Raiatea than at
Tahiti, but there is no direct evidence for this and the specimen agrees with Tahiti birds.
Georg Forster’s watercolour depicts a bird with a very long bill. It seems probable that this is a n
ow-extinct
race from Raiatea. Nothing more is known about it. Mees (1991) located a unique skin of this species in
Bremen, which was collected by Garrett between 1870 and 1873.
The Millerbird A. familiaris was so named because its principal prey was several species of ‘miller’ moths
Agrotis sp. (Munro 1944), including on Laysan the endemic and similarly extinct Laysan Miller Moth Agrotis
laysanensis (Butler & Usinger 1963). There were two Millerbird races, the nominate on Laysan and A. f. kingi
on nearby Nihoa Island (which is still extant).
Fisher (1906) described the Millerbird as one of the most abundant of the endemic land birds on Laysan,
and found it very active in the cool of the morning or in late afternoon. Bailey (1956) found the Millerbird in
1913, and added that his party were probably the last to see the bird alive, for they were gone when Wetmore
(1925b) visited Laysan in 1923. Rabbits were introduced to Laysan in 1904. They began to destroy the vege-
tation, and by 1920 it had nearly all gone (Bailey 1956). The loss of habitat and associated food sources
probably caused the Millerbirds’ extinction. They also suffered an increase in egg predation from the Laysan
Finch Telespiza cantans and other birds, plus competition for nesting habitat, once the vegetation had been
destroyed.
Habits
Munro (1944) eloquently described the bird:
Greyish-brown on upper parts, wings and tail darker and underparts lighter, the little miller bird slipped about
among the half dried grass almost unseen . . . It was found all over the island where there was vegetation. It
hunted its food among the grass and shrubbery. It came into the laboratory through the window and hunted
moths among the rafters under the roof while we worked . . . Its food seemed to be entirely insects, moths, flies
and small beetles. It swallowed large moths whole, wings and all. It had a rather nice little song, a deep harsh
note a little hard; but it was the end of the breeding season and we may not have heard it at its best. We found a
number of nests. One in the top of a shrub had two eggs. It was three inches in diameter with a hollow in the top
about two inches across and one and a half inches deep, built of grass, down and feathers. We found a number of
nests with either two eggs or two chicks . . . Eggs were b luish-white with brown spots . . . Some of the eggs
instead of being spotted had a black blotch on the large end.
The blotch referred to here is in fact a well-known oological phenomenon, in which spots ‘migrate’ to form
such a blotch or blotches at the large end.
Pycnonotidae (Bulbuls)
The bulbuls are widely distributed and occur across the majority of Africa and tropical Asia. There are
about 24 genera with around 130 species. Medium-sized birds, they occur in a wide range of habitats, espe-
cially rainforest and open-forested areas. In general, bulbuls are adaptable and can tolerate habitat
disturbance, but some forest species are threatened with habitat loss. Bulbuls occur on a number of smaller
Indian Ocean islands and on Madagascar, but only one species is known to have become extinct.
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Habits
The Chatham Islands Fernbird probably inhabited dense vegetation at the edges of swamps and tidal inlets,
where it fed on insects (Oliver 1955; Gill & Martinson 1991; Higgins et al. 2006). Travers (1872) gave the best
description:
I only found this bird on Mangare [sic], where it is not uncommon. Its peculiar habit of hopping from one point
of concealment to another renders it difficult to secure. It has a peculiar whistle, very like that which a man
would use in order to attract the attention of another at some distance, and although I knew that I was alone on
the island, I frequently stopped mechanically on hearing the note of this bird, under the momentary impression
that some other person was whistling to me. It always uses the same cry as Sphenœacus punctatus [South Island
Fernbird Bowdleria punctata]. It is solitary in its habits and appears to live exclusively on insects.
Olson (1990) described its distinctive skeletal morphology, which showed that it had a robust pelvis and
legs, with comparatively short wings and a reduced sternal keel. In other Bowdleria and some other passerine
genera, the feet are used in foraging, especially for removing vegetation and detritus to expose the inverte-
brate prey. The Snares Island Fernbird B. cordata is known to use this technique when foraging in tussock
grass, on the forest floor and in penguin nests (Best 1979). Olson also remarked that the peculiar modified
tail of B. rufescens is reminiscent of that of the Australian Orthonyx logrunners. Their rectrices are similarly
stiff and free of barbs at the tip; the tail is used as a prop while the bird scratches at the ground beneath.
The discovery of the Bismarck Thicketbird extends the range of the genus some 965km north-west from its
nearest congeners on the New Hebrides. The Bismarck bird was collected only once, on December 18, 1958,
when two birds were secured, and after days of searching another pair were briefly seen. There have been
few surveys in the high montane forests of New Britain where it occurred, and being a cryptic, skulking
species best identified by its call, it may still persist. However, recent surveys have failed to find the species
(BirdLife International 2011), and it may be extinct. The reasons for its disappearance are not clear.
Habits
Little is known of the bird. The specimens were taken in montane forest at approximately 1,700m on lime-
stone karst. Its call was a single note and the stomach contents consisted of insects and a snail (Gilliard
1960). Gilliard recalled the fate of the first bird to be collected:
[The native hunter Tesako] was creeping through a clump of bamboo forest when he spied this bird alone on
the papery floor under the bamboos. It hopped to the base of a slender bamboo, where it clung to the vertical shaft
like a wren, then it hopped to a low part of another bamboo and hung sideways facing Tesako, who then shot it.
The rediscovery of the nominate L ong-legged Thicketbird M. r. rufa on Viti Levu on 1974 received much
publicity, as it was previously known from four specimens collected between 1890 and 1894. Recent surveys
have located it in at least five sites, and its status is not currently considered as critical as once believed
(BirdLife International 2011). A subspecies, M. r. cluniei, was discovered on June 10, 1974 on Vanua Levu
(Kinsky 1975). This bird is still known only from the single type specimen, a male collected in the Delanacau
Mountains. It has not been positively seen since, but there was one unconfirmed sighting in 1990. Vanua
Levu has large populations of introduced Indian Mongoose Herpestes auropunctatus and Black Rat Rattus
rattus, which could prove to be a serious threat to the predominantly terrestrial thicketbirds, but the Vitu
Levu population survives alongside these potentially lethal predators. For this reason, the disappearance or
extreme rarity of the Vanua Levu subspecies remains a mystery.
Habits
Little is known about the Vanua Levu subspecies, but on Viti Levu they prefer primary forest with dense
understorey adjacent to streams and creeks, at an elevation between 200m and 800m. They forage close to
the ground and on the forest floor in pairs or family groups, and are unobtrusive and difficult to locate
except by the presence of their loud, melodic song (BirdLife International 2011).
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The White-winged Apalis A. chariessa is divided into two subspecies; nominate A. c. chariessa from the lower
Tana River, Kenya, and A. c. macphersoni from Tanzania, southern Malawi, and northern Mozambique (King
1981). The latter is declining over most of its range due to deforestation, whereas the northern population
has only been recorded once in the 20th century, in 1961 (Britton 1980; Oyugi & Amutete 1999). The forests
of Mitole have been cleared since the 1960s, and unless another population exists elsewhere, it is likely that
the Northern White-winged Apalis is now extinct.
Habits
Little is known about the nominate race, but A. c. macphersoni occurs in tall, dense humid forest, being most
abundant on forest edges (Dinesen et al. 1993). It forages almost exclusively in the high canopy, where it
gleans twigs and leaves for insects.
The Western population of Turner’s Eromemela has a patchy distribution and is known from five localities
in south-east DR Congo, with one record from south-west Uganda (Chapin 1953). It has not been recorded
at either locality since the 1970s, and the Congo range has now been deforested (BirdLife International
2011). No further reports are known and it might well be extinct as a result.
Habits
Little was recorded about the bird’s habits. It was an inhabitant of tropical moist lowland and m
id-altitudinal
montane forests, and preferred the canopy of large trees, especially m id-size and mature species such as
Croton megalocarpus (Otienio et al. 2011).
Zosteropidae (White-eyes)
The white-eyes are small, typically nectarivorous warblers which are usually green, olive and yellow with a
contrasting white eye-ring. They have slender bills and brush-tipped tongues. White-eyes are adaptable
birds, and have colonised a large number of remote islands and archipelagos, principally in the Indian
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Ocean, Gulf of Guinea and the western Pacific. Island endemicity is high. Some island populations are seri-
ously threatened by habitat destruction and introduced mammalian predators.
The Chestnut-flanked White-eye Zosterops semiflava was thought to have been either closely related to or a
subspecies of the Mayotte White-eye Z. mayottensis (Dickinson 2003), especially as it was somewhat similar
in coloration. Using mtDNA, Warren et al. (2006) have shown that Chestnut-flanked White-eye was not
closely related to the Mayotte bird at all, but was instead part of an earlier invasion of white-eyes from Asia
(as opposed to a later one from Africa, from where the Mayotte White-eye and the other Seychelles endemic
white-eye, Z. modestus, were derived).
Edward Newton’s account (1867b) of the discovery of this bird is brief:
While I was skinning . . . Mr. Nevill went out with my gun, and returned shortly with examples of the
‘Tourterelle rouge’, ‘Pigeon hollandais’, and ‘Serin’ . . . The ‘Serin’ proved to be a second new species of
Zosterops, which I have called Z. semiflava. Mr. Nevill saw a flock of some dozen or so, from which he killed a
couple. I was too busy to go out again on this day, so I did not see the species alive.
Newton was told that the Chestnut-flanked White-eye occurred on Silhouette, Praslin and La Digue, but
this was never confirmed. He was sure he saw one on Mahé, and the Marion-Dufresne expedition probably
observed the species on South-east Island (Lionnet 1980, 1984a; Skerrett et al. 2001); otherwise they were
only ever encountered on Marianne. The last specimens were taken in 1892 by Abbott (Lionnet 1984a).
Vesey-Fitzgerald (1940), who resided on the islands for quite a number of years, was unable to find it, or
report any sightings, and it presumably became extinct between 1892 and 1900. The extinction of the
Chestnut-flanked White-eye was due to deforestation, as native forest made way for coconut plantations
(Skerrett et al. 2001), competition with introduced birds, and predation by the Black Rat Rattus rattus.
Nothing was recorded about its habits, but it was probably gregarious, living in small flocks.
The Bridled White-eye is divided into two subspecies; nominate Z. c. conspicillatus on Guam, and Z. c.
saypani, which is sometimes treated as a full species, on Saipan, Tinian and Aguijan, where it is abundant.
The Guam population was equally common before the arrival of the Brown Tree Snake Boiga irregularis in
the 1940s (Seale 1901), after which numbers dramatically crashed. They still occurred in restricted areas in
southern Guam during the 1963–68 bird censuses, but were confined to the cliff regions and associated
forest of the northern plateau by the 1970s (Drahos 2006). Super-typhoon Pamela hit the area on May 19–21,
1976; this devastated the remaining habitat of the white-eye.
A survey in 1981 by the United States Fish and Wildlife Service recorded approximately 2,200 individ-
uals, the white-eye having disappeared from 98% of its original range. By 1982 the bird was restricted to the
Pajon Basin in the extreme north of Guam, where fewer than 50 birds remained. This population was soon
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overcome by the depredations of the Brown Tree Snake, as the last observation was made just a year later in
June 1983. The bird is now extinct (Savidge 1987).
Habits
The Guam Bridled White-eye was widespread over the island and occurred in almost all habitats including
mature limestone forest (Baker 1951), where it foraged in the larger Ficus and Guettarda trees, scrubby
secondary growth, and grasslands and foothills. It also utilised beach strand zones (King 1962) and wetland
swamps. It was predominantly insectivorous, but very occasionally took fruit and nectar, and formed small
flocks while foraging in the upper canopy, making low, chipping contact calls (Jenkins 1983).
There was no territorial behaviour and the birds appeared to nest year-round. The nests were rarely
observed, but Hartert (1898b) found some within 1–2m of the ground, while another was 8 feet (2.4m) up in
a Leucaena leucocephala shrub; it was constructed from fine fibres and rootlets, woven into a hanging basket
(Jenkins 1983).
This bird, once common on the island, was described by Gould (1855) from a specimen collected in 1853 by
John MacGillivray. Extinction apparently occurred after rats escaped from the wreck of the SS Makambo in
1918. Sharland (1929) could not find the bird in 1928, and Hindwood (1940) conducted an enquiry and
searched for it diligently in 1936, but without success.
Habits
Little was recorded about the habits of
this white-eye. It occurred in small
flocks and inhabited lowland forests
and palm glades (Higgins et al. 2006).
In 1913, Bell (in Hindwood 1940) noted
that the ‘Big Grinnells’ occurred in
their thousands and were highly
destructive to fruit and crops, and also
to other birds, whose eggs they
‘sucked’ (predated). They bred during
November and December. Hull (1909)
recorded that the nest was a cup-
shaped structure in a fork, built of
roots and grass lined with finer mate-
rials, and placed among the masses of
fibre clothing the undersides of kentia
palms, or in shrubs overgrown with Robust Silvereye
vines. Zosterops strenua
The eggs, like those of all
white-
eyes, were pale blue, and the
clutch size was two or three. Hull also
found the white-eye in great numbers,
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and remarked that ‘at Lord Howe Island, its powerful song makes music all day long in the palm-glades
and on the wooded hillsides’.
The White-breasted W hite-eye began to decline due to a combination of competition from the Silvereye
Zosterops lateralis, which was introduced in 1904 (North 1904), and the accidental introduction of Black Rats
Rattus rattus in the mid-1940s (Garnet & Cowley 2000), which are serious predators of birds, their eggs and
young. The clearance of forest further impacted the population and it was reduced to just 50 birds by 1962.
In 1986 the establishment of the Norfolk Island National Park was made to protect this bird and other
species, but by this time, the White-breasted White-eye was probably effectively extinct.
Only four were seen in 1978 (Schodde et al. 1983), with a few sightings in the 1980s and 1990s (Moore
1999). An observation of a single bird was made in 2000 (Garnet & Cowley 2000), while a bird seen in 2005 is
the last unconfirmed report (van Balen 2008). Later observations may be misidentifications.
Habits
The bird flourished only in native forest free of invasive weeds, and foraged in the canopy of tall shrubs and
trees (Schodde et al. 1983). The White-breasted White-eye preferred remnant rainforest, with the canopy
dominated by native Hibiscus Lagunaria patersonia, Ironwood Sophora howinsula and other trees, emergent
Norfolk Island Pine Araucaria heterophylla and creepers (Higgins et al. 2006). It also occurred in residents’
gardens in the early years, and Hull (1909) records the birds around his dwelling:
This handsome Silver-eye [sic] is very plentiful in the vicinity of dwellings, and especially favours the
fruit-gardens with its presence. It has a loud and not very musical note when in flocks, and a number of birds
arguing together make a noise similar to a mob of quarrelling house-sparrows. Solitary birds, however, occasion-
ally indulge in a long-sustained liquid song, very pleasing to the ear.
Its diet was varied, comprising fruits, berries, nectar and insects. The breeding season lasted from October
to December, and a cup-shaped nest, constructed almost entirely from dried grasses, was situated in dense
vegetation, including Spanish moss hanging from pine trees (Hull 1909). Between three and six eggs were
laid; these were a uniform pale blue.
The Mukojima White-eye has been variously described as a bulbul, a babbler and an Old World warbler,
and most often included in the honeyeaters, or Meliphagidae. However, it is now clear that it is a member of
the Zosteropidae (van Balen 2008).
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After its discovery by Baron von Kittlitz in 1828 (Kittlitz 1831) on his voyage round the world, it was not
recorded again until 1889 when it was found by Holst (in Seebohm 1890a), who sent Seebohm 12 specimens.
The Mukojima White-eye probably disappeared due to the clearance of almost all of the subtropical forest
on the Ogasawara Islands, and it has not been seen since the early 1940s. The bird was previously thought
to have been confined to the Mukojima Group, but Suzuki & Morioka (2005) have shown that it may also
have inhabited Chichijima. The surviving race, A. f. hahashima, remains common in the Hahajima group to
the south.
Habits
This white-eye formerly inhabited subtropical forest, occurring in large flocks, and was regularly hunted
for food. After this forest type was almost completely destroyed, it adapted to live in low secondary forest,
forest edge, bushes, plantations and gardens (Brazil 1991). It bred from March to June, with one (or usually
two) eggs being laid in tree forks, but occasionally in tree cavities. It was frugivorous, but also took flowers
and probably insects.
Regulidae (Kinglets)
The kinglets were once placed in the family Sylviidae, but have now been given familial status. The
Regulidae contains one genus, with six species distributed in Europe, Asia, North Africa, Madeira, Taiwan,
and North America. They are the smallest passerines, and are characterised by having a distinctive crown
patch, the centre of which can be erected during display. The kinglets are not considered threatened, but one
island subspecies is extinct.
The Ruby-crowned Kinglet R. calendula ranges across North America, breeding as far north as Alaska and
British Columbia and wintering in the southern states and Mexico. In addition to two migratory mainland
races, a sedentary race, R. c. obscurus, formerly occurred on Guadalupe.
This bird was confined to the last surviving cypress grove, at the northern summit of the 1,220m ridge, in
1906. It was thought to be extinct by 1953, but Howell & Cade (1953) discovered five male birds, and
reported on the dire circumstances surrounding the birds’ habitat. The cypress grove comprised old trees
with no regeneration, as all of the seedlings had been eaten by goats (Greenway 1967).
The continued ravages of goats and predation by cats had reduced the population to critically low
numbers by the 1990s. Recent surveys have failed to locate the bird, and it is probably extinct (Martens &
Päckert 2006). This tiny bird had managed to survive the h uman-induced catastrophic events that elimi-
nated almost all of the other Guadalupe land birds a century before, but it too has now become a victim of
human encroachment.
Habits
The Guadalupe Ruby-crowned Kinglet nested in March in large cypress and pine groves at high elevations.
The nests were c up-shaped, well-hidden and hanging or placed on a tree branch. While foraging, the birds
moved along branches in a series of short hops or bursts of flight, constantly active, and with characteristic
wing-flicking (Martens & Päckert 2006). The kinglet was predominantly insectivorous, but also took fruit
and seeds.
Thayer & Bangs (1908) describe the birds discovered by the collector Brown, who poignantly recalled the
song of the Guadalupe Ruby-crowned Kinglet, the only sound resonating from the Guadalupe forest:
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Mr. Brown found the species restricted to the large cypress wood, and in small numbers, noting in all about
thirty-five individuals. The breeding season was passed, as young on the wing were seen. The males, however,
were still singing, and Mr. Brown characterizes the song as ‘indescribably sweet; in fact I have seldom heard its
equal, and given as it always is in the silent gloomy depths of the cypress woods of Guadaloupe, it is ever to be
remembered’.
Troglodytidae (Wrens)
The wrens have their greatest diversity in the New World. They are generally small, inconspicuous birds
that skulk in dense vegetation. The family is complex and some generic relationships are in need of further
study. At present, the Troglodytidae is divided into 20 genera with about 80 species. They are widely distrib-
uted and occur on some islands and island archipelagos. Island populations have suffered from
anthropogenic changes, and three subspecies have become extinct. Deforestation has threatened a number
of continental species and, as a result, one may have disappeared.
The case of the San Benedicto Rock Wren is one of the few documented avian extinctions in which a natural
catastrophe was entirely to blame. According to Townsend (1890) and Anthony (1898b), the San Benedicto
Rock Wren and the regional race of Common Raven Corvus corax clarionensis were the only resident land
birds on the island, with the latter author describing the wren as ‘abundant’. It is said to have bred in May,
and was quite common on the volcanic island until the eruption of a new vent, El Boquerón, on August 1,
1952.
Just two weeks later, the entire island and offshore pinnacles were covered in ash and pumice, to a depth
of up to 3m. More eruptions took place in November and December, but by March 9, 1953, the volcano was
dormant (Brattstrom & Howell 1956). However, a new 300m high crater, Volcán Bárcena, had completely
obliterated the wren’s habitat. Nothing was recorded about the San Benedicto Rock Wren’s habits.
Bewick’s Wren T. bewickii ranges over much of the US and Mexico, including some offshore islands, and is
very variable, with nearly twenty geographical races being recognised. Howell (1917) found the San
Clemente race abundant in 1917, as did Grinnell & Miller (1944), who noted that it bred in dense thickets of
thorn scrub and cactus. However, it had disappeared by 1968 as a result of habitat destruction by goats and
sheep (Jones & Diamond 1976).
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A single Bewick’s Wren seen on the island in 1973 and captured and photographed in 1974 did not belong
to this race; it was evidently a vagrant of another subspecies.
The last sight record of this race of Bewick’s Wren T. bewickii was by Henry Kaeding on March 22, 1897
(Kaeding 1905), the last specimens having been collected by Anthony in May 1892. Several subsequent
surveys have failed to find it (Greenway 1967). It was rare in 1875 when the type was collected, and the
population may never have been large. Shortly after Palmer (in Ridgway 1876b) collected the first spec-
imen, goats were introduced to Guadalupe. They virtually eliminated the sagebrush, the yellow blossoms
of which were a favourite food (Bryant 1887); unfortunately this was a vital foraging habitat for the wrens.
Kaeding (1905) stated categorically that the bird’s extinction was due to destruction of the underbrush. He
found the birds only in a very small area of dying pine trees Pinus radiata binata at the northern edge of the
915m high ridge.
In May 1892, Anthony and Streator (Anthony 1901) collected a single specimen in this grove and two in
open country on the hillside below. Kaeding (1905) recorded them as nearly extinct by March 1897, and they
were not positively seen again.
Only Palmer, Bryant, Kaeding and Anthony ever saw the bird alive. The sagebrush had been completely
destroyed by 1906, and the bird was definitely extinct by that date (Thayer & Bangs 1908). Greenway (1967)
thought that cats might have been partly to blame for its demise, but by his admission these animals had no
effect on the numbers of the Guadalupe Rock Wren Salpinctes obsoletus guadeloupensis, which is tame and
still common on the island. Over-collecting may have also been a reason for the bird’s decline.
Habits
Palmer (in Ridgway 1876b) reported that the bird lived in scrub, rarely in trees, and fed on a small black
insect which occurred on the blossoms of the white-leafed sagebrush Senecio palmeri which covered the
mountainside. He also mentioned that:
Their motions are very quick; their general habits restless, impatient and shy. Their almost incessant activity,
together with shyness, renders them difficult to secure.
The nest and eggs were never recorded. Bryant (1887) noted that they took refuge among the dry branches
of fallen trees:
The birds were timid rather than shy, being alarmed by the crushing of dry branches as I worked my way amid
dense windfalls of pines; where they were found, they fled into the thickest parts. When all was quiet they would
cautiously approach until within a few feet of me, seemingly prompted by curiosity. A frightened female uttered
a few ‘twit twits’ of alarm, but with this exception they were utterly silent.
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When this race was described in 1866 the birds were said to be common, but by 1878 Ober (in Lawrence
1878c) found that the birds were scarce on Martinique:
An inhabitant of the woods; I have not seen it near houses or sugar mills, only in the forests of the hills, and
along the borders of streams where the bushes are thick.
Nothing more than this is known of the wren’s habits.
Five other subspecies of House Wren occur in the West Indies. The St. Vincent T. a. musicus and St. Lucia
Wren T. a. mesoleucus Wrens are rare; the Guadeloupe House Wren T. a. guadeloupensis was rediscovered in
1969 (Kroodsma & Brewer 2005), but is now considered probably extinct (see below); the form on Dominica,
T. a. rufescens, is common, and that on Grenada, T. a. grenadensis, relatively so. The extinction of the
Martinique race is all the more puzzling as each of these islands seems to have been subjected to the same
pressures of human population and introduced predators. On St. Lucia, the range of the predatory native
boa Constrictor orophias coincides with that of the wren and other ground-nesting birds, and apparently
controls introduced mongooses (Greenway 1967). But the snake does not occur on Grenada, where the wren
is still common. This interesting ecological relationship awaits further analysis.
The Guadeloupe House Wren has had a gloomy, checkered history. It was thought to be extinct in 1914, the
last specimen being taken near the village of Sainte Rose, Basse Terre (Bond 1950), but three territorial males
were seen and their voices recorded in 1969 at Cacao, northern Basse Terre; another small colony was
observed and their songs recorded in 1973 at Basse Terre (Barlow 1978; Kroodsma & Brewer 2005). Barlow
continued searching for wrens in 1973 and located two individuals in ‘dense bush’ on 25 May, but upon
returning the next day, a bulldozer had extended a road 350m through the bird’s territories. Barlow saw
another pair at the end of the new
road, and the following day, using
play-back of the wren’s song,
observed a few birds off the old
and new roads. There have been no
more reports since then, and the
species might now be extinct
(Levesque & Mathurin 2008). If it
does survive, the numbers must be
critically low. Deforestation
appears to be at least partly respon-
sible for the Guadeloupe House
Wren’s disappearance.
Habits
Little was recorded. Barlow saw
birds in cut-over forest with an
understorey of tree ferns and
deciduous scrub. Ober (in
Lawrence 1878d) observed the
wrens ‘only in the second growth
of the hills and in wood in the flat Guadeloupe House Wren
portion of the island’. Males were Troglodytes aedon guadeloupensis
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singing from territories in May, and a pair was seen in the same month (Barlow 1978), so breeding probably
took place in May to June.
The Scaly-breasted Thrasher M. fuscus occurs in the Lesser Antilles, where five subspecies have been
described. They are generally considered not threatened but some races are in decline. The Barbados race
was first collected in 1903, again in 1905, observed in 1933, and four were netted in 1985 (Arendt 2006). It
was considered very rare in 1990 (Evans 1990), and Buden (1992a, b) failed to locate it in 1991.
The race was described as distinct only in 1993 (Buden 1993a). Feilden (1889) found no trace of the bird in
1889, and concluded that it was no more than a straggler to the island. Regardless of its subspecific status, it
has not been definitely located since 1990. Barbados has suffered almost complete deforestation and little
regeneration has taken place. The same situation affected the Antiguan population, but secondary forest
growth was utilised by the thrashers. With virtually no suitable habitat left, the Barbados race is now prob-
ably extinct.
Habits
Little was recorded about the Barbados bird. It was supposedly restricted to dry coastal scrub and possibly
mangroves on the island, but like other Scaly-breasted Thrasher subspecies it probably occurred in a wide
variety of habitats. The diet comprised fruit, insects, and lizards, but it was also a predator of other birds’
eggs and young (Arendt 2006).
This species was endemic to Cozumel Island, where it was common, but a series of devastating hurricanes
seriously depleted the population. Hurricane Gilbert hit the island in September 1988, after which only a
few sight records were made, the last of which was in 1995. The bird was then thought to be extinct
(Macouzet & Escalante Pliego 2001). However, it did persist, as two birds were seen in June 2004. Two
further hurricanes, Emily and Wilma, hit Cozumel in 2005 (Curry et al. 2006), and the bird has not been seen
since, despite an extensive survey in December 2006. Unconfirmed sightings in 2007 appear to be based on
misidentification, and the Cozumel Thrasher might now be extinct. The idiotic introduction of the
bird-predating Boa Constrictor Boa constrictor in 1971 by a film company (Martínez-Morales & Cuarón 1999)
and feral cats must have also impacted on the population.
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Habits
This thrasher was a skulking bird that foraged on or near to the ground, which made it vulnerable to intro-
duced predators. Recent observations occurred in semi-deciduous and deciduous forest lacking scrubby
understorey (Macouzet & Escalante Pliego 2001), but it formerly inhabited scrubby woodland and thick
undergrowth bordering open fields, and forest edge (Howell & Webb 1995). It had a complex, scratchy,
warbling song, often singing from conspicuous perches, and the breeding season is recorded as being from
May to July (BirdLife International 2011).
Sturnidae (Starlings)
The starlings are a wide-ranging group of birds restricted to the Old World that has been divided into 30
genera. They are extremely varied in colour and morphology, and some species are strikingly sexually
dichromatic. Their greatest diversity lies in the Indo-Malayan and Afrotropical regions, but some genera,
e.g. Aplonis, have populated a number of remote Pacific islands and archipelagos. Two genera once occurred
on the Mascarenes in the Indian Ocean. The island populations in particular have declined due to human
interference, and a number of species are now extinct.
The Réunion Crested Starling was discovered in 1669. Just a few years later, Dubois (1674) in 1671–72 gave
the first account:
Huppes or Callendres with a white tuft on the head, the rest of the plumage white and grey, a long beak and feet
like a raptor; they are a little smaller than baby pigeons. It’s another good game [bird] when fat.
A number of specimens were sent to Europe during the 1770s, where its outward appearance caused its
taxonomic status to be in doubt. It was considered to be a relative of the Hoopoe Upupa epops, but Murie
(1874) examined its osteology in great detail, and confirmed its relationship to the starlings. A recent mito-
chondrial DNA analysis (Zuccon et al. 2008) has shown that Fregilupus is an isolated lineage, only distantly
related to some South-east Asian Sturnus starlings, although its origins lie in South-east Asia (Cheke &
Hume 2008; Hume 2014b).
The bird was evidently common and adaptable on Réunion, and out-survived all of the other large bird
species by nearly a century and a half. However, at the beginning of the 19th century, as more and more of
Réunion’s forests were turned over to agriculture, the starling was persecuted for the damage it did to
crops. Levaillant (1807), citing information from an anonymous Réunion resident, makes note of this:
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[the Réunion Crested Starling] lives in large flocks in Bourbon [Réunion] where it frequents humid places
and marshes, and causes big damage to coffee trees.
Cordemoy’s testimony
It must have still been reasonably common until the 1840s, as new specimens were relatively easy to obtain;
no less than seven were sent to Italian museums in 1844 (Legendre 1929). Eugène Jacob de Cordemoy wrote
to Manders (1911), and left the last detailed note about the starling:
I have known the bird you ask me about since childhood, namely the Fregilupus varius . . . which has in fact
entirely disappeared . . . When I was a boy this bird lived in the forests of the interior of the island and never set
foot nor wing in towns or inhabited places. It remained faithful to the forests where it bred, which it enlivened
with its clear notes. I used to hunt it then at an age when one is pitiless. I can see it now, a little larger than the
white blackbird [Réunion Cuckoo-shrike Coracina newtoni], with a white crest on the head in the case of the
male, the wings a blackish-grey on the upper surface, the beak and feet yellowish. By no means shy, it was not
frightened even by the sound of firearms, and after a regular slaughter one went off with dozens of these poor
victims in one’s g ame-bag.
After ten years spent in Paris I did not find a single one in the forests where formerly they flew about in
flocks. All ruthlessly destroyed. I shall never forgive myself for the part, slight though it was, which I took in the
matter. I lost my taste for sport and the best bag would not tempt me . . . We will now consider the feeding habits
of this bird. Having raised several in the aviary, I can risk talking about it though I never saw one feeding in the
wild state. In my aviary its food consisted of bananas, potatoes, and choux-choux [Chayote Sechium edule,
boiled]. But when left to its own instincts, it must, like the other winged denizens of the forest, have eaten
insects as is done by its companion in the forests, the Bourbon Blackbird . . . [Réunion Bulbul Hypsipetes
borbonicus], and as is the habit of most fruit-eating birds.
Cheke (1987) examined the literature and concluded that the time Cordemoy was referring to was the late
1840s or early 1850s, and that the bird was still common at that time. It had survived alongside rats, cats and
a host of anthropogenic changes, yet it disappeared within the following decade; it was certainly gone by
1860.
Brasil (1912) suggested that disease or a parasite was the culprit, which may have been contracted from
introduced Common Mynas Acroditheres tristris, though these birds had been living on Réunion on and off
since 1759 (Cheke & Hume 2008), but this does not appear likely. More damaging was severe deforestation,
which accelerated from the 1830s, especially as coffee plantations began to give way to sugar cane. The
emancipation of the slave population from 1848 also resulted in severe degradation of the comparatively
untouched mountain cirques.
However, further research has shown that the Réunion Crested Starling was in steep decline as early as
1817, by which time it was already restricted in distribution (Hume 2014b). The bird was considered a pest
in the early 1800s, and, being confiding and easily approached, it is likely that over-hunting finished off the
birds.
Habits
The Réunion Crested Starling had strong jaw musculature for gaping, robust pelvic elements, and large feet
and claws, which suggests that it was an opportunistic forager in all types of forest, and clearly capable of
utilising the forest understorey and canopy as well as ground substrates (Hume 2014b). It was omnivorous,
as are most starlings, feeding on fruit, seeds and animal matter. It probably nested in tree cavities. The
Réunion Crested Starling was not only able to survive in captivity, but also the journey by boat to Mauritius,
as Desjardins (in Milne-Edwards & Oustalet 1893; Hume & Walters 2012) received four live birds from his
friend Sauzier in May 1835 and noted that they would ‘eat anything’, and that two escaped into the
Mauritian forest and might populate it. Vinson (1877) provided a little extra information about the habits of
the bird (translation from Hume 2014b):
Now these daughters of the wood, when they were numerous, flew in flocks and went thus in the rain forests,
while deviating little from one another, as good companions or as nymphs taking a bath: they lived on berries,
seeds and insects, and the créoles, disgusted by the latter fact, held them for an impure game. Sometimes, coming
from the woods to the littoral [coast], always flying and leaping from tree to tree, branch to branch, they often
alighted in swarms on coffee trees in bloom, and there was in the past the testimony of an inhabitant of the Island
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of Bourbon, said the naturalist Levaillant, that they caused big damage in coffee trees by making the flowers fall
prematurely. But it is not the white flowers of coffee that the hoopoes were searching for and thus behaving so, it
was for the caterpillars and insects that devoured them; and in this they made an important service to the silvi-
culture of the Island of Bourbon and the rich coffee plantations, with which this land was then covered, the
golden age of the country!
These observations suggest that the Réunion Crested Starling undertook vertical migration on mountainous
Réunion, descending to the lowlands from March to November, and returning to the mountain forests to
breed during the austral summer (November–June) (Hume 2014b). The spectacular crest was thought to
have been rigid, but Vinson (1877) gave the following information confirming its erectile capabilities (trans-
lation from Hume 2014b):
But what especially is a marvel of simplicity and elegance is the beautiful white crest composed of feathers of
various lengths but higher in the middle, very flexible, barbs disunited, mobile, which curl forward as in a crest
of a cockatoo. When anxious, happy, surprised or angry, the bird erects them to his liking.
The discovery of subfossil remains of a large starling on Rodrigues by Rev. H. H. Slater during the Transit of
Venus Expedition (Günther & Newton 1874) added yet another extinct bird to the growing list of lost
Mascarene species. Günther & Newton noted that the skull of this starling was very similar to that of the
Réunion Crested Starling Fregilupus varius, but it differed in several features, particularly in having a more
developed supraoccipital ridge. This suggests that the neck muscles were stronger, as were those of the bill.
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The upper bill is shorter and less curved, less slender at the tip than in Fregilupus, and the lower bill is
stouter. These noticeable differences warrant generic separation. Günther & Newton remarked that:
These features indicate that Necropsar had to use greater force than Fregilupus in searching and perhaps
digging for its food, and that it possessed in an eminent degree the capability found in several species of Starlings
and Crows, of removing objects or forcing an entrance in their search for food, by first inserting their wedge-
shaped bill, and then opening the mandibles. The greater strength of the bill would also be fully accounted for, if
Necropsar be the bird described in the old manuscript previously mentioned as feeding on eggs and dead
tortoises.
Tafforet’s observations
The manuscript in question is one by Julien Tafforet, a Réunionese mariner who was accidently marooned
on Rodrigues for nine months in 1725–26 (North-Coombes 1971). Tafforet had been sent to survey the island
to report its potential for French occupation, and he was an astute and careful observer. Unlike François
Leguat (1708), who had no boat and was unable to reach the lagoon islets, which in some cases are quite
large and were originally forested, Tafforet visited them all. On Île Gombrani, one of the large southern
islets, he made the only known observation of the Rodrigues Starling (translation from Hume (2014b), taken
from the original manuscript):
There is a small bird that is not very widespread, because it is not on the mainland; one sees it on the islet du Mât
[= Île Gombrani], which is south of the main island; and I believe that it is on this island because of the birds of
prey on the mainland, and also to live with more facility on the eggs of the fishing birds which lay there, because
they do not eat anything other than eggs or a few turtles dead of hunger that they know well enough to tear.
These birds are a little larger than a blackbird [referring to the extant endemic bulbul, Hypsipetes borbonicus]
and have a white plumage, a part of the wings and tail black, a yellow beak as well as the legs, and have a
wonderful song; I say a song, though they have several, and all different, and each is most beautiful. We fed some
with finely-chopped cooked meat that they ate in preference to seeds of forest trees.
Habits
Little is known, but the robust limbs and strong gaping jaws suggest that it was a terrestrial foraging species.
It may have preyed on the diverse snail and invertebrate fauna of the island, as well as scavenged among
the seabird colonies and tortoise and turtle breeding areas.
Réunion and Rodrigues once harboured endemic starlings that are represented by physical remains and
early accounts. However, no starling was ever reported from Mauritius, which is situated between the other
two Mascarene islands. Around the turn of the 20th century, the amateur natural historian Etienne Thirioux
collected a large number of subfossil remains from caves and rock overhangs around the Moka range in
central Mauritius; Hume (2014b) identified some of these as belonging to a new genus and species of star-
ling. It differed from Fregilupus and Necropsar in the great development of the processus extensorius of the
mandible, which would have made the bill extremely strong, and in characters of the post-cranial skeleton.
As the Mauritius Starling was not mentioned during the Dutch occupation of the island from 1598 to
1710, it may have disappeared before 1598. The Black Rat Rattus rattus is now known to have arrived on
Mauritius during the 14th century (Hume 2013, 2014a), so it is likely that the ground-inhabiting Mauritius
Starling was quickly exterminated by these capable predators.
Habits
The Mauritius Starling occurred in the semi-dry evergreen forests of the island, and probably nested in tree
hollows. It was a powerful gaping species, and probably foraged on the ground. It had a bill similar in
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structure to that of the Eurasian Starling Sturnus vulgaris, which inserts its bill into soft substrates and forces
it open to reveal food items. Like the Rodrigues Starling Necropsar rodericanus, the Mauritian bird was
predominantly carnivorous, possibly feeding on small geckos, skinks and invertebrates, raiding the nests of
forest birds, and seasonally scavenging among turtle, tortoise and seabird breeding colonies (Hume 2014b).
Both races of the Tasman Starling A. fusca – the nominate on Norfolk Island and A. f. hulliana on Lord Howe
– are now extinct. Gould’s (1836b) description of the Norfolk Island bird is brief, and erroneously states that
the type was collected by Captain Sturt on the Murrumbidgee River, on his expedition to the interior of
Australia. The specimen was apparently presented by Sturt to the Zoological Society. It is unclear how
Gould’s misunderstanding regarding the place of origin of the specimen arose.
The starling was last observed in 1923 (Garnett 1993) and disappeared shortly after. Rats cannot be
blamed as they were for the extinction of the Lord Howe race (see below), but the Norfolk Island birds had
a preference for fruiting crops, and they were persecuted accordingly. Hull (1909) described them attacking
bananas:
It is a bold and noisy marauder, creating havoc amongst the banana plantations and orchards. Its soft, slaty-grey
plumage, darker in the male than in the female, is somewhat at variance with its bright orange-red eyes; and its
assertive manner, attitudes, and loud challenging notes are not in keeping with its sober coat. I have often
watched a pair attacking a bunch of bananas hanging to ripen under the verandah of the house where I was
staying on Norfolk Island. The male would utter a few calls from an adjacent p ine-tree, and then dart on to the
iron roof, making a great clatter as he alighted. Then, whistling a sharp staccato note at short intervals, he would
drop on the bananas, rip open the ripest, and swallow large pieces of fruit, uttering satisfied notes between
mouthfuls. The female would follow, with less noise and assurance, and in a few minutes the empty banana skin
would only remain. A sudden movement on my part would send the birds back to the p ine-tree where they
scolded for a while, and then returned to scoop out another banana.
It is likely that habitat destruction, along with competition with introduced birds, especially European
Blackbird Turdus merula, Song Thrush Turdus philomelos and Eurasian Starling Sturnus vulgaris (Garnett &
Crowley 2000), were responsible for its extinction.
Habits
The Norfolk Island Starling inhabited rainforest dominated by Norfolk Island Pine Araucaria heterophylla
(Smithers & Disney 1969), but it was also associated with plantations and orchards. They were generally
frugivorous, but also took invertebrates including snails, and may have raided other birds’ nests, as did the
birds on Lord Howe Island (Hindwood 1940). The breeding season was from September to November, and
February to March. Their nest was a slightly open structure made of small twigs and dry grass, placed in the
trunk of a dead tree fern, quite close to the ground. Eggs were b
luish-green, sparsely freckled and blotched
with pale red, concentrated on the larger end (Hull 1909). There was apparently variation in size and shape
of the eggs, and the usual clutch was four, but sometimes five were laid.
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When Bell visited Lord Howe Island in 1913–14 (in Hindwood 1940) he found the starling in ‘thousands’
and remarked that it was highly destructive to fruit and other crops, and to other birds, whose eggs it
‘sucked’ (predated). Like many of Lord Howe Island’s birds, the grounding of the SS Makambo on Ned’s
Beach in June 1918 marked the end for the starling, as Black Rats Rattus rattus escaped and overran the
island. A local resident, Alan McCulloch, recorded the disaster that took place (in Hindwood 1940):
Within two years this paradise of birds has become a wilderness, and the quietness of death reigns where all was
melody. One cannot see how the happy conditions are to be restored. The very few birds remaining are unable to
breed, being either destroyed upon their nest or driven from them by rats, and their eggs eaten. One can scarcely
imagine a greater calamity in the bird world than this tragedy which has overtaken the avifauna of Lord Howe
Island.
It is all too clear why the birds disappeared.
Habits
The starling was known locally as the Cudgimaruk (based on the call note) or Red-eyes (Hindwood 1940).
Its habits were similar to those of the Norfolk Island Starling A. f. fusca, but it apparently preferred to nest in
holes in trees, and higher up than the nominate form. They were forest-dwellers, but would visit gardens,
and foraged in pairs or flocks (Hull 1909).
Edgar Waite, who wrote a journal about a visit to Lord Howe Island in 1902 (in Hutton 1991), described
the starling when it was still numerous:
. . . the commonest bird is the Cujermerook, a kind of starling, which possesses many of the habits of true star-
lings. It is a great pest in the garden, destroying enormous quantities of peaches & other fruits. At the same time
it is the tamest & cheekiest of birds, allowing itself to be caught & feeding out of the hand directly afterwards. I
cannot imagine birds anywhere being more tolerant of man than on Lord Howe Island. The Cujermerook is a
most inquisitive bird; standing beneath a tree where a flock is feeding they will hop to the branches nearest &
crane their necks in a most ludicrous manner. So unsophisticated are they that almost the whole flock may be
caught one at a time & the cries of the bird in the noose only excite their curiosity more and bring reinforcements
from neighbouring fruit trees.
The only known specimens of this bird were collected by Baron von Kittlitz in 1827 (Hume 2002), during his
ornithological tour of the south-west Pacific. He took the first specimen on December 15, another on
December 21 and a juvenile on December 29, in mountain woods near Lelu. A total of six were collected,
although only three are mentioned, after which Kittlitz departed (on December 30).
Finsch (1881) did not find the species on his visit to Kosrae in 1880, and made the following note:
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It no doubt exists on Kuschai [Kosrae], just as it did when Kittlitz visited the island. Nobody has reached the
mountains in the interior since Kittlitz’s time; and it is strictly a mountain bird. When I was on Kuschai the
path from one side of the coast was no longer used by the natives, the reason being the great and rapid decrease in
their number, only a few hundred of them now surviving (I believe not more than three hundred and fifty all
told). Kittlitz only got two specimens, which I saw in St. Petersburg. It is a very distinct species.
The native population of Kosrae was victimised by whaling crews after European discovery of the island,
when supplies and female companionship was obtained by bartering with the chiefs. The first missionary
station was established in 1852, and by 1870 almost the entire population (or what was left of them) had
become converted to Christianity (Ritter & Ritter 1982).
Japanese collectors, and W. F. Coultas on the Whitney South Seas Expedition in the 1930s, searched for
months for the bird and failed to find it (Greenway 1967). It must have disappeared during the intervening
years. There are large populations of rats on the island – this was even the case when Kittlitz first arrived in
1827 (Kittlitz 1835) – which are thought to have escaped from whaling ships. These rats may have preyed on
the birds, but the Micronesian Starling Aplonis opaca, a smaller species, is still quite common in the lowlands,
even though it suffers from rat depredations (Feare & Craig 1998; Hume 2002). It is more likely that the
Kosrae Starling could not tolerate forest disturbance, and disappeared as a result.
Habits
Little is known of the habits of Aplonis corvina. Kittlitz (1832–33 [1832]) found it only in the deepest moun-
tain forest regions, where it was rare, solitary and avoided humans. Kittlitz (1835, translated in Hume 2002)
stated:
This species lives deep in the wooded region in the centre of the island and is just about the only bird occasionally
to be seen here. It eats small animals, larger insects and lizards etc., which it swallows whole. It also seems occa-
sionally to eat fruit, the stones of which have been found in its stomach. It [the stomach] is relatively smaller and
more muscular than the other species. The mating call of this solitary bird is loud and consists of a single, often
repeated tone. The young (probably in their first year) have very different feathers, so that one is led to believe
that they are a completely different sort of bird. They are all over yellowy white with black/brown speckles, espe-
cially on their upper half. The iris is light brown. I would doubt if they belonged to this species if I hadn’t once
shot a black bird that had white feathers still visible from the transition period. Both sexes have the same plumage,
but the female seems smaller, and its black feathers are less shiny, although the males also differ here, probably
due to their age.
Kittlitz (1832–33 [1832]) further stated that it fed on crickets.
Much speculation has surrounded this unique specimen (see Olson 1986c). Sharpe (1890) suggested that it
was the same as Forster’s drawing of Bay Thrush Aplonis (‘Turdus’) ulietensis (see p. 318), and therefore the
type of that species. However, Stresemann (1949) found that this was not so, as the descriptions do not agree
at all. Olson (1986c) examined manuscript notes made by the naturalist Andrew Bloxam, and concluded
that the type of mavornata had been collected on Mauke in the Cook Islands, by Bloxam in 1825. Olson
showed, by reference to a hitherto overlooked manuscript by Bloxam, that he had collected three specimens
on Mauke in that year on his way back from Hawaii. One of these was a starling, which Bloxam named
Sturnus Mautiensis and described as follows:
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The Huahine Starling was the easternmost of the Aplonis starlings in the Pacific. It is known from a single
tarsometatarsus, which is longer than that of the largest extant Aplonis, the Samoan Starling A. atrifusca, but
differed in some characters (Steadman 1989c).
The Fa’ahia subfossil material has an age range between 750 and c.1250 bp, so the Huahine Starling disap-
peared at an early period of Polynesian colonisation of the island, probably due to deforestation,
over-hunting and predation by the Pacific Rat Rattus exulans. It is very likely that a number of other Aplonis
occurred in the Society Islands, the Cook Islands and possibly in the Tuamotu and Marquesas Islands as
well (Steadman 2006a). The widely distributed Polynesian Starling A. tabuensis tolerates anthropogenic
changes quite readily (Feare & Craig 1998), but it appears that the Aplonis island endemics are extremely
susceptible to changes.
Steadman (1989c) suggests (with some authority) that the mysterious Bird of Raiatea ‘Turdus’ ulietensis,
known only from a painting by Georg Forster executed in 1774, is an Aplonis species related to the Huahine
bird. I agree with this hypothesis.
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The Bay Thrush is one of many mysterious birds that was noted just once and disappeared before anything
was recorded about it. It is known only from a painting by Georg Forster, and from descriptions by J. R.
Forster and Latham of a female specimen, now lost, taken during Captain Cook’s second voyage. The spec-
imen was collected on Raiatea in late May 1774. The locals were hostile when Cook called at the island again
in 1777 on his last voyage, so little or no exploration was possible. Certainly no more specimens of the bird
were seen. Andrew Garrett searched for it in 1850, but was unable to find any (Greenway 1967).
The unique specimen eventually became part of the Banksian collection, where Latham (1783) saw it. He
gave a brief description:
Size of the Song Thrush: eight inches and a half. Bill an inch and a quarter, notched at the tip, and of a reddish-
pearl colour: general colour of the plumage rufous brown: quills edged with dusky: tail rounded in shape and
dusky: legs dusky black. Inhabits Ulietea [= Raiatea]. In the collection of Sir Joseph Banks.
Much has been made of Latham’s mention of a notch at the tip of the bill. It was not described by J. R.
Forster, or illustrated in the painting by Georg Forster, so its taxonomic value must be approached with
great caution.
Taxonomic confusion
It was Tristram (1879), Seebohm (1881) and Sharpe (1890, 1906a) who caused much of the confusion
surrounding the bird. Tristram thought that it was closely related to the New Caledonia Thrush ‘Merula
xanthopus’ (nowadays considered a race of Island Thrush Turdus poliocephalus), which may have been the
reason why Henry Seebohm had the bird reconstructed ‘thrush-like’ in his Catalogue of Birds (Seebohm
1881). Forster’s plate was copied by Keulemans for Seebohm’s work, possibly on the instructions of
Seebohm, and was thus biased toward looking like a thrush. This published plate, can, however, be disre-
garded as it bears little or no resemblance to the original.
Sharpe (1890, 1906a) twice suggested that it was the same as the erstwhile Mysterious Starling (now
known as Mauke Starling Aplonis mavornata, see p. 316), but Wiglesworth (1892) noted that the measure-
ments differed, and Kinnear (in Stresemann 1949) showed that there were too many discrepancies between
the starling skin and Forster’s painting for Sharpe to be correct. Olson (1986c) has shown that the prove-
nance of the Mysterious Starling (Mauke) and the Bay Thrush (Raiatea) are totally different.
The affinities of the Bay Thrush are unclear. Forster’s manuscript name for the bird, Turdus badius, was an
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arbitrary one, for he states quite clearly that the bird resembles no known genus, though it has characters in
common with several. Forster does not seem to have believed it to really be a Turdus. Greenway (1967)
thought that the s cale-covered tarsi and forked tongue are also uncharacteristic of thrushes. The cilia on the
tongue seem to suggest the brush-tongue of a honeyeater, and Holyoak (pers. comm) thought that the bird
may well have been one of these birds. However, considering the distribution of endemic species of Aplonis
in the Pacific (e.g. A. mavornata and Huahine Starling A. diluvialis), Steadman (1989c) suggested that the
‘Bay Thrush’ was likely to have been an Aplonis as well, and that attribution is followed here.
Habits
J. R. Forster made the only comments about the bird’s habits. Known as E boona-nou-nou by the locals, it
occurred among thickets in deep valleys. The bird had a soft, fluting voice. Forster also mentions that the
wing was short and the claws on the hind toe were powerful, and twice as large as the rest. This may be
indicative of terrestrial habits.
Turdidae (Thrushes)
The thrushes are widely distributed throughout the world, including remote island archipelagos, and have
been divided into 21 genera with about 136 species. Some of these are under taxonomic review and may
warrant separate familial placement. They are small to medium-sized birds that predominantly forage on
the ground. A number of species are migratory, but most island forms are sedentary. Their terrestrial habits
have made them extremely vulnerable to introduced predators, especially rats, and habitat destruction has
resulted in the probable extinction of two African and South American subspecies. M
osquito-borne diseases
have also contributed to the almost complete extinction of the endemic Myadestes Hawaiian thrushes.
Kittlitz’s Thrush was discovered by Baron F. H. von Kittlitz in June 1828 on Peel Island, now known as
Chichijima, the largest of the Ogasawara Islands. Kittlitz described the bird as common in the coastal
woods, usually keeping close to the ground. He collected four specimens; the bird was never seen again.
In 1830, Peel Island became a port of call for whaling vessels, with the ships being careened onshore. Rats
must have escaped during this time and it appears that Kittlitz’s Thrush, being a probable g round-nester,
succumbed rapidly to these vermin. A small settlement was established on the island, and by 1854 pigs,
goats, cats and dogs were introduced. The able collector P. A. Holst searched for the thrush in 1889, as did
Alan Owston’s Japanese collectors in 1899. Momiyama and Hachisuka also searched between 1920 and
1930, but all were unsuccessful (Greenway 1967). The similarly terrestrial Doctorbird Turdus poliocephalus
vinitinctus (see p. 322) became extinct within a few years of rats invading Lord Howe Island, so it is conceiv-
able that Kittlitz’s Thrush disappeared some time during the 1830s, for the same reason.
Description 25.5–27.5cm (10–10.25in). Resembles Z. h. eichhorni of the Bismarcks, but the wing is much longer, and rump more
rufescent; bill blackish, base of lower bill pale; iris dark brown; legs and feet light horn.
The Russet-tailed Thrush includes four races, present in eastern Australia, New Guinea, the Bismarcks and
on Choiseul in the Solomon Islands (Clement & Hathaway 2000). The Choiseul Russet-tailed Thrush is
known from only one specimen, a female collected on January 13, 1904 by A. S. Meek (Meek also collected
the only known specimens of the Choiseul Ground Pigeon Microgoura meeki; see p. 176). The thrush was
never reported again, despite a recent extensive search (Boseto & Pikacha 2016). However, a juvenile spec-
imen of an indeterminate subspecies of Russet-tailed Thrush Zoothera heinei was recently collected in
lowland forest on neighbouring Santa Isabel Island in the Solomons, and considered to be a range extension
of the species (Joyce et al. 2014). This specimen may represent a new taxon, or it may indicate that the
Choiseul subspecies occurs on at least one other island in the Solomons.
These thrushes are terrestrial birds that forage in leaf-litter for invertebrates; this would have made the
Choiseul birds vulnerable to introduced predators. Therefore the introduction of cats, which became feral
and numerous in the forests, appears to be responsible for exterminating the population, which probably
disappeared some time after 1904. Nothing else is known about it.
The Isle of Pines was formerly home to an endemic race of Cuban Solitaire M. elisabeth, the nominate race of
which race still survives on Cuba. Bangs & Zappey (1905) described the Isle of Pines race as very rare, and
occurring only in the densest forests, where, because of its shy habits, it was very hard to shoot. It had a
long, ringing call which could be heard at a great distance, and was ‘almost startling’ in the still forests. It
occurred in the vicinity of the Paso Piedras (at sea level), which connects the southern portion of the Isle of
Pines with the northern, and was said to occur in other places on the island, but apparently only one spec-
imen was ever collected. The reasons for its disappearance are unknown.
According to Greenway (1967), Bloxam found the Oahu Thrush common on the sides of the hills in 1825,
when he visited Oahu as naturalist on HMS Blonde. Munro (1944) gave the best account of the bird:
It was collected and described in a manuscript by Bloxam, who was naturalist on the HMS Blonde when Lord
Byron brought the bodies of King Liholiho and Queen Kamamalu to Honolulu from England in 1823 [they had
died of measles while on a visit to Britain]. While in the islands Bloxam collected a number of specimens of
Hawaiian birds, and among them was the Oahu Thrush. The bird became extinct, the specimens were lost, and
there is no known specimen existing in the world at this time. Bloxam and his companions spent a night on the
east side of Oahu. After an uncomfortable night they made an early start up the Nuuanu Pali back to Honolulu.
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George Munro (1944) believed that the decline in this bird began in 1923, when Lanai City was built and
human populations increased. Bird diseases, endemic in imported poultry and transmitted by mosquitoes,
were probably responsible for the bird’s extinction. His description of the decline of this bird is moving:
It inhabited all the present forest, frequenting the low trees and under-brush and . . . did not seek the highest
dead trees to broadcast its few call notes which replaced the beautiful song of the other thrushes. From 1911 to
1923 this bird was under my observation as I frequently rode the bridle trails of the forest. It was at that time a
common bird and its call notes could be heard constantly, especially in the north and south ends of the small
Lanai forest. It declined from 1923 when the population of Lanai increased and the town of Lanai City was built.
The people brought bird diseases with their poultry and these, evidently carried by mosquitoes, were fatal to the
native bird population. I watched its decline till 1931. The few times I have been through the Lanai forest since
1931 the thrushes’ notes have been conspicuously absent. There is little or no hope of this bird’s survival as there
is of other native Lanai birds.
Munro said that the bird’s food was berries and insects. Although he searched diligently for nests from 1913
onward, he never found one.
The Molokai Thrush formerly occurred on the central Hawaiian Islands of Lanai, Molokai, Maui and prob-
ably Oahu. It was considered common in the 19th century, but declined rapidly, almost certainly due to
deforestation, the introduction of disease-carrying mosquitoes and perhaps rat predation of eggs and
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young. The montane m osquito-free refugia afforded on Hawaii are not available on Molokai, and pigs have
facilitated mosquito distribution by opening up the most remote forested areas. The oloma’o has not been
officially recorded since 1980, although unconfirmed reports suggest it survived until the end of the decade
(Wakalee & Fancy 1999). It is probably now extinct.
Reports of an oloma’o on Maui in the 1850s are probably genuine, and have been confirmed by the
discovery of subfossil remains (James & Olson 1991). This would have almost certainly been a separate
subspecies. The Maui bird may have survived until the end of the 19th century, as few ornithological
surveys had been undertaken before then to record them. It probably disappeared for the same reasons as
its congeners on other islands.
Habits
The Molokai Thrush was a shy bird, with its presence revealed only by its loud and melodious song. In the
late 19th century it occurred in forests from the lowlands upwards (Wakalee & Fancy 1999), but it became a
bird of the montane forest canopy as its range shrank in the 20th century. It was primarily a frugivore, but
also took insects.
The Kama’o was common in moist forests at sea level as well as mountain forests during the late 19th
century, and was considered to be much more numerous than the smaller Puaiohi (or Small Kaua’i Thrush)
M. palmeri (Perkins 1903). By the early 20th century, the Kama’o was in drastic decline, and by 1928 it was
restricted to montane forest above 1,100m in the Alaka’i Wilderness Preserve. There it continued to decline;
during 1968–1973 its population was estimated to be around 340 individuals. By 1981, the numbers were
down to 20 or so, with the last probable sighting in 1989 (Wakelee & Fancy 1999). Despite a few uncon-
firmed reports, intensive surveys in the 1990s failed to find it. The Kama’o is almost certainly extinct.
Its probable extinction can be blamed on habitat destruction, mosquito-borne diseases, and predation by
and competition with introduced animals. The Puaiohi has managed to survive in small numbers, and has
been successfully reintroduced into areas where it formerly occurred. Unlike the Kama’o, its future seems
secure.
Habits
The Kama’o preferred open canopy Ohia Metrosideros polymorpha and Olapa Cheirodendron sp. forests, with
a dense fern- and e piphyte-rich understorey. It often perched motionless on low branches, where it would
reveal its presence by wing quivering, and by its song (Perkins 1903). The song was varied and complex,
described as a melody of fluting notes, liquid warbles, buzzy trills, and gurgling whistles (Wakelee & Fancy
1999). The call was a harsh braak, with an alternate h igh-pitched note. It fed primarily on fruits and berries,
especially the fruit of the ’ie’ie Freycinetia arborea, and also on insects.
Description 22.9cm (9in). Russet-brown upperparts; slightly more russet on uppertail-coverts; head, nape, lores and ear-coverts
dark brown, wings and tail brown, with a russet o uter-web to most of the wing feathers; underparts dark vinous brown, shading
into brown on chin, upper throat, thighs, centre of abdomen and undertail-coverts; bill yellow; legs, feet and claws pale yellow.
The Doctorbird was common on Lord Howe Island before 1909 (Hull 1909), but by 1913 (Bell in Hindwood
1940) it was in decline due to human interference, and probable predation from cats, dogs and feral pigs.
Goats were also introduced, and would have reduced the areas of suitable habitat by o ver-grazing. The
species disappeared soon after June 1918, when the SS Makombo was shipwrecked off Lord Howe Island,
and Black Rats Rattus rattus overran the island. Being ground-nesting birds, it is not surprising that they
disappeared so quickly.
An unsuccessful attempt was made to introduce the Norfolk Island nominate race, T. p. poliocephalus,
now also extinct, to Lord Howe Island (Greenway 1967; see below). Sixty-eight Norfolk Island birds were
introduced in two transfers, in 1925 and 1935, but they were quickly wiped out by rats (Tennyson &
Martinson 2007).
Habits
The Doctorbird foraged in open paddocks and in forested areas, turning over leaf-litter, and even flipping
over dry cow-pats when searching for invertebrate prey (Hutton 1991). It moved around in large flocks,
gathering together at sunset, and sometimes even frequented the islanders’ homes (Bell in Hindwood 1940).
Hull (1909) was on Lord Howe Island in October 1907, and recorded the bird’s habits in great detail:
A very common and exceedingly tame species peculiar to Lord Howe Island, the ‘Doctor Bird’ is seen everywhere
scratching among the dead leaves with the industry of a barnyard fowl, or perched on the low shrubs, emitting
its sharp whistling chirp. The locals’ name derives from a sharp double knocking sound when it was alarmed.
Though so tame that one can approach quietly within a short distance of a few feet, any sudden movements, or
loud noise, would send it scolding away for a short flight, but it soon stopped to reconnoitre, curiously watching
the intruder. It is also very suspicious, and will desert a nest either when building, with eggs, or even with
young if touched by man.
The nest is a large, loosely built structure, open, shallow, cup-shaped, outwardly of woven fibre from the sheaths
of palm fronds, and dead leaves, lined with dry grasses, placed at the foot of a palm amid the dead fallen fronds,
on top of a stump, or in a mass of intergrown vines a few feet from the ground.
He mentions that two or three eggs were laid; these were pale greenish-grey with reddish markings scat-
tered over the whole surface.
The Norfolk Island Thrush was an adaptable bird, even inhabiting gardens adjacent to remnant rainforest,
but it gradually disappeared as the forests were cleared. Gould (1865) was informed that it had become
scarce around the settlements:
When Norfolk Island was first made a penal settlement, this bird was doubtless very common there; but I have
reason to believe it has now become scarce, having been partially extirpated by the Government officers and
convicts who tenanted this beautiful island for so many years.
A combination of feral cats and rat predation, plus competition with the European Song Thrush T. philo-
melos, and interbreeding with the European Blackbird T. merula (which yielded sterile young), finally caused
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its extinction (Higgins et al. 2006). Although there are reports of it surviving until the late 1970s, the last
confirmed record was in 1975.
Habits
Hull (1909) stated that the habits of the Norfolk Island Thrush were similar to those of the Doctorbird T. p.
vinitinctus on Lord Howe Island, foraging in leaf litter for invertebrates, but it was also partial to the fruit of
the introduced guava. Its nesting preferences, however, were different, as it constructed its nest from
between 2–6m off the ground. It often used the introduced lemon trees. There were two breeding seasons,
one from August to December, the other March to May, with two eggs laid in the former, and four in the
latter.
The Maré Thrush, a race of the widespread Island Thrush T. poliocephalus, has not been recorded since 1939
and is now probably extinct. Its disappearance can be almost certainly attributed to anthropogenic changes
to the island. Virtually nothing was recorded about it, but Layard & Tristram (1879) were informed that it
frequented dense bush, and visited native plantations at certain seasons; it was extremely shy and difficult
to approach. Sharp (1900a) noted that the alarm note was ‘like a Blackbird’s siss siss’.
Bond (1950) described the Grand Cayman Thrush as rare, restricted to the eastern and north-eastern parts
of the island, and last seen in the summer of 1938 by C. B. Lewis (Johnston 1969; Bradley 2000; Bradley &
Rey-Millet 2013). It was the only full species endemic to the Cayman Islands, and had been declining for
some time before the final sighting (Bangs 1916). Bangs had based this information on a collection of birds
made by W. W. Brown, who had shot thirteen Grand Cayman Thrushes between April and July, 1913.
Bird-killer Brown
Wilmot Wood Brown Jr. was a professional collector, and widely acknowledged by ornithologists of his day
for obtaining large numbers of birds, sometimes thousands in a single expedition. For example, in their
study of the critically endangered Worthen’s Sparrow Spizella wortheni, Webster & Orr (1954) named a now
extinct subspecies, S. w. browni, in honour of him for supplying so many bird specimens to museums in the
USA. He supplied skins to many institutions including the US National Museum of Natural History,
Washington, DC, American Museum of Natural History, New York, Field Museum of Natural History,
Chicago, and Museum of Comparative Zoology, Harvard (e.g. Thayer & Bangs 1907a; Bangs 1916). Brown
indiscriminately collected birds wherever he was sent, and had no qualms about shooting large numbers of
critically endangered species, some of which would never be seen again.
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His record of extermination reads like an avian book of doom: Puerto Rican Parakeet Psittacara maugei (p.
185), one taken in 1892 from the last few on Mona Island, not recorded again; Culebra Island Amazon Parrot
Amazona vittata gracilipes (p. 186), three taken in 1889, not recorded again; Guadalupe Flicker Colaptes cafer
rufipileus (p. 214), 12 skins and 6 clutches of eggs taken in 1906 from a population of 40, not recorded again;
Slender-billed Grackle Quiscalus palustris (p. 319), a series taken in 1910, not recorded again; McGregor’s
House Finch Carpodarcus mexicanus mcgregori (p. 315), five specimens taken in 1906, when the species was
recorded as being nearly extinct (Thayer & Bangs 1907); and Grand Cayman Oriole Icterus leucopteryx bairdi
(p. 319), 17 taken in 1911, even when he reported it as ‘one of the rarest birds he had ever hunted for’ (Bangs
1916). Brown covered all of Grand Cayman to obtain Grand Cayman Thrush Turdus ravidus, taking 13 in
1911, including the last specimen ever collected on 8 June, and found the thrush extremely rare and in only
two remote patches of woodland (Bangs 1916). Brown remarked that in the case of the Grand Cayman
Thrush, he was careful to have left enough birds to perpetuate the population (Bangs 1916), a statement
which was soon to become a hollow fallacy!
The ornithologist, T. M. Savage English, who resided on Grand Cayman for three years, wrote a letter to
W. L. Sclater on May 19 1916, then curator at the Bird Room, Natural History Museum, London, complaining
about Brown’s abhorrent activities. This unfortunately was never acted upon:
Dear Mr Sclater
Many thanks for sending me Mr Bangs’ paper on the results of the activities of “Birdkiller” Brown – as the
Cayman Islanders christened him.
After a somewhat stormy interview with the commissioner (since dead) I did manage to get Brown’s permit to
collect “for scientific purposes” limited to one district (particularly disliked by the commissioner) in the I. of Gd.
Cayman – some miles to the E. of my property here – but he had a free hand as to numbers and was said to have
taken “thousands” of skins.
Is it not time that such a thing as the practical extermination of Mimocichla [Turdus] ravida – simply to put
money into the pockets of a foreigner – should be made impossible in any British colony.
I have neither made nor tried to make one halfpenny from the birds of Cayman, nor am I responsible for the death
of a single one of them, even as an egg, so I think I may express myself freely on the subject.
The Grand Cayman Thrush appears to have bred only within dry forest on limestone (Bradley 2000),
much of which was destroyed during the 19th century, hence the bird’s rarity, but it also occurred in
mangrove (Bradley & Rey-Millet 2013). Bunyon Whittaker (in Bradley 2000; see below) reported that he
never saw it again after the 1932 hurricane, but C. B. Lewis observed it for the last time between Battle Hill
and Winter’s Land at the east end of the island (the present Furtherland Farm area) in the summer of 1938,
when he led a part of naturalists during the Oxford Expedition (Johnston 1969; Bradley 2000).
Habits
Johnston (1969) interviewed the oldest resident on Grand Cayman, then aged 72, who provided informa-
tion on the bird from his youth. Johnston wrote:
At that time the thrushes were, according to Mr. Whittaker, conspicuous, noisy, and common where timber was
being cut about ¾ mile inland from Northside. This area, known locally as ‘mountainous cliff’ and “bullrush
walk,” contains the highest ridges on the island, some 60 feet above sea level, and was originally heavily wooded
with old, large trees of mahogany Swietenia mahugoni, cedar Cedrela odorata, ironwood Gymnunthes lucida,
and red birch Bursera timaruba . . . Many of the large trees have now been logged out. Mr. Whittaker believed
the thrushes preferred the densest woods, not occurring in clearings.
Savage English (1916) recorded an individual that occurred in a tangle of dense knife-edged coral-rock,
swamp and mangroves, with patches here and there of the poisonous manchineel tree, and of climbing
cactus. He also published the only record of the thrush’s vocalisations, and stated:
Its song was very subdued, recalling the warble of a Budgerigar. Its manners and general appearance, apart from
its colour, were not unlike those of a blackbird, and the second time it showed itself it flew across the road, giving
a ‘thrush’ chatter as it flew. This, like its song, was only just audible.
D. W. Johnston searched for the bird between 1965 and 1967; Patricia Bradley has been searching for it since
1982. It is now certainly extinct.
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Pieper (1985) briefly mentioned subfossil remains of a number of bird species that once occurred on Madeira
and Porto Santo. He identified three species of extinct, flightless rails, a flightless Coturnix quail, and a
thrush Turdus sp. Pieper compared the tarsometatarsus of the Madeira Thrush with that of the Eurasian
Blackbird T. merula, and noted that the Madeira bird was larger and had greatly lengthened hindlimbs.
Bowdich (1825) was the only person to have possibly recorded the Madeira Thrush when it was still extant.
He shot an individual (now lost) on Porto Santo in 1823, and remarked that:
The back and belly are brown, with patches of yellow, the wings and tail brown; the beak is strong, and of
a brown colour, except the first half of the lower mandible, which is yellow.
It is clearly evident from the long hindlimbs that the Madeira Thrush was a ground-inhabiting species,
and foraged in the laurel forests that formerly covered the islands. It may have nested on or close to the
ground, and would have been extremely vulnerable to introduced mammals, as well as to deforestation.
House mice Mus musculus were introduced by the first human colonists around 1000 AD (Rando et al. 2014)
and would have reached plague proportions a short time later. This pest, along with rats and cats, would
have been a serious predator and competitor. If Bowdich did encounter the thrush in 1823, it had certainly
disappeared by the time Lowe (1853) surveyed Madeira and Porto Santo in the early 1850s.
Ploceidae (Fodies)
The fodies are small relatives of the weaver birds, and are endemic to Indian Ocean islands. They occur
in Madagascar, the Seychelles including Aldabra, the Comoros, and the Mascarene Islands. Fodies are
generally forest, woodland or scrub inhabitants, but one species, the Madagascar Fody Foudia madagas-
cariensis, is extremely adaptable and has been introduced to almost all of the smaller Indian Ocean
islands, where it occurs in both native forest and around human settlements. Fodies are vulnerable to rat
predation, and coupled with forest destruction, the continued existence of some island populations
remains in doubt.
The Madagascar Fody F. madagascariensis has now been introduced to a number of other islands, where it
sometimes competes with native species. Several races of the Red-headed Fody F. eminentissima inhabit the
forests of Eastern Madagascar, the Comoros and Aldabra; the Mauritius Fody F. rubra is confined to
Mauritius where it is extremely rare; a yellow species F. flavicans occurs on Rodrigues; and there is a
dull-yellow bird F. sechellarum on the Seychelles (Moreau 1960). In addition, a species formerly occurred on
Réunion.
Cheke & Rocamora (2014) suggest that a second species of fody once inhabited the Seychelles, alongside
the yellow F. seychellarum. It was described as having a red head and throat, with the rest of the plumage
brown. If the bird was an endemic, it disappeared shortly after settlement by humans in the late 1700s, most
probably a victim of introduced black rats Rattus rattus.
The Réunion Fody was first mentioned by Gabriel Dellon (1685), who described their inherent tameness
when faced with human habitation:
While the sparrows are no larger in Mascareigne [=Réunion] than in other countries, their numbers render
them inconvenient. They strip sown fields and the houses are full of them, as ours are with flies. One often sees
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them fall into pots and platters, and burning their wings on
fires, which are lit out of doors, the sun providing enough heat Réunion Fody
for [even] the coolest houses. Foudia delloni
Dubois (1674) in 1671–72 also noted that the vast numbers of
‘sparrows’ were causing damage to the inhabitants’ crops:
There are, besides, in the Isle a quantity of other birds, which
wou’d take too long to describe, contenting myself with naming
the principal, & particularly the sparrows, which here are so
thick, and in such numbers that they bring great damage to the
island, eating a large part of the cereal that is planted, without
it being possible to destroy them because there are too many.
Several cereal harvests would be made during the year if it were
not for these sparrows, because of which one is restricted to a
single harvest, in the period they go nesting in the mountains.
These sparrows have plumage like those in Europe, except that
the males, when breeding, have the throat, head and top of the
wings the colour of fire. All the birds of the Island have each
their season at different times, being six months in the flat
country & six months in the mountains, from whence returning
they are very fat and good to eat.
This behaviour is much like that of the Madagascar Fody,
rather than the other Foudia species, which are generally
insectivorous. However, there are reports that the Rodrigues Fody behaved in a similar way, before the
introduction of the Madagascar form in the 1850s (Cheke & Hume 2008).
Decline
Black Rats Rattus rattus were absent from Réunion when Dubois made his observations, but shortly after-
wards in 1676 they had already reached plague proportions. Oceanic island fodies are notoriously
susceptible to rat predation (e.g. Safford 1997), so it is likely that the Réunion population crashed once these
vermin had become established. Moreau (1960) suggested that the Réunion Fody occurred in the lowland
belt of evergreen forest up to an elevation of 300m, which would have been the one of the first forest types
cleared for agriculture. Whatever the reason for its disappearance, the Réunion Fody was certainly extinct
by the end of the 17th century.
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The Star Finch N. ruficauda is divided into three subspecies, the easternmost of which, N. r. ruficauda, may
have disappeared recently. Its original range is poorly documented, but it is known to have occupied central
Queensland and northern New South Wales (Holmes 1996; Higgins et al. 2006).
The Eastern Star Finch began to decline in the 19th and 20th centuries, primarily due to habitat alteration
caused by livestock over-grazing and trampling (Garnett & Crowley 2000). As the subspecies became rarer,
it was sought-after for the aviculture trade. Thousands appear to have been captured for this market during
the 1930s, but the effects were probably only temporary; trapping was outlawed in the 1960s (Holmes 1998).
Drought may have also been significant in this bird’s decline. Surveys to find them in central Queensland in
1993–94 and 1996–97 (Holmes 1996, 1998) were unsuccessful, and it might well now be extinct. Captive
birds of various subspecies survive, but it is not certain if any of these represent the eastern form.
Habits
The Eastern Star Finch occurred in pairs or flocks, rarely up to 50 individuals, and probably nested in loose
colonies (Higgins et al. 2006). It was a sedentary bird, preferring grasslands and grassy woodlands adjacent
to fresh water (Holmes 1996), but in earlier times it foraged in suburban areas, including roadsides, gardens
(Baldwin 1975) and even in the streets and yards of townships (Cayley 1932; Marshall 1932).
Diet comprised various seeds and insects, and the nest was a bottle-shaped construction, usually
between 3m and 9m above the ground. Its call was frequent and often loud, especially when in flight
(Higgins et al. 2006).
Fringillidae (Finches)
The majority of the true finches occur in the Northern Hemisphere, but genera are also found in South and
Central America, and on Pacific island archipelagos. There are four distinct subfamilies: Fringillinae (fring-
illine finches), Carduelinae (cardueline finches), Drepanidinae (Hawaiian finches), and Euphoniinae
(euphonias), which are further divided into about 33 genera (not including fossil taxa), with around 138
species. Some of these generic placements are tentative and await further analysis. Fringillid finches that are
endemic to islands have suffered severe losses from anthropogenic changes, and none more so than the
Hawaiian finches. Taking into account the number of new taxa that has been described from subfossil
remains, those that have left no trace at all, and the high percentage of historical losses, perhaps more than
100 Hawaiian finch species have become extinct. Two species also once occurred in the Canary Islands, and
one was almost flightless.
The Makawehi Finch is s o-named because subfossil remains were found in the Makawehi Dunes, Kaua’i.
The extant Laysan Finch Telespiza cantans and Nihoa Finch T. ultima were long regarded as races of the same
species, but it is now clear from the fossil record that they were once sympatric on some if not all of the main
islands of the group. Fossils of T. ultima have been found on islands to the east and west of T. persecutrix,
indicating that these forms are likely to be distinct species rather than subspecies, and both T. ultima and T.
cantans occurred on Molokai, along with the Maui Nui Finch T. ypsilon (James & Olson 1991; see below).
Bill-size – ranging from smallest to largest, Maui Nui Finch T. ypsilon; Makawehi Finch T. persecutrix; Nihoa
Finch T. ultima; and Laysan Finch T. cantans – would have provided sufficient niche segregation for these
otherwise comparatively similar finches to maximise utilisation of food sources. The Laysan and Nihoa
birds are predominantly terrestrial, and survive today because they inhabit islands in the remote north-west
chain. The main island populations would have been subject to major anthropogenic changes, and the
introduction of the Pacific Rat Rattus exulans, which is a known predator of birds, must have been cata-
strophic. It is unlikely that the main island populations survived to the historic period.
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The Makawehi Finch was closely related to the two extant species, but it had a smaller bill than either
(James & Olson 1991). If its ecology was the same and it was equally terrestrial, it would have disappeared
rapidly after colonisation of the Hawaiian Islands by the Polynesians.
The Maui Nui Finch has been found as a subfossil on Maui and Molokai, which suggests that it once
occurred on the greater island of Maui Nui (the combined islands of Maui, Molokai, Lanai and Kaho’olawe;
a combination of sea level rises, erosion and subsidence around 200,000 years ago saw the islands separate;
Ziegler 2002).
This species had the smallest bill of the genus. It was heavily preyed on by the Molokai Stilt Owl
Grallistrix geleches, its bones occurring abundantly in their pellets (James & Olson 1991). This infers that the
species was an adaptable one, and survived in the harsh, arid environment at coastal Ilio Point on Molokai.
It appears to have been widely distributed on the island, as other subfossil remains were discovered at a
cave site at about 305m above sea level. A similar but much s maller-billed bird was found at a much higher
altitude of 808m; this may represent another distinct Telespiza species (James & Olson 1991).
The Lanai ’Alauahio was still common at the start of the 20th century in all parts of the Lanai forest, from
600m to the highest point on the island (Perkins 1903), but like many birds of Lanai, it declined rapidly. It
was scarce long before the last pair was seen in March 1937 (Munro 1944). Munro could offer no explanation
for the extinction, but stressed that it was not due to the importation of foreign birds and their potential
diseases, for almost none had been introduced for a very long time. Deforestation and the activities of feral
livestock may have been at least partly responsible for its demise.
Habits
Its habits were much like those of other subspecies. It made short flights from tree to tree, and fed mainly on
small invertebrates in the bark of trunks and branches, but it would habitually feed on the ground or at the
bases of trees. It had a chipping call and Perkins (1903) described its song as vigorous and pretty. He saw it
rise up straight in the air, singing as some Hawaiian thrushes do. Perkins collected young birds in July 1894.
Munro (1944) found what he believed to be a nest of this species on 19 April 1913, which had been blown
out of a tree. It was different from other nests, a neat little ball compactly built of very fine grass stems and
skeleton leaves. Other nests of this species were more ragged on the outside.
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This species was quite common in the early years of the 20th century in the forests of Molokai, above 450m,
but expeditions in 1980 and 1988 to Mount Olokui, Molokai (where original forest still survived) (Scott et al.
1986; Reynolds & Snetsinger 2001), and more recently in 2004, failed to find any. It was last seen in the
Kamakou Preserve, Ohialele Plateau, in 1963 (Collar et al. 1994). The Kakawahie disappeared due to a
combination of habitat destruction, rat predation, and mosquito-borne avian diseases.
In the 1890s Munro had found the bird common in the upper boggy forest of the island and in 1907
collected a series. It declined steadily between then and 1944, when Munro (1944) wrote that the Kakawahie
was so reduced in numbers that there seemed little chance of saving it. Its last refuge was the high plateau
between the Wailau and Pelekunu Valleys.
Richards (1946) reported the dearth of birds on the island, observing only the commonest surviving
species, the Amakihi Chlorodrepanis virens wilsoni, but even this species was scarce on Molokai:
In the heights of the East End mountains . . . I saw what looked like an amakihi . . . I didn’t go on any hikes with
bird hunting in mind, but the silence struck me as odd. All I could hear was cricket-chirping and rarely, the
chirp of the small green bird I was unable to identify.
Other species of Paroreomyza still survive on Hawaii and Maui, but in greatly reduced numbers.
Habits
The Kakawahie foraged on trunks and branches of the Koa Acacia koa, working both on the upper and
underside of horizontal limbs, where they fed on insects. Very occasionally nectar was also taken. The nest
was constructed from moss neatly woven together and lined with dry moss and disintegrated ’Ie’ie
Freycinetia arborea leaves. One was located about 4.6m above the ground in an ohia tree. Bryan (1908) gave a
useful account of the bird:
In habits the Kakawahie resembles [other] species of the genus to which it belongs, and from which it differs in
color so widely. They prefer to feed over the trunks and branches of the trees. Here they secure the insects that
make up almost the whole of their diet. However, they will be seen in the tops of the tallest trees, but apparently
paying little or no attention to their flowers. In short, they are persistent and sturdy entomologists, always
active and alert, but strange to say, they seldom, if ever, take insects on the wing. At intervals moths are taken of
such size they are compelled to hold them under their feet and pull them to pieces so as to devour them piecemeal,
much after the fashion of the common chickadee . . .
The Kakawahie, like his cousins, is full of curiosity. The sound of one making one’s way through the woods is
sure to attract the little resident to the scene, when uttering their never varied ‘chirk’, they will come close
enough to the person to take in every detail of his makeup in wide-eyed inquisitiveness. Once satisfied that their
show of authority has no intimidating effect, they will resume their feeding close to the observer. One can thus
study their movements at close range. I have often watched them under the most favorable circumstances, for an
hour or more at a time, but have never seen them paying the slightest attention to the nectar-bearing flowers
about them. Occasionally they go down in the shrubs to within a foot or so of the ground, and it is probable that
on rare occasions they do alight on the ground, although I have never seen them do so.
The bird had a loud chirp, and could be attracted by an imitation of it. The call was said to resemble chip-
ping wood, hence the bird’s name Kakawahie, which means ‘to break up firewood’.
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The Oahu ’Alauahio was considered common in the last decade of the 19th century, despite having disap-
peared from the mountains in the neighbourhood of Honolulu (Perkins 1903), and was found in both the
Wai’anae and Ko’olau mountain ranges. Subfossil remains have shown it was also once an inhabitant of the
lowlands (James & Olson 1991). There was a gradual decline ever since, with only occasional reports of its
existence in recent decades. The last confirmed observation was of two birds on December 12, 1985 on the
Poamoho Ridge Trail (Bremer 1986), and recent searches for the species have failed to find it (Pratt 2010). It
appears that deforestation, introduced predators and mosquito-borne diseases are the most likely causes of
the Oahu ’Alauahio’s extinction.
Habits
This bird foraged on the trunks and branches of tall trees and fed on beetles, spiders, slugs, caterpillars and
moths. Perkins (1903) noted that they had a special technique for collecting a particular species of carabid
beetle:
The Oahuan species has the knack of obtaining one of the most remarkable of the Hawaiian Carabid beetles in
quantities, to judge from the remains found in the bird’s stomach, yet for the entomologist it is difficult to obtain
even a single specimen. I have little doubt that it obtains these from the interior of dead branches of lofty Koa
trees, practically inaccessible to the insect collector, since they are generally found in those shot when feeding in
this situation.
A single nest with two eggs was collected in late January, 1901 (Berger 1972). The call was a loud cherk.
Munro (1944) was the only man who ever described a living Lanai Hookbill, and although Perkins’s
description was not accepted as a valid taxon, Munro believed that this bird was a valid species. It inhabited
the forests of Akoko Euphorbia lorifolia that formerly covered the lowland plains on Lanai. Munro collected
the type specimen while it was feeding on the large fruit of an Akoko tree. Thousands of acres of Akoko
forest originally covered these plains (Snetsinger et al. 1998), which are now planted with pineapples. Akoko
trees still occur in the higher forests, and it was Munro’s belief that a few birds still survived there.
Perkins (1919), in his description, quoted Munro’s observations:
On March 17th, 1916, further up the same valley, where it is very densely wooded, I heard two or three birds
calling to one another, the cry being less sweet and not so loud as that of the Ou [’O’u Psittacirostra psittacea],
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. It looked
smaller than an Ou and more active, but less so than Chlorodrepanis [Amakihi]. The form of its bill could not
be made out, but it was not that of the latter. On Aug. 12th, 1918, in a patch of dry forest on the s outh-west side
of the mountain, at about the same elevation as that where the original specimen was obtained, I saw another
bird, and was near enough to note the light colouring round the eye, but not the form of the beak. Some of the
notes were like those of Psittacirostra, but others new to me, especially a low squeak or whistle, and it was too
small for that bird, not so t hick-set, and with a very short tail. So I feel sure it was the other.
Perkins (1919) continued:
As so few specimens have been seen by so skilled a collector, the bird must be a great rarity, but its discoverer
hopes that it may increase in numbers, as the forest is now rigidly protected and rapidly recovering. When I
collected on Lanai in 1893 and subsequently the forest was in a deplorable condition, being rapidly destroyed by
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Habits
Nothing was recorded about the bird’s habits, except that the type was collected while feeding on Akoko
fruit. Its stomach contained a few native berries. James et al. (1989) suggest that the bill of D. munroi was not
designed for strong biting or crushing, and that it may have been frugivorous, or possibly a predator of
terrestrial molluscs.
The Greater ’Amakihi inhabited the mountain rainforest on the windward slope of Hawaii behind Hilo,
near the Wailuku River, between 152 and 1,220m (Perkins 1903). When first discovered in 1892 it was very
rare and hard to find. Palmer (in Munro 1944) found it frequenting high trees and masses of creepers in the
densest parts of the forest. Perkins collected a series in December 1895, Henshaw (1902) obtained some
more in 1900, and the last was taken in 1901 by A. M. Woolcott (Stattersfield et al. 1998; Fuller 2000). Its
native forest was destroyed and replaced by sugar-cane (Amadon 1950), which is probably the main reason
for the bird’s extinction.
Habits
The Greater ’Amakihi was a shy and active bird, with a high, clear call, and a song with two or three loud
notes at the end, most similar to that of the extant Common ’Amakihi H. virens. It had a strong bill, which
was used to extract insect prey from the dense masses of ’ie’ie Freycinetia arborea covering Ohia Metrosideros
polymorpha trees. Perkins (1903) found that stomach contents were small Paratrigonidium forest crickets,
carabid beetles, spiders and caterpillars. The bird may also have taken nectar occasionally.
Munro (1944) made the following comment:
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The Hawaiians did not know the bird and had no native name for it, so presumably it was always rare. It fed on
crickets and a small caterpillar which occurred on the dry shoots of the ieie plant. The call was low and plaintive,
but audible from a long way off somewhat resembling an old native’s imitation of the Mamo call.
The nomenclature of this group of birds is extremely complicated. For a detailed review see Olson & James
(1995), Pratt (2005, 2010) and recommendations of the AOU (1998). Four species of ’Akialoa are known
historically from the Hawaiian Islands, all now probably extinct. Although no species was recorded histori-
cally from Molokai or Maui, the subfossil record has established that ’Akialoas probably referable to H.
lanaiensis once occurred on both islands (James & Olson 1991). A further two new species have been
described from subfossil remains.
After being considered fairly common in the early 1890s (Perkins 1903), the Kaua’i ’Akialoa declined
rapidly, being considered rare by the end of the century (Henshaw 1902), and thought to be nearly extinct
by 1944 (Munro 1944). Munro found it in all parts of the forests of Kaua’i, from the high plateau down to the
forest edges near the coast, but he did not see it after 1920. However, he mentioned that Walter Donagho
had reported one on the upper plateau in October 1941.
Munro believed that the birds were very susceptible to mosquito-borne diseases, and were exposed to
these by their habit of coming to the forest edge and to low elevations. He reported individuals with
tumours and cancers, and Perkins (1903) found birds with tapeworms. Richardson & Bowles (1964) redis-
covered a population in 1960, describing it as a rare species in the undisturbed forest of the Alaka’i Swamp.
This population must have disappeared extremely rapidly, as the last well-documented report was in 1965
(Huber 1966). Despite intensive searches and persistent rumours of its continued existence, no further
authenticated sightings have taken place, and the bird must now be considered extinct (Sincock et al. 1984).
Habits
The Kaua’i ’Akialoa was an active bird, hopping along tree branches, or moving w oodpecker-like on the
trunks of trees (Perkins 1903). It did not fly much, mainly just short darts from tree to tree, and it often sang
as it flew. The tubular b
rush-tipped tongue, common to all ’akialoas and nukupu’us, was used to extract
nectar from the flowers of lobelias and various flowering trees. The bird also foraged for insects on the
trunks and branches of trees, especially on Koa Acacia koa.
Their song was like that of a linnet but a little louder, and the birds answered each other with a chirp
(Munro 1944). When Munro studied them in 1891, they sang very little during January and February, but
the song was noticeable in March and April. He also heard a different call in April, which he took to be a
breeding call. Munro (in Wilson & Evans 1890–99) described the bird’s habits as follows:
This bird is much more common and enjoys a wider range than the Nukupu’u, which bird it much resembles in
habits. It seems to inhabit the whole forest-region of Kaua’i; its food consists of insects, their eggs and larvae, and
we have also seen them sucking honey from the Lehua flowers . . . I have seen these birds from the branches in the
tops to the roots of the trees, probing into holes and under the bark, where they find a harvest of cockroaches’
eggs, beetles and grubs; on one occasion I saw one alight on the ground and insert its bill amongst mats of dead
leaves and bits of wood; have also seen them collecting insects from the bases of the leaves of the halapipi tree;
have not often seen them feeding on honey.
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The Maui Nui Akialoa was extremely rare when first discovered, and is known from just three specimens,
taken in 1892 on Lanai. It was seen on perhaps only two more occasions, the first when Perkins saw one in
1894 (Perkins 1903). He considered that this bird was part of the same brood that had been collected two
years before in 1892:
I saw but a single example. This was evidently an adult male, its plumage appearing quite brightly yellow, and
unlike any of the figures in Mr Rothschild’s work [1893a]. There is no doubt that his figure of the adult bird, if
really taken from an adult, represents the bird in its non-breeding stage, for in January, when I saw the one above
mentioned, all of the adult birds on Lanai were in the fullest and most perfect plumage. It was extremely tame, at
times not five yards distant, hunting for insects along the trunk and large limbs of a partly fallen Ohia, which
overhung the edge of a precipitous cliff. As if killed, would necessarily have fallen in the brush far below, or have
lodged in the shrubbery on the side of the cliff, being without a dog I forbore to shoot, and when after some
minutes it flew off, it was seen no more. It is probable that this was really a survivor of the brood obtained by Mr
Rothschild’s collectors, since Wolstenholme, who discovered the bird, informed me that all of their specimens
were obtained in the same spot and practically at the same time. Certainly the bird seen by me was quite alone,
and this at a time when mature birds were paired, and it may even be feared that it is the possible sole living
representative of its species.
Munro (1944), who knew the island’s birds well, may have seen one:
I watched for it for 20 years and on only one occasion saw a bird that might reasonably be supposed to be this
species. It was across a small valley from my position, moving up the steep hillside, flying from tree to tree. It
was more yellow in colour than any other Lanai bird, about the size and with the action of an Akialoa, but I could
not say for certain that it was this bird. I made two collections of the forest landshells of Lanai, two botanical
collections, and hunted goats all through the forest; I explored for water in its valleys and rode its bridle trails
scores of times, but I never again saw a bird I even remotely thought might be an Akialoa.
It appears that forest destruction was the primary reason for its disappearance. Olson & James (1982, 1995)
and James & Olson (1991) discovered subfossil remains of ’akialoas, probably of this species, on the islands
of Molokai and Maui, but these populations disappeared before being scientifically described.
The Lesser ’Akialoa was discovered and first collected on the third voyage of Captain Cook in 1779 near
Kealakekua Bay. Munro (1944) reported that in 1891 it was common in Kona and widely distributed over
the rest of the Hawaiian forests. It was still moderately common in 1894–95, when Perkins (1903) found it
between elevations of 460m and 2,130m, distributed through the continuous forest. The population
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collapsed extremely rapidly, however, and the last specimens were taken at Pa’auilo, Hamakua District, in
September 1903 (Banko 1979).
In 1936 and 1937, Munro (1944) did not see any in the Kau, Kona or Kohala Mountains. The last uncon-
firmed and probably doubtful sighting was in 1940 by Chester E. Blacow (Baldwin 1941). There have been
many searches for it since without success (Greenway 1967). The rapid disappearance of this bird, going
from being considered common to being extinct in less than a decade, is presumably due to m
osquito-borne
avian diseases, accelerated by habitat destruction.
Habits
Perkins (1903) noted that nectar comprised the bulk of the diet, but the birds also fed on insects, and he
suggested that the genus was evolving towards insectivory. The Lesser ’Akialoa obtained its food by
probing for insects in bark crevices, under lichens and in the leaf-bases of the ’ie’ie vine Freycinetia arborea,
and was fond of nectar from lobelias, Koa Acacia koa and Ohia Metrosideros spp. Perkins commented that
when feeding on nectar the birds seemed to sing more freely, and suggested that nectar stimulated singing.
Perkins also found a nest, apparently the only one recorded, at the end of June. It was well-concealed near
the extremity of one of the largest branches of a Koa tree. It contained one chick, but a recently fledged bird
was nearby.
Wilson & Evans (1890–99) made further comments on the habits of the bird:
This species – peculiar, so far as my observations go, to the Island of Hawaii – occupies the lower forest-zone
from about 1100 to 2500 feet, and is most plentiful among the tall ohia trees. Like its larger relative [Kaua’i
’Akialoa H. (e). stejnegeri], it prefers decayed timber in which to search for its food, and invariably chooses a
rotten or half-dead tree for its hunting ground, no doubt on account of its slender bill, which requires soft mate-
rial to work upon. It is also very partial to the great tree-ferns which in the forests of Hawaii reach a height of
more than 30 feet, and, as the sombre colour of its plumage is very nearly that of their foliage, it is most difficult
to observe, and is at the same time more quiet and unobtrusive in its habits than any other member of the genus;
in fact, had it not been for its clear and characteristic call-note, I doubt whether I should have noticed it at all.
Munro (1944), rather poignantly, remarked:
With many birds going over and over the flowers each will get but a small taste from each flower and that might
perhaps cause their haste. But again the birds at this time seem care-free and dash about and sing joyously when
not seeking food. It is really a scene of merry-making; a scene I fear will never be replaced.
This Oahu endemic was considered rare when first discovered and, after the first specimens were collected
in 1837, was thought not to have been recorded again. However, Perkins believed he saw a pair in October
1892 (Perkins 1903) and stated:
. . . on one occasion, I saw a pair of this species, the one chasing the other over a narrow ridge high up in the
mountains. The leading bird passed over this ridge and down into a deep gulch on the other side, squeaking as it
flew, the other alighted in an Ohia tree on top of the ridge about 10 yards in front of me. The latter when I shot
dropped over the steep ridge. After much searching I was unable to find it. I distinctly noticed the sombre
plumage of the upper parts, which in no way approached the yellower colour of the other species with which I
was then familiar.
For several days he hunted round Waolani, and spent a night in the open on the ridge, but he never again
saw anything that looked like an ’akialoa.
There were two reported sightings in 1937 and 1940, both of which are considered to be of questionable
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accuracy (Greenway 1967). The species was long thought to be known only from the holotype in the Berlin
Museum, collected in the Nu’uanu Valley in January 1837 by Ferdinand Deppe. Olson & James (1995)
located another specimen in the Philadelphia Academy, taken at the same time by J. K. Townsend.
This was a species with a very long, decurved bill, not modified for hammering as in the Nukupu’u
Hemignathus lucidus or Akiapola’au H. munroi. The lingual trough of the mandible, the excavated part of the
bill through which the tongue can pass, is much less excavated in Hemignathus upupirostris, which indicates
that the tongue was short, not long and tubular (James & Olson 1991). This may have been associated with a
different diet; perhaps this bird did not predominantly feed on nectar as in other ’akialoas. Nothing else is
known about the bird. As the subfossil remains are known from coastal sites on Kaua’i and Oahu, the
Hoopoe-billed ’Akialoa appears to have been a victim of Polynesian habitat alteration, which was much
more severe in the lowlands.
The species was very rare in 1898, and even then probably confined to the Alaka’i Wilderness Preserve, but
it was easy to collect owing to its habit of accompanying little flocks of ’Akikiki Oreomystis bairdi, whose
chirping attracted attention.
Munro (1944) described the bird as tame and unafraid, hardly seeming to notice the approach of humans.
It was thought extinct until rediscovered in 1960, when it was observed only once (Richardson & Bowles
1964). It was possibly sighted in 1995 (Pratt & Pyle 2000), but all other surveys have failed to find it, and it
seems very likely that this bird is now extinct (Reynolds & Snetsinger 2001; Pratt 2010). Its extinction can be
attributed to forest clearance, opening of the understorey by cattle and pigs, and m osquito-borne avian
diseases.
Habits
This was a tree-creeping species, searching the trunks and branches of Ohia Metrosideros polymorpha and
Koa Acacia koa for insect prey. The incredible bill – with the upper bill being almost twice as long as the
lower – was used as a dual purpose tool. The lower bill was used for hammering at loose bark or to widen
larval burrows to expose prey, which was then hooked out with the long, thin upper bill. The tapping sound
made while hammering was audible for some distance. The bird would also forage among leaves for cater-
pillars, and very occasionally took nectar. It had a clear and distinct call kee-wit, while Perkins (1903)
described its song as a short trill and ‘fairly lavish’.
This race of Nukupu’u was believed to be common in the lower forest belt until about 1860 (Perkins 1903),
but it had totally vanished by about 1893, and nobody has seen it since. Very few specimens were ever
collected, and none at all after January 1837, when Deppe took specimens in the Nu’uanu Valley, behind
Honolulu. Wilson & Evans (1890–99) provides an interesting summary:
While Deppe was in the Sandwich [Hawaiian] Islands he was joined by the still better known American natu-
ralist Townsend . . . In his work Townsend states that he and Deppe, in January 1837, hired a house in the
Nuano Valley [Nu’uanu Valley], five miles from that town, with the object of collecting birds and plants, and
we may well suppose that this species was found by them in that district. On Townsend’s return he sent several
specimens of birds collected by him to Audubon, then in this country, and among them two of the present
species, which were acquired by the late Sir William Jardine, at the sale of whose collection, in 1886, they were
bought for the Museum of the University of Cambridge. On one of them being submitted, at my request, to
Professor Cabanis for comparison with the type at Berlin, that eminent authority declared the two to be specifi-
cally identical. This result was the more satisfactory since I myself was unable to meet with the species, and later
explorers have been no more fortunate, so that there is reason to fear that it has become extinct. It was undoubt-
edly peculiar to the island of Oahu, where Deppe informed Lichtenstein that it frequented the p lantain-blossoms
in considerable numbers.
Habits
Perkins (1903) managed to acquire some meagre information from the locals about this bird. It was almost
exclusively associated with the Koa Acacia koa, though the birds collected by Deppe in 1837 frequented
plantain flowers. Nothing else is known, and the belt of forest behind Honolulu and elsewhere is now
completely destroyed.
Perkins (1903) found the Maui Nukupu’u to be more numerous in the early 1890s than the Kaua’i Nukupu’u
H. l. hanapepe, but restricted to the Haleakala mountain forest. Few ornithologists visited Maui in subse-
quent years, and the population declined over the next century. Greenway (1967) thought that as
considerable forest still remained on the eastern and southern slopes of Haleakala, perhaps a few birds
might still survive. There have been a number of sightings, but the last was of a single male seen in 1995
(Reynolds et al. 1995). Despite intensive surveys since, no birds have been recorded, and it is now almost
certainly extinct.
Habits
Perkins (1895, 1903) noted that the Maui and Kaua’i Nukupu’us scarcely differed in habits or song. The call
was a loud kee-wit, repeated once or twice. This call was repeatedly uttered, the birds pausing in their
feeding for this purpose. Food consisted of arthropods, found by probing bark, and the stomachs of
collected specimens contained spiders, wood-burrowing larvae of moths and beetles, and adult beetles.
Small pieces of lava were also swallowed, apparently to assist in grinding up of the hard shells of beetles.
Perkins (1903) recorded it only rarely sucking the nectar of the flowers of the Ohia Metrosideros polymorpha.
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The dual-purpose bills of the Nukupu’u H. lucidus and Akiapola’au H. munroi are truly amazing adapta-
tions for foraging, but the discovery of the Giant S
cimitar-billed Nukupu’u takes this to extremes. This bird
was a giant, much larger than any other Hemignathus; it had a long, decurved bill, the maxilla (upper bill)
being fl at-sided, deep and narrow, and the mandible (lower bill) approximately one third of its length. The
upper bill was reminiscent of the ’akialoas, but the short mandible was nearly straight, as in H. munroi, but
less short and stout. The ’akialoas and nukupu’us foraged by hammering and twisting off bark and lichens
on trees, or probing into epiphytes, bark crevices or insect burrows (Pratt 2005). The specialist bill of the
Scimitar-billed Nukupu’u may have had a different function. James & Olson (2003) hypothesise that the
bird might have been more terrestrial, probing into g round-level vegetation or using the long upper bill to
move leaf litter, in the manner of the New Zealand Weka Gallirallus australis or Kiwi Apteryx sp. If it was a
terrestrial forager, there can be no doubt that it disappeared soon after the arrival of humans and their
commensal animals.
A unique specimen of this species, of indeterminate race, was collected on Hawaii by the US Exploring
Expedition in 1840 or 1841 (Olson & James 1995). The specimen is in poor condition, with the tip of the
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maxilla missing. Olson & James quote the original account of Peale (1848), which almost certainly refers to
the same specimen:
There is another species very much like this, still undescribed; it has a very slender bill, the under mandible not
being larger than the upper, and little more than half its length; it differs in other respects, but the only specimen
obtained by the Expedition, is too much mutilated to venture specific characters for it: the indication is made to
call the attention of future ornithologists visiting that island.
The Hawaii population probably disappeared rapidly after European contact, but Olson & James note that
on June 29, 1971, a bird most closely resembling the Maui Nukupu’u H. l. affinis was possibly observed in
the Kohala Mountains (van Riper 1982). There have been no other reports, and the bird is now presumably
extinct.
The discovery of this specimen now shows that the Nukupu’u H. lucidus was sympatric with the
Akiapola’au H. munroi on Hawaii. Whether their ranges overlapped however, will never be known. A
subfossil maxilla, almost certainly referable to H. lucidus, has been recovered from sand dune deposits on
Molokai, which extends the range of the species to all of the main islands except Lanai (James & Olson 1991;
Olson & James 1991).
the birds could be construed as belonging to separate genera (James & Olson 1991). The Pololei S hovel-billed
Finch V. neophasis had a larger, straighter and deeper upper bill than that of V. mirandus, and there were a
number of other structural differences in the post-cranial skeleton. Its ecology is equally perplexing.
The Hawaiian gapers had sturdy, straight or decurved, very long bills with elongate retroarticular processes
(a flange of bone that projects backwards from the posterior part of the mandible). A similar condition
occurs in a number of other Hawaiian finch genera, but the bill of Aidemedia differed in some other aspects
(James & Olson 1991).
‘Gaping birds’, which include members of the Sturnidae and Icteridae, are so named because of their
feeding strategy (Beecher 1951). They habitually insert the bill into a substrate and spread their beak apart
using strong muscular actions. For example, the Eurasian Starling Sturnus vulgaris and Western Meadowlark
Sturnella neglecta feed by gaping in a variety of substrates including bark, fruit, flowers and soil. Because of
the similarity in bill morphology, the gapers of Hawaii probably had an equivalent feeding technique
(James & Olson 1991).
The Straight-billed Gaper was sympatric with the Sickle-billed Gaper on Oahu (see below), and differed
in the shape of the bill. The bill of the S
traight-billed Gaper was most similar to that of the extinct Greater
’Amakihi Hemignathus sagittirostris (p. 292) but longer and narrower. All of the gapers disappeared after the
Polynesian colonisation of the Hawaiian Islands, but before the arrival of Europeans, doubtless due to
habitat destruction and predation by the introduced Polynesian Rat Rattus exulans.
The Sickle-billed Gaper had a long, narrow and decurved bill, although the degree of curvature was much
less than in the ’akialoas (see p. 333). James & Olson (1991), with some degree of caution, suggest that the
bill differences seen between the S ickle-billed Gaper A. zanclops and S
traight-billed Gaper A. chascax could
be males and females of the same, highly dimorphic species. They compare these differences with the
extinct Huia Heterolocha acutirostris of New Zealand (see p. 267), which exhibited extreme sexual dimor-
phism, the male having a short, straight bill and the female a long, curved one. If James & Olson are correct,
the gapers of Oahu would be unique among the Hawaiian finches in this respect.
The Maui Nui Gaper is distinguished from the two species on Oahu by its shorter, very straight bill, and
flattened dorsal surface on the maxilla. The bill is similar to that of meadowlarks of the genus Sturnella,
which also have a long, straight bill and flattened dorsal surface, but this is due entirely to convergence
(James & Olson 1991).
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It is interesting to note that this species occurred on two islands, Maui and Molokai, so it is likely that the
Maui and Molokai populations of A. lutetiae became separated due to rising sea-level rather than the popu-
lation of one island colonising the other by crossing a channel. This was also the case with the Maui Nui
Finch Telespiza ypsilon (see p. 329) and Maui Nui ’Akialoa Hemignathus (ellisianus) lanaiensis (see p. 334). It is
also probable, based on biogeography, that A. lutetiae may have once occurred on Lanai, the middle island
of the Maui Nui group, though subfossils have not been found.
Once considered to be subspecies of the extant Hawaii ’Akepa Loxops coccineus, recent mitochondrial DNA
studies have shown that the Oahu and Maui ’Akepas are distinct and warrant full specific status (Pratt
2010).
Perkins (in Munro 1944) may have seen the last Oahu ’Akepas. The bird had previously been collected
by Wolstenholme (after whom the bird was named) for Rothschild on May 20, 1893 (Rothschild 1893c). This
was the last known specimen, but Perkins saw a pair in 1900 in forest in the Wahiawa district. The Oahu
‘Akepa was not seen again.
The Oahu ’Akepa appears to have suffered the same fate as so many Hawaiian finches – destruction of
habitat, predation by introduced mammals, and vulnerability to avian diseases. Despite rumours that it still
persists, it is now almost certainly extinct (Lepson & Freed 1997).
Habits
This bird was commonly seen throughout the 19th century, though it appears to have been restricted to
dense mountain forests, particularly associated with Koa Acacia koa and Ohia Metrosideros polymorpha. It
was an active bird, gleaning caterpillars and spiders from leaves and sometimes from bark, the bird having
a specialised cross-bill for the purpose. The lower bill was asymmetrical, variably crossing the upper bill to
the right or to the left depending on the individual. The bill was also used to open buds for insects, and
nectar was occasionally taken.
The Maui ’Akepa was, in recent times, confined to the mountain of Haleakala in the east of the island. It was
considered common in the 19th century, but declined as the forests were cut and livestock opened up the
understorey. The increase in altitude of mosquitoes carrying deadly avian diseases appears to have been a
significant factor in its disappearance, and it is possible that the successful rearing of young was impaired
by competition with the introduced Japanese White-eye Zosterops japonicus, a scenario affecting the extant
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Hawaii ’Akepa today (Camp et al. 2009). The last record of the Maui bird was in 1988 (Engilis 1990; Pratt
2010). Despite intensive surveys in suitable remaining forest there have been no further sightings.
Habits
The Maui ’Akepa occurred in Koa Acacia koa forest, but preferred Ohia Metrosideros polymorpha for foraging.
The bird had a tubular, nectar-sucking tongue, but nectarivory was only occasionally practised.
Perkins (1903) only once saw a nest, high up in an Ohia tree, at the extremity of a horizontal branch. The
pair was collecting nesting material from the ground, and taking the woolly down of tree ferns, probably as
lining for the nest.
The Greater Koa Finch had a restricted range, in the Koa Acacia koa forests between 1,220m and 1,520m on
the western and southern slopes of Mauna Loa, in the Kona and Kau Districts (Olson 1999a). The species
was first found by Rothschild’s collector, Palmer, on September 26, 1891 at Puulehua in Kona, when he and
George Munro collected a series of specimens (Munro 1944). Despite the restricted distribution of the bird it
was considered common (Perkins 1903), and possibly the entire population could be observed when the
Koa was in fruit (Greenway 1967).
Although there was a reported sighting in 1937 by Walter Donagho (in Munro 1944), none has definitely
been seen since Perkins collected a series in March 1896 in Kau (Perkins 1903; Olson 1999a). Henshaw (1902)
failed to find it a few years later. The speed at which this species disappeared – in less than a decade – is
remarkable. The loss of habitat, and competition, predation and disturbance from introduced alien species,
are all likely to have caused a population crash, and avian disease contracted from mosquitoes must have
been significant; but these factors do not fully explain the rapidity of the bird’s decline. It is probable that
Polynesian destruction of lowland Koa forests had already forced the birds into less suitable marginal
habitat (James & Price 2008), making them more susceptible to extinction after Europeans arrived on the
island.
Interestingly, Olson (1999a) suggests that the persistence of Rhodacanthis and Kona Grosbeak Chloridops
kona (see p. 344) into the late 19th century was more to do with the decimation by European diseases of the
native Hawaiian human population than for any other reason. The disease outbreaks occurred following
contact with members of Captain Cook’s expedition in 1778 and 1779. The Kona District was the most
intensely cultivated area on Hawaii, but from the late 18th century (and for the next 100 years) there was a
period of respite from human interference, during which time the highland forests recovered. However,
this would change drastically once a new wave of settlers from Europe arrived.
Habits
The Greater Koa Finch may have undertaken some elevational movement in response to food availability,
as it periodically occurred at about 900m and at 1,200m (Olson 1999a). The bird was easily located by its
song, which was distinctive and comprised several whistled, fluting notes, the last ones prolonged; the
song ‘seemed to flood the whole surrounding forest’ (Munro 1944). It fed on the green beans of the Koa, and
sometimes the birds’ stomachs would be stuffed with nothing else. They were also insectivorous, and fed
on caterpillars and other insects and their larvae.
Perkins (1903) saw males collecting nesting material from the ground, which they carried to the tallest
Koa trees, and only when the caterpillars had stripped the leaves could he see largish nests that were almost
certainly those of the Greater Koa Finch:
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The Koa Finch is the largest and most beautiful of all the Hawaiian Finches. It frequents the tallest and most
leafy acacias, both when growing on the roughest l ava-flows and in the grassy openings in the forest. It belongs
entirely to the upper forest, and is probably most numerous at about 4,000 feet. Its peculiar whistle, though not
very loud, is very clear, and can be heard for a considerable distance. If imitated closely it will readily answer,
and sometimes, after fruitless hunting for hours without even hearing a sound from this bird, a whistle has been
immediately responded to. At other times a distant bird has been drawn close by the imitation of its whistle and
easily secured. It devours the beans of the acacia, and these it swallows in very large pieces . . . I have seen both
male and female feeding the f ull-grown young, and as I could find nothing but the large pieces of koa bean in the
latter, I conclude that the young are fed on pieces similar to those swallowed by the parents, without mastication.
The young male soon acquires the peculiar whistle, for I have shot one in almost perfect song in quite immature
plumage and with the skull still cartilaginous. It does not restrict itself to the koa bean, but varies its diet by
feeding on lepidopterous larvae . . . When it has been feeding on the koa beans its bill is often much stained with
their green juice and green fragments.
The status of the Lesser Koa Finch has caused much debate, as the bird was observed and collected only
over a period of a few days in the autumn of 1891, and never seen again (Olson 1999a). Rothchild’s collector,
Robert Palmer, along with George Munro, obtained a few in the same region, Kona, where they obtained
the Greater Koa Finch R. palmeri, but neither collector considered the birds distinct; only Munro noted that
they differed in size and coloration. Although Perkins (1903) and Henshaw (1902) searched the area care-
fully in subsequent years the bird was never seen again.
Several authors doubted the validity of the Lesser Koa Finch, but Amadon (1950) and James & Olson
(2005) considered them distinct, and this now seems to be the consensus (e.g. Pratt 2005). The discovery of
subfossil remains of Rhodacanthis on Maui, where a large and a small species have been found, has shown
that sympatric species did occur elsewhere (James & Olson 1991; James & Price 2008).
Fortunate collection
With some foresight, Perkins (1903) noted that the collecting of the Lesser Koa Finch was purely down to
chance, and it is fortunate that Palmer and Monro were in the right place at the right time:
Munro informed me that he and Palmer saw in all a score or two of Rhodacanthis, amongst which were two or
three of the yellow-headed species. In 1892 in a longer stay in exactly the same locality, I saw several score of the
larger bird, and in two subsequent visits, the latter of which was made for the special purpose in investigating
this matter, certainly some hundreds were examined with the naked eye or glasses, as the case required. On none
of these occasions did I see a single one of the y ellow-headed bird, nor did I hear any song or call note that could
be referred to a species unknown to me.
Habits
Nothing was recorded about the birds other than they were found in association with the Greater Koa Finch
in large Koa trees, and their stomach contents comprised seeds of that tree. Munro (1944) expressed regret
that, at the time of collection, he failed to realise that the bird was different from the Greater Koa Finch;
otherwise he would have made notes on its habits.
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Originally thought to represent a population of Greater Koa Finch R. palmeri (James & Olson 1991), the
Scissor-billed Koa Finch has distinct characters that warrant specific recognition. It differs from Greater Koa
Finch in bill structure, and from Lesser Koa Finch R. flaviceps in having a larger skull and jaws (James &
Olson 2005). The bill was robust, with strong muscle attachments which presumably allowed the bird to
slice into the tough, fibrous pods of Koa Acacia koa, which was the main food item of the koa finches on
Hawaii (Perkins 1903).
The habitat where the subfossil remains were found on Kaua’i contained a number of leguminous plants,
including Kanaloa Kanaloa kahoolawensis and Koai’a Acacia koaia (Burney et al. 2001); these were probably
utilised for food, with the birds also feeding on caterpillars and berries (James & Price 2008). The birds prob-
ably disappeared as a result of deforestation and o ver-hunting following the arrival of the Polynesians, but
prior to the arrival of Europeans in the 1770s.
The Oahu Koa Finch is known from subfossil material from both Oahu and Maui. James & Olson (2005)
suspect that the Maui population represents a distinct taxon but refrain from describing it, pending the
discovery of further diagnostic material. This was a smaller koa finch, approximately the size of R. flaviceps,
but it differed in the structure of the bill, being more similar to the larger R. palmeri and R. forfex (the latter of
which it would have been sympatric with on Maui).
As in the case of R. forfex on Kaua’i, the habitat surrounding the fossil locality on Oahu comprised various
legumes, but the Koa Acacia koa was comparatively scarce, suggesting that in the lowlands at least other
leguminous plants were preferred. The Oahu Koa Finch disappeared during the period following
Polynesian occupation.
The Kona Grosbeak had, in historic times at least, a tiny distribution. It appears to have been restricted to an
area of about 10.3km2 on the slopes of Mauna Loa, between 1,100m and 1,700m but probably not much
below 1,400m (Olson 1999a, 2014). It was discovered by Scott B. Wilson (1888) on June 21, 1887, when he
obtained two specimens. Henry Palmer (in Munro 1944) collected it again in September 1891. A specimen
taken in September 1892 by Robert Perkins (not in 1894 as often cited, e.g. Munro 1944) was the last known
(Banko 1986; Olson 1999a). Olson (2014) suggests that populations of the Kona Grosbeak were probably
always low in number, and at the time of its discovery, by which time the birds had been further reduced by
introduced disease, would have comprised a few scattered groups and lone individuals. Even in 1887,
Wilson, followed by Perkins in 1892, considered it rare and highly localised (Wilson & Evans 1890–99;
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Perkins 1893; Walther & Hume 2016); therefore its historical existence lasted for only five years and six
months (Olson 1999a).
Munro (1944) believed that a few might still survive at very high altitudes on the lava flows, but this was
unlikely, and the bird has probably been extinct since the end of the 19th century. Like the koa finches, the
rapidity of this bird’s extinction is difficult to explain. Its disappearance must be attributed to anthropo-
genic changes, especially the introduction of mosquitoes and associated avian diseases.
Habits
Those ornithologists who saw the Kona Grosbeak found it in Naio or Bastard Sandalwood trees Myoporum
sandwicense, where the birds fed almost exclusively on the dry seeds (Olson 1999a). An osteological analysis
has shown that the skull and jaws of the Kona Grosbeak were highly adapted for crushing and cutting in
order to feed on the excessively hard seeds of Naio (Olson 2014); the cutting edge (or tomium) of the rostrum
was pressed against an anvil-like pad on the opposing mandibular surface with tremendous force. Munro
(1944) noted that the birds seldom strayed from the more recent lava flows, where there was no under-
growth and the trees were of medium-size. Olson (2014) suggests that the limited distribution of the Kona
Grosbeak was due to its total reliance on specific but restricted young ‘clinker’ lava flows (large, rugged
masses of boulder-rich lava) on the drier leeward side of the island, where the extreme environmental
conditions caused scrubby Naio trees to dominate and produce year-round fruits. The specialised Kona
Grosbeak was the only species capable of utilising this food source, and was replaced elsewhere on the Big
Island and elsewhere in Hawaii by the smaller and more numerous Wahi Grosbeak Chloridops wahi (see p.
345); it is likely that the Wahi Grosbeak was the direct ancestor of C. kona (Olson 2014). Therefore the Kona
Grosbeak was continually forced to move to younger flows when conditions through erosion, soil develop-
ment and vegetation succession no longer provided suitable habitat (Olson 2014).
According to Perkins (1903), they were sluggish birds that had a short sweet song, but they were most
often heard to squeak. Perkins (1895) remarked that:
It is a dull, sluggish, solitary bird, and very silent – its whole existence may be summed up in the words ‘to eat’.
Its food consists of the seeds of the fruit of the aaka [Myoporum sandwicense] . . . and as these are very minute,
its whole time seems to be taken up cracking the extremely hard shells of this fruit, for which its extremely
powerful beak and heavy head have been developed. I think there must have been hundreds of the small white
kernels in those that I examined. The incessant cracking of the fruits when one of these birds is feeding, the noise
of which can heard for a considerable distance, renders the bird much easier to get than it otherwise would be. It
is mostly found on the roughest lava, but also wanders into the open spaces in the forest. I never heard it sing (I
once mistook the young Rhodacanthis’ [one of the koa finches] song for that of the Chloridops [Kona
Grosbeak], but my boy informed me that he had heard it once, and that its song was not like that of the
Rhodacanthis. Only once did I see it display any real activity, when a male and female were in active pursuit of
one another amongst the sandal-trees. Its beak is nearly always very dirty, with a brown substance adherent to
it, which must be derived from the sandal-nuts.
Adult birds were seen taking endemic looper caterpillars (Scotorythra sp.) (Perkins 1903), which were prob-
ably an important food source when feeding the young (Olson 1999a, 2014).
The Kona Grosbeak C. kona was thought to have been the only member of its genus, but recent discoveries
in the fossil record have shown that large-billed finches of the genus Chloridops once occurred throughout
the Hawaiian Islands. The Wahi Grosbeak occurred on Kauai, Oahu and Maui, and is likely to have been the
commonest member of the genus, and probably occurred on all of the large islands of the Hawaiian
Archipelago (Olson 2014). It was so-named because of its ability to crack hard seeds; the Hawaiian name
wahi means ‘to cleave or break’. It was smaller in size than C. kona, but the bill was similar in structure, only
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smaller and shorter (James & Olson 1991). Olson (2014) suggests that the Wahi Grosbeak may have fed on
the hard seeds of Zanthoxylon, a genus that occurs throughout the Hawaiian Islands, and the 8-mm seeds
could have been accommodated by the jaw apparatus. It occurred sympatrically on Oahu with the largest
of the Chloridops, C. regiskongi, so presumably foraged in different types of forest or on a different-sized class
of food. The Polynesian destruction of habitat was almost certainly the reason for its extinction, as it does
not appear to have survived into historic times.
Another probable grosbeak species has been recovered from fossil localities on Maui. The Maui spec-
imen, which might be a small individual of C. wahi, is known from a single mandible, but pending the
discovery of further material, may prove to be a distinct species (James & Olson 1991). A series of jaw
remains found on Kaua’i are now known to be referable to C. wahi, and not a distinct taxon as was formerly
believed (Olson & James 1982; Hume & Walters 2012).
This was a huge grosbeak with a massive bill, larger than that of any other known finch-like member of the
Hawaiian finches or Drepanidini. The structure of the bill differed considerably from that of other Chloridops
species, and the bird may well warrant separate generic status (James & Olson 1991). James & Olson
concluded that it could in fact represent a sister taxon to the Rhodacanthis koa finches.
The great size of the bill alludes to its ecology and no doubt it was a granivore, crushing the hard seeds of
trees and shrubs with its massive jaws. All of the subfossil remains found are from Barber’s Point, a lowland
coastal site on Oahu. If this species was endemic to the coastal dry forest, it would have disappeared rapidly
after Polynesian habitat destruction, which was most severe in the lowlands.
James & Olson (1991) considered the Mauka Grosbeak generically distinct from Chloridops in a number of
characters, most notably by the large nasal openings of the maxilla. The Mauka Grosbeak was equal in size
with the largest of the Hawaiian finches, the Greater Koa Finch Rhodacanthis palmeri, but the bill was
comparatively short and broad. It appears to have been one of the few montane endemics, as there is no
evidence of it occurring in the rich lowland fossil sites on Maui.
The early extinction of the Mauka Grosbeak is somewhat puzzling considering that other species of
Hawaiian finches and the ’o’os survived until comparatively recent times in the montane forests.
The ’O’u formerly occurred throughout the Hawaiian Islands, and it was considered one of the commonest
of the Hawaiian finches (Perkins 1903; Snetzinger et al. 1998). However, Perkins only saw a single pair in
March 1893 on Oahu, shortly after which it disappeared. Rothschild (1905a) considered the Oahu popula-
tion distinct (as P. p. deppei), but, along with P. p. oppidiana, once considered to be a distinct race from
Molokai, it has been relegated to the synonymy of P. psittacea.
Perkins (1903) noted that rats were out-competing the ’O’u for its main food source, the fruit of the ’ie’ie
Freycinetia arborea, and that once the forests were opened up by pigs, cattle and goats, the ’O’u disappeared
from places where it was previously numerous. In addition, the bird’s habit of migrating to low-elevation
forest exposed it to avian diseases spread by mosquitoes (Snetzinger et al. 1998). Each island population of
’O’u rapidly disappeared; it was extirpated on Maui by 1901; on Molokai by 1907; and by 1931 on Lanai
(Pyle & Pyle 2009).
Last strongholds
The last strongholds of the species were in the Alaka’i Wilderness Preserve on Kaua’i, and the Upper Waikea
Forest Preserve on Hawaii, but the populations continued to dwindle. A population of some 400 birds
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survived on Hawaii into the 1980s, but a lava flow from Mauna Loa destroyed much of the Upper Waikea
forest, and the last individual was recorded in 1987 (Conant et al. 1998; Snetsinger et al. 1998). On Kaua’i
there were approximately 60 birds in the early 1970s, which declined to only two in 1989, with the bird seen
for the last time on February 17 of that year (Pyle & Pyle 2009). There have been a number of unconfirmed
sightings since 1995, but the species is now likely to be extinct (Pratt 2005, 2010).
Habits
The ’O’u was an adept long-distance traveller, and probably island-hopped on a regular basis, evidenced
from the absence of speciation in each island population. This was unusual compared with other Hawaiian
finches, and may be explained by a regular m ass-dispersion event following a steep population rise in good
years; thus gene flow was maintained between the populations. It appears to have been restricted to
mid-elevation wet and mesic Ohia Metrosideros polymorpha forest, between 800 and 1,900m, but more
commonly between 1,200–1,500m (Scott et al. 1986). It fed predominantly on the fruit and bracts of the ’ie’ie,
and there was considerable altitudinal seasonal movement to locate fresh food sources.
The young were fed almost exclusively on caterpillars (Perkins 1903), and the ’O’u would travel great
distances to obtain this prey. The song was rather like that of a canary, with whistles, trills and warbles
(Snetzinger et al. 1998), beginning with three or four distinct and penetrating whistled notes. Perkins noted
that they would sometimes sing in concert on the wing.
The Cone-billed Finch is very rare in the Hawaiian fossil record and known from only two maxillae found
on Kaua’i, the oldest Hawaiian Island in the group. The genus differs from all other Hawaiian finch genera
in having the bill cone-shaped, rather than arched (James & Olson 1991).
The bill of the C
one-billed Finch was rather elongate and narrow as well as being c one-shaped, and it is
most similar to the recently extinct Po’ouli Melamprosops phaeosoma (see p. 352), but it differs in a number of
other characters. The Cone-billed Finch subfossil remains were found in pellet deposits in caves of the
Kaua’i Stilt Owl Grallistrix auceps (see p. 226), which was probably the finch’s main predator.
The Ridge-billed Finch differed from the Cone-billed Finch and all other Hawaiian finches in having two
converging ridges on the dorsal surface of the maxilla (James & Olson 1991). The bill is somewhat reminis-
cent of that of Molothrus cowbirds, but this resemblance is due entirely to convergence.
The Ridge-billed Finch has been identified in subfossil cave deposits on the central Hawaiian islands of
Maui, Molokai and Oahu, which makes it the most widely distributed of all fossil Hawaiian finch genera.
As there appears to be no differentiation between each island population, it may have been a regular
island-hopping species like the ’O’u Psittirostra psittacea (see p. 347). Despite the wide distribution of the
Ridge-billed Finch, it disappeared before European settlers arrived, probably as a result of habitat destruc-
tion and predation by introduced mammals.
Specimens Specimens are in Cambridge, England; Cambridge, Massachusetts; Honolulu; Leiden; New York; Paris; Tring;
and Vienna.
Status Extinct. Last seen in July 1898.
Range Hawaii, Hawaiian Islands.
Description 20cm (8in). Medium-sized with a very long, curved bill. Generally glossy black; bright yellow feathers on bend of
wing, rump and upper- and u ndertail-coverts; primary and u
nderwing-coverts white; four central tail-feathers black, rest
brown with patch of dull white near tip; bill black; legs and feet b lackish-brown. Juvenile browner, yellow areas reduced and
may have been absent in younger birds (Olson & Hume 2009).
The Mamo was described by Latham on the basis of two specimens in the Leverian Museum, said to be a
male and a female, which are now probably in the Vienna Museum (Pelzeln 1873). Latham erred in stating
that the bird came from the ‘Friendly Islands’ (i.e. Tonga), and writers such as Vieillot and G. R. Gray
repeated this error.
The Mamo’s yellow feathers were used in Hawaiian feather-work, and were the most highly prized of
all. It was a very tame bird, and easily captured by a method similar to that used by boys on Kaua’i to catch
finches for Townsend (1839) in 1835. This involved holding up a flower; as the bird dipped its bill into the
blossom the boy closed his finger over the flower and trapped the bird by its bill. Despite this intense
hunting, Munro (1944) was of the opinion that native capture for food and feathers would have had little
effect on its numbers. It is more probable that destruction of the forest and perhaps disease was responsible
for the bird’s ultimate demise.
Until the 1880s the Mamo was still relatively common, and Perkins (1903) was told that a local hunter
could shoot up to 12 in a day, but there may have been some confusion with the similarly coloured Hawaii
’O’o Moho nobilis (see p. 286). Perkins considered it extremely rare in the 1890s, and Wilson & Evans (1890–
99) and Wilson (1891) described the difficulties of obtaining a specimen:
Of this extremely rare and apparently extinct species I obtained two specimens from a collection which was
formed by the late Mr. Mills of Hilo in Hawaii, some thirty years or more ago . . . I could obtain no certain infor-
mation of examples having been observed since those in the Mills collection were procured – about 1859, though
while staying at Olaa in the district of Puna in Hawaii, where Mr. Mills secured them, I was assured by the
natives that the bird still existed, and at the time of my visit (October) had, together with the O’o, migrated to
the mountains, which is barely possible.
Habits
The Mamo was probably dependent on the native lobelias, where it took nectar (Palmer’s captive bird
sucked sugar and water readily), and Henshaw (1902) saw them actively hunting for insects in the tops of
tall Ohia trees. The showy b
lack-and-yellow Mamo was said to forage with the similarly striking black, red
and grey ’Ula-’ai-Hawane Ciridops anna (Pratt 2002; see p. 354), which must have made an impressive sight.
Perkins was given an imitation of the call by old bird-catchers, a single long plaintive note that so closely
resembled that of the Black Mamo Drepanis funerea, with which he was familiar, he had no doubt of its
correctness.
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The Black Mamo was one of the last of the Hawaiian finches to be described whilst still extant. It was
discovered on June 18, 1893 by Perkins, but was to succumb quickly to the same processes that had devas-
tated so many other members of its family. While describing the species, Alfred Newton (Newton 1893)
wrote with some foresight:
Its sombre plumage and the sad fate that too probably awaits the species induce me to propose for it the name of
Drepanis funerea.
Perkins (1895), as usual, gave excellent field notes about the Black Mamo, and quotes from his diary provide
an insight into a period just prior to the time when so many of the Hawaiian finches became extinct. Despite
all of his efforts, the Black Mamo was the only Hawaiian Finch that Perkins collected that was new to
science. He records taking a second specimen in 1893:
June 27th: The specimen obtained, I watched for some time before shooting. It was thrusting its bill under the
wet moss which covered the treetrunks, in search of insects [Perkins later was to recant this statement]. I
could tell that the long tongue
was being darted in and out,
and that so rapidly that it
appeared like a liquid streak,
the eye not being able to distin-
guish each separate movement.
It had not so silvery an appear-
ance as is familiar in our own
woodpeckers.
Perkins was told by an old
native woman in the Pelekunu
Valley that the local name was
Oonukuuma (‘the ’O’o with the
sucking beak’), and commented
that ‘like most other birds,
which have been thrust back to
the furthest depths of the
forest’, the Black Mamo was
very tame and would come
close to the observer out of
curiosity, even perching
directly overhead.
Perkins probably obtained
six specimens in 1893 (Bryan
1908), and the following year
Theodore Meyer, a resident of
Molokai, secured another six in
about six months of continuous
collecting. He found no more,
and believed the species to be
extinct. In May 1907, Alanson Black Mamo Drepanis funerea
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Bryan (Bryan 1908) set out to search other localities, and after a journey involving much hardship, found
some birds. He indicated that the bird was quite inquisitive and could be called down to an imitation of its
call. He secured a total of three specimens, all males. In March and April 1908 George Munro (1944) searched
for, but failed to find it. A survey in 1936 produced only a rumour that one had been seen in the Wailau
Valley a few years before. Munro (1944) claimed that William Donagho searched much of the remaining
forests of Molokai in 1937 and found no trace of it. Munro suggested that it might still survive on the plateau
between the Waileu and Pelekumu valleys, the only likely place that could support a population. However,
Richardson (1949) found no trace of it after an extensive survey in this area.
Habits
The Black Mamo was a very tame and inquisitive bird of underbrush and low forest trees, feeding on nectar
of arborescent lobelias. Captured specimens had heads smeared with lobelia pollen. They also probed wet
moss for water, and it appears that they never took insect prey. The long, curved bill was perfectly adapted
for probing into flowers, and Perkins (1903) noted the difficulties that this species had when trying to preen
itself:
. . . they will sit quietly preening their feathers, when they have a very comical appearance, much stretching of
the neck being necessary to enable them to reach the fore parts of the body with the tip of their long beaks.
According to Perkins (1895) it had a loud cry of remarkable clearness and a milder note which was probably
similar to that of the Mamo D. pacifica. Bryan (1908) described this call as hoa, and said that this was a name
also given to the bird by the natives.
Before collecting the last known specimens, Bryan (1908) made some important observations of the
bird’s habits:
This bird . . . confined its range to the undergrowth, several times coming down to within three or four feet of the
ground. At no time did it make a long flight or alight on the top of the trees. As it was raining all the while the
bird was especially active in preening and shaking its feathers. The trees and vines were everywhere covered with
thick wet moss, and although the bird hopped about from branch to branch, carefully inspecting each limb, I did
not see it catch any insects, or even probe into the moss. Hopping from tree to tree, it worked its way around the
head of the little side valley, up which it had come in answer to my call, to where a large purpleflowered lobelia
was in profuse blossom, and began to feed. The ease and grace with which the feat was accomplished was indeed
interesting, and left no doubt in my mind as to one of the probable causes of the remarkable development of the
tongue and bill. The tongue was inserted with great precision, up to the nostrils, in the flower, while the bird
balanced itself on the branches, assuming almost every imaginable attitude in its operations. In all three of the
birds secured, the crown was smeared with the sticky purplish white pollen of this lobelia.
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The ’Apapane H. sanguinea occurs on all of the main Hawaiian Islands; it is one of the few Hawaiian finches
that has maintained reasonable numbers, despite being susceptible to the same problems that have deci-
mated so many of its congeners. A closely related but distinct species, H. fraithii, once occurred on the remote
island of Laysan, which differed in having a shorter bill and overall paler coloration.
From 1890, Laysan was exploited for the guano deposited by the large colonies of breeding seabirds. By
about 1904 this had become unprofitable, and a Max Schlemmer decided to introduce rabbits, intending to
start a rabbit-meat canning business. This did not flourish, but by 1923 the rabbits had almost totally
destroyed the native vegetation, leaving a waste of sand with only an occasional tree (Bailey 1942). In that
year the rabbits were eradicated and the vegetation began to make a remarkable recovery, but it was too late
to save the Laysan ’Apapane. Although Dill & Bryan (1912) had reported a population of some 300 in 1911,
by 1923 only three remained; these were seen by members (including Alexander Wetmore) of the Tanager
expedition (Wetmore 1925b). A remarkable piece of film footage by Donald Dickey shows one of these birds
singing from a limestone pinnacle.
A few days later, a three-day gale swept the island, causing a spectacular sandstorm in which the last
three Laysan ’Apapane died. Although a number of searches have been made, no birds have been seen
since.
Habits
Fisher (1906) reported that the Laysan ’Apapane was:
Found all over the island, but is most abundant in the interior among the tall grass and low bushes, bordering
the open stretches near the lagoon, where all the land birds seem fond of congregating. Its favourite nesting place
is in the same area, and the proximity of broad patches, acres in fact, of a prostrate succulent portulacea with
yellow and a sesuvium with pink flowers has many attractions for the honey-eaters. Here they may be found
throughout the day walking around after small insects or drinking honey from the blossoms . . . It is not
uncommon to see one go from flower to flower and insert its bill between the petals of a nearly blown bud.
The nest was constructed in the middle of a grass tussock; made of fine grasses and rootlets, it was lined
with the brown down of young albatrosses. Nests containing one to four eggs were found between May 16
and 23, 1902 (Fisher 1903, 1906).
Munro (1944) noted that the birds actively sought water, and seemed to suffer from the lack of it more
than the other endemic Laysan birds. The only natural permanent water on the island was a seep of brackish
water, but after heavy rains large areas would be flooded (Bailey 1942). The ’Apapane fed on nectar and
insects – particularly caterpillars and adults of the large miller moth, which was common on the island.
When eating these moths it would hold them in its left foot and delicately pick at them, discarding the
wings and other hard parts.
Description 14cm (5.5in). Adult male has broad black mask extending behind eye; crown grey merging to dark brown on back;
scapulars and w ing- coverts dark rufous-
brown; primaries and secondaries dark brown; rump and uppertail-
coverts
rufous-brown; chin black; throat and breast white shading to g
reyish-white on breast; belly white, merging with deep cinnamon
undertail-coverts; bill glossy black; iris medium brown; legs and feet dark pink–brown; foot pads yellowish. Female similar but
duller, with smaller black mask.
The Po’ouli was discovered as recently as 1973, in the Ko’olau Forest Reserve on the north-eastern flanks of
the Haleakala volcano on Maui, at an altitude of 1,980m (Pratt 2005). Because of the distinctiveness of this
taxon, Pratt (1992) thought that it may not have been a Hawaiian finch at all, but some other type of
passerine; however, Fleischer et al. (2001) have shown that the Po’ouli is indeed a Hawaiian finch, although
it is phylogenetically unique, with its affinities to other members of the group uncertain.
On its discovery, the Po’ouli population numbered around 200 birds. This declined sharply over the next
decade, and by 1985 was estimated to have dropped by 90%. By 1995, the species was down to just five or
seven individuals, and by two years later, only three; one male and two females (Baker 2001).
Each had a definite, separate territory in the Hanawi Natural Area Reserve and the neighbouring
Haleakala National Park. The birds were no longer in contact, so an intensive effort to translocate a female
to the male’s territory was put in place in 2002 (Groombridge et al. 2004). After some months of logistical
preparation, a female was captured and released within the male’s home range, but the bird returned to its
own territory within one day. On September 9, 2004, one of the last three individuals was captured and
taken into captivity in the Maui Bird Conservation Center, Olinda, but it died on November 28, just two
months later (VanderWerf et al. 2006). Tissue samples were taken from the captive male in the rather forlorn
hope that the species may one day be cloned, which seems most unlikely. The last two wild birds disap-
peared in 2003 and 2004, with the last known individual dying in captivity, and as no other population has
been found since 1998, the likelihood that the Po’ouli still survives is extremely remote.
Why efforts to save the species were left for so long when its plight had been known for two decades has
yet to be adequately explained (see Donald et al. 2010).
Habits
Although the Po’ouli was restricted to h igh-elevation forests in historic times, the discovery of subfossil
remains in lower-elevation, drier habitats on Maui (James & Olson 1991; Reilly 1998) suggests that this
range contraction was a result of Polynesian habitat destruction. It was a cryptic bird, foraging on trunks
and branches of trees, where it gleaned invertebrates, especially snails, from the bark, lichens and moss
mats (Mountainspring et al. 1990).
The tongue of the Po’ouli was spoon-shaped, robust and unique amongst Drepanidini (Bock 1978; Pratt
1992), and appears to have been an adaptation for extracting the soft parts of Succinea snails (Bock 1978), a
choice food item that it foraged for in understorey vegetation (Baldwin & Casey 1983). It was often associ-
ated with Maui Parrotbill Pseudonestor xanthophrys and Maui ‘Alauahio Paroreomyza montana in mixed
feeding flocks within its home territory (Groombridge 2009).
The nest was an open cup made from twigs and bound together with moss, built in the stems of Ohia
Metrosideros polymorpha (Engelis et al. 1996). The eggs were speckled brown-grey on a whitish background.
The song was described as a quiet mixture of chittering notes, and the call a loud chirk, often repeated in a
short series.
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Almost nothing is known about this bird. The Hawaiian name means ‘the red bird that feeds on the Hawane
palm’. It was collected only five times, the last specimen being taken by Palmer on 20 February 1892 near
the headwaters of the Awini River on Mount Kohala. This specimen was originally preserved in spirit, and
its stomach contained undigested food (Olson 2012a).
Perkins (1903) believed that it was formerly widespread, having been found on the windward side of the
island in the Hilo District and the Kohala Mountains, and on the leeward side in the Kona district, but there
is no evidence that it ever occurred in the Kona District. Olson (2012a) has shown that it may have had only
a patchy distribution, and that it was inextricably linked with loulu palms Pritchardia sp., which are found
in clumps in forest (Perkins 1903). The severe decline in loulu palms due to deforestation, especially in the
lowlands, and seed-predation by the Pacific Rat Rattus exulans and pigs, drastically reduced the habitat of
the ‘Ula-‘ai-Hawane. The last surviving birds were forced into less suitable, marginal habitats, and had
almost certainly disappeared by the late 1890s. However, Munro (1944) thought that he saw one in 1937, but
he could not be sure.
Olson (2012a) has shown that probably four of the five known specimens were part of the James Mills
collection, all taken from the area above Hilo, Hawaii. Based on circumstantial evidence, Olson (1992b) had
previously considered that two of the specimens were taken on Molokai, but he no longer considers this to
be the case (Olson 2012a). A Ciridops species did once occur on Molokai, however, as subfossil remains have
been discovered, but it may prove to be conspecific with the ‘Ula-‘ai-Hawane (James & Olson 1991; Olson
1992b).
Of the five specimens, there are three plumages; one is scarlet, black and grey, one is predominantly
green, grey and brown, and one is red and black with much brown intermingled (Pratt 2005; Olson & Hume
2009; Olson 2012b). The green-plumaged specimen is almost certainly an adult female (and not an imma-
ture as suggested by Pratt 2005). This means the ‘Ula-‘ai-Hawane differed from other red-and-black
Hawaiian finches (Himatione, Palmeria, Vestiaria and Drepanis) in being sexually dichromatic (Olson & Hume
2009; Olson 2012a).
James & Olson (1991) and Olson (2012a) examined the skeletal morphology of Ciridops and noted that the
pelvis was broad and robust, as were the tibiotarsus and tarsometatarsus, much more so than in other red-
and-black Hawaiian finches. It also had comparatively large feet. Analysis of the musculature showed that
in comparison with ‘I’iwi Vestiaria coccinea and Apapane Himatione sanguinea, the muscles were markedly
better developed (Olson 2012a).
Habits
Very little was recorded about the habits of the ‘Ula-‘ai-Hawane. Emerson (1894), who presumably obtained
his information from native hunters, stated:
There was, I am told, another red-feathered bird called ula-ai-hawane, a beautiful thing in scarlet, wild and shy,
a great fighter, a bird rarely taken by the hunter.
Olson (2012a) compared the skeletal morphology of Ciridops with other bird genera and concluded that two
New Zealand species, the Whitehead Mohoua albicilla and Yellowhead M. ochrocephala, were the best
analogues. They have similar strong legs and feet, and feed in litter and debris that collects in the forks of
trees. The birds achieve this by gripping with one foot, and, using the tail as prop, scratch vigorously with
the other foot (Soper 1976; Olson 2012a). Olson (2012a) suggest that the robust hindlimbs and large feet of
Ciridops were an adaptation for foraging in the matted mass in the axils of loulu palm fronds, in which the
legs were used to move tough material to obtain food. The birds did not eat the fruit and blossoms of the
palm itself, but consumed invertebrates and small seeds occurring in the palm crowns. The stomach
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contents of the pickled specimen contained various invertebrate remains, including moths, spiders, plant
and bark lice and their eggs, fruit flies and small seeds.
The discovery of Ciridops species on islands other than Hawaii in the main Hawaiian group has shown
again how inadequate the historical record can be in determining the supposed biogeographical ranges of
taxa. The Kaua’i Palmcreeper was a smaller bird than C. anna of Hawaii, with a narrower, longer bill and
less robust legs, but still stouter than the other red-and-black Hawaiian finches (James & Olson 1991).
Because it did not possess the degree of characters exhibited in C. anna, it was presumably less derived from
other Hawaiian finch genera than its Hawaii congener, and James & Olson suggest that it possibly provides
a ‘missing link’ between C. anna and the rest of the red-and-black clade.
The Kaua’i Palmcreeper disappeared before the arrival of Europeans, no doubt a victim of Polynesian
habitat destruction and perhaps o ver-hunting for its feathers (which were probably red and black). The
subfossil remains were discovered in a Kaua’i S tilt Owl Grallistrix auceps pellet (see p. 226), which confirms
that they formed part of the diet of these raptorial birds.
Another series of Ciridops subfossil remains has been discovered at the coastal site of Barber’s Point,
Oahu. These bones are similar to those of C. anna but are slightly shorter in the bill and legs and the bill was
shorter and wider than C. tenax (James & Olson 1991). Further diagnostic material is needed before these
differences are sufficient to warrant treating the extinct palmcreeper population of Oahu as a separate
species.
The only known specimens of this finch came from Chichijima, or Peel Island, but it is possible that they
also occurred on Hahajima, where they may have survived until the 1880s (or so the naturalist A. P. Holst
was told by local residents Seebohm 1890a). There is some reasoning to doubt this statement, however, as
the bird was never reported there in the 1850s, when Mathew C. Perry surveyed the island (Perry 1856).
The types were collected in June 1827 during the voyage of HMS Blossom. Kittlitz (1832) visited the larger
Bonin Islands the following year, during which he collected additional specimens. According to Greenway
(1967), William Stimpson found none on Chichijima in 1854, and nor has anyone since. Stimpson was a
naturalist on the Rodgers-Ringgold North Pacific Exploring and Surveying Expedition, and although he
failed to locate the finch, he did report goats, sheep, dogs, cats and rats, which would surely have decimated
a predominantly terrestrial finch. The island had been used by whalers for careening ships since the 1830s;
the Bonin Islands Finch disappeared sometime between the visit of Kittlitz in 1828 and that of Stimpson in
1854.
There is considerable variation in the surviving skins, which show size and plumage differences, beyond
that explicable solely by sexual dimorphism. Although it is known for certain that specimens were taken on
Chichijima, it is equally possible that some were collected on other islands, and they may represent different
subspecies or even species (see Fuller 2000).
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Habits
Kittlitz (1832–33) recorded the only known details about the bird:
This bird lives on Bonin-Sima [= Chichijima], alone or in pairs, in the forest near the coast. It is not common,
but likes to hide, although of a phlegmatic nature and not shy. Usually it is seen running on the ground, only
seldom high in the trees. Its call is a single soft, very pure, high piping note, given sometimes shorter, sometimes
longer, sometimes singly or sometimes repeated. In its muscular crop and spacious gullet I found only small
fruits and berries.
There is considerable variation in the morphology of the House Finch C. mexicanus, with differences in body
and bill size, wing, tail and tarsus length, and degree of red in the male plumage. A number of subspecies
occur in the south-western United States, of which some are doubtfully distinct. McGregor’s House Finch
represents one such example of variation in plumage. McGregor (1901), in discussing dichromatism in the
genus Carpodacus, where yellow feathers frequently replace red ones, remarked:
Among the finches from the San Benito Islands . . . I have never seen two alike. The colors range from bright
crimson through orange to lemon yellow with all manner of variations resulting from combinations of these
colors and their shades or tints. It is impossible to take any example and say it is the typical coloration.
After having been considered numerous in 1895, the birds disappeared extremely rapidly. Kaeding (1905)
reported that the bird was practically extinct; it was last recorded in 1938 (Boswall 1978). The reason for its
disappearance is not clear, but cats were present on the island (Anthony 1925), and would have been a real
threat to the birds. San Benito was an unforgiving place, especially for a terrestrial finch, and Anthony
(1897) remarked:
There is little vegetation on this island, which is little more than a reef less than two miles in extent, and it is
rather surprising that a species of this genus should be found there at all.
Collar & Newton (2010) suggest that over-hunting by specimen-collectors was the primary cause of its
extinction.
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Habits
Almost nothing seems to have been written about this bird. It was found on San Benito and Isla Cedros;
McGregor (1898) found the birds distributed all over these two islands, but they were extremely shy and
difficult to collect. He was too late to find eggs, but collected three young birds. The nests were ‘miscella-
neous’ constructions of bark, twigs and fibre.
The Trías Greenfinch was related to the Goldfinch Carduelis carduelis and Greenfinch C. chloris, but differed
in its long, robust legs, which were probably an adaptation for terrestrial habitation (Alcover & Florit 1987).
It had a large, robust bill and probably fed on large seeds. These adaptations could evolve only in the
absence of predatory terrestrial mammals. The bird no doubt disappeared rapidly after humans arrived on
the islands, bringing cats and rats, but it may have survived into the historical period. Nothing else is
known about it.
The Slender-billed Greenfinch was sympatric on Tenerife with the Common Chaffinch Fringilla coelebs and
Blue Chaffinch F. teydea; differences in bill structure suggest that it occupied a different niche and had
different food requirements. Morphometrics obtained from the wing elements show that it was probably a
weakly flying species, and the long tarsi, which were equivalent in size with those of the Trías Greenfinch C.
triasi, indicate that it was a terrestrial species, perhaps foraging and nesting entirely on the ground.
The subfossil material was found in an area associated with the dense herb layer of the laurel forests, so
presumably the Slender-billed Greenfinch, with its long bill, foraged for seeds and perhaps invertebrates on
the ground (Rando et al. 2010). Reasons for its extinction are unclear but the arrival of humans and
commensal mammals would have proved devastating to a predominantly terrestrial finch. It may have
survived into the historical period, to become a victim of the introduced Black Rat Rattus rattus and Brown
Rat R. norvegicus.
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Endemic to the island of St Lucia, Semper’s Warbler was briefly described by Sclater (1876) on the basis of
specimens sent to him by the Rev. J. E. Semper, a clergyman resident on St. Lucia.
Although it may have once been common, it was very rare throughout the 20th century, and there have
been only five or six records since the 1920s. Bond (1950) stated that the last specimen was taken on the
summit of Piton Flore in 1934 by Stanley John. It was sighted between Piton Lacaombe and Piton Canaries
in March 1947, also by Stanley John. Although the last confirmed sighting was in 1961 (Keith 1997), uncon-
firmed reports were made as recently as 2003. A number of visitors to the island have searched diligently for
it over the years, without success (Collar et al. 1992).
It is believed that predation by the mongoose has been an important factor in the bird’s disappearance.
The mongoose was introduced to the island in 1884 to control the F er-de-Lance snake Bothrops atrox, but
both Bond (1962) and Wingate (1969) noted that the snake was still numerous in some areas of the island,
suggesting that mongooses were uncommon there and not the main threat to the bird. Towle & Towle (1991)
believed habitat destruction to be a major factor, much of the forest having been cleared for agriculture.
Habits
Rather little is known of the bird. Semper’s Warbler was found in virgin forest, where it foraged in the
undergrowth, close to the ground. Skutch (in Griscom & Sprunt 1957) stated:
Semper’s Warbler [is] a very plain nondescript species with dark grey upperparts and whitish underparts.
During many weeks exploration in the island I have never seen, nor even heard, this bird, which is probably on
the verge of extinction, although it was apparently not uncommon many years ago. Semper’s Warbler is known
to inhabit forest undergrowth and may nest on or very near the ground like the majority of tropical warblers, and
consequently has suffered from the depredations of the mongoose. Nothing is known of its song, but it is said to
chatter when alarmed.
The Bahama Yellowthroat G. rostrata is sometimes considered conspecific with the Common Yellowthroat
G. trichas, Belding’s Yellowthroat G. beldingi and Altamira Yellowthroat G. flavovelata; the genus is in need of
taxonomic revision. The nominate race of G. rostrata was restricted to New Providence in the Bahamas, and
appears to have been quite scarce even when first described (Bryant 1867). It was recorded as rare in the
1990s (Raffaele et al. 1998), and there appear to be no recent reports on its status. The Bahamas have suffered
from deforestation due to land development and agriculture, and the introduction of numerous mamma-
lian predators has impacted on many bird populations.
The New Providence birds might well now be extinct. However, another subspecies, G. r. tanneri, occurs
on the Abaco islands, and this form is considered by some authorities to be conspecific with the New
Providence species.
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Habits
This bird was an inhabitant of arid lowland scrub and bracken, building its nest close to the ground in dense
vegetation or tree stumps (Curson et al. 1994). Stomach contents of a male contained the remains of an anole
lizard Anolis sp., but invertebrates were usually taken.
The Jamaican Oriole Icterus leucopteryx occurs on Jamaica and San Andrés Island (off Colombia); it is not
considered threatened. However, the Grand Cayman subspecies I.1. bairdi appears to have been affected by
severe deforestation. It was reported to be extremely rare in 1911 by the collector W. W. Brown (Bangs 1916),
who still saw fit to take 17 specimens. By contrast, Savage English (1916) found it not uncommon in the
north of the island. It was observed in 1938 by C. Bernard Lewis during the Oxford Expedition (Johnston et
al. 1971), but not recorded by Johnston (1969) during three thorough expeditions in 1965, 1966 and 1967.
However, Patricia Bradley (Bradley 2000; Bradley & Rey-Millet 2013) was shown photographs and field
notes by a local resident, Bernard St Aubyn, who had photographed it in his George Town garden in May
1967. Unfortunately the photographs and field notes were subsequently destroyed (P. Bradley pers. comm.
2012). The Grand Cayman Oriole is now certainly extinct.
Habits
Brown (in Bangs 1916) found it scattered here and there at wide intervals over the island, and a nest made of
palm fibres and attached to a hemp palm leaf about at a height of 18m from the ground was found in late
May; the nest contained three dependent young. Savage English (1916) gave the final account:
Icterus bairdi, one of the Troupials, is by no means uncommon in the north of Grand Cayman and is usually seen
in small parties of five or six birds, hunting for food in the crowns of Coconut and Thrinax palm trees. These
parties are presumably families, for, though three years was not long enough to enable the writer to find a nest,
there can be little doubt as to this bird’s breeding somewhere on the island . . .
Habits
Haemig (2010) discovered new information about the ecology of the bird, noting that the grackle nested in
emergent aquatic vegetation, in cultivated areas and also in towns, but whether it was dependent on a
marsh habitat is not known. It was abundant, and formed flocks, which did considerable damage to maize
crops.
Three eggs, supposedly to be of this species, were collected by A. Forrer at Mazatlan, West Mexico, but
the locality is not within the range of Q. palustris, so they are likely to belong to another species.
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The Seaside Sparrow Ammodramus maritimus ranges from Massachusetts along the Atlantic and Caribbean
coasts of the United States to Texas and the mouth of the Rio Grande. The species is sedentary, and nine
subspecies are recognised, some of very limited distribution. The Dusky Seaside Sparrow is one of the
saddest of recent extinctions, and a bitter indictment of modern civilisation. It had one of the most restricted
ranges, if not the most restricted, of any North American bird.
Pesticide problems
Its decline was probably due to the spraying of the salt marshes in which it lived with DDT to control
mosquitoes; this took place from 1942 to 1953. By 1957 this had caused the sparrow population to crash by
at least 70%. Ingestion of even tiny amounts of DDT can cause reproductive failure in birds, and this was the
root cause of the decline. Spraying was discontinued after 1953, and the area impounded by earthen dykes
thrown up round the perimeter of the salt marshes, with the result that they were covered with 10–30cm of
fresh water. Naturally this altered the vegetation markedly, and the rushes in which the sparrows tradition-
ally nested began to disappear in the lower parts of the marsh. Delaney et al (1981) recorded:
On the St. John’s marsh, drainage, housing developments, and highway construction all contributed to habitat
loss. Extensive wildfires during the dry season (December–April 1973–1976) also contributed to the population
decline.
Habits
The Dusky Seaside Sparrow was restricted to salt marshes, where the birds foraged for insects, marine
invertebrates and seeds on the ground or in marsh vegetation, sometimes probing in the mud. The nest was
an open cup, usually built among tidal reeds and spartina grasses; between two and five eggs were laid. The
call was described as a raspy buzz.
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Worthen’s Sparrow Spizella wortheni is one of the least known and most endangered birds of the United
States. When Webster & Orr began their studies in the 1950s, the Zacatecas race, browni, was reasonably
common but restricted in distribution. A single bird was collected in 1942, which had hitherto been unrec-
ognised, four pairs were recorded in 1952, at least eight in 1954, with the final observation made in August
1961, when two immatures were collected (Wege et al. 1993). Searches in 1991 were unsuccessful. The disap-
pearance of this sparrow was a result of habitat loss to agriculture, overgrazing and erosion from cattle
herding, and a decline in native herbivores which possibly maintained the bird’s habitat (Behrstock et al.
1997).
Habits
The Zacatecas Worthen’s Sparrow was found in grassy pasture sparsely dotted with small mesquite Prosopis
up to a metre tall, Juniper Juniperus deppeana, and a few other shrubby species at an altitude of between
2,350m and 2,450m (Webster 1954; Webster & Orr 1954). At least eight pairs were recorded in 1952 (Webster
& Orr 1954), with an average territory of 20–25 hectares per singing male, in a 200-hectare tract of habitat
(Webster 1954, 1958). The males were singing from waist-high bushes. Three specimens were taken in
breeding condition in June, and the two immature specimens were taken in August. The peak period for
breeding activity was between June and July, when singing males, pairs behaving territorially, and nest-
building was observed (Wege et al. 1993).
The Spotted Towhee P. maculatus is widely distributed across the eastern United States. A subspecies once
occurred on the island of Guadalupe. The last specimens were collected in 1855 (Bryant 1887) and one was
sighted in September 1896 (Gaylord 1897). Thoburn (1899) was probably the last person to see one, in June
1897. Howell & Cade (1954) searched for it in vain, confirming that the subspecies had vanished.
It was probably never a common bird, and was recorded as uncommon by Palmer (in Ridgway 1876b).
All the scrub and undergrowth in the cypress groves where the birds lived was destroyed by goats, at a time
when the birds were already in serious decline. Feral cats were also introduced, potentially lethal to any
ground-nesting bird, and there seems little reason to doubt that these two factors were primarily respon-
sible for the towhee’s extinction.
Habits
The Guadalupe Towhee occurred in groves of cypress on the plateau below Mount Augusta, and in later
years appeared to be confined to that area. Earlier, Palmer in 1875 had found the birds generally in woods,
and frequenting brushwood, fallen logs, fences etc., rather than trees, suggesting they were found
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elsewhere on the island (Ridgway 1876b; Greenway 1967). The diet comprised seeds and insects. The nest
and eggs were never described, but it probably nested close to the ground. The song included a unique trill,
not recorded in the nominate on the mainland, which resembled that of a Western Bluebird Sialia mexicana
(Greenway 1967).
The Bermuda Towhee was similar to the Eastern Towhee Pipilio erythrophthalmus, but differed in having a
heavier bill, the maxilla and mandible being wider, reduced wing and pectoral girdle, and more robust legs
(Olson & Wingate 2012). Olson & Wingate further noted that the sternum was shorter and wider, with a
much smaller keel than the mainland bird. As towhees are ground-dwelling inhabitants that occur in forest
understorey and scrub (Rising 2011), it is likely that the Bermuda Towhee once inhabited a similar habitat
on Bermuda. The reduction of the sternum and wings and an increase in robustness in the limbs suggest
that the Bermuda Towhee was weakly volant, but still capable of flight. Similar ratios of pectoral, pelvic,
wing and leg elements are present in the Mascarene starlings, for example, (Hume 2014b), all of which were
capable of flight.
The Bermuda Towhee was described just once in life in 1609–1610 by an exiled seaman, William Strachey
(in Olson & Wingate 2012). He stated:
Sparrowes fat and plumpe like a Bunting, bigger then ours.
Extinction
Pigs were already on Bermuda when Strachey encountered the towhee in 1609–1610 (Olson & Wingate
2012), and rats and cats arrived around 1614 (Sterrer et al. 2004). Land clearance also began in earnest, so it is
no surprise that the Bermuda Towhee disappeared extremely rapidly, at least by 1625, and especially as it
was a probable ground-nester and feeder.
The fossil record has shown that the Bermuda Towhee was one of the longest-surviving passerines on a
small island, and it had remained relatively morphologically unchanged for 400,000 years (Olson & Wingate
2012). It was present during an interglacial with associated high-magnitude sea-level rise, when Bermuda
Bermuda Towhee
Pipilio naufragus
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was reduced to a few small islets (Olson & Hearty 2009), and it survived the sea-level fluctuations that have
beset the Bermuda platform ever since. The case of the Bermuda Towhee is a tragic example of vulnerability
in an insular species; upon the arrival of human beings and their commensal animals, extinction can over-
come a species in less than a decade.
The Santa Barbara Song Sparrow was a race of the Song Sparrow M. melodia, one of several occurring on the
islands off the coast of California. Cookman (1930), although remarking on the island being well-forested,
recorded that feral pigs and cats had become established. It is likely that pig-induced habitat change and cat
predation were the main causes of extinction. According to Garrett & Dunn (1981), the sparrow was last
seen in 1960.
Habits
These sparrows occurred in brush cover on the slopes and ravines, and in long coarse grass. Wright &
Snyder (1913) remarked that it was distributed all over the island, wherever bushes provided cover. Grinnell
& Miller (1944) found it still abundant, and added that bushes were used for nesting.
The Rufous-crowned Sparrow A. ruficeps occurs across the south-western United States and Mexico,
including islands off the coast of Baja California. Its distribution, however, is discontinuous, with a number
of isolated populations considered subspecifically distinct. The Todos Santos race occurred on the tiny
Todos Santos Islands, off Baja California, where it was considered common in the early 20th century.
However, it declined due to habitat destruction by rabbits, and the introduction of cats. It was terrestrial in
its habits, which made it particularly susceptible to these predators.
This bird has not been recorded since the 1970s. Mellink (1992) made a thorough search for the species in
1991, using bird-call recordings, but without success. It is probably now extinct (Donlan et al. 2000).
Habits
The birds foraged on or near the ground or in low branches and low shrubs. They would peck or scratch at
leaf litter to obtain their food, which comprised insects and seeds, and often walk or hop under low shrubs
or dense grasses, only flying when absolutely necessary.
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The Antioquia Brush-finch is a poorly known species. It was scientifically described in 2007 from three
museum specimens from Antioquia, Colombia (Donegan 2007). The specimens were previously labelled as
the closely related Slaty B rush-finch A. schistaceus. The three skins were all collected in the 20th century;
only one includes a collection date, 1971. All were collected on the Llano de Ovejas, a small plateau in
forests between 2,400 and 2,800m.
Subsequent ornithological surveys in 2008 and 2009 in Antioquia have failed to find the species (BirdLife
International 2011), and it might be now extinct. The type locality has suffered major deforestation due to
cattle pasture development and commercial flower cultivation, and unless another population exists else-
where, the likelihood of the bird’s survival is remote. However, another brush-finch, the Pale-headed
Brush-finch Atlapetes pallidiceps, was not seen for 29 years prior to its rediscovery in November 1998 (Agreda
et al. 1999; see p. 509), so there is some hope for the Antioquia birds.
Cayman Brac, along with Grand Cayman and Little Cayman, is situated between Cuba and Jamaica in the
West Indies. Cayman Brac was thought to have an avifauna similar to the other Cayman Islands, apart from
the endemic subspecies of Red-legged Thrush Turdus plumbeus coryi (Bradley & Rey Millet 2013). However,
Patton (1966) discovered thousands of vertebrate subfossil remains in a limestone cave called Patton’s
Fissure, located in the village of Spot Bay, Cayman Brac. Among the remains were two species of Melopyrrha
bullfinches, one referable to the Grand Cayman Bullfinch M. nigra taylori, which is now found only on
Grand Cayman (Bradley 2000; Bradley & Rey Millet 2013), and a larger species, the Cayman Brac Bullfinch
M. latirostris, which is now extinct. The Cayman Brac Bullfinch is known from at least 14 individuals, based
on the number of jaw elements; the maxilla and mandible were larger than the Grand Cayman Bullfinch,
especially in the width of the maxilla and depth of the mandible (Steadman & Morgan 1985). The maxilla,
quadrate and mandible of the Cayman Brac Bullfinch are broader and more massive than in other West
Indian finches, so it may have fed on a mixed diet of fruit and seeds.
Steadman & Morgan suggest that M. latirostris evolved from a population of the Grand Cayman bird, or
its immediate progenitor, which became isolated on Cayman Brac. The Grand Cayman Bullfinch recolo-
nised Cayman Brac after the establishment of M. latirostris, and may have hybridised with it, before
disappearing again at some time during the Holocene. No bullfinches occur on Cayman Brac today.
The Cayman Brac Bullfinch may have survived into the historic period and disappeared due to defor-
estation. Steadman & Morgan further suggest that M. latirostris may have been genetically swamped out by
interbreeding after the more recent arrival of M. n. taylori.
Thraupidae (Tanagers)
The tanagers are a diverse group of birds occurring in North and South America, with the greatest diversity
in South America. There are more than 100 genera with about 240 species, but the taxonomic placement of a
number of genera may be incorrect. They are small to m edium-sized birds, many being brightly coloured,
but the family includes the dull b lackish-brown Galápagos finches, made famous by Charles Darwin and
his theory of natural selection. Many species are poorly known, and the forest- and island-inhabiting tana-
gers are threatened by habitat destruction and introduced mammalian predators. Two species and four
subspecies have become extinct, including one subspecies of Galápagos finch.
The Large Ground Finch is divided into two subspecies, the nominate, which is endemic to Floreana and
San Cristóbal, and G. m. strenua, which occurs on several of the other islands. The nominate had the largest
bill of any of the Galápagos finches, and was slightly larger than G. m. strenua. It was first collected by
Charles Darwin and others from Floreana and San Cristóbal during the voyage of the Beagle in September
1835, and was described by Gould (Gould 1837b) from seven specimens. But Darwin was inconsistent in his
specimen labelling, and his locality data-labels were considered entirely unreliable (Swarth 1931; Hellmayr
1938). However, Frank Sulloway (Sulloway 1982) examined manuscripts and specimens of Galápagos
finches collected by Captain Fitzroy, Harry Fuller, who had accompanied Darwin on James Island, and
Syms Covington, Darwins’s servant. Unlike Darwin, they all recorded the provenance of their specimens,
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including G. m. magnirostris, and collected specimens of this form on both Floreana and San Cristóbal, thus
confirming its provenance.
Extinction
Steadman (1986) provides strong evidence to suggest that Darwin’s Large Ground Finch became extinct
shortly after the visit of Charles Darwin in 1835, which constitutes its only historic record. After humans
settled on the island in 1832, the island was rapidly altered, especially in the lowlands, which was almost
certainly the preferred habitat of the finch. Furthermore, the introduction of donkeys, cattle and goats
resulted in a scarcity of Opuntia cacti in the lowlands (Townsend 1930; Dawson 1962; Hamann 1975), which
would have affected the finch’s nesting sites and food supply (Steadman 1986); introduced dogs, cats and
rats would have been serious predators. Darwin appears to have collected the birds when still relatively
common, but reports from visitors to Floreana in 1838, 1846 and 1852 failed to find it (Lack 1947; Sulloway
1982; Steadman 1986). It probably became extinct within a decade or two after 1835 (Steadman 1986). On
San Cristóbal, Darwin’s Large Ground Finch has not been recorded since the visit of the Beagle in 1835
(Sulloway 1982). As the island was not settled until a decade or so later, it may have survived there a little
longer than it did on Floreana (Steadman 1986).
Bowman (1961) recorded collecting a single specimen of G. m. magnirostris on Floreana in September
1957, but Sulloway (1982) showed that Bowman had in fact encountered a small population of G. m. strenua
that had recently colonised the island.
Another undescribed large ground finch, the Sharp-beaked Ground Finch Geospiza nebulosa ssp. became
extinct on Floreana. This may have been an endemic (Steadman 1986); it had the largest bill of any of the G.
nebulosa races. Three skins were collected by the Beagle crew in 1835, two by Darwin (one now lost) and one
by Fitzroy, and the fourth and last was taken in 1852 (Sulloway 1982). Steadman (1986) also collected
subfossil remains. Sulloway (1982) and Steadman (1986) suggest that the Floreana Sharp-beaked Ground
Finch disappeared due to a combination of habitat destruction, predation by introduced predators, and
competition from other Darwin’s finches, which invaded the habitat of G. nebulosa once the forests were
cleared in the highlands.
Habits
Almost nothing is known about the habits of Darwin’s Large Ground Finch. Steadman (1986) found that
subfossil remains of this bird were the most abundant of all ground-finch material in the Floreana fossil
record, making up 75% of the total finch fauna, which is testament to its former abundance. It was an
extremely tame bird, and Darwin saw a small boy killing them with a stick near a well on Floreana
(Greenway 1967). Darwin (in Gould 1841) records that ‘he seldom saw them in the upper and damp regions’,
inferring that they usually inhabited the arid lowlands, which is supported by fossil evidence (Steadman
1986). Steadman further suggests that Darwin’s Large Ground Finch relied heavily on the seeds and fruit of
the endemic variety of Floreana cactus Opuntia megasperma var. megasperma. The fruit and seeds of O.
megasperma are the toughest of all Galápagos Opuntia, therefore only the massive bill of the finch could
utilise this food source.
The Puerto Rican Bullfinch Loxigilla portoricensis occurs on Puerto Rico, where it is considered not threat-
ened. A large, distinct race, L. (p). grandis, given full specific status by some authors (Garrido & Wiley 2003),
once inhabited the island of St Kitts, but is now extinct.
Known locally as the Mountain Blacksmith, it was not uncommon in the forests of Mount Misery in 1880,
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when Ober collected the type series (Lawrence 1882). However, it was nearly extinct by 1920, with the last
known specimen being taken by Paul Bartsch on July 26, 1929 (Olson 1984).
Bond (1950) considered the bird to have been exterminated by the introduced African Green Monkey
Chlorocebus aethiops, but a number of other races of the bullfinch survive on other West Indian islands, and
on Barbados L. p. barbadensis survives alongside the same monkey. Raffaele (1977) pointed out that Bond’s
suggestion was most unlikely, as monkeys have been wild on St Kitts for some 300 years, being established
from about 1700 to the present day (McGuire 1974). The monkeys flourish in the mountain ravines, but are
rare high up in the mountains where the bullfinch occurred. It seems very unlikely that these monkeys
would have caused the extinction of the bullfinch at so late a date, and furthermore, they would be expected
to have had more effect on the distribution of bird species in the lower mountain forests, with which they
apparently coexist.
Sympatry
Olson (1984) suggested that the St Kitts Bullfinch was probably not just confined to St Kitts in the past, and
occurred elsewhere in the archipelago including the neighbouring islands of Nevis and St Eustatius, which
would have been joined to St Kitts during the lower sea-levels of the last glacial. Although no subfossil
remains of the St Kitts Bullfinch have been found on Nevis and St Eustatius, Olson was eventually proved
correct as a rostrum of this species, or a closely related one, was discovered on Barbuda in a prehistoric
fossil locality (Steadman et al. 1997). Olson further suggested that the restriction of L. (p). grandis to the high
mountain forests might have been due to habitat destruction of the lower forests. The Lesser Antillean
Bullfinch L. noctis arrived on St Kitts at some point during the late 1800s, and apparently inhabited the
lower forest belt, while the St Kitts Bullfinch was confined to the mountain forests (Garrido & Wiley 2003).
The suggestion that the presence of the smaller L. noctis was detrimental (Raffaele 1977) appears to be
unfounded, however, as their former presence on Barbuda shows that both the St Kitts Bullfinch and the
Lesser Antillean Bullfinch were once naturally sympatric in the northern Lesser Antilles (Steadman et al.
1997).
Before the discovery by Olson (1984) of the last specimen taken in 1929, Raffaele (1977) hypothesised that
the bird was exterminated by two disastrous hurricanes, which hit St Kitts in 1899. Although inaccurate,
severe deforestation such as has taken place on St. Kitts does exacerbate the effects of hurricanes on bird
populations, and it may have reduced the already declining population. There seems little doubt that the
bird is now extinct, since Raffaele in 1972, Browning in 1977 and Steadman in 1982 all failed to find it (Olson
1984).
Habits
Nothing is known of the habits of the St Kitts Bullfinch. However, it may have been similar to the Puerto
Rican Bullfinch, which is secretive and found in dense forests. It is an opportunistic feeder, taking seeds,
fruits, insects and spiders.
The Eastern Chat Tanager is endemic to the island of Hispaniola, where it separates into four subspecies.
The races C. f. neibae in the Cordillera Central and C. f. tertius (formerly considered a separate species) in
southern Haiti and south-west Dominican Republic are both locally common but declining. The Gonâve
Island population C. f. abbotti is probably extinct (see below), as is the Samaná Peninsula race, C. f. frugivorus,
which has not been reliably recorded since 1982 (Raffaele et al. 1998). The prime reason for its disappearance
is deforestation due to logging and conversion of the forest to agriculture.
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Habits
The Samaná Eastern Chat Tanager was a predominantly terrestrial species, occurring in broadleaved forest
and dense understorey thickets, especially in ravines and adjacent to water. It inhabited forest in areas
below 600m, which has made it particularly vulnerable to the impact of deforestation. Breeding apparently
occurred in May to July (Raffaele et al. 1998), and the diet comprised a variety of invertebrates. The tana-
ger’s large, strong feet were adapted for searching through the leaf litter (Latta et al. 2006). The call was
described as a low slurred or whistled swerp, swerp, chip, chip, chip . . ., sometimes accelerated into a chatter
(Hilty 2011).
This race of the Eastern Chat Tanager C. frugivorus was restricted to Gonâve Island off Haiti, where it was
formerly common (Danforth 1929), but the population has not been recorded in recent years, despite exten-
sive searches (Latta et al. 2006). It is now probably extinct. The island has suffered severe deforestation, and
little suitable habitat remains.
Habits
The birds inhabited semi-arid scrub in areas below 600m. Danforth (1929) found the birds difficult to
observe in the dense undergrowth, but not shy, and they could be located by their song, which he described
as a rather buzzy, whistled wee–chee–chee–chee–chee. The stomach contents of two collected males comprised
mainly insects, including moths, ants, spiders and a cockroach, plus a few seeds.
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The accounts
Megapodiidae (Megapodes)
Anderson’s description of this bird is brief and virtually unidentifiable: ‘Fusca nigraque; pedibus nudis [brown
and black]. New Caledonia’. Gray (1862a) added:
The name given above (with the very short specific characters) was found among others in the manuscripts of
Anderson, who was assistant surgeon during the third voyage of the famous circumnavigator Cook. Though the
description is so short and concise, I am, however, induced to suppose that it can only be referred to a species of
Megapodius; certainly it cannot be reconciled with any of the present known birds from New Caledonia. I refer
to it in the hope that this slight indication may lead to its being searched for by collectors . . .
Gray (Brenchley 1873) placed M. andersoni in the synonymy of the extinct New Caledonian Wood Rail
Tricholimnas lafresnayanus (see p. 109). Balouet & Olson (1989) mention, but do not overly support,
Anderson’s account based on the available evidence, but recent work (Steadman 2006a) has shown that
another (as yet undescribed) megapode once occurred alongside the extinct P ile-
builder Megapode
Megapodius molistructor (see p. 39). The possibility exists that Anderson saw either one of these.
Cheeseman (1891) visited the Kermadec Islands in August 1887 and, in reporting on the avifauna, mentioned
that no endemic terrestrial species of bird is known to inhabit the islands, the land birds of which all seem to
be shared with New Zealand. This contrasts greatly with Norfolk Island, which is the same distance from
the New Zealand. This may be because successive volcanic eruptions repeatedly wiped out the fauna,
necessitating constant repopulation and precluding endemism.
Cheeseman
Following discussions with a Mr. Johnson, who resided on Sunday Island for about 15 years prior to 1887,
Cheeseman (1891) wrote that:
. . . when he [Johnson] lived on Sunday Island, which was prior to the eruption of 1876, a bird inhabited the
floor of the large crater which made mounds of sand and decayed leaves 2ft. or 3ft. high, laying its eggs in the
mounds. He was in the habit of visiting the mounds for the sake of the eggs and young birds, and has frequently
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taken five or six of the latter from the same nest at one time. The eruption of 1876 covered the floor of the crater
with a deposit of mud very similar to that thrown out by the eruption of Tarawera, and apparently killed out the
species, for it has not been seen since. The evidence, such as it is, seems to point to the former existence of a
species of Megapodius. It is worth mention that in the crater-basin of Niuafou, one of the Tongan Islands, and
which is not further removed from Sunday Island than the mainland of New Zealand, a species of Megapodius
has long been known to exist.
Buller (1905), based on Cheeseman’s work, considered this to provide evidence for the existence of the
Tongan Megapode Megapodius pritchardi in the New Zealand area. This, however, is untenable on zoogeo-
graphic grounds, and the habits of the two birds seem to have been different the Tongan Megapode on
Niuafo’ou lays in burrows, while the Sunday Island bird described by Cheeseman was said to have built
mounds of sand.
Lister (1911) referred to Cheeseman’s account, and commented:
We may remark that the statement that five or six young birds were taken from the same nest at one time is
hardly in accordance with the habits of the genus, for the eggs being laid at some intervals the young ones are not
of the same age, and leave the mounds to feed themselves as soon as they are hatched. The statement would be
more appropriate to the young of the [Pacific] Grey Duck (Anas superciliosa) which frequents the island. It is
the mound-building habit which, as Mr. Cheeseman says, ‘seems to point’ to the existence of a Megapode on
Sunday Island.
Lister went on:
I have had the good fortune to meet at the Natural History Museum [in London] Mr. Iredale, who has recently
resided on the Kermadecs with the object of studying their fauna. He assures me that he was not able to obtain
any confirmation of the report of the existence of a megapode on Sunday Island which was given to Mr.
Cheeseman, and finds that the successors of his informant are not inclined to regard that report as worthy of very
serious consideration.
Nonetheless, the strong possibility exists that a megapode formerly inhabited Sunday Island; the subfossil
record shows that many, if not most, major Melanesian and western Pacific island groups were once popu-
lated by megapodes (Steadman 2006a).
This species has puzzled ornithologists for more than a century. The feather cannot be identified with any
other Argusianus, but it may or may not represent a distinct species. The feather was procured from A. D.
Bartlett, the superintendent of the Zoological Gardens, Regent’s Park. It was presented to the then British
Museum by his son Edward Bartlett; that is as much as we know about the origin of the feather, though
Wood (1871) said that it was found among a box of loose feathers of the Great Argus A. argus. Delacour
(1951a) stressed that there was little doubt that this elaborate feather belonged to a still-undiscovered
species of Argusianus, and did not represent an aberration.
(1983) concluded that the Two-banded Argus had poor aerodynamic properties, and therefore the species
was probably flightless (though Parkes 1992 disputes this). Delacour (1951a) suggested Java as the prove-
nance, as this represents a large gap in the range of the Great Argus (which occurs on Peninsular Malaysia
and on Sumatra). Davison (1983) disagreed, as the avifauna of Java was well-known long before the
discovery date of A. bipunctatus, and he hypothesised, without supporting evidence, that the bird probably
came from the island of Tioman. Regardless of these fanciful interpretations, the feather’s provenance
remains completely unknown, although recently Davison & McGowan (2009) came to the conclusion that
the most likely explanation of the feather is that it is an anomalous growth from a Great Argus, as Parkes
(1922) had suggested.
Little can be said about a brown duck collected on Newfoundland. No bird that matches the descriptions
occurs there now, and McAtee (1963) was unable to identify it with any known species. Pennant (1785)
gives no reference for his brief description:
Duck with a large bluish bill: head and neck of a very pale brown: lower part of the last, and breast, of the same
colour, edged with rust-colour: wings cinereous grey: speculum blue, tip’t with white: tail and legs dusky:
Inhabits Newfoundland.
Latham (1824) adds little more:
This has a large, bluish bill; head and neck very pale brown; lower part of the last and breast the same, the
feathers edged with r ust-colour; wings cinereous grey; speculum blue, tipped with white; tail and legs dusky.
Inhabits Newfoundland.
Stephens (1824) provides some extra detail, but on rather spurious grounds:
Inhabits Newfoundland. 16 inches; beak large, stout at its base, bluish, with its tip black; the nostrils are placed
near the end; the head and neck are very pale brown; the lower part of the last, and the breast the same; the
feathers tipped with white; the tail and legs dusky.
Nothing else is known of this bird, yet, in view of the extinction of the Labrador Duck Camptorhynchus labra-
dorius (see p. 60), the Newfoundland Duck may have existed but disappeared even earlier.
Barrow (1807) was the only person to mention the St Paul Island Duck. He described it as small and brown,
and not much larger than a thrush. He also stated that it was the favourite food of the five sealers then living
on the island (Bourne et al. 1983). These sealers remained on the island for several years, and as nothing
more was heard of the duck, one must presume that they killed them all. Bourne et al. (1983) point out that
by the time of Barrow’s visit in February, northern winter visiting ducks should have already been in
breeding condition, so that the description of the St Paul bird as ‘small and brown’ suggests a resident
island population without sexually dimorphic plumage. No fossil material has yet been found, so it is
uncertain if this population differed from the Amsterdam Island Duck (see p. 54). Olson and Jouventin
(1996) suggest that the Amsterdam and St Paul Island ducks may have had a common ancestor, but if the
birds on St Paul were flightless, as was the duck on Amsterdam Island, they would have evolved
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flightlessness independently. This would have almost certainly resulted in the St Paul Island duck being
morphologically distinct from the Amsterdam Island population.
Cathartidae (Vultures)
The case history of this bird is remarkable, and its status still remains a mystery (e.g. Allen 1871).
Bartram
In the course of his travels along the St John’s River, above Lake George, Florida, William Bartram described
a number of birds. All of them are well-known birds in the area today, except for the Painted Vulture, which
nobody has subsequently seen. Bartram described his mysterious bird as follows:
. . . near the size of a turkey buzzard, but his wings are much shorter . . . the head and neck bare of feathers nearly
down to the stomach, where the feathers begin to cover the skin, and soon became long and of a soft texture,
forming a ruff or tippet . . . the bare skin on the neck appears loose and wrinkled, which is of a deep bright yellow
colour, intermixed with coral-red; the hinder part of the neck is nearly covered with short, stiff hair; and the skin
of this part of the neck is of a dun_purple colour, gradually becoming red as it approaches the yellow of the sides
and foreparts. The crown of the head is red, there are lobed lappets of a reddish orange colour, which lay on the
base of the upper mandible . . . The plumage of the bird is generally white or cream colour, except the quill
feathers of the wings and two or three rows of the coverts, which are of a beautiful dark brown, the tail which is
large and white is tipped with this dark brown or black; the legs and feet of a clear white, the eye is encircled with
a gold coloured iris, the pupil black.
Description 25–30 cm (9.8–11.8in). Head, neck and breast rich chestnut; mantle brownish-olive; chin and throat whitish; rest of
upperparts and tail dark olive-brown; belly, flanks and to undertail-coverts dark brown with fine, buffy barring; bill green or
yellowish-green, with blue-grey base; iris red; legs and feet olive. Differed from nominate in being smaller, being more heavily
barred with buff on the abdomen, and having shorter wings and tarsus.
The Red-necked Rail Rallina tricolor is an elusive medium-sized rail that occurs in eastern Indonesia,
Australia, and New Guinea and its outlying islands (Ripley 1977; Taylor & van Perlo 1998). It has been
divided into six subspecies, but there appears to be much individual variation and range overlap. The
Bismarck Islands subspecies was first described in 1925 from a male collected on New Ireland (Stresemann
1925), and the species was considered rare there. Two specimens were also collected on New Hanover
Island in 1944 (Mayr 1949; Ripley 1977), and these are the last known (Mayr & Diamond 2001). This elusive
bird may still exist in the Bismarck archipelago, but the lack of recent records makes its continued survival
somewhat precarious. Ripley (1977) noted that the Bismarck Islands Red-necked Rail had shorter wings
and tarsus than the nominate, and suggested that it was in the process of becoming flightless. This would
have made it more vulnerable to human disturbance and predation by introduced mammals.
The unique type specimen of this bird was obtained along with a number of others during a voyage to the
Solomon Islands prior to 1869 by the yacht Chance, owned by a Mr J. A. Buttray. The bird was probably
personally collected by John Brazier, a conchologist who also collected birds, which he preserved in spirits.
He made several expeditions to the Bismarcks (Sharpe 1906a). The Solomon Islands collection was described
by Sclater (1869a), at which time the ornithology of the islands was poorly known.
It subsequently transpired that a number of the birds were of species known only from New Ireland,
where, it was presumed, the yacht must have called during the course of the voyage (Mayr 1933b). It is
possible therefore that the rail may have come from there. If this is correct, the Solomon Islands B
are-eyed
Rail would be synonymised under the nominate G. p. plumbeiventris (BirdLife International 2011), a move
that has been followed by several authorities (e.g. Sharp 1894; Dickinson 2003). But as no further specimens
exist, and the provenance could well be the Solomon Islands after all, Taylor & van Perlo (1998) conclude
that intactus may represent a valid but overlooked subspecies. Despite its somewhat different appearance to
other Bare-eyed Rails, Dutson (2011) mentions the alternative possibility that it might be a vagrant.
The isolated Kermadec Islands are situated about 1,000km northeast of North Island, New Zealand.
Although remote, Raoul (also called Sunday) Island once harboured some probable endemic landbirds,
including a megapode Megapodius sp., a Hemiphaga pigeon (see p. 189) and a rail Hypotaenidia philippensis
ssp., all now extinct (Sorenson 1964; Worthy & Brassey 2000). Cheeseman (1891) interviewed an island resi-
dent, Roy Bell, who made the only comment about the rail:
Striped Rail (Rallus philippensis, L.). Sunday Island, vicinity of the lagoon in Denham Bay, but by no means
common.
Whatever species or subspecies Bell was describing, it was not recorded by Iredale in 1908 after a 10-month
stay on the island (Iredale 1912), and it had certainly disappeared by 1944 (Sorenson 1964). Cheeseman
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(1891) was told that cats had exterminated the Raoul Island Pigeon Hemiphaga novaeseelandiae ssp., and no
doubt the rails would have been equally vulnerable to these predators. The Spotless Crake Hypotaenidia
tabuensis plumbea was recorded as a breeding species at Denham Bay around 1909 and 1911, but rats were
taking eggs (Oliver 1912).
There is a possibility that the Raoul Island Banded Rail was a population of Buff-banded Rail Hypotaenidia
philippensis (Iredale 1912), as this species has colonised a vast number of islands throughout the Pacific
(Ripley 1977; Taylor & van Perlo 1998). However, it is just as likely to have been an endemic species that was
rapidly wiped out once humans colonised the island. Only with the discovery of fossil remains can the true
affinities of the Raoul Island Rail be determined.
Olson (2006) reported the existence of a drawing of a Hypotaenidia rail, made during the Malaspina
Expedition in 1793. The expedition was poorly known, primarily due to Alejandro Malaspina being impris-
oned and ultimately exiled after he returned from the voyage to Spain (see Olson 2006 for details). Two
ships, the Descubierta and Atrevida, departed from Cádiz, Spain, on 30 June 1789, and returned in September
1794. The expedition included scientists and draughtsmen.
Four drawings were made by an anonymous artist during the expedition’s visit to Vava’u in the Kingdom
of Tonga, of which two are the only known representations of now-extinct species; there are also brief notes
describing coloration. One of the two drawings depicts the Pacific Islands Eclectus Parrot Eclectus infectus
(see p. 198), whereas the other illustrates a probably flightless Hypotaenidia rail. Olson compared the latter
drawing with the Banded Rail H. philippensis, a species that still occurs on the island, and concluded that the
Vava’u bird had more robust legs and shorter wings, characters associated with flightlessness.
A large number of rails occurred throughout the Pacific Islands, and Kirchman & Steadman (2005)
described an extinct, flightless species from Eua, also in Tonga, Hypotaenidia vekamatolu (see p. 108). The
Vava’u bird was presumably closely related to the Eua bird and derived from the same ancestral stock.
Olson (2006) noted that the Vava’u Rail differed from Hypotaenidia philippensis in proportions of the skel-
eton, and in being greyer with a much brighter red beak. The rails presumably became extinct on Vava’u at
some point after 1793 due to habitat destruction and the introduction of predatory mammals.
To all intents and purposes, Latham’s description is the only information we have on this bird. Gray (1862b)
had no idea what it was, and merely stated ‘What is the Dark Rail (Rallus tenebrosus, Lath. MSS)?’. A ‘Mr.
Francillon’ whom Latham mentions in his description may possibly be the entomologist John Francillon
(1744–1816). Latham’s information was evidently obtained from him, but the circumstances behind the
description seem to be unknown.
No one else seems to have mentioned such a bird, including Sharpe (1894). The Spotless Crake is a rare
vagrant to Norfolk Island, so it is possible that the original specimen was a vagrant; indeed, Ripley (1977)
tentatively placed this bird within the nominate race of Spotless Crake, P. tabuensis tabuensis. However, it
could equally represent a distinct but now-extinct subspecies from the island.
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As on Réunion Island in the Mascarenes, there are a number of reports describing a bird called a Poul Ble
(Poules Bleues) from Mahé in the Seychelles, which are probably referable to an endemic but now-extinct
Porphyrio swamphen. It was first mentioned in 1768 by the Marion-Dufresne expedition, which comprised
two ships, La Digue and La Curieuse (Lionet 1980). An expedition party explored the northwest of Mahé, and
noted (in Lionet 1980, 1985; Skerrett et al. 2001):
There are in these rocks many giant tortoises, goats and also the ‘poules bleues’, of which we killed one; they are
like a large chicken, with blue feathers, the beak flat, wide and red and the same colour feet.
The Poul Ble apparently had a reputation for ferocity as attested by Pierre Poivre, Governor of Mauritius,
who wrote to the Commandant of the Seychelles in 1770 (in Lionet 1985; Skerrett et al. 2001):
If you could send me some pigeons, doves and other birds peculiar to your island, it would give me great
pleasure, but don’t send me any poules bleues for they are ‘mal faisantes’.
The meaning of ‘mal faisantes’ is not known (Lionet 1985), but Skerrett et al (2001) thought this statement
was in reference to the aggressive nature of the birds and that they would ‘make short work of any cage
they were put in’. There are place names referring to the bird on Mahé, e.g. Anse aux Poules Bleues, which
suggests that it was once abundant, but no physical remains have yet been found to resolve its affinities,
despite palaeontological searches (Hume 2004). The birds presumably died out due to over-hunting, defor-
estation and predation by introduced animals.
Another supposed Seychelles rail called the ‘poule pintade’ (‘spotted rail’?) was mentioned in the early
literature (Lionnet 1984b), which Cheke (2013b) suggests may have represented a Gallirallus (= Hypotaenidia,
following recent taxonomic revisions) banded rail. Seeing that only one vagrant specimen of banded rail
has ever been reported in the Indian Ocean, collected in 1863 (Benson 1970), and that Hypotaenidia is an
otherwise exclusively Australasian-Pacific genus, this assignation is probably entirely erroneous. The orig-
inal description is vague and may not even be referable to a rail at all.
Habits
Little known, but one Brayer de Barré in 1775 (in Lionet 1985) described the habitat of the birds (translation
by JPH):
In the northwest of the Seychelles Islands [Mahé], there is a plain watered by the rivers, which is a half-mile
wide. The land is good for cultivation, and there are lots of surrounding woods. The rocks are full of tortoises,
goats, and poules bleues.
Nothing is known about this pigeon other than Latham’s (1823) description:
18 inches. Head, neck and breast purple; nape glossed with green; chin and throat white; the rest of the body
cinnamon colour. Inhabits Howe’s Island [Vava’u Islands].
The description is reminiscent of the wide-ranging Metallic Pigeon C. vitiensis, but as no subspecies
answering to this description appears to inhabit Tonga today, it might represent an extinct race that formerly
inhabited the islands. Races of C. vitiensis do occur in Fiji and Samoa, so there is a possibility that the Tonga
locality is in error.
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Governor Philip (1789) commented that he saw on Lord Howe Island ‘very large pigeons in great plenty’.
Gray (1862b) questioned what they might be. In 1844–47, a Dr Foulis (in Hindwood 1940) made a brief
comment which may be referable to this bird:
There are a few birds that belong to the island, the only valuable kind being a large blue pigeon which is tolerably
numerous.
Nothing more is known about them. Biogeographically, it is reasonable to assume that a race of the New
Zealand Pigeon Hemiphaga novaeseelandiae occupied Lord Howe Island in the past, and disappeared due to
over-hunting and predation by introduced mammals. Any discovery of subfossil material will elucidate its
relationships, but until that time its status remains in doubt.
Moseley (1879) was the only person to mention a black parrot on the Admiralty Islands. He stated:
I saw on the main island a scarlet and black Parrot or Cockatoo of some kind, which flew out of some high trees on
the seashore, screaming loudly, like a Cockatoo. The bird was wary, and I could not get a shot at it. It reminded
me at the time of the rare Dasyptilus pequetti [Pesquet’s Parrot Psittrichas fulgidus] of New Guinea; it was
of about that size.
Pesquet’s Parrot is a bird of the central highlands of New Guinea (Juniper & Parr 1998). If an isolated popu-
lation of such a bird formerly occurred on the Admiralty Islands it was almost certainly specifically or at
least subspecifically distinct. Nothing more is known about it.
Kotzebue’s description is a brief one. It stated merely that the bird was the size of a sparrow, red and green
in colour with a red tail four times as long as the body. At face value this sounds like an exceptional species,
related to the Papuan Lorikeet Charmosyna papou, in an isolated and now-extinct population. The Papuan
Lorikeet, which is confined to New Guinea (Juniper & Parr 1998), is a red and green bird with a long, yellow-
tipped tail that is nearly twice as long as the body. No other lorikeet has a tail of comparable length. No
Charmosyna occurs on Samoa today, but one species, the Red-throated Lorikeet C. amabilis, lives on Fiji
(though it is short-tailed and now, incidentally, extremely rare, with the most recent sighting in 2001;
BirdLife International 2011), and there is no good reason why another should not have occurred on Samoa.
If the description is correct, the bird cannot be identified with any known species, and is probably specifi-
cally distinct.
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The validity of this subspecies has been open to question, as many of the surviving museum specimens
have clipped wings and clipped tails, indicating that they are captive birds; moreover, the lack of uniformity
in the coloration of the female specimens suggests hybridisation by cross-breeding. However, Forshaw
(1989) doubted that it was an aberration as a result of captivity. If it was a genuine subspecies, it may have
occurred on islands at the northern or eastern end of the Banda Sea.
Salvadori (1891a) thought that Latham’s R ose-fronted Parrot was ‘probably a young bird’, but based on
coloration it is more likely to be an adult female. When comparing the specimen, which is in poor condition,
with other Psittacula, it differs from all other members of the genus. Unless it is an aberrant individual, the
Rose-fronted Parakeet may well represent a valid but now-extinct species. Nothing more is known about it.
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The identity of this parrot remains a mystery. Linnaeus (1766) synonymised this name with the extinct
Mascarene Parrot Mascarinus mascarin, but the descriptions totally disagree. It was described from a spec-
imen seen alive by Hasselquist, of uncertain origin. Latham (1822) describes the bird as:
Size of a Jay. Bill black, the feathers round the base of it black, rough, and beset with hairs; space round the eye
white; irides yellow; crown variegated cinereous and black; upperparts of the neck and wings black; belly and
thighs cinereous, marked with transverse hoary lines; tail wholly ash-coloured, cuneiform; legs tuberculated,
black; toes the same; claws crooked and black. Inhabits Africa. The only one who has described this is Hasselquist,
from whom Linnaeus had his account.
Because Linnaeus associated the Obscure Parrot with the Mascarene Parrot, albeit in the synonymy, some
authors have suggested that the Mascarene Islands of Mauritius or Réunion were probably the provenance.
There is no evidence for this whatsoever.
The Jamaican Yellow-headed Macaw is known from a specimen shot in about 1765 by a Mr. Odell, and from
a contemporary painting dated the same year (Fisher & Warr 2003). It was seen by a Dr Robinson, who sent
a description to Gosse (1847). Gosse wrote:
If this be not the Tricolor of Le Vaillant [Cuban Macaw], which is the only Macaw I am aware of marked with a
yellow nape, it is probably undescribed. The two descriptions do not, certainly, agree exactly; yet still I cannot
but think the bird seen by Robinson, whose description I give below, to be this very rare species. Of the present
specimen the Doctor says, ‘This bird I saw stuffed. The legs and tail were wanting. It seemed less [in size] than
the common Red and Blue Macaw [Scarlet Macaw Ara macao]. By what I can judge from this sample, this bird
has never yet been figured or described. Sir Henry Moore, the late Lieutenant Governor, often assured me that
the Jamaican Macaw was very different from any he had ever seen. The subject now before us was shot in the
mountains of Hanover parish, about ten miles east of Lucea, by Mr. Odell’.
Gosse was uncertain whether the bird was a new species, or conspecific with A. tricolor of Cuba. In spite of
the evident differences in the descriptions, the Jamaican species was repeatedly united with A. tricolor until
given specific status (Rothschild 1905b). Even Clark (1905d) united the two, though he queried this in a
footnote.
Lack (1976) rejected the validity of A. gossei on the basis that neither Sloane nor Browne, early writers on
the island, mentioned such a bird, although he admitted that the description was plausible. But there is no
reason to assume that every early visitor would have visited every part of the island or have noticed every
item of the avifauna. Greenway (1967) had already pointed out that Browne might no mention of an
endemic red macaw in Jamaica, though he indicated that the ‘common red macaw’ was imported from the
mainland. A recently discovered illustration of a probable Scarlet Macaw Ara macao, painted on Jamaica in
1765 (Turvey 2010), confirms that this mainland species at least had been introduced onto the island.
However, if A. gossei was very rare and confined to mountains in the interior, Browne may easily have failed
to observe it. Thus there seems to be reasonable evidence that at least one other endemic red macaw, along
with Ara tricolor of Cuba, once occurred in the West Indies. Whether it justified specific status is now impos-
sible to ascertain.
Status Known only from Brisson’s 1760 description, and perhaps Edwards’s plate of 1751.
Range Perhaps occurred on Martinique, West Indies.
The Brown-throated Parakeet Aratinga pertinax is a polytypic species ranging from Panama through
northern South America and a number of islands off the coast of Venezuela (Juniper & Parr 1998). About 14
races are generally recognised. Linnaeus’s name aeruginosus, still in use for one of the subspecies, was based
on Edwards’s 1751 plate of a bird allegedly from the West Indies, but the plate does not agree with the bird
currently known by this name (of northern Colombia and north-west Venezuela). Based on Edwards’s
description, the Martinique bird was generally similar to the nominate B rown-throated Parakeet, differing
mainly in the brown of the neck turning to spots on the breast.
Edwards’s description and plate do not agree precisely with any B rown-throated Parakeet race, though
it is obviously very closely related. Edward’s definite comment about the brown of the neck falling ‘in spots
on the breast’ is particularly hard to reconcile.
Linnaean problems
The application of the name aeruginosus has posed a number of nomenclatural problems. Peters (1937
[1931–48]) comments:
Berlepsch and Hartert [1902] . . . substitute Cumaná, Venezuela as the type locality and Brabourne and Chubb
[1912] . . . designate Cayenne. None of these authors could have compared their birds with pl. 177 of Edwards,
the exclusive basis of Linnaeus’ name, which does not agree with birds from north-eastern Venezuela or Cayenne.
Peters uses the name aeruginosa for birds from the arid tropical zone of Colombia, western Venezuela and
north-western Venezuela, but Edwards’s plate does not agree with birds from these areas either. Clark
(1905b) commented on the evidence for a species of parakeet on Martinique:
Under the name of ‘La Perruche de la Martinique’, Brisson gives a description of a bird which covers perfectly
the well-known Conurus aeruginosus of the mainland, referring to Edwards (1751) for a figure of the species.
It is on this plate and description that Linnaeus’s [Psittacus] aeruginosus is founded. Brisson says the bird is
found in Martinique, and also in various places on the mainland, while Edwards gives for it the rather vague
habitat of the ‘West Indies’. Very likely this is a case of a continental species wrongly credited to the Antilles . . .
but we have no evidence to show that this is the case, and it is not at all improbable that the Conurus of
Martinique was a brown-throated species similar to C. aeruginosus.
Thus Linnaeus’s name aeruginosus may well have been based on birds from Martinique, but nomenclatural
confusion might be avoided by employing the name martinica used by Brisson (1760).
Both Brisson’s (1760) and perhaps Edwards’s (1751) accounts might refer to a B rown-throated Parakeet
subspecies that once occurred on Martinique. It was not recorded again, and the reasons for its disappear-
ance are unknown. No parakeet survives on Martinique today.
No Aratinga occurs on Dominica now, but there is ample evidence that one did so formerly. Frederic Albion
Ober travelled extensively, but particularly in the Lesser Antilles, where he collected from 1876 to 1878 and
again in 1880, discovering 22 new species of birds and adding many types to the collections of the
Smithsonian Institution. He seems to have had a particular affection for Dominica, which he visited several
times. His collections from Dominica were written up by George Lawrence, with liberal quotes from Ober’s
own notes. His account (Lawrence 1878a) of a parakeet on the island is as follows:
‘Parrot’. Not abundant. This bird, about the size of our Northern Carolina Parrot, but more robust, is very shy,
keeping mainly to the higher mountains; sometimes descending to the inner valleys, to feed upon the wild
guavas. It is sometimes captured by being wing-broken, and takes kindly to confinement, but unlike its larger
brother, the Cicero [Imperial Amazon Amazona imperialis], does not learn to talk. It congregates in small
flocks. It is oftener shot in the months between September and February. A very beneficent law of Dominica
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prohibits the shooting of Parrots, Ciceros, Ramiers &c. in any other months, thus ensuring protection during
the breeding season.
Ober did not obtain a specimen.
Clark (1905b) briefly discussed the evidence for a parakeet on Dominica:
There appears to be no description extant of the species which formerly inhabited Dominica, but it seems to have
been exterminated at an early date. Ober’s remarks doubtless had reference to Amazona bouqueti [i.e. the
Red-necked Amazon Amazona arausiaca], and not to any species of Conurus [i.e. Aratinga].
Although it is curious that Ober did not mention the Red-necked Amazon, which is also endemic to
Dominica, his comments cannot refer to it, as his comparison of the bird to a Carolina Parakeet Conuropsis
carolinensis unquestionably implies a parakeet rather than an amazon parrot. If this bird did exist, it must
have disappeared soon after Ober observed it in the 1870s.
Little is known of this bird, for which Clark (1905b) provides a summary:
Hughes (1750) under ‘The Parakite’ says: ‘This is of the frugivorous kind, and about the bigness of a thrush,
having a longer and more crooked bill. It feeds on all manner of berries, popaws, and ripe plantains, residing
chiefly in inaccessible gullies. The bird borrows its name from its resemblance in make, but not in plumage, to
the small green parakite’. Sir Robert Schomburgk (1848) includes ‘Psittacus passerinus’ in his list of the birds of
the island, but no parrakeet is mentioned by Ligon (1673).
As other Aratinga parakeets occur in the West Indies, and subspecies of Brown-throated Parakeet A. pertinax
occur in the southern island chain of the Lesser Antilles (Juniper & Parr 1998) and perhaps formerly on
Martinique (see p. 380), there seems little reason to doubt that a species or subspecies of Aratinga once
inhabited Barbados. It must have disappeared between 1750 and 1840, but the reasons for its extinction are
unclear.
Trochilidae (Hummingbirds)
Bell’s Woodnymph Thalurania belli Verrill
Thalurania belli Verrill (no date = 24 October 1905)
Status Known only from Verrill’s description. Possibly represents an extinct species.
Range Dominica, West Indies.
Verrill (’1905’) appears to be the only person who witnessed a hummingbird on Dominica other than the
Blue-headed Hummingbird Cyanophaia bicolor, a species that still occurs on the island. He described the
adult male as follows:
Upperparts rich iridescent metallic green, becoming deep peacock blue or v erditer-green on forehead and crown;
coppery on shoulders and deep bluish or emerald green on rump: scapulars, upperwing- coverts and
uppertail-coverts, deep peacock or bluish green. Wings metallic purple or steel blue, the outer web of outer
primary narrowly edged with white or pale ash grey. Basal portion of tail dull copper green, the outer half deep
steel blue with violet reflections. The three outer feathers on each side broadly tipped with white and the outer-
most feather white at base also. Lower parts uniform snow white more or less washed with greyish on flanks and
sides. Flanks and sides beneath wings spotted with isolated bright green feathers. Ear-coverts and loral region
deep velvety black in marked contrast to green of occiput. Bill dusky black with lower mandible slightly lighter
near base.
The adult female was similar but duller and more uniform in colour. He mentions that the nest was a beau-
tiful structure of felted t ree-fern down, ornamented with spider webs, beetle elytra, and bits of b
lue-green
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lichen, usually saddled to a slender branch or twig up to 10m from the ground, and preferably overhanging
some mountain stream.
The bird was confined to the highest mountains, at an elevation of above 600m. It was very locally
distributed and strictly confined to forest, occurring in the deepest and darkest areas and never visiting
sunny glades as did all other species. It tended to occur in the upper part of the tree canopy particularly in
the tangled network of lianas and orchids. Cory (1918) considered that if the description of the ‘type spec-
imen’ was correct, the bird was most likely an aberrant B lue-headed Hummingbird; however, Verrill’s very
detailed description seems to be from life, and it is unclear if any type specimen was ever designated.
Moreover, he seems to have observed a population that almost certainly no longer exists in the mountain
forests of Dominica. If the species is valid, the reasons for its extinction are unclear.
Halcyonidae (Kingfishers)
This species was one of the few species mentioned by Latham that was not subsequently described by
Gmelin, and has been overlooked or ignored by all subsequent authors. There appears to be no kingfisher
that approaches Latham’s 1781 description. The main description is Black-capped Kingfisher Halcyon
pileata. Latham states:
(Var A [i.e. H. pileata]) The bill is large, and of a bright red: the head and hind part of the neck are black: the back,
the tail, and middle of the wings, are of a deep glossy violet blue: the shoulders and ends of the wings are black:
the throat, fore part of the neck, and breast, are white, and encircling the neck at the lower part, near the back: the
belly is of a pale rufous colour: legs pale red.
(Var B) Length ten inches. Bill thick at the base, two inches and a half in length, and of a red colour; the general
colour of the upper parts of the body is black, mixed with ferruginous: the feathers of the head longish: the quills
and tail are blue green, with the inner webs and shafts dark-coloured: the throat and breast are white; on the last
some of the feathers are tinged with pale green: the belly ferruginous brown: legs red. Inhabits the South Sea, but
what part uncertain.
The two descriptions are very different, and Latham’s ‘Var. B’ is likely to be a distinct species. It may repre-
sent a now-extinct kingfisher from somewhere in the South Pacific.
The Ryukyu Islands stretch from 225km s outh-west of Okinawa to 290km from the northern tip of Taiwan.
The Miyako Kingfisher was collected on February 5, 1887 on M iyako-jima by Y. Tashiro (Kuroda 1919) and
has not been seen since. The islands have been searched carefully, particularly Orii (Greenway 1967), but no
birds of this species have been found.
There is great doubt as to its validity (Forshaw 1985); Brazil (2009) points out that Miyako-jima is a highly
383
unlikely place for a kingfisher to live, and suggests that the type was in all likelihood a vagrant (possibly not
found on Miyako-jima at all), with the taxon representing (at best) a Ryukyu race of Micronesian Kingfisher
H. cinnamominus.
The horny sheath of the bill is missing in the type, so the bill coloration is unknown, and the supposed
difference in the primaries being longer in proportion to the secondaries is almost certainly an artefact of
preservation (Greenway 1967); otherwise the bird is similar to the Guam subspecies of Micronesian
Kingfisher H. c. cinnamominus.
If there was once a valid population on the Ryukyu Islands of this bird it has certainly gone now, due to
widespread deforestation and drainage of wetlands. Nothing was recorded about its habits.
Galbulidae (Jacamars)
The type and only specimen of this race of Brown Jacamar B. lugubris was collected in 1923 by Samuel
Klages (Todd 1943). Originally described as a full species, Mayr (1957) relegated Klages’s Jacamar to a
subspecies, while Meyer de Schauensee (1966) confirmed that at the time of his writing it was still known
only from the type. Its taxonomic position and its validity await confirmation. If it is a valid taxon, it may
now be extinct.
Troglodytidae (Wrens)
The Common or Winter Wren Troglodytes troglodytes is the only member of the Troglodytidae to occur in the
Old World. Some 35 races are recognised, and many more have been described. The species ranges across
the entire Holarctic, with many races being confined to single islands.
The status of T. t. orii is disputed. Paynter & Vaurie (1960) relegated it to a synonym of T. t. mosukei, with
the comment that it required confirmation. The race is not mentioned at all by Austin & Kuroda (1953). A
large part of the island was cleared for an airfield and military installations during World War II. Regardless
of whether it was a valid subspecies, the population is certainly now extinct. Nothing was recorded about
its habits.
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Timaliidae (Babblers)
Delacour (1921) explained that a male and a female Red-billed Leiothrix Leiothrix lutea were sent to him
from China in two separate consignments. The male arrived in February (c.1921) and was at first thought to
be an aberrant specimen, but closer examination and the arrival of the female two months later made
Delacour believe that he had a new species. He pointed out that the differences between astleyi and L. lutea
are much the same as the differences between two races of the Silver-eared Mesia Leiothrix a. argentauris and
L. a. laurina. Delacour did not say who Astley was, but presumably he was the dealer from whom the speci-
mens were obtained. If this is a genuine taxon, it may have disappeared through o
ver-trapping.
Fringillidae (Finches)
Bowdich (1825) was the only person ever to have encountered the Madeira Finch. He observed it in 1823
and remarked:
I saw another and more curious bird, but I doubt if it is a native of the island. The outline of the beak most resem-
bles that of the widow-bird, (vidua, Cuv.) but the commissure is situated like that of the grakle, (gracula, Cuv.)
immediately beneath the nostril, and forms a much deeper angle; it evidently belongs to the goniaphea, between
fringilla and corythrus.
Bowdich describes it further in a footnote:
The upper mandible closes over the lower, and the middle toe is longer than the others; the whole bird is black,
with the exception of the head, which is azure. G. leucocephala.
The Madeira Finch was figured in Bowdich’s volume, but unfortunately a large part of his collection was
lost en route to Europe (Pieper 1985), which probably included a finch specimen. No subfossil remains have
yet been identified, so the relationships of the Madeira Finch remain a mystery. Extant in 1823, it was never
reported again, so it is likely that human-induced changes and the introduction of predatory mammals
exterminated the birds. Lowe (1853) did not record it during his survey of Madeira and Porto Santo, so it
almost certainly had disappeared between 1823 and the time of Lowe’s visit in the early 1850s.
Cardinalidae (Cardinals)
This is an enigmatic bird that has puzzled most authors. Paynter (1970) noted that:
Based on a unique specimen, is generally conceded to be a Spiza, but whether it represents an extinct species or a
hybrid is unknown.
and as late as 1983, the AOU Checklist (1983) still said that ‘its peculiarities cannot be accounted for by
hybridism or apparently by individual variation’.
Coues (in Sharpe 1888) suggested that it might be a hybrid female S. americana × male Blue Grosbeak
Passerina caerulea. The c olour-variant hypothesis appears to be the most likely; it might be that Townsend’s
type, in lacking normal carotenoid pigments (Parkes 1985) and through the foxing of the now aged spec-
imen (see Fuller 2000), contrive to give a more distinctive plumage than the specimen actually merits.
A recent study has shown that the coloration is more complex (Hume & van Grouw 2016). If Townsend’s
Bunting was a female Dickcissel as suggested by Parkes (1985), the diagnostic brown crown and cheeks
should still be present, and in Townsend’s male attribution, it should have exhibited the black throat patch,
unless juvenile. Holt (2002–03) considered the bird to be a juvenile male Dickcissel, but this is not supported
by the lack of a full black throat patch, and juvenile males are brown above, not grey. Furthermore, there is
black speckling (eumelanin) concentrated on the sides of the upper breast, no dark streaking on the flanks,
and the upperparts are grey in the Townsend’s Bunting specimen, a pattern that is not found in any plumage
of Dickcissel. Only a mtDNA analysis will resolve this long-overdue conundrum.
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Data-deficient taxa
This appendix includes those taxa that lack sufficient information for them to be placed with confidence in
any category. Some known from a single specimen or just a few specimens may be extinct, but it is perhaps
as likely that they are not. A number of these birds may be relocated one day, as they occur in remote, rarely
visited places where few ornithological surveys have taken place. The birds that have defied rediscovery
are often cryptic forest inhabitants that are always going to be difficult to find, especially if they are
nocturnal; no doubt the owls and nightjars are the most likely of these ‘lost’ birds to come back from the
dead. Others are in a state of taxonomic flux and may be synonymous with extant populations. Until further
information becomes available, the status of these enigmatic data-deficient birds remains unclear.
Although the type of this tinamou was collected by M. A. Carriker on June 9, 1943 at Ayacucho, Colombia, a
second specimen referable to this species was collected nearly 150 years before, in 1786 (although this is
now lost). The status of the Magdalena Tinamou remains in doubt. It is generally thought to be extinct;
however, a captive individual survived until the 1990s (Donegan et al. 2003a), and the species is possibly
extant in the wild, with reported sightings in the 1970s, 1980s, and 2008. If it survives, it is critically
endangered.
Almost nothing is known about this bird, which is presumed to have inhabited forested lowlands. The
type, which is probably an immature male, was collected at an altitude of 150m, while the second (lost)
specimen, an adult male, was taken at 500m. The foothill forests in the Magdalena River Valley have been
almost completely deforested, but a recent survey (Donegan et al. 2003a, b) found remnant foothill forests
on the western slope of the Cordillera Oriental and also on the eastern slope of the Serranía de San Lucas.
The bird may still survive.
The Magdalena Tinamou is considered by some authorities to be a subspecies of the similar R ed-legged
Tinamou C. erythropus, while others have lumped all Red-legged Tinamou subspecies into the nominate
form. Pending further studies, the Magdalena Tinamou has been dropped from the 2008 IUCN Red List
(BirdLife International 2011).
This elusive hawk was recorded as being ‘not uncommon’ on Car Nicobar in the last years of the 19th
century (Gurney 1898; Butler 1899). It was recorded in 1901 (Richmond 1902), and after a lapse of almost 80
years it was supposedly seen again in 1977 (Abdulali 1978). This last record is unconfirmed, however; and it
was not recorded by an extensive survey in 1993–95 (Sankaran 1998). The main reason for its disappearance
is the destruction of habitat.
There is a second race in the species, A. b. obsoletus from Katchall and Camorta Islands. This is very rare,
and almost as elusive as the Car Nicobar nominate.
Habits
The Car Nicobar Sparrowhawk was a bird that kept almost exclusively to the tops of trees, and was
extremely shy. Stomach contents contained lizards and insects. Breeding appears to have occurred in
September and February or March, suggesting the species was double-brooded, and the nest was located
about 12m from the ground at the end of a large branch in a huge Ficus tree. Butler (1899; see also Gurney
1898) made the following observation on the bird:
Mr. Butler writes that they are ‘not uncommon in forest on Car Nicobar, keeping almost exclusively to the tops
of high trees; continually utters a shrill little double cry, exactly like Astur badius [Shikra] Young birds are
extremely chestnut in colour. The one I send had one or two filaments of nest-down still hanging to it, proving
this to be the first plumage acquired . . . Young birds have a trick of fluttering on a bough like a broken-legged
bird . . . In September I noticed several r ufous-crowned young birds probably bred in March or April, and at the
same time both adult cocks killed were in a state of breeding.
Baillon’s Crake Porzana pusilla has a vast distribution from Eurasia and Africa to Australia and a host of
islands in between. Although not globally threatened, some of its races are very poorly known and their
status is uncertain. The Bornean subspecies was collected just once, on June 25, 1912, and has not been
recorded since. The reasons for its disappearance are not known, but there is a possibility that it might still
survive. However, Borneo has suffered severe deforestation, so the likelihood of relocating the birds is
becoming increasingly remote. Nothing is known of its habits.
small white spots along edge of inner web; chin and throat whitish; foreneck and breast more grey; bill shorter. Immature lacked
rufous on head and bright olive margins to feathers of upperparts; slight barring on underparts.
The Flores subspecies of Lewin’s Rail Lewinia pectoralis was first collected by the naturalist A. H. Everett on
November 1896 in Manggarai district, west Flores (Hartert 1898b), with another five specimens, including a
juvenile and 10 clutches of eggs, taken by a priest and naturalist, Jilis A. J. Verheijen, in 1958–59 (Mees 2006;
Trainor & Verbelen 2016). All of the Verheijen’s specimens were taken within 20km of the town of Ruteng in
the Manggarai district (Trainor & Verbelen 2016). It was thought that as the birds were last recorded in
1958–59, the Flores rail could possibly be extinct (Paynter 1963; Hume & Walters 2012). However, another
specimen, a juvenile male, was taken 52km west of Ruteng on February 1, 1974 by another naturalist-priest,
Erwin Schmutz (Mees 2006). Since then, searches for it have been unsuccessful and its status remains in
doubt, but this elusive bird may still exist (Trainor & Verbelen 2016). The reason for its disappearance or
rarity is not known.
Habits
The skin specimens and egg clutches of Flores Rail were taken in rice fields, agricultural land, vegetated
stream channels, secondary forest, dry gardens and in scrub from 650–1120m in altitude (Trainor & Verbelen
2016). The clutches were taken in February, March, April, June, September, November and December
(Verheijen 1964; Mees 2006), so the birds presumably bred year round.
The Slender-billed Curlew was once numerous, and as recently as the early 1960s hundreds could be
counted on its traditional migration routes. It migrated from its breeding grounds in southwest Siberia
through north Kazakhstan and other eastern European countries to the Mediterranean, where it over-
wintered generally in Atlantic Morocco and Tunisia (See Gretton 1996). Its disappearance can be attributed
to intense hunting pressure on its Mediterranean wintering grounds, and habitat destruction in both
Kazakh steppe and winter range (Gils & Wiersma 1996). There are very few recent records. Birds were
regular visitors to the well-watched wintering site at Merja Zerga, Morocco, but they have not been recorded
there since February 1995 (Donald et al. 2010). An individual was seen in May 1998 at Druridge Bay,
Northumberland, UK (Cleeves 2002, Gretton et al. 2002; the only UK record of Slender-billed Curlew,
though it was and remains controversial. The record was removed from the British List in 2014; see Collinson
et al. 2014) and a few other observations have been made elsewhere, but many need verification (Donald et
al. 2010; del Hoyo & Collar 2014). Authenticated records were made in April 1999 in Greece, and February
and August 1999 in Oman, while the last accepted record is of a bird seen in April 2001 in Hungary (Oláh
and Pigniczki 2010). There have been other unconfirmed reports, but they are becoming more infrequent
(see Crockford 2009; Donald et al. 2010; Kirwan et al. 2015). If the Slender-billed Curlew still survives its
numbers must be critically low, and its habit of long migration, exposure to hunting, plus habitat destruc-
tion in its breeding and wintering grounds renders the long-term survival of this bird virtually impossible.
The Slender-billed Curlew is the Eurasian equivalent of the extinct Eskimo Curlew Numenius borealis (see p.
148), and is almost certainly doomed to the same fate, if it has not disappeared already.
Habits
The Slender-billed Curlew occurred in a variety of habitats including saltmarsh, coastal brackish lagoons,
estuaries, and steppe grasslands during migration, and wintered in coastal bays with saltmarshes and
inland marshes (Gretton 1996). It fed on a variety of invertebrates, including earthworms, molluscs, snails,
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small crustaceans and small insects. Little was recorded about its breeding habits, but it apparently bred in
small colonies in dense vegetation on dry areas within bogs. The nest was constructed from grass, and up to
four eggs were laid. Eggs were recorded once in May 1914 (Ushakov 1916), and a communal nesting area
was found in 1924 (Ushakov 1925). The common name of ‘curlew’ is derived from the soft cour-lee call given
in flight, and was similar to that of the Eurasian Curlew, but shorter and higher-pitched. The alarm call was
a rapid kew-ee.
This lory has not been recorded since the type material was collected. The specimens are labelled from the
Nenusas Islands, which are now deforested and the locals do not know of such a bird; but in any case the
specimens appear to be immature birds of the Talaud subspecies E. h. talautensis, and could have been
traded or even possibly blown to the Nenusas, which are only an hour’s flight for a bird from Talaud (Collar
et al. 2001).
The Brown-throated Conure is a widespread species, with 14 subspecies occurring in Panama, northern
South America, and islands off the northern coast of Venezuela (Forshaw 1989). The Sinú valley subspecies
is of unclear taxonomical position; it is known from just two specimens, and has not been seen since 1949.
Almost nothing was recorded about the habits of this bird. The reason for its disappearance is unclear.
The area has been in the grip of armed conflict for many years, and the survival of this subspecies cannot be
confirmed.
The Red-throated Lorikeet was considered common when it was first collected in Ovalau on June 17, 1875
(Ramsey 1875), yet it must have been comparatively scarce on other islands, as other collectors failed to
mention it (see Swinnerton & Maljkovic 2002). However, twelve specimens were collected in one week in
1925 on Viti Levu by the Whitney South Seas Expedition (Watling 2000). It has gradually disappeared from
each of the four Fijian islands on which it occurred in recent years.
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The last stronghold was on Mount Tomaniivi on Viti Levu, where the bird was observed during the
1970s; W. Beckon filmed and photographed the bird between 1975 and 1978 on the Nadarivatu Plateau. The
last confirmed sighting was on August 12, 1993 at around 800m, when three birds were seen feeding with a
Collared Lory Phigys solitarius (Watling 2000). Apart from one possible sighting in 2001 in the same locality,
there have been no further reports, despite long-term surveys (Swinnerton & Maljkovic 2002).
The Red-throated Lorikeet has suffered from continued habitat destruction, and perhaps from predation
by introduced Black Rats Rattus rattus, which have benefited from anthropogenic changes (Watling 2000);
the chances of the lorikeet being rediscovered are becoming increasingly remote. The bird is presently
considered to be critically endangered (BirdLife International 2011), and it may already have gone.
Habits
This lorikeet favours mature o ld-growth native forest above an altitude of 500m, but it probably occurred in
all suitable habitats down to sea level prior to serious deforestation on the islands (Swinnerton & Maljkovic
2002). The Red-throated Lorikeet foraged in small flocks high in the canopy, where it fed on nectar and
pollen from flowering trees, especially Metrosideros collina and Erythrina subumbrans (Clunie 1979). As a
result, it was likely to have undertaken seasonal migration in search of food (Clunie 1999). The birds often
fed in the company of other species, and were best located by their short, h igh-pitched squeaks (Forshaw
1989).
Little was recorded about the Bahia R ufous-vented Ground Cuckoo. The species occurs in lowland ever-
green and higher-ground forests, canebrakes and thickets. Diet comprises varied insects, arthropods, small
vertebrates, and some seeds and fruit. It is a terrestrial species foraging on or close to the ground. The Bahia
race N. g. maximiliani disappeared as a result of severe deforestation, but the nominate subspecies still
survives.
The taxonomy and type locality data of the Rufous-vented Ground Cuckoo complex are confused. The
type specimen of Bahia R ufous-vented Ground Cuckoo was collected in Bahia, which has now also been
designated the type locality of the species N. geoffroyi (Raposo et al. 2009); if this is correct the Bahia race N.
g. maximiliani becomes a junior synonym of the nominate, and no longer valid.
The Greater Crested Coua is known from only one skin collected in 1950; some doubt has been placed on its
validity, and it has been considered a distinct species, a geographical variation and even a hybrid. However,
Goodman et al. (1997) suggest that it differs so considerably from nominate Crested Coua Coua cristata cris-
tata that it probably merits specific recognition. The type locality is now heavily degraded, and deforestation
and over-hunting are probably responsible for its disappearance. Nothing else is known about the bird.
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The nominate race of Sulawesi Masked Owl, T. r. rosenbergii, occurs on Sulawesi and some adjacent islands,
but is poorly known. It appears to be widespread on Sulawesi and has managed to adapt to human-disturbed
areas (König et al. 2008). The Peleng subspecies, endemic to Peleng in the Banggai Archipelago, is known
from one individual collected on August 22, 1938; the bird was never recorded again. Peleng has suffered
severe deforestation, but due to the ability of the nominate race to survive in deforested areas, the Peleng
Masked Owl may still be extant. Calls thought to possibly come from this form were heard in Western
Peleng in March 2009 (Rheindt et al. 2010).
The nominate race of Lesser Masked Owl, T. s. sororcula, which occurs on Tanimbar in the Lesser Sundas
(and which may prove a distinct species; Collar et al. 2001), is known from just three specimens and is
considered endangered (König et al. 2008). The type and only specimen of the Buru subspecies, a female,
was collected on Kayeli in October 1898 (Hartert 1900). Apart from a few unreliable records, this bird has
not been seen since 1921. Nothing was recorded about its habits or why it disappeared, but it probably
occurred in lowland rainforest on Buru.
This owl may have nested in hollow trees (König et al. 2008), which would make it extremely vulnerable
to deforestation. These forests are under immediate threat of complete destruction due to commercial
logging, so if the owl still survives its long-term future is extremely bleak.
The taxonomy of the bay owls is extremely confused, with a number of subspecies described, some of which
are dubious. Seven races of Oriental Bay Owl Phodilus badius are generally recognised, ranging from Sikkim
through the Indian subcontinent and South-east Asia, with three subspecies endemic to islands; P. b. parvus
of Belitung Island, Sumatra, the doubtfully distinct P. b. riverae of Samar Island, Philippines, and P. b. arixu-
thus of Bunguran Island in the Natuna archipelago (Bruce 1999).
The reasons for the possible disappearance of the Natuna subspecies are unclear, but owls are notori-
ously cryptic, and Bunguran is not commonly visited by ornithologists, so a population may still survive.
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The only specimen of the Siau Scops Owl was collected in 1866 by Jonkheer R. von Duyvenbode on the
small volcanic island of Siau, north of Sulawesi, Indonesia. The nominate Sulawesi Scops Owl O. m.
manadensis is considered common on Sulawesi, and other subspecies occur on Peleng, Sula Island and
Tukangbesi Island. Deforestation is severe on Siau, and much of what remains comprises secondary growth
(Collar et al. 2001), so the Siau Scops Owl probably disappeared as a result. A recent survey failed to locate
the birds, but an unconfirmed call was apparently reported (Hunowi 2006).
The Puerto Rico Screech Owl M. nudipes is endemic to Puerto Rico and considered common on that island
(Stotz et al. 1996); a race of this species, M. n. newtoni, considered doubtfully valid by some authorities, was
endemic to the nearby Vieques, Culebra and adjacent Virgin Islands. It was last reported in 1936 on St Croix,
but there were unconfirmed reports in 1985 that is still survives on Guana Island, a small cay in the British
Virgin Islands. If it does survive, it is extremely rare. Destruction of forest to make way for plantations
appears to be the cause of its disappearance (Marks et al. 1999).
Habits
This owl preferred forests with large hollowed trees, and would call y ear-round, mostly at dawn, and from
dense foliage. Its local name in the Virgin Islands is ‘cuckoo-bird’, derived from the loud c oo-coo call. The
diet comprised mainly large insects, particularly orthopterans, but also small birds, geckos and small
rodents.
Little is known about the origin of the sole specimen of this species. Todd (1920) says merely that explora-
tion in Cayenne and the lower Amazon by Samuel M. Klages had ‘brought to light’ a number of apparently
new birds. The Cayenne Nightjar was one of them, with the type taken on 24 April 1917, but nothing is
known about it besides its appearance. It may be extinct, though Cleere (1999) thinks this unlikely.
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The Itombwe Nightjar is known from a single specimen, an adult female, collected in August 1955 at
Melenge in the Itombwe Mountains (Louette 1990). However, sound recordings taken in 1996 in Itombwe
may be of this species (BirdLife International 2011). Identical sound recordings were taken in Gabon in 1985,
northern Congo in 1996 and south-east Cameroon in 1997 ( Dowsett-Lemaire & Dowsett 2000), which indi-
cates that the species may be more widespread.
The type was taken in transitional forest between lowland and montane forested areas at an elevation of
1,280m (Louette 1990). The territorial call is probably a series of short tchoc notes (Cleere 2010). If it still
survives the numbers must be very small, and the continued clearance of its forested habitat due to agricul-
ture and for livestock grazing makes the long-term survival of this elusive species doubtful.
The only known specimen of the Nechisar Nightjar was collected on the side of a dirt road in 1990 (Safford
et al. 1995). The body had been destroyed, but a single wing and some tail feathers were retrieved which
constituted a new species, distinct from any other nightjar. It has not been seen again. A report of a possible
sighting in 2009 awaits confirmation (Cleere 2010). Nothing else is known about it, except that the specimen
was found in open, short grassland on the Nechisar Plains at an altitude of 1,200m. The Nechisar Plains are
under threat of destruction from firewood collecting, the extraction of construction material and
over-grazing (BirdLife International 2011). As nightjars are cryptic, nocturnal species, the Nechisar Nightjar
may still be extant.
This mysterious species is known from one adult female collected on September 7, 1929 in cold, arid sand
hills covered with low scrub at 1,230m in Xinjiang, China. It has not been seen since. There are some doubts
as to its validity, but it occurs alongside the Eurasian Nightjar C. europaeus plumipes, and is considered
smaller in size (Cleere 2010), with marked differences in wing-patterning (Vaurie 1960a, Leader 2009).
Nightjars are notoriously cryptic birds, so the chances of this species being rediscovered are reasonable.
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This distinctive nightjar is known from one specimen, an adult female, collected on August 21, 1939 and
never recorded again. It was found on the lowland coastal flats near Mount Panie, north-western New
Caledonia, but nothing else is known about it. It was originally considered a subspecies of the W
hite-throated
Nightjar E. mystacalis, a widespread species found in eastern Australia and vagrant to New Guinea (Cleere
2010), but it is now considered a distinct species.
This enigmatic bird was formerly known only from the type specimen, an adult male, caught at Tongué,
Païta, near Nouméa on the night of April 11, 1880, having flown into a bedroom, probably attracted by the
light (Layard & Layard 1881). The captor of the specimen took it to Théodore Savés, a French naturalist and
traveller, who then gave it to the Layards. The bird was unknown to both the local population and to colo-
nists (Layard & Layard 1882). A second specimen (a young bird), dated 1915, was recently discovered in an
Italian museum by Carlo Violani (BirdLife International 2011). Other reports include a dead individual
found in the 1950s, one shot in 1960, and a possible sighting in 1998 (Ekstrom et al. 2002), which hint that
some may still survive. If it still exists, the population must be extremely rare and restricted to the most
remote and inaccessible forests (Tobias & Ekstrom 2002).
Habits
Very little is known about the bird. It was nocturnal, probably sitting upright on branches or on the ground
during the day, and fed on beetles (Layard & Layard 1881), other insects and perhaps small vertebrates. It
had rather long robust legs, strong feet and short rounded wings, which suggests that it may have been in
part a terrestrial species, similar to the New Zealand Owlet-nightjar Aegotheles novaezealandiae (see p. 235).
discovered in 1974 (Benson 1974). It is only known from one specimen, and the type locality has already
been destroyed. No more records are known. If it still exists, it must be critically endangered, and there is
little hope for its future.
However, there is some doubt as to the validity of this taxon, and its status needs reassessment (Safford
& Hawkins 2013).
Habits
Nothing known, but the nominate occurs in dry scrub, savanna woodland and margins of wet evergreen
forest, where it forages for insects, small amphibians and lizards. It is not dependent on water.
There are 14 recognised subspecies of the Variable Kingfisher C. lepidus, ranging from the Philippines to the
Solomon Islands. Little is known about the Variable Kingfisher in general, but the species is not considered
endangered (Woodall 2001). Only one of the races, the Malaita Variable Kingfisher, has not been seen in
recent years, and it may have become extinct. Almost nothing is known about it, but Malaita is a very diffi-
cult island to access, so a population may still survive. It was seen frequently in 1997, but no details were
published (G. Dutson in litt. 2011).
The taxonomy of the cave swiftlets is confused and still in need of clarification. Whitehead’s Swiftlet is
divided into two subspecies, both confined to mountains, the nominate A. w. whiteheadi on Mount Data,
Luzon, and A. w. origenis on the Mountains of Apo, Kitanglad and Matutum, Mindanao (Chantler 1999;
Chantler & Driessens 2000). These swiftlets are extremely difficult to identify and easily confused with
other genera and species, even in the hand.
The nominate Luzon species was collected only once (Ogilvie-Grant 1895) and has not been recorded
again. It might be extinct, but swiftlets are elusive birds, and occur in inaccessible regions, so some may still
survive. The Mindanao birds are considered endangered due to restricted range and habitat loss.
Deforestation appears to be the primary reason for the Luzon birds’ disappearance; Mount Data has been
almost completely cleared of forest. If the bird still survives, it must be critically endangered.
Habits
Whitehead’s Swiftlet favours subtropical or tropical moist montane forests above 1,200 m elevation. It may
utilise hollow trees while roosting, and netted individuals of A. w. origenis on Mount Kitanglad (on
Mindanao) may have been taken on c liff-faces (BirdLife International 2011). A nest of the Luzon race was
taken in July 1895; otherwise nothing is known of these birds.
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The validity of the Sierra Leone Black Swift remains in doubt, mainly due to the lack of specimen material.
Nine races of African Black Swift Apus barbatus are recognised, ranging discontinuously from Liberia
through Congo and Kenya south to South Africa, and on Madagascar (Chantler 1999; Chantler & Driessens
2000). The taxonomy of the species is in need of clarification. The Sierra Leone race A. b. glanvillei may be
conspecific with the race A. b. sladeniae (see below), itself a poorly known subspecies. If the Sierra Leone
Black Swift is a valid subspecies, it may be extinct. The reasons for its disappearance are unknown, but
deforestation is a likely cause, especially as they nest in hollow trees.
The Fernando Pó Swift has sometimes been considered a race of African Black Swift A. barbatus, but Dowsett
& Forbes-Watson (1993) accorded it specific status. It is a poorly known taxon, recorded less than 10 times
and from very few specimens (Collar & Stuart 1985), with the last confirmed taken in 1903 (Pérez del Val
1996). Since then there have been no confirmed records. A recent sighting of around 10 birds in January 1998
at Moca in Monte Alen National Park (Mills & Dean 2007) remains unconfirmed, but if correct, this would
be the first record for mainland Equatorial Guinea (Dowsett-Lemaire & Dowsett 1999). It occurred over a
wide area, including the Obudu Plateau in southeast Nigeria, Mount Kupé in western Cameroon, Bioko
Island, and Mount Moco in Angola (Chantler 1999). Habitat destruction occurs over the entire range of the
Fernando Pó Swift, so its future, if indeed it is still extant, is far from secure.
The type of this hummingbird was obtained by Jules Bourcier, an expert on hummingbirds, who seems to
have exchanged it with Gould, who had also published intensively on hummingbirds (Gould 1861). The
specimen was held in the Gould Collection (now at Tring), and another was in the Elliot Collection, with
both collected before 1852. Warren (1966) pointed out that the Gould specimen agrees exactly in make up
with other Bourcier skins and that therefore this must be Bourcier’s original type, retained by Gould in
exchange for other birds. This was a not uncommon practice with Gould.
Graves (1999c) studied the two specimens in detail, and concluded that the Coppery Thorntail could not
be considered a hybrid, an immature specimen or a variant of some kind, and must therefore be regarded as
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a valid species. The provenance may be in error, however, as trade in 19th-century skins often provided
unreliable locality data.
Habits
Nothing was recorded about the bird, but it may have occurred in primary and secondary forest in the
Amazonian lowlands and foothills. If this was the case, it would have been severely threatened by habitat
destruction.
The Turquoise-throated Puffleg is known from six skins, but only the type has locality data, with the others
being Bogotá trade specimens. The type locality is now almost entirely destroyed. There was an uncon-
firmed sighting in the Chillo valley near Quito in 1976, but searches in the 1980s failed to locate the species
(Collar et al. 1992; BirdLife International 2011).
There is some dispute as to the validity of this species; Fjeldså & Krabbe (1990) considered it valid, while
Graves (1996) thought it a possible hybrid between Glowing Puffleg E. vestita and a second undetermined
Eriocnemis species.
Habits
Little was recorded about this bird. The type was collected between 2,100 and 2,300m in a ravine in forest on
the steep-sided slopes in the arid upper Guaillabamba (Fjeldså & Krabbe 1990).
The White-chested Tinkerbird is known from only one specimen, collected in 1964 at Mayau in north-western
Zambia, and further searches to relocate the bird have proved unsuccessful (Dowsett & Dowsett-Lemaire
1993). Collar & Fishpool (2006) suggest that it might survive in restricted areas other than the type locality,
that it was a possible straggler from elsewhere, or that the White-chested Tinkerbird replaces its congener,
the Yellow-rumped Tinkerbird P. bilineatus, in Angola and DR Congo. These adjoining countries are in need
of thorough ornithological research, but such actions are hampered by their unstable political nature.
At present, the status of this bird remains unresolved. It has variously been considered an aberrant
Yellow-rumped Tinkerbird (Short & Horne 2002), or, more recently and with sound reasoning, a valid
species (Collar & Fishpool 2006).
Habits
The White-chested Tinkerbird appears to favour dense, evergreen Cryptosepalum thicket, and large areas of
this forest type dominated by C. pseudotaxus still occur in n
orth-west Zambia and neighbouring Angola.
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The bird has a large, comparatively robust bill, which may be an adaptation for feeding on the berries of
mistletoes (Collar & Fishpool 2006).
The Kinglet Calyptura is the smallest member of the cotinga family, and is (or was) restricted to a tiny area
in Rio de Janeiro in the foothills of the Serra do Mar (Kirwan & Green 2011). A number of specimens were
taken during the mid-19th century, which suggests that it was fairly common, but the population crashed
soon afterwards and it was last collected around 1890. Deforestation due to gold and diamond mining, plus
an increase in coffee plantations, were the primary reasons for its disappearance. It was not found during
surveys made in 1981–82 and was feared possibly extinct (Collar et al. 1992), but surprisingly the bird was
rediscovered in October 1996, when two birds were observed in the Serra dos Órgãos (Pacheco & Fonseca
2001). Unfortunately there have been no subsequent reports, and the present status of this species remains
unknown. If the Kinglet Capyptura survives, its numbers must be perilously low.
Habits
The Kinglet Calyptura occurred in primary forest and abandoned clearings in secondary forest, and actu-
ally appeared to favour the latter (Pacheco & Fonseca 2001), where it foraged for small berries, seeds and
insects (Snow 1982). It was usually found in pairs, and had a surprisingly loud call that was described as
‘brief, raucous and disagreeable’ (Collar et al. 1992; Kirwan & Green 2011).
This species almost certainly still exists, but more recent evidence of its existence has yet to be reported. It
was described from a single individual found dead in May 1984 by Don Smith at Sanganeb lighthouse,
north-east of Port Sudan, Sudan. The lighthouse is situated on a small island about 30km off the coast of
Port Sudan, so the true provenance of the swallow is difficult to establish. Smith was certain he saw other
individuals, and thought the bird likely to occur in the Red Sea hills of Sudan or Eritrea, or possibly in the
coastal hills of western Saudi Arabia north of Jedda (Madge & Redman 1989). Possible sightings of cliff
swallows, which may be referable to this species or perhaps an undescribed taxon, have been made at Lake
Langano and in Awash National Park in the Rift Valley, Ethiopia (BirdLife International 2011). The Red Sea
Swallow has not been relocated some 37 years after discovery, and its status remains shrouded in mystery.
399
The White-eyed River Martin was first discovered in January 1968 in a winter roost at Bung (or Beung)
Boraphet Lake reservoir along with swallows and other migrant birds (Kitti 1968). Over the next few years,
a number of specimens were taken for the aviculture trade between November and February, one was
photographed alive in December 1968, and two collected from the same locality were exhibited in Bangkok
Zoo in 1971 (Sophason & Dobias 1984). The last confirmed record is an observation of six birds flying low
over the water on February 3, 1978 (King & Kanwanich 1978). Apart from an unconfirmed report of juve-
niles perched in trees on an island lake in Bung Boraphet in January 1980, none has been seen since, despite
extensive searches (Sophason & Dobias 1984; Turner & Rose 1989; Collar et al. 2001).
The disappearance of the White-eyed River Martin is attributed to over-hunting for food and the pet
trade, plus the destruction of the reed beds on which the bird depended in winter. However, it may still
survive, as the martin’s summer distribution remains unknown and there have been rumours that breeding
populations persist in river valleys in south-western China and Cambodia (Dickinson 1986). However,
thirty years has now passed since the last confirmed sighting and the status of this mysterious and elusive
species remains unknown. If it is still extant, its numbers must be perilously small.
Habits
Little is known. The martins were migratory birds, roosting in reed beds in winter, and their flight was
described as graceful and buoyant (Turner & Rose 1989). The stomach of one individual contained the
remains of a beetle (Kitti 1968). The White-eyed River Martin may have been nocturnal or at least crepus-
cular, as its eyes, feet and claws were comparatively large, inferring that it may have been a cave, rock or
tree-hollow nesting species that emerged only at night (Rasmussen, in Tobias 2000).
The Black-browed Babbler is an enigmatic bird that was briefly mentioned by Büttikofer (1895b), and
figured by Smythies (1940), with some doubt as to its status. Mees (1995) confirmed its validity and noted
that the type and only known specimen was obtained by the collector C. Schwaner some time between 1843
and 1848; he almost certainly encountered the bird in lowland forest in South Kalimantan in southern
Borneo. It has not been recorded since.
South Kalimantan has suffered severe deforestation since the mid-19th century, most recently for
oil-palm and rubber plantations, and the major fires that occurred in 1997–1998 may have removed any
hope of its continued existence (Collar & Robson 2007).
Habits
Nothing is known about this bird, but the comparatively short and stout tarsi suggest that it may have been
a more arboreal species than other Malacocincla babblers.
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The Sangihe White-eye appears to have always been a rare species. Until recently it was known only from
the type specimen, collected during an expedition in 1888 in a coconut plantation near Manganitu, a coastal
town (Blasius 1888b). However, Manganitu was actually the base of this expedition, and it is more likely
that the specimen was taken in forest on the steep slopes of Gunung Sahendaruman (Rasmussen et al. 2000).
The bird was not seen again until August 1996, when it was observed on two consecutive days (Riley
1997). Subsequent sightings took place on November 10, 1996, once in November 1998, and three sound
recordings of an individual bird were made between February 16 and 19, 1999 in a small remnant patch of
forest in Gunung Sahendaruman and adjacent Sahengbalira (Wardill & Hunowu 1998; Rasmussen et al.
2000; Riley 2002). Despite recent surveys of this locality, the white-eye has not been recorded since. Sangihe
has now been almost totally deforested, and current agricultural and forestry developments threaten even
the last remaining scraps of forest. If the Sangihe White-eye survives, its long-term future is extremely
bleak.
Habits
This bird was associated with primary broadleaved forest, often with a high density of screw pine Pandanus
spp., at between 750m and 1,000m. It was a bird of the mid-storey to upper canopy (BirdLife International
2011), and primarily insectivorous, gleaning insects from leaves; it may have also taken fruit. The contact
call was described as thinner and higher-pitched than that of the closely related Black-crowned W
hite-eye
Z. atrifrons of Sulawesi (Riley 1997).
The specific status of the Whiskered Wren Thryothorus mystacalis is in need of review, as it is sometimes
considered conspecific with the Moustached Wren T. genibarbis. The Whiskered Wren is divided into eight
races, all of which are vulnerable due to severe forest destruction (BirdLife International 2011). The
Colombian Whiskered Wren T. m. macrurus is known from few specimens, and has not been recorded in
recent years. It may be extinct. However, the nominate Whiskered Wren T. m. macrurus is difficult to observe
in dense undergrowth, and best located by its song (Ridgely & Tudor 1994), so there is a possibility that the
Colombian subspecies still survives. If it is still extant, the numbers must be extremely low. Little is known
about the Colombian Whiskered Wren, but the nominate prefers dense undergrowth in humid forests
(Kroodsmar & Brewer 2005).
401
Rueck’s Blue Flycatcher is known from four skins, two obtained from Malaysia as trade skins, the other two
collected around 1917–1918 in secondary lowland forests at Tuntungan and Delitua in the lowland forests
of northern Sumatra (Robinson & Kloss 1919). The exact distribution remains in doubt, and their season of
collection suggests they may have been migrant birds (Collar et al. 2001).
The forests at Tuntungan and Delitua are on the outskirts of Medan City and severely depleted, so if this
species was sedentary, it is likely to have now disappeared. Its morphology suggests some relationship with
the Hainan Blue Flycatcher C. hainanus of China and South-east Asia (van Marle & Voous 1988).
The Black-lored Waxbill is known from very few specimens, the first taken in 1927 at Kiabo on the banks of
the Lualaba River (Chapin 1928), another in 1948 in Mabwe, one in 1949 at Sombe and another in Kaleka in
1950, the last known (Collar & Stuart 1985).
The Black-lored Waxbill had a restricted distribution. It occurred in small flocks in swampy, level grassy
plains with tall grasses and bushes (Chapin 1928; Collar & Stuart 1985). The range of this species has been
poorly surveyed and if it still exists it might persist in the Upemba National Park (Collar & Stuart 1985).
The Red Crossbill Loxia curvirostra has a vast range, occurring in northern spruce forests in Europe, Asia and
North America. It has been divided into a large number of subspecies, most of which are not endangered.
One, the Newfoundland Red Crossbill L. c. percna, was endemic to Newfoundland, Canada, where it was
occasionally sympatric with visiting populations of the larger White-winged Crossbill Loxia leucoptera
(Environment Canada 2006). It had co-evolved with the Black Spruce Picea mariana (Benkman 1993),
although it also fed on the seeds of other trees, and had a more robust bill than other North American
subspecies; it was able to survive year-round on Newfoundland due to the retention of seeds in the cones of
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the Black Spruce during the winter (Benkman 1993; Parchman & Benkman 2002). Furthermore, the closed
cones and thinner scales of Black Spruce on Newfoundland, as opposed to the thicker scales and more open
cones of mainland trees, required more robust jaws to be exploited (Parchman & Benkman 2002). The cone
structure of the spruce and the jaw adaptation exhibited in the Newfoundland birds had evolved in the
absence of American Red Squirrels Tamiasciurus hudsonicus (Benkman 1993).
Decline
The Pine Marten Martes americana had been over-hunted on Newfoundland, so in an effort to restore the
population, the Newfoundland Wildlife Service introduced the American Red Squirrel in 1963 and 1964 as a
food source (Dodds 1983). In a short time, the squirrels had reached extremely high densities and started
out-competing the crossbills for spruce seeds (Benkman 1993). The crossbill was considered common prior
to the squirrel introduction (Austin 1968), but severe declines were noted in the population during the 1970s
(Pimm 1990); the last confirmed observation on Newfoundland was in 2002 (COSEWIC 2004). Some Red
Crossbills were observed breeding in 2005 and 2006 but it is uncertain if these were of the Newfoundland
race (Environment Canada 2006). More recent observations may also be of mainland birds. There is some
hope that a small population persists on squirrel-free islands off Newfoundland.
Habits
The Newfoundland Red Crossbill was known locally as the ‘Spruce Mope’, in reference to their slow move-
ments when feeding in the tops of spruce trees. They foraged in flocks, often with White-winged Crossbills,
and were generally associated with Black Spruce forest, but also fed on the seeds of other tree species and
on invertebrates.
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This section covers taxa known only from inadequate illustrations and written accounts from untrained
observers, or from descriptions or illustrations where the original specimen or specimens are now lost. The
term ‘invalid taxa’ is used for convenience only, as the names are not invalid in terms of nomenclature. The
primary basis for including these birds is that they sometimes appear in lists of extinct or hypothetical
birds. Many are unidentifiable, whereas others have proved to be multiple descriptions of the same species,
or even in some cases completely invented birds. Where skins exist, molecular and morphological work has
shown them to be hybrid forms, aberrant individuals, or artefacts. No doubt some of these taxa are genu-
inely extinct species, but their correct status can now never be determined.
Now considered a synonym of Ornate Tinamou Nothoprocta ornata branickii (Krabbe & Schulenberg 2005).
Considered a synonym of the nominate race of Solitary Tinamou Tinamus s. solitarius (Amaral & Silveira
2004).
Known only from hearsay comments by Sclater (1862) and Heuglin (1859) and an unpublished picture by
Levaillant. Now considered a synonym of Ornate Tinamou Nothoprocta ornata branickii (Krabb & Schulenberg
2005).
Described from a wax cast taken from an egg in the Museo de La Plata (Lydekker 1894b), which lacks diag-
nostic characters. This wax cast is at Tring).
May represent a n ow-extinct species, but at present best regarded as a nomen dubium, mainly because the
holotypical specimens (eggs only) lack determinable characters.
May represent a n ow-extinct species, but at present best regarded as a nomen dubium, mainly because the
holotypical specimens (eggs only) lack determinable characters.
Gilbert (1789) reported ‘peacocks’ on Lord Howe Island, which have been interpreted as representing a
species of megapode. Five eggs and two chicks of a megapode were supposedly collected on Lord Howe
Island by H. H. Romilly, and listed in Seebohm’s c.1895 catalogue. However, these were actually collected
on New Hope Island (Niuafo’ou), not Lord Howe Island, and are specimens of the extant Tongan Megapode
M. pritchardii (Lister 1911).
Known only from Sonnerat’s description, from which all others derive. Unidentifiable and now rejected
(Mayr 1963).
Known only from the accounts of Stork (1767) and Le Conte (1854).
405
Known only from Linnaeus’s description, which he took from a depiction on a Chinese dish. Perhaps a
fabulous bird, a phoenix, or some kind of domestic fowl.
Considered synonymous with nominate Grey Partridge Perdix perdix perdix (Violani et al. 1988; McGowan
1994).
Known only from the types. Believed to be a hybrid between Gambel’s Quail Lophortyx gambelii and
Douglas’s Quail L. douglasii (Peters 1934 [1931–48]).
A population of Northern Bobwhite C. virginianus, now extinct on the island. Known only from the type
collected in 1888. Aldrich (1946a) examined a large series of Colinus quails and concluded that individual
variation was intense, and that C. v. insularis fell within the range of the Florida race, C. v. floridanus.
Known only from eggs and immature birds. Conspecific with B. c. asiatica or B. c. hutchinsii.
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Known from a single specimen, now lost. Probably a leucistic Bewick’s Swan C. bewickii (Kear, in Scott
1972).
Formerly believed to be a full species, but now considered a morph or mutation of the Mute Swan Cygnus
olor (Kear in Scott 1972).
Known only from Forskål’s description. Synonymised under Ferruginous Duck Aythya nyroca by Salvadori
(1895).
Known only from S. G. Gmelin’s description. Synonymised under Ferruginous Duck Aythya nyroca by
Salvadori (1895).
Known from various early accounts. Synonymised under Ferruginous Duck Aythya nyroca by Salvadori
(1895).
407
Known only from the inadequate descriptions of Pennant and Latham. Considered to be conspecific with
the Common Eider Somateria mollissima by Salvadori (1895).
Known only from Sonnerat’s account and depiction. Lysaght (1952, 1956a) showed that Sonnerat was
unscrupulous in using other people’s material, and the account and description was almost certainly
plagiarised.
Known only from Molina’s description. Hellmayr & Conover (1948) state that owing to ‘the incomplete
diagnosis, together with the absence of any definite locality’, the name cannot be identified with any extant
species, and suggest that Molina was probably describing Magellanic Spheniscus magellanicus or Humboldt
S. humboldti penguins.
Considered to be either a dark morph of the Black-capped Petrel Pterodroma hasitata, a population of the
Jamaican Petrel Pterodroma caribbaea (see p. 67), or a mistaken identification of Audubon’s Shearwater
Puffinus lherminieri (Collar et al. 1992; BirdLife International 2011).
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Known from a water colour of about 1792 held in the Mitchell Library in Sydney, Australia. Considered to
be synonymous with the Black-winged Petrel Pterodroma nigripennis (Jouanin & Mougin 1979).
Known only from Solander’s description. G. R. Gray (1871) included P. agilis as a doubtful synonym of his
own P. phillipii, the Mount Pitt Petrel (see p. 68).
409
Known only from the type, in Tring, collected by Rev. T. Powell. Generally considered to be a melanistic
Polynesian Storm-petrel Nesofregetta fuliginosa, but some authorities (e.g. Clements 2000) give this form
subspecific status. If so, this subspecies is critically endangered, if not already extinct.
Known only from a right ulna, but misidentified. Cowles (1987) concluded that it actually belonged to a
Whimbrel Numenius phaeopus, a common migrant to the Mascarenes.
Known only from Molina’s description. Deautier & Steullet (1929) considered that this description agreed
with the Great White Egret Ardea alba egretta (Gmelin), whereas Hellmayr & Conover (1948) consider it be
an invented bird.
Known only from Molina’s description. Never identified, but the name cyanocephala was formerly used
incorrectly for a race of the Black-crowned Night Heron Nycticorax nycticorax obscurus, and for the Whistling
Heron Syrigma sibilatrix. Hellmayr & Conover (1948) dismissed the Blue-headed Heron as ‘indeterminable
or even fictitious’.
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Unidentifiable.
Known only from Molina’s description. Never identified, and ignored by Sharpe (1898) and Hellmayr &
Conover (1948).
Known from the descriptions of Pennant and Latham. Considered unidentifiable by McAtee (1963).
Known only from Catesby’s description, from which the other accounts derive. Never identified, but may
represent a n
ow-extinct form. Sharpe (1898) identified Linnaeus’s Ardea aequinoctialis as the white morph of
the Reddish Egret Egretta rufescens.
Known only from Catesby’s description. Ridgway (1878) treated it as fictitious, or the same as the Great
Blue Heron Ardea herodias.
Now considered synonymous with the Australian Pelican Pelecanus conspicillatus (Worthy 1998).
411
Known from a number of specimens from tropical Africa. Usually dismissed as an aberration ( Ogilve-Grant
1898; Bannerman 1930a).
Known only from Pennant’s brief description. Unidentifiable (Hellmayr & Conover 1948).
Known only from bones collected in New Zealand, and said to be larger than other New Zealand species
(Forbes 1892d). Falls within the range of variation of Great Cormorant P. carbo (Worthy & Holdaway 2002).
Known only from Audubon’s description (1849), based on specimens seen by Townsend. Considered to be
Pelagic Cormorant P. pelagicus (Ogilvie-Grant 1898).
Known only from Audubon’s description (1849), based on specimens seen by Townsend. Considered to be
Pelagic Cormorant P. pelagicus (Ogilvie-Grant 1898).
Known only from Pallas’s description, based on Steller’s notes. There is little evidence to suggest that
Steller’s ‘white sea raven’ was even a cormorant. Probably unidentifiable. Pallas (1811) thought Steller was
referring to a Gannet Sula sp.
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Known only from the account of Sonnerat. Almost certainly a dreamed-up bird.
Known only from early descriptions. Synonymised under Egyptian Vulture Neophron percnopterus
(Strickland 1855).
Known only from Gesner’s description, from which all others cited. Unidentifiable.
Known only from Levaillant’s description. Synonymised under the Eurasian Black Vulture Aegypius mona-
chus (Sharpe 1874).
This has generally been considered as either a subspecies (‘niger’) of Steller’s Sea Eagle Haliaeetus pelagicus
(for example, F erguson-Lees & Christie 2005), or a r ange-specific dark morph. Not seen since the 1950s, a
female hatched in captivity in 2001 exhibited this coloration, but both parents were regularly coloured
Steller’s Sea Eagles (Kaiser 2010). This effectively settled the question.
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Known only from the description of Rafinesque. Considered to be an aberrant Bald Eagle Haliaeetus leuco-
cephalus (Rhoads 1911), which make no sense on zoogeographical grounds. Unidentifiable.
Latham (1822) considered the White Eagle to be merely a colour morph of the Golden Eagle. It has not been
seen since the 18th century.
Known only from the descriptions of Pennant and Latham. Strickland (1855) synonymised the B
lack-cheeked
Eagle with the North American race of Osprey Pandion haliaetus carolinensis.
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Strickland (1855) placed the Louisiana White Eagle in the synonymy of the Bald Eagle Haliaeetus leucoceph-
alus, with a note that Du Pratz was inaccurate.
Known only from Latham’s description of the lost type, formerly in the Bullock Collection. Strickland (1855)
synonymised it with Bonelli’s Eagle Aquila fasciata.
Generally believed to be an abnormal specimen of Red-tailed Hawk Buteo jamaicensis harlani (Ridgway 1884,
1885a).
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Status Extinct; known only from bones and possibly one very brief account. Date of extinction c.17th century. Now shown to be
referable to the extant Réunion Harrier.
Range Mauritius, Mascarene Islands.
Subfossil bones found in the Mare aux Songes, Mauritius, were considered by Newton & Gadow (1893) to
be similar to the Black Sparrowhawk Accipiter melanoleucus of South Africa in size and relative proportions,
but they retained the specific alphonsi on biogeographical grounds. Mourer-Chauviré et al. (2004), however,
have not only shown that the subfossil remains are referable to Circus not Accipiter, but they are indistin-
guishable from the extant Réunion Harrier Circus maillardi; the two birds were actually conspecific. Réunion
would have had a distinct lack of open areas in the past, so the Réunion Harrier has evolved short, broad
wings, short tarsi and long claws (Ferguson-Lees & Christie 2001), adaptations for hunting birds in thick
forest, a similar evolutionary trait exhibited by the Hawaiian Harrier Circus dossenus (see p. 90). The harrier
was possibly mentioned on Mauritius in 1602 (Cheke & Hume 2008), but never again, and probably disap-
peared as a result of persecution and deforestation.
Carter (2007) showed it represented a hybrid population, which has been extirpated since 2000 (perhaps
earlier) due to hybridisation with Black Kite Milvus migrans (see also Johnson et al. 2005, Hille & Collar
2009).
Known only from the original description with no surviving specimens. Considered referable to the Black
Caracara Daptrius ater (Hellmayr 1921).
Known only from the plate in Frisch, on which the descriptions of Brisson and Latham are based.
Unidentifiable.
Known only from the description by Henry Salt following his voyage to Abyssinia. Unidentifiable.
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Known only from Piller’s description. Strickland (1855) placed it in the synonymy of the Hen Harrier Circus
cyaneus.
Known only from a description in Albertus Magnus. Strickland (1855) placed it in the synonymy of Saker
Falcon Falco cherrug.
Known from several early accounts, but it is unclear whether they all refer to the same bird. Unidentifiable.
Known only from Forster’s description. Synonymised under the Gyrfalcon Falco rusticolus (Hellmayr &
Conover 1948).
417
Known only from Levaillant’s description. Considered unidentifiable (Hartert & Neumann 1914).
Known only from Pennant’s description. Probably based on a misidentification of a Canada Goose Branta
canadensis, or other large geese, large grouse or cranes.
Based on the account of Peter Mundy. Probably referable to a juvenile Mauritius Red Rail Aphanapteryx
bonasia (Olson 1977b) (see p. 126).
Known only from the type. Now considered a melanistic colour aberration of Buff-banded Rail Hypotaenidia
philippensis, based on unpublished genetic work and recent analysis (R. Dekker in litt. 2008; Hume & van
Grouw 2014), and removed from the 2010 IUCN Red.
Known from one specimen, obtained from the zoological dealer Leadbeater, of doubtful origin. BirdLife
International (2011) removed this species from their list of extinct species as the type is believed to be a
badly prepared specimen of G rey-necked Wood Rail A. cajanea (Meyer de Schauensee 1966), or a possible
subspecies of it (Taylor & van Perlo 1998).
Based only on the account of Leguat (1708), who almost certainly confused it with a flamingo Phoenicopterus
sp. (Cheke & Hume 2008).
Known only from a single element with an insufficient description (Olson 1977b). Synonymised under
Gallirallus australis (Gill et al. 2010).
Known only from subfossil material, which has never been properly characterised or illustrated (see Olson
1977b). Synonymised under Gallirallus australis (Gill et al. 2010).
Known only from an inadequate description in King (1784), and now a synonym of Hawaiian Crake
Zapornia sandwichensis (see p. 120).
Known only from the plates of Miller and Georg Forster. The name Rallus ecaudatus is already preoccupied
(Lysaght 1953); this is now synonymised with Hawaiian Crake Zapornia sandwichensis (see p. 120)
Known only from the type in the Museum of Comparative Zoology at Harvard, formerly preserved in
alcohol. Now considered a juvenile Lord Howe Wood Rail Hypotaenidia sylvestris (Olson 1992a).
419
Based on the extremely poor probable rendition of Aphanapteryx bonasia in Herbert (1634; see Olson 1977b).
Known only from Colenso’s account. Now thought to be a misidentification of the New Zealand Grebe
Poliocephalus rufopectus.
Considered probably extinct (Dickinson et al. 1991). However, now synonymised under Grus antigone sharpii
(Dickinson 2003).
Known only from Audubon’s description, as published by Nuttall. Ridgway (1878) placed it in the
synonymy of the Great White Heron Ardea occidentalis, which is now considered to be a white morph of the
Great Blue Heron Ardea herodias.
420
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Known only from Marco Polo’s description, as translated by Yule. Synonymised under Siberian Crane Grus
leucogeranus (Blaauw 1897).
Known only from the type specimen, collected in November 1840. Considered to be an immature Shore
Plover T. novaeseelandiae (Buller 1887–88) or invalid (BirdLife International 2011).
Unidentifiable.
Known only from Müller’s account, perhaps based on an earlier work by Ólafsen (1774–75). Never identi-
fied, and almost certainly unidentifiable.
Known only from Hartert’s comments on Krustenstern’s plate. Probably fictitious, based on a mixture of
species.
421
Now considered to probably represent a hybrid between Curlew Sandpiper Calidris ferruginea and
Sharp-tailed Sandpiper C. acuminata (Cox 1989, 1990).
DNA analysis has shown that Cox’s Sandpiper is a hybrid Curlew Sandpiper C. ferruginea x Pectoral
Sandpiper C. melanotos (Christidis et al. 1996).
Originally known only from Pallas’s description, taken from Steller’s manuscript. Synonymous with the
extant Aleutian Tern Onychoprion aleuticus.
Known only from several early accounts and Daubenton’s plate in the Planchees Enluminées. Inadequate
description and difficult to determine whether more than one species is involved.
422
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Description based on an aberrant individual, or possibly confused with Great Northern Diver Gavia immer.
Much ink has been spilt over the White Dodo (or Solitaire) of Réunion, a species even given specific status
in a relatively recent monograph on the pigeons (Gibbs et al. 2001), and yet no physical evidence of any kind
has been found to substantiate its existence. The supposed illustrations of white dodos have no provenance
data whatsoever, and are likely to represent a leucistic bird from Mauritius (Hume & Cheke 2004), or are
entirely imaginary. The accounts of white birds on Réunion, which some authorities continue to associate
with dodos, are almost certainly referable to the now extinct Réunion Ibis Threskiornis solitarius
(Mourer-Chauviré et al. 1995a, b) (see p. 76). Large numbers of bird subfossil remains have now been recov-
ered from Réunion, and not a single dodo bone has been identified (Mourer-Chauviré et al. 1999). Until
evidence to the contrary is made available, the White Dodo of Réunion must remain as mythical as the
griffin or phoenix.
Known only from the description by Hernandez, on which all subsequent accounts are based. Considered
fictitious (Salvadori 1893) or possibly a domestic pigeon (Smellie 1812).
Known only from the plate in Temminck & Knip (1811). The provenance data for this supposed species is
doubtful, and the bird itself is probably fictitious (Salvadori 1893).
423
Known only from Sonnerat’s description, the basis of Scopoli’s name Columba cinerea. Almost certainly
referable to Madagascar Turtle Dove Nesoenas picturata (Walden 1877 [1875]), and considered doubtful
(Wagler 1827).
Known only from Daubenton’s plate, on which Latham’s (1790) description was based. Almost certainly
fictitious (e.g. Wagler 1827; Salvadori 1893).
Known only from Miller’s plate and inadequate description. Never identified, and considered a doubtful
species (Wagler 1827; Bonaparte 1854a).
Known only from Pallas’s inadequate second-hand description, based on Gmelin’s manuscript.
424
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Because of the ambiguity surrounding Latham’s (1801b) description under Columba norfolciensis of three
quite distinct birds, the name was formally suppressed (ICZN 2010; Forshaw 2016), and the name is no
longer considered valid.
Known only from the descriptions of Linnaeus (1766) and Möhring (1752), both based on Seba’s (1734)
plate. It is uncertain that this bird even represents a pigeon, as some authors referred it to Cuculidae
(cuckoos).
Unidentifiable.
Apparently known only from Levaillant’s description. Considered fictitious (Sundervall in Layard 1867).
Known only from a picture collected by Scottish explorer Sir Alexander Burnes (also known as Bokhara
Burnes). Uncertain if it even represents a columbid, and some authors suggest the image is a sandgrouse.
425
Known only from Levaillant’s plate and description, from which all other accounts derive. Considered a
fictitious bird (Salvadori 1893).
Known only from Latham’s description. Never satisfactorily identified. With such a supposedly vast range,
it defies belief that no other specimen was obtained or described. Probably represents the European Turtle
Dove Streptopelia turtur.
Known only from Daubenton’s plate and Buffon’s description. Never satisfactorily identified. No other
Macropygia occurs in Africa, but provenance uncertain.
Known only from Albin’s plate and description. Provenance uncertain and the plate differs from the
description.
426
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Known only from King’s statement that a ‘large white pigeon’ was found in the Hawaiian Islands. No
species of Columbidae has ever been recorded from Hawaii (Olson & James 1991), so the identification or
provenance are in error.
Known only from Latham’s description. Considered indistinct (Stephens 1819) and an unidentifiable Treron
(Salvadori 1895).
Known only from Latham’s description. There is no evidence that Latham even had a specimen. Possibly a
description of New Zealand Pigeon H. novaeseelandiae (Salvadori 1893), or the locality is in error (Gray
1862b).
Known only from Percy Earl’s inadequate description. The provenance may be in error (Gray 1862b).
427
Known only from Temminck’s description, based on a captive bird in Amsterdam. Provenance uncertain,
and now unidentifiable.
Known only from Albin’s description and plate. Considered conspecific with European Turtle Dove
Streptopelia turtur (Salvadori 1893).
Known only from Latham’s description, but Gmelin’s name, which was based on it, has been used, possibly
erroneously, for the Thick-billed Green Pigeon Treron nipalensis. Unidentifiable.
Known only from Latham’s description, and doubtfully distinct. Salvadori (1893) listed this bird as an
unidentified form, and Peters (1937 [1931–48]) said that it is ‘doubtless’ the same as the Polynesian Ground
Dove A. erythropterus.
428
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Appendix 2: Doubtful and invalid taxa
Known only from Gosse’s brief and inadequate account and a contemporary illustration.
Known only from Fermin’s description, from which the others derive. No such bird has been reported since.
Known only from Tschudi’s description of the type, formerly in the Museum of Neuchâtel, but now appar-
ently lost. Referred to the synonymy of the M
aroon-chested Ground Dove Claravis mondetoura (Taczanowski
1884–86; Ridgway 1916).
Known only from Hartlaub’s description, based on a specimen received by him in 1883. Salvadori (1893)
dismissed the specimen as probably an example of Tambourine Dove Turtur tympanistria in which the
metallic spots were missing.
Known only from Pelzeln’s lost type with an abnormally developed bill, Latham’s apparently normal spec-
imen (also lost), and another specimen in Liverpool, which lacks the bill. Considered to be an aberration
and thus synonymised under Nestor productus (see p. 189).
429
Known only from a very brief statement by Scarlett (1955a): ‘many bones of a small form [of Kaka] have
been recovered’. Holdaway & Worthy (1993) have shown that this subfossil material is referable to the
extant South Island Kaka Nestor meridionalis.
Known only from the type in Tring, purchased from the Masséna Collection. Although Salvadori (1891a)
considered this species valid, it is generally considered to be a variation of the Kaka Nestor meridionalis
(Buller 1905).
Known only from the unique type, a male in the Leiden Museum. It was collected by Laglaize on March 13,
1876, and purchased from A. A. Bruijn in 1878. Peters (1937 [1931–48]) explained that Ernst Mayr had
suggested to him that this bird was either a hybrid Rajah Lory Chalcopsitta insignis x Yellow-streaked Lory
C. scintillata or an intermediate form, closer to the former.
Known only from Lesson’s description, based on the unique type, now lost. Considered doubtful (Salvadori
1891a) or unidentifiable (Peters 1937 [1931–48]).
Known only from Edwards’s plate and description. For a long time the name was used for the Red Lory Eos
rubra. Considered dubious by Finsch (1868). Coloration considered to match a nutritionally deficient Red
Lory (Low 2011).
430
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Never identified.
Known only from Levaillant’s plate and description, based on a specimen, now lost, formerly in the collec-
tion of Raye de Breukelvaert in Amsterdam. Provenance in doubt.
Known only from the descriptions of Brisson, Daubenton and Levaillant, apparently all based on a spec-
imen (now lost) in the collection of the Abbé Aubry. Salvadori (1891a) put this bird in the synonymy of
Red-and-blue Lory Eos histrio.
Known only from the original types, two living specimens brought by Walter Goodfellow from Obi in the
Moluccas. The specimens were apparently never preserved and are now lost. Holyoak (1970) and Forshaw
(1989) considered it to be a juvenile Red Lory Eos bornea, and Mees (1982) stated it was an invalid species.
Known only from Latham’s description based on a specimen, now lost, formerly in the Leverian Museum.
Synonymised with Violet-necked Lory Eos squamata ricinata (Salvadori 1891a).
431
Known only from a passing comment by E. L. Layard, almost certainly referring to Trichoglossus haematodus
massena, the race of the Rainbow Lorikeet in the New Hebrides.
Known only from Pennant’s description. Salvadori (1891a) included it in the synonymy of Trichoglossus
novaehollandiae, now called T. haematodus moluccanus, the north Australian form of Rainbow Lorikeet.
Known only from the type collected by Alfred Russel Wallace, an adult male that Salvadori (1891a) consid-
ered either a hybrid Rainbow Lorikeet T. [haematodus] nigrogularis x Yellow-streaked Lory Chalcopsitta
sintillata, or an immature of something else.
Known from the type in Paris, and a specimen from the Gould Collection, now in Tring. The type is of
unknown locality; the Gould specimen came from Melbourne and was assigned to the same species by
Finsch (1868). Salvadori (1891a) examined this second specimen, and concluded that it was a hybrid
Rainbow Lorikeet T. h. moluccanus x Musk Lorikeet Glossopsittacus concinnus.
Now considered to be immature of the race T. h. nigrogularis (Juniper & Parr 1998).
Known only from Levaillant’s plate and inadequate description. Synonymised by Salvadori (1891a) with
Purple-naped Lory Lorius domicella.
432
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Known only from Levaillant’s description and plate, based on three specimens, now lost. Considered a
variety of the Purple-naped Lory Lorius domicella (Salvadori 1891a).
Known only from the description and plate of George Edwards, based on a specimen, now lost, in the
possession of Sir Hans Sloane. Considered a questionable species (Salvadori 1891a).
Known only from the description of Latham, based on a specimen, now lost, formerly in the collection of a
Mr. Harrison. Finsch (1868) and Salvadori (1891a) considered this species to be an artefact, composed of a
red lory specimen with feathers of a jay attached.
Known only from Seba’s description, presumably from a specimen in his collection, from which all others
derive. Finsch (1868) and Salvadori (1891a) considered it an imaginary bird.
433
Known only from Albin’s description, based on a live specimen, now lost, which he saw in London.
Unidentifiable.
Known only from Meyen’s description. Finsch (1868) thought the bird was probably the Halmahera race of
the Moluccan King Parrot Alisterus amboinensis hypophonius, or a variety of the Chattering Lory Lorius
garrulus that had been brought as a captive bird to China, where no such parrot is native.
Known only from the unique type in Tring. Believed it to be an aberrant specimen of Lorius domicella
(Forshaw 1971).
Known only from the unique type, a female, considered by Forshaw (1971) to be an aberrant specimen of
the Purple-bellied Lory Lorius hypoinochrous.
Generally considered to be the immature plumage of the Blue Lory Vini peruviana (Finsch 1868; Townsend
and Wetmore 1919).
Known only from the description of Bontius. Salvadori (1891a) stated that both Wagler and G.R. Gray iden-
tify it with the Papuan Lorikeet Charmosyna papuensis.
Known only from the unique type, a female, in the Museum of Genoa. Salvadori (1891a) considered it a
possible accidental variety of Red-cheeked Parrot Geoffroyus geoffroyi pucherani.
Known only from Levaillant’s plate and description. According to Salvadori (1891a) both G.R. Gray and
Otto Finsch considered this to be a variety of Red-cheeked Parrot Geoffroyus geoffroyi, and Wagler (1832)
thought it was a variety of the Senegal Parrot Poicephalus senegalus. Locality confused.
Known only from the unique type of unknown locality. Forshaw (1989) and Collar (2006) examined it and
believed it to be an aberrant specimen of Muller’s Parrot T. sumatranus.
435
Known from the descriptions of Shaw and Latham, probably independently derived. Salvadori (1891a)
remarked that Shaw’s description seemed to apply to a specimen of the King Parrot Alisterus scapularis, and
that Latham’s description differed and could not be identified.
Known only from Latham’s description. Finsch (1868) believed it to be a variety of a species of rosella
Platycercus, a genus of brightly coloured Australian parrots, or possibly a king parrot Alisterus sp.
Unidentifiable.
Known only from the type, shot a few miles north of Dalby, Darling Downs, Queensland, by a Mr. Shaw. It
was paired with a female King Parrot Alisterus scapularis. Peters (1937 [1931–48]) identified it as a hybrid
King Parrot x Red-winged Parrot Aprosmictus erythropterus, while Ramsay (1878) thought it was simply an
aberrant King Parrot.
Known only from the type, a female, now in Tring. Peters (1937 [1931–48]) considered that it represented a
specimen of Barnard’s Parrot Barnardius barnardi in which the yellow pigment is largely lacking.
Known only from a male and female which Salvadori saw alive in the London Zoo. The male is now in
Tring. Now considered a hybrid Crimson Rosella P. elegans x Eastern Rosella P. eximius (Wells 1992).
436
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Known from two specimens in Tring. Peters (1937 [1931–48]) concluded that it is an aberration of Eastern
Rosella P. eximius.
Known from only two specimens in the Australian Museum. Considered to be a hybrid between the
Crimson Rosella Platycercus elegans and the Pale-headed Rosella P. adscitus (Peters 1937 [1931–48]).
Known only from Latham’s description. Finsch (1868) considered the Chlorotic Parrot to be a variety of
some species of the genus Platycercus. Unidentifiable.
Known from Levaillant’s plate and description. Usually considered to be a lutino (yellow variety) of the
Rose-ringed Parakeet Psittacula krameri.
Known only from a specimen, formerly in the Leverian Museum but now lost, on which Latham’s descrip-
tion was based. Unidentifiable.
Known only from an inadequate reference by Russ (1871), probably in error for the Paradise Parrot Psephotus
pulcherrimus.
437
Apparently known only from Bechstein’s description. Considered to be a Platycercus (Salvadori 1891a).
Known only from Kuhl’s description, based on a specimen (now lost) that was formerly in the Bullock
Collection. This bird is now unidentifiable. Gray (1859a) and Finsch (1868) considered it conspecific with
the Black-fronted Parrot Cyanoramphus zealandicus of Tahiti (see p. 172).
Known only from Kuhl’s description of the lost type, formerly in the Bullock Collection. Both Finsch (1868)
and Salvadori (1891b) cite this form as a synonym of C. erythronotus, which itself is a synonym of the
Black-fronted Parrot C. zealandicus (see p. 196). Kuhl described it as resembling Red-fronted Parakeet C.
novaezelandiae in all respects, except that it has a brown forehead.
Known only from Reichenow’s description. This may be a synonym of Malherbe’s Parakeet Cyanoramphus
malherbi, (which itself was considered a doubtful taxon, until recently).
Formerly occurred in the North Canterbury area of New Zealand. Status never satisfactorily determined.
Finsch (1875) said that the form was similar to the small race of R
ed-fronted Parakeet on the Auckland
Islands, C. novaezelandiae aucklandicus.
Known only from Latham’s description. Shaw (1811), Vieillot (1817, 1823), Kuhl (1820a), Wagler (1832),
Finsch (1868) and Salvadori (1891a) all repeat Latham’s account without coming to any conclusion as to
what this bird could be. Unidentifiable.
438
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Appendix 2: Doubtful and invalid taxa
Known only from Virey’s account (in Sonnini 1799–1805). May have represented a Cyanoramphus, but
Virey’s account also suggest that the bird might have been a Rosella Platycercus sp., a group now confined to
Australia and Tasmania.
Known only from Latham’s description, based upon a captive bird of uncertain origin. Salvadori (1891a)
suggested that it is a young Palm Lorikeet Charmosyna palmarum.
Known only from Latham’s description. Salvadori (1891a) stated that ‘Dr. Finsch thinks this is an artificial
specimen’.
Known only from Shaw & Nodder’s description, from which others derive. Finsch (1868) and Salvadori
(1891a) believed that it represented an ‘albino’ (or lutino – a yellow form) of some species of Neophema.
439
Apparently known only from a very old Japanese drawing, on which the various descriptions are based.
According to Salvadori (1891a) both Wagler and Gray identify the bird with Charmosyna papuensis (Papuan
Lorikeet Charmosyna papou).
Known only from Marcgrave’s description, which has been the subject of discussion but never satisfactorily
identified. Provenance uncertain.
Known only from descriptions derived from Aldrovandus. Generally considered to be a variant of the
African Grey Parrot Psittacus erithacus.
Probably known only from Aldrovandus’s description, from which all others derive. Salvadori (1891a)
dismissed it as ‘a fabulous bird’.
440
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Appendix 2: Doubtful and invalid taxa
Apparently known only from Daubenton’s plate, and Buffon’s description, based on a specimen in the Jardin
des Plantes, Paris, but presumably no longer extant. Salvadori (1891a) considered it a doubtful species.
Unidentifiable.
Known only from Latham’s description of a specimen formerly in the British Museum. Unidentifiable.
Based on two male specimens from West Africa. One was from Lord Derby’s collection, which had died in
captivity in 1867; the other was identical in every way. Now considered to be an immature Brown-necked
Parrot Poicephalus robustus fuscicollis (Bannerman 1931; Peters 1937 [1931–48]).
Known only from Heuglin’s inadequate description. Finsch (1868) and Salvadori (1891a) both suggested
that it was conspecific with the Senegal Parrot Poicephalus senegalus.
442
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Appendix 2: Doubtful and invalid taxa
Known only from Alexander’s description of the type, now in Tring. Bannerman (1931) concluded that the
type falls within the range of the West African race of Brown-necked Parrot Poicephalus robustus fuscicollis;
all the supposed characters of P. kintampoensis occur in examples of it.
Both Hartlaub (1879) and Salvadori (1891a) considered this bird to be probably the same as the Black-
collared Lovebird Agapornis swinderniana.
Known only from the descriptions of Latham and Levaillant, from which the others derive. Salvadori
(1891a) remarked that ‘Dr Finsch suspects that this is an artificial bird’.
Known from two specimens. It is now considered to be an aberration of Loriculus philippensis apicalis (Peters
1937 [1931–48]; Forshaw 1989; Juniper & Parr 1998).
Known only from Edwards’s description. Salvadori (1891a) placed it in the synonymy of the Ring-necked
Parakeet Psittacula krameri.
443
Known only from Latham’s description. Salvadori (1891a) called this bird a lutino (yellow) variety of the
Plum-headed Parakeet Psittacula cyanocephala.
Known only from the descriptions and plates of Edwards and Levaillant. Salvadori (1891a) called this bird
a lutino (yellow) variety of the Plum-headed Parakeet Psittacula cyanocephala.
This parakeet was known from a few specimens and its status was surrounded in mystery. It is now consid-
ered to be a captive-bred hybrid between the Slaty-
headed Parakeet Psittacula himalayana and the
Plum-headed Parakeet P. cyanocephala (Rasmussen & Collar 1999a, b).
Known only from Rothschild’s (1905c) account based on Dubois’s (1674) description of the Réunion Parrot
(see p. 205), in which Rothschild completely confused the provenance (Hume 2007).
The Violet Macaw is known only from Rothschild’s (1905b) description, supposedly based on an earlier,
untraceable, reference. There is no other evidence to support the existence of this name, and as no physical
remains of any Anodorhynchus macaws have ever been discovered on the West Indies – only the genus Ara is
known to have colonised the islands. Lenoble (2015) has now shown that Rothschild (1905) erroneously
based the name Anodorhynchus purpurascens on the Guadeloupe Amazon Amazona violacea.
Known only from Bouton’s inadequate account. No other evidence is known. A painting of a blue and
yellow macaw, alongside a dodo, executed by Roelandt Savery in 1626, has sometimes been considered to
represent this species. However, there is no indication at all of the provenance of the specimen upon which
Savery’s painting is based.
Known only from De Rochefort’s inadequate description. Provenance data unreliable. Considered a
synonym of A. martinica (Greenway 1967), itself a dubious species (see above).
Based on one hearsay observation only (see Greenway 1967). Only one species of Jamaican macaw, the
Jamaican Yellow-headed Macaw A. gossei (see p. 380) is known from credible evidence. That three other
large macaws, the Red-headed Macaw A. erythrocephala, the Jamaican Blue-and-Yellow Macaw Ara sp. and
Albin’s Macaw Ara. sp. (see below) supposedly occurred sympatrically on the island, all described by orni-
thologists without a shred of physical evidence, epitomises the over-zealous passion often associated with
extinct birds.
445
Known only from the 1791 account of Thomas Atwood (Atwood 1791). While on Dominica, Atwood
mentions two parrots, a ‘Mackaw’ which was larger than the ‘Common Parrot’, and that both the macaw
and parrot had green and yellow plumage. This description is conceivably referable to the two extant
Dominican Amazona parrots, the Imperial Amazon A. imperialis and the Red-necked Amazon A. arausiaca. If
so, the Imperial Amazon, which is larger than the Red-fronted, was probably the ‘Mackaw’ of Atwood,
while the Red-fronted Amazon was the ‘common parrot’.
Known only from Albin’s (1738) description. Almost certainly referable to the R
ed-and-green Macaw Ara
chloropterus. Albin describes the commonly practised transportation of macaws around the West Indies:
‘They are commonly brought from Jamaica, and other parts of the West Indies, and bear a good price, being
commonly sold for ten guineas’. No doubt this was such an import.
Known only from Buffon’s and other early accounts. Records for Hispaniola are in error (see Walters 1995;
Olson 2005).
Based on a good description by Latham (1822) but almost certainly referable to a colour variety of Scarlet
Macaw Ara macao (Finsch 1867).
446
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Appendix 2: Doubtful and invalid taxa
Known only from the accounts of Marcgrave and Edwards, the latter based on a specimen in the possession
of Lord Carpenter. Considered to be a description of the Chestnut-fronted Macaw Ara severus.
Known only from Azara’s description. Salvadori (1891a) suggested that this was a variety of the
Scarlet-fronted Parakeet A. wagleri frontata, or possibly the Golden-capped Parakeet A. auricapillus.
This supposed subspecies is known from one individual. Juniper & Parr (1998) regard it as an aberrant A.
leucophthalma callogenys.
Known only from the type, of unknown origin, now in Tring. Peters (1937 [1931–48]) considered it a hybrid,
as did Forshaw (1989), who thought it was between the Burrowing Parakeet Cyanoliseus patagonus and
Scarlet-fronted Parakeet Aratinga wagleri.
It seems that Masséna & Souancé never mentioned a Parrot under the above name, and no description of it
has been given either by Bonaparte or anybody else (Salvadori 1891a). Finsch (1868) says that Bonaparte
took the name from a Souancé manuscript.
447
Known from four specimens (according to Peters 1937 [1931–48]) or three specimens (according to Forshaw
1989), all from western Matto Grosso. Now considered to be an aberrant G reen-cheeked Parakeet Pyrrhura
molinae sordida (Juniper & Parr 1998).
Known only from Levaillant’s description and plate. Wagler (1832), Gray, Reichenow (according to
Salvadori 1891a) and Phipps (1989) all believed it to be the young of the Monk Parrot Myiopsitta monachus.
Known only from Latham’s description. May represent a Forpus sp., otherwise unidentifiable.
This description has been generally identified with the Golden-winged Parakeet Brotogeris chrysoptera, but
Salvadori (1891a) rightly points out that the body colour more resembles the Canary-winged Parakeet B.
versicolorus. Provenance locality is also doubtful.
Known only from Miller’s plate, from which other descriptions derive. The origin of his information is
unknown. Salvadori (1891a) thought it was probably an artificial bird.
Unidentifiable but Finsch (1868) suggests a variety of Yellow-billed Parrot Amazona collaria.
Known only from the original description of Aldrovandus, from which all others derive. Provenance in
doubt.
Known only from Reichenow’s description and coloured plate. The sole specimen, a cagebird, seems to
have disappeared. Peters (1937 [1931–48]), following Salvadori (1891a), synonymised A. hecki with race
lilacina of the R
ed-lored Parrot Amazona autumnalis.
Known from Levaillant’s description and plate, and possibly also from Daubenton’s plate. Salvadori (1891a)
identified it as a variety of the Cuban Parrot Amazona leucocephala.
449
Known only from the plate and description of Catesby, based on a live specimen he saw in Carolina.
Identified by Salvadori (1891a) as a leucistic specimen of Cuban Parrot Amazona leucocephalus.
The evidence for this species is spurious at best, and was considered extremely doubtful by Salvadori
(1891a).
Never identified.
Known only from Levaillant’s plate and figure, on which the other references are based. Never satisfactorily
identified and now unidentifiable.
Doubts have been expressed as to the validity of this supposed subspecies (Mees 1964). It represented a
cave-dwelling population, and is poorly known; the birds have not been reported since the 1960s.
Known only from Vieillot’s description. Not mentioned by the AOU (1957), and Ridgway (1914) cited the
species with a query in the synonymy of the American Wood Owl Ciccaba virgata virgata. Unidentifiable.
Known only from Latham’s description of a specimen, now lost. Not identifiable with any known species
(Peters 1940 [1931–48).
Known only from Vieillot’s description. Never identified, but the name was formerly used for the Spotted
Eagle Owl Bubo africanus.
Known only from the unique type, a female with traces of downy juvenile feathers on the head. Peters (1940
[1931–48]) doubted its validity, and Vaurie (1960b) merged this race and another in Bubo bubo kiautschensis.
Known only from Pennant’s description. McAtee (1963) considered it synonymous with the Snowy Owl
Bubo scandiaca.
451
Known only from the type, now lost, stated to have been in the Würtzburg Museum. Considered unidentifi-
able (Peters 1940 [1931–48]).
Known only from Latham’s description. Peters (1940 [1931–48]) stated that is was practically
unidentifiable.
Known only from the type, an adult male. Bond (1951) and Traylor (1952) considered this bird synonymous
with Rufescent Screech Owl Otus ingens.
Known only from the type. Considered a synonym of Plain Nightjar Caprimulgus ludovicianus by Grant &
Mackworth-Praed (1947) and Mackworth-Praed & Grant (1952), and not included in a recent monograph
on the group (Cleere 2010).
452
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Known from three specimens, one in the Elliot Collection, a second in the Salvin-Godman Collection, and a
third in the Berlepsch Collection. Peters (1945 [1931–48]) considered it to be a hybrid Black-throated Mango
Anthracothorax nigricollis x Ruby Topaz Chrysolampis mosquitus.
Known from the type, an immature male collected by Longuemare, and considered an immature Amethyst
Woodstar C. amethystina (Schuchmann 1999).
Known only from Bogotá trade skins. Considered a melanistic bird of doubtful origin (Schuchmann 1999).
Known only from the type. Now considered a melanistic specimen of an unknown hermit (Hinkelmann
1999).
Known only from the type. Affinities uncertain, generally regarded as conspecific with the G
rey-chinned
Hermit P. griseogularis (Peters 1945 [1931–48]).
453
Known only from the type, an imperfect specimen. Considered a synonym of P. zonura (Peters 1945 [1931–
48]), which itself is now a subspecies of Grey-chinned Hermit P. griseogularis.
Known from two specimens, an immature male collected by Buckley, and an adult male collected in 1898.
Now considered to represent an aberrant Sparkling Violet-ear C. coruscans (Schuchmann 1999).
Known only from a specimen in the Elliot Collection. Now considered a hybrid B lack-throated Mango
Anthracothorax nigricollis x Ruby Topaz Chrysolampis mosquitus (Peters 1945 [1931–48]).
Known only from two specimens of uncertain origin, no longer extant. Now placed in the synonymy of the
Black-throated Mango Anthracothorax nigricollis (Peters 1945 [1931–48]).
Known only from the type. Generally considered to be an artefact (Simon & Hellmayr 1908).
Known only from Floericke’s account, based on specimens in his collection, now lost.
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Known only from the unique type, a Bogotá trade skin. Now regarded as a hybrid between the Festive
Coquette Lophornis chalybeus and the Spangled Coquette Lophornis stictolophus (Schuchmann 1999).
Known only from a unique specimen in the Elliot Collection. Considered an aberrant B
lue-chinned Sapphire
Chlorostilbon notatus (Schuchmann 1999).
Known only from two specimens, both ‘Bahia’ trade skins. Considered an aberrant B lue-chinned Sapphire
Chlorostilbon notatus or a hybrid C. notata x White-chinned Sapphire Hylocharis cyanus (Schuchmann 1999).
Known only from the type, believed to be an adult male, a former mount in the Gould Collection. Considered
to be a probable aberrant Green-tailed Emerald C. alice (Schuchmann 1999).
Known only from the type series, and considered a synonym of the eastern Brazilian race of G
littering-bellied
Emerald, Chlorostilbon aureoventris pucherani (Cory 1918; Peters 1945 [1931–48]).
Known only from the unique specimen, an adult male, in the Berlepsch collection, which was a Bogotá
trade skin. Probably an aberrant Short-tailed Emerald C. poortmani (Schuchmann 1999).
455
Known only from the type in the Museum Heine. Probably an aberrant S
hort-tailed Emerald C. poortmani
(Schuchmann 1999), which would make the locality of Peru in error.
Known only from the type, believed to be an adult male, collected by Gould’s correspondent, Reeves. Now
regarded as a hybrid Glittering-bellied Emerald Chlorostilbon aureoventris x Amethyst Woodstar Calliphlox
amethystina (Butler 1931; Peters 1945 [1931–48]; Graves 1999b).
Known only from a unique specimen in the Vienna Museum, collected by Natterer. According to Berlioz
(1938) this is probably a hybrid between an Amethyst Woodstar Calliphlox amethystina and another undeter-
mined species.
Known only from Bogotá trade skins. Considered to be a hybrid (Berlioz 1937), the most recent assignation
being between the Fork-tailed Woodnymph Thalurania furcata x G
olden-tailed Sapphire Chrysuronia oenone
(Schuchmann 1999).
Known only from two Bahia trade skins. Considered a hybrid, with parents unknown (Meyer de Schauensee
1966, 1970).
Known only from the unique type. Thought to be the female of the Emerald Woodnymph Augasma smarag-
dinea (Simon & Hellmayr 1908), itself considered to be a hybrid (see below). Berlioz (in Peters 1945 [1931–48])
considered it a hybrid.
457
Known only from an adult male in the Berlepsch collection, and one other specimen in the Nehrkorn
Collection. Meyer de Schauensee (1966) considered the type to be a hybrid Long-tailed Sylph Aglaiocercus
kingi x P
urple-backed Thornbill Ramphomicron microrhynchum.
Known only from three specimens. Berlioz (1938) considered the bird to be a hybrid W hite-chinned
Sapphire Hylocharis cyanus x Glittering-bellied Emerald Chlorostilbon aureoventris pucherani.
Known only from the descriptions of du Tertre and Bancroft, of uncertain origin
Known only from the type series in the Rothschild Collection, and considered to be a variety of the
Red-billed Streamertail Trochilus polytmus.
Known only from the unique type collected by Beske. Considered a possible hybrid White-throated
Hummingbird Leucochloris albicollis x G
littering-bellied Emerald Chlorostilbon aureoventris pucherani (Berlioz
1938; Schuchmann 1999).
458
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Known only from Reichenbach’s plate. Placed in the synonymy of the White-bellied Hummingbird Amazilia
chionogaster (Salvin 1892).
Known only from a Bogotá trade skin in the Paris Museum and another in the R ousseau-Decelle collection,
Paris. Considered a probable aberrant Glittering-throated Emerald A. fimbriata (Schuchmann 1999).
Known only from the type, an adult male from Colombia (probably a Bogotá trade skin) in the Gould
Collection. Simon (1910) considered it a hybrid Versicoloured Emerald Amazilia versicolor x Golden-tailed
Sapphire Chrysuronia oenone.
Known only from a Bogotá trade skin, believed to be a female. Considered a probable melanistic female of
Andean Emerald A. franciae (Meyer de Schauensee 1966; Schuchmann 1999).
Known only from the type, which is an artefact, with the body of a Ruby-topaz Hummingbird Chrysolampis
mosquitus and the head of a R
ed-bellied Emerald Chlorostilbon gibsoni (Simon & Hellmayr 1908; Simon 1921).
Known only from the type, an adult male collected by de Oca. Considered to be a hybrid Azure-crowned
Hummingbird Amazilia cyanocephala x Berylline Hummingbird A. beryllina (Berlioz 1932).
459
Known only from de Oca’s description and plate. Generally placed in the synonomy of Amazilia ocai (Peters
1945 [1931–48]; see above).
Florence van Rossem’s Hummingbird Amazilia florenceae (van Rossem & Hachisuka)
Saucerottia florenceae van Rossem & Hachisuka 1938, 407 (Rancho Santa Barbara, 20 miles n
orth-west of Guirocoba, Sonora)
Status Doubtful taxon.
Putative range Sonora, Mexico.
Known only from the unique type, a female. Now considered to be a hybrid (Peters 1945 [1931–48]).
Known from only two specimens. This bird is believed to be an extreme example of variation occurring
throughout the western Mexican range of the Violet-crowned Hummingbird Amazilia violiceps (Weller &
Schuchmann 1997).
Apparently known only from the co-types in the Elliot Collection. Simon & Hellmayr (1908) considered the
‘type’ of neglectus to be an artefact composed of the body of the Versicoloured Emerald Amazilia versicolor
and the head of a White-chinned Sapphire Hylocharis cyanus.
Known only from an adult male specimen held in New York. Status and affinities uncertain (Peters 1945
[1931–48]; Meyer de Schauensee 1966; Sibley & Monroe 1990).
Known only from the type, an adult male. Considered to be a hybrid between Cinnamon Hummingbird
Amazilia rutila and Rufous-tailed Hummingbird A. tzacatl (Graves 2003).
460
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Known only from the type, and possibly another specimen from the Gould Collection. Simon (1897)
believed the bird to be a melanistic variety of the G reen-bellied Hummingbird A. viridigaster and it was
placed in the synonymy of A. viridigaster by Peters (1945 [1931–48]). Weller (2001) considered it to be a mela-
nistic form of the Indigo-capped Hummingbird A. cyanifrons, rather than A. viridigaster.
Apparently known from two males. Cory (1918) listed it as a melanistic variety of G reen-
bellied
Hummingbird A. viridigaster, and Peters (1945 [1931–48]) also synonymised it with A. viridigaster.
Known only from specimens in the Simon collection. Status and affinities uncertain.
Known from two adult male specimens in the Gould Collection, one from Popayán, the second a Bogotá
trade skin. Elliot (1878) considered it to be a dark variety of the Brown Inca Coeligena wilsoni, while Berlioz
(1936a) considered it to be a hybrid Black Inca C. prunellei x Bronze Inca C. coeligena columbiana, which was
confirmed by Graves (2001).
Known from a dozen or more Bogotá trade skins. Believed to be a hybrid Collared Inca Coeligena torquata x
Buff-winged Starfrontlet C. lutetiae (Berlioz 1936a; Schuchmann 1999).
Known only from an adult male Bogotá trade skin held in Tring. Considered to be a hybrid between the
Collared Inca Coeligena torquata and the Mountain Velvetbreast Lafresnaya lafresnayi (Schuchmann 1999).
461
Known only from the unique type in the Royal Albert Memorial Museum, Exeter, of unknown locality.
Graves (1991) considered it to be a light-damaged specimen of Sword-billed Hummingbird Ensifera ensifera.
Known only from the type held at Tring, and possibly another in the Simon collection. Considered a prob-
able melanistic Amethyst-throated Sun-angel Heliangelus amethysticollis (Fjeldså & Krabbe 1990;
Schuchmann 1999).
Known only from Bogotá trade skins. Considered an aberrant Amethyst-throated Sun-angel Heliangelus
amethysticollis of race clarisse (Peters 1945 [1931–48], Fjeldså & Krabbe 1990).
Known only from the type held at Tring, a male collected by Henry Whitely and given by him to Boucard.
According to Simon & Hellmayr (1908), the type of Henry’s S un-angel is a specimen of Gorgeted S
un-angel
H. strophianus with the central rectrices tinged with bronze green.
Known only from two specimens of uncertain origin, held at Tring. Graves (2001) concluded that it was a
colour variant of Gorgeted Sun-angel H. strophianus.
Known only from Bogotá trade skins, and a specimen from Antioquia. Graves (1990) determined that this is
a hybrid Amethyst-throated Sun-angel Heliangelus amethysticollis x
Copper-
bellied Puffleg Eriocnemis
cupreoventris.
462
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Known from Bogotá trade skins. Only the type, an adult male, is listed by Salvin (1892). Graves (1990) deter-
mined that this is a hybrid Amethyst-throated Sun-angel Heliangelus amethysticollis x C
opper-bellied Puffleg
Eriocnemis cupreoventris.
Known only from the type. Berlioz (1936b) suggests that it is a hybrid of ambiguous character, perhaps
between the Tourmaline Sun-angel Heliangelus exortis and the Greenish Puffleg Haplophaedia aureliae. Status
uncertain according to Schuchmann (1999).
Known only from the type, an adult male Bogotá trade skin. Considered a probable hybrid Heliangelus sp. x
Eriocnemis sp. or P
urple-backed Thornbill Ramphomicron microrhynchum (Meyer de Schauensee 1966).
Known only from the type, a Bogotá trade skin. Generally believed to be an immature female of Rothschild’s
Sun-angel H. rothschildi (Simon & Hellmayr 1908), itself a dubious species.
Known only from Bogotá trade skins. Considered a probable hybrid Heliangelus sp. x Eriocnemis sp. or
Purple-backed Thornbill Ramphomicron microrhynchum (Meyer de Schauensee 1966).
Known only from the type, a Bogotá trade skin, which Simon (in Peters 1945 [1931–48]) suggested may be a
hybrid Amethyst-throated Sun-angel Heliangelus amethysticollis of race clarisse x Buff-tailed Coronet
Boissoneaua flavescens.
463
Known only from one adult male specimen. Graves (1996) proved that it was a hybrid Black-breasted
Puffleg E. nigrivestis x Sapphire-vented Puffleg E. luciani.
Known only from three Bogotá trade skins, which differ somewhat. Possibly a hybrid Eriocnemis x Coeligena
(Schuchmann 1999).
Known only from the type and another skin, presumed to be of this species, in the Gould Collection, now in
Tring. Graves (1998) concluded that the type is a melanistic specimen of the C opper-bellied Puffleg
Eriocnemis cupreoventris, but did not comment on the Gould specimen.
Known from three specimens, all from the Gould collection, an adult male, an adult unsexed skin, and a
‘variety’. Peters (1945 [1931–48]) placed it in the synonymy of the Copper-bellied Puffleg Eriocnemis
cupreoventris.
Known only from the type, considered by Peters (1945 [1931–48]) to be a synonym of the Glowing Puffleg
Eriocnemis vestitus.
Known only from the type, an adult male Bogotá trade skin. Schuchmann et al. (2000) considered it a prob-
able hybrid Glowing Puffleg E. vestitus x Copper-bellied Puffleg E. cupreoventris.
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Known only from Benvenuti’s description of a specimen in his collection. Never satisfactorily identified,
but Salvin (1892) regarded it as a synonym of the Glowing Puffleg E. vestitus.
Known only from the type, said to be a juvenile male, collected by Mossa. Graves (1999b) concluded that it
was a hybrid Black-tailed Trainbearer Lesbia victoriae x Rainbow-bearded Thornbill Chalcostigma herrani.
Known only from the type, an unsexed specimen. Meyer de Schauensee (1966) believed Orton’s Comet to
represent a hybrid Black-tailed Trainbearer Lesbia victoriae x Purple-backed Thornbill Ramphomicron micro-
rhynchum, and this identification has been supported by Graves (1997a).
Known only from three specimens, a male and a female from Illalo, Ecuador, and a Bogotá trade skin.
Considered a probable a hybrid L
ong-tailed Sylph Aglaiocercus kingi x Black-tailed Trainbearer Lesbia victo-
riae (Schuchmann 1999).
Known only from Boucard’s description, tacked on to the end of his description of the Tyrian Metaltail
Metallura tyrianthina. Cory (1918) placed this species in the synonymy of M. tyrianthina with a query.
Known only from the type. Generally considered to be a hybrid of the Black-chinned Hummingbird
Archilochus alexandri with either Anna’s Hummingbird Calypte anna or Costa’s Hummingbird C. costae
(Thayer & Bangs 1907; Schuchmann 1999).
465
Known only from the type, a male. Now considered to be a hybrid Scintillant Hummingbird Selasphorus
scintilla x Volcano Hummingbird S. flammula (Carriker 1910; McCarthy 2006).
Known only from an adult male collected by Floresi, in the Loddiges Collection. The bird is generally
considered to be a hybrid between Allen’s Hummingbird Selasphorus sasin and Anna’s Hummingbird
Calypte anna (e.g. Schuchmann 1999).
Apparently known only from two adult females shot by H. G. Rising on July 2, 1896. Considered to fall
within the range of individual variation of the Bumblebee Hummingbird A. heloisa (Bangs 1927).
Apparently known only from the type, but possibly from other specimens at Tring. Meyer de Schauensee
(1966) thought the Decorated Woodstar was probably a hybrid W
hite-bellied Woodstar Chaetocercus mulsant
x Gorgeted Woodstar C. heliodor, which was upheld by Graves (1997b).
Known only from the type, a male. Meyer de Schauensee (1966) and Schuchmann (1999) considered C.
harterti to be a hybrid, but did not specify any parentage, although the latter mentions it under Esmeraldus
Woodstar C. berlepschi.
Known only from Gould’s description. The type appears to be lost. Salvin (1892) synonymised H. splendens
with Violet-fronted Brilliant Heliodoxa leadbeateri.
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Known only from Boucard’s description of a specimen in his collection. Peters (1945 [1931–48]) placed it in
the synonymy of the Green-backed Firecrown Sephanoides sephanoides.
Known only from Molina’s description. Salvin (1892) used Molina’s name for the bird now called the
Green-backed Firecrown Sephanoides sephaniodes. Usually synonymised with S. sephaniodes with a query, but
considered unidentifiable.
Known only from the type. Peters (1945 [1931–48]) relegated Penard’s Hummingbird to a synonym of the
Rufous-breasted Hermit Glaucis hirsutus.
Salvador Mountain Gem Lampornis viridipallens connectens Dickey & van Rossem
Lampornis viridipallens connectens Dickey & van Rossem 1929, p. 209 (Los Esesmiles, Dept. Chalatenango, El Salvador)
Status Invalid taxon.
Putative range El Salvador.
Known only from the type. Monroe (1963) carried out a detailed revision of the group and considered it
invalid.
467
Known only from Levaillant’s description, based on two specimens collected by Dombey, and then in the
Jardin des Plantes, but now lost. Provenance and species considered doubtful (Sclater 1857).
Known only from Kuhl’s description, based on the plates of Daubenton and Levaillant. Considered a prob-
able artefact (Peters 1945 [1931–48])
Known only from Brisson’s description. Unidentifiable (see Whistler & Kinnear 1935).
Known only from the type, a young bird. Mayr (1957) suggested it was a subspecies of Black Scimitarbill
Rhinopomastus aterrimus, but it is now generally assumed to be inseparable from R. a. notatus from east
Sudan, Ethiopia and north-west Somalia.
Known only from Bertoni’s description. Probably based on memory (Peters 1948 [1931–48]).
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Known only from the type, an adult male, which Zimmer & Mayr (1943) considered to be a hybrid
Red-necked Araçari Pteroglossus bitorquatus sturmii x Ivory-billed Araçari P. azara. Short & Horne (2001)
considered it to be a hybrid between P. azara mariae and Lettered Araçari P. inscriptus humboldti.
Known only from the type. Friedmann (1958) doubted its validity, and it is now considered a synonym of
Lettered Araçari Pteroglossus inscriptus humboldti (Short & Horne 2002).
Known only from the plates and descriptions of Albin and Aldrovandus. Never identified with any known
species. Peters (1930) considered it an artefact.
Known only from Edwards’s plate and description, on which Linnaeus’s name was based. Peters (1930)
claimed that Edwards’s plate is not identifiable, and that apart from the white throat the bird could be the
Keel-billed Toucan R. sulfuratus brevicarinatus.
Known only from Wagler’s description. This bird has been generally synonymised with the common Curl-
crested Araçari Pteroglossus beauharnaesii.
469
Known only from the type. Generally considered to be a young specimen of the Least Honeyguide Indicator
exilis (Friedmann 1954).
Known only from the type, a young female, perhaps merely the young of some other species, but never
identified.
Known only from the type. Locality data may be in error, thus Short (1982) suggested that C. immaculatus
might be an aberrant specimen of the Chestnut Woodpecker Celeus elegans.
Known only from the description of Pennant, based on a specimen sent to him by Dr. Garden of Charlestown,
South Carolina. Unidentified.
Known only from J. E. Gray’s description from a drawing. Peters (1948 [1931–48]) considered it to be an
aberrant specimen of White-bellied Woodpecker D. javensis javensis.
Known only from Albin’s description and plate, from which the others derive. Unidentifiable and probably
a dreamed-up bird.
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Known only from the type in Brussels. Peters (1948 [1931–48]) thought it was a hybrid Yellow-tufted
Woodpecker M. cruentatus x Red-fronted Woodpecker M. rubrifrons, the latter nowadays being considered a
colour-morph of the former (Short 1982).
Almost certainly synonymous with the San Salvador race of West Indian Woodpecker, M. s. nyeanus, which
is a variable form; the supposed differences of the Grand Bahama taxon fall within the range of variation of
those from San Salvador (Winkler & Christie 2002). The population disappeared in the 1950s.
This nuthatch was described by Latham (1790) from one or two paintings in the collection of Lady Impey
(Fisher & Warr 2003), which may or may not have been based on actual specimens. Its provenance may
have been Siam (Thailand) (Fisher & Warr 2003), or it is a species closely related to birds from south-west
Iran (Dickinson 2006). However, Mlíkovský (2007) suggests that it may represent a now-extinct taxon from
the Zagros Mountains, western Iran. Nothing more can be said about it.
Known only from Swainson’s description. Unidentifiable (Cory & Hellmayr 1927).
Known only from the type. It is now considered to be a probable hybrid Tropical Kingbird Tyrannus melan-
cholicus x Varied Flycatcher Empidonomus varius.
Known only from a solitary Bogotá trade skin. Cory & Hellmayr (1927) believed it to be an aberrant spec-
imen of Lawrence’s Tody-tyrant Todirostrum sylvia superciliare, and Traylor (1979) regarded it as a synonym
of that form.
Apparently known only from the type, collected by B. Hermann and presented to the National Museum of
Chile. Gigoux & Looser (1930) consider Phillipi’s Spinetail to be an artefact, with the body of the Cape Horn
Sedge Wren Cistothorus platensis hornensis and the tail of Des Murs’s Spinetail Sylviorthorhynchus desmursii.
Known only from the type. Olrog (1963) treated it as a synonym of the Chilean S
harp-billed Canastero
Asthenes pyrrholeuca sordida.
A. sclateri was originally known only from Cabanis’s description, based on the type, and one other spec-
imen in the Berlin Museum; a third specimen, formerly in Tring, now appears to be lost. A. sclateri was split
from the closely related A. punensis, but following SACC (2007) these two are reunited, with the name A.
sclateri taking precedence. Therefore, the Puno Canastero is widespread and not considered endangered.
Known only from the type taken in Minas Gerais. Both Pinto (1952) and Meyer de Schauensee (1966)
suggested that this bird might be a race of the White-throated Woodcreeper Xiphocolaptes albicollis, but it is
now treated as a subspecies of Moustached Woodcreeper X. falcirostris (Collar et al. 1992; Marantz et al.
2003).
Known only from the type. Status uncertain, but may well still be extant. Placed in the synonymy of
Moustached Woodcreeper X. falcirostris by Hellmayr (1925), but considered by Pinto & Camargo (1961) and
Marantz et al. (2003) to be either a race of the White-throated Woodcreeper X. albicollis or possibly a distinct
species.
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Known only from the unique type. Regarded as a melanistic specimen of the B
uff-faced Scrub Wren
Sericornis perspicillatus (Beehler 1978; Mayr et al. 1986).
Known only from the type, now lost. Regarded as an immature of the C
hestnut-rumped Heath Wren
Calamanthus pyrrhopygius Parker (1984b).
Known only from the type in Tring. Considered an artefact composed of the head of the Black Drongo
Dicrurus macrocercus and the body of the Black-headed C
uckoo-shrike Coracina melanoptera (Sharpe 1879).
Known from a single specimen collected in 1931. Probably a vagrant of the Australian nominate race of
Cicadabird.
Known only from Pennant’s description. Coues (1878) considered that it was not a shrike, and that it did not
originate in North America. McAtee (1963) stated that it was unidentifiable.
Known only from the type in Washington, D. C. Ridgway (1904) and Hellmayr (1935) considered this to be
a hybrid between the Yellow-throated Vireo Vireo flavifrons and the B
lue-headed Vireo V. solitarius.
473
Known only from the plate of Daubenton and the descriptions of Buffon and Pennant. McAtee (1963)
considered it a partial albino, and unidentifiable. Clearly not an Old World oriole and if anywhere perhaps
best placed with the icterids.
Known only from two specimens, both collected by Stalker. Believed to be a hybrid between two
Australasian Figbird races, S. vieillotti vieilloti and S. v. flaviventris (Ford 1975).
Known only from Forster’s description. Todd (1984) has shown that it was not a Pomarea, but a Polynesian
Triller Lalage maculosa tabuensis.
Known from two recently fledged specimens collected in 1931, and considered doubtful by Goodman et al.
(1997). Molecular analysis has now shown the Short-toed Nuthatch Vanga is a juvenile White-throated
Oxylabes Oxylabes madagascariensis (Fjeldså et al. 2013), which was previously classified as a babbler, but is
now placed within the endemic Madagascar family Bernieriidae.
Known only from the type. Considered by dos Anjos et al. (2009) to be an abnormal immature specimen of
the Australian Raven Corvus coronoides.
474
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Sharpe (1877) placed C. fallax in the synonymy of the Slender-billed Crow Corvus enca.
Both Sharpe (1877) and Mathews (1930) put C. modestus in the synonymy of C. violaceus, which is itself now
regarded as a subspecies of the Slender-billed Crow Corvus enca.
This is now known to be a colour morph of the North Atlantic subspecies of Common Raven Corvus corax
varius, which once occurred on the Faroe Islands. It is not albinistic as often stated, but a population of birds
exhibiting leucism (van Grouw 2014). The Faeroese White-speckled Common Raven was considered a pest
by the Faroe Islanders, and regularly hunted, but once they began declining to dangerously low numbers,
the remaining birds were sought after by collectors (Droste 1869). The speckled birds were reduced to a
handful at the beginning of the 20th century; the last confirmed specimen was an individual shot in 1902,
and the unique gene that caused the extensive white plumage in these birds is now also extinct (van Grouw
2014).
Known only from the type of unknown locality, an adult male in Washington D. C., obtained from A.
Boucard. Generally considered to be a hybrid King Bird of Paradise C. regius x Magnificent Bird of Paradise
Diphyllodes magnificus (Fuller 1995).
Known from the type, an adult male, collected by Walter Goodfellow in August 1906 and now at Tring, and
one other specimen in Berlin. Generally considered to be a hybrid King Bird of Paradise C. regius x
Magnificent Bird of Paradise Diphyllodes magnificus (Fuller 1995).
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Known from nine specimens in Berlin, New York, Tring and Paris. Considered to be a hybrid Magnificent
Riflebird Ptiloris magnificus x Twelve-wired Bird of Paradise Seleucidis melanoleucus (Fuller 1995).
Known only from the type in the Leiden Museum, an adult male from the Arfak Mountains of north-west
New Guinea. Considered to be a hybrid Magnificent Riflebird Ptiloris magnificus x T
welve-wired Bird of
Paradise Seleucidis melanoleucus (Fuller 1995).
Known from about 24 specimens. Considered to be a hybrid Magnificent Bird of Paradise Diphyllodes
magnificus x King Bird of Paradise Cicinnurus regius (Fuller 1979, 1995).
Known only from the type, now in the Rothschild collection in New York. Considered to be a hybrid Black
Sicklebill Epimachus fastuosus x Arfak Astrapia Astrapia nigra (Fuller 1995).
Known only from four specimens, all in New York. Considered to be a hybrid Lesser Bird of Paradise
Paradisea minor x Twelve-wired Bird of Paradise, Seleucidis melanoleucus (Stresemann 1930; Fuller 1995).
Known only from three specimens, in Dresden, Leiden and New York. Considered to be a hybrid Superb
Bird of Paradise Lophorina superba x Magnificent Bird of Paradise Diphyllodes magnificus (Fuller 1995).
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Known only from an adult male collected by Captain Neptune B. Blood on September 20, 1944, now in
Sydney. Considered to be a hybrid Raggiana Bird of Paradise Paradisaea raggiana x Blue Bird of Paradise P.
rudolphi (Fuller 1979, 1995).
Known from the type and possibly another specimen in the Berlin Museum, and four others in New York.
Considered to be a hybrid between the Greater Bird of Paradise Paradisaea apoda or Raggiana Bird of
Paradise P. raggiana and the Emperor Bird of Paradise P. guilielmi (Fuller 1979, 1995).
Known only from the type, in Paris. Considered to be a hybrid Lesser Bird of Paradise Paradisaea minor
finschi or Raggiana Bird of Paradise P. raggiana with Emperor Bird of Paradise P. guilielmi (Fuller 1995).
Known only from the types. Considered by Stresemann (1930) and Fuller (1995) to be a hybrid Lesser Bird
of Paradise Paradisea minor finschi x Raggiana Bird of Paradise P. raggiana augustaevictoriae.
Known only from the type. Considered to be a hybrid between two races of the Greater Bird of Paradise,
Paradisea apoda novaeguineae and P. a. salvadorii (Fuller 1995).
Known from a specimen in New York and another in Paris. Considered to be a hybrid Western Parotia
Parotia sefilata and Superb Bird of Paradise Lophorina superba (Stresemann 1930).
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Considered to be an intergeneric hybrid, Magnificent Riflebird Ptiloris magnificus x Superb Bird of Paradise
Lophorina superb (Frith & Frith 2010).
Considered to be an intergeneric hybrid, Black Sicklebill Epimachus fastosus x Arfak Astrapia Astrapia nigra
(Frith & Frith 2010).
Now considered to be an intergeneric hybrid Lesser Bird of Paradise Paradisaea minor x Magnificent
Riflebird, Ptiloris magnificus (Frith & Frith 2010).
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Considered to be an intergeneric hybrid, Lesser Bird of Paradise Paradisaea minor x Magnificent Bird of
Paradise Diphyllodes magnificus (Frith & Frith 2010).
Unidentifiable. Known only from Lesson’s description, from a type now lost. Generally considered to be
identical with the Kioea Chaetoptila angustipluma (see p. 289), but the description does not agree at all, and
the provenance is in extreme doubt.
Known only from the descriptions of Brünnich, Latham and Pennant, based on Ström. Synonymised under
Great Tit Parus major (Gadow 1883).
Known only from a single specimen, originally one of four. Sometimes listed as extinct but Kirwan & Grieve
(2010) suggest the type locality is in error, its range being much more widespread in Iran, and that the very
similar extant subspecies P. a. gaddi should be synonomised under P. a. phaeonotus.
This supposed subspecies is known from only one specimen collected in 1937, but the Blue-wattled Bulbul
Pycnontus nieuwenhuisii itself may be of hybrid origin and so an invalid taxon (Williams 2002).
Known only from the type, a female. Considered an aberrant Tristram’s Warbler S. deserticola maroccana, or a
hybrid S. d. maroccana x Desert Warbler S. nana deserti (Bairlein 2006).
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Known only from Aububon’s description and plate of a specimen obtained in June 1812. From the fact that
a number of his drawings of birds obtained about this time were later destroyed, it has been considered that
the published plate may have been based to some extent on memory. Mayr et al. (1986) relegated it to the
synonymy of the Golden-crowned Kinglet R. satrapa.
Known only from the type, collected by George Vanderbilt. Now known to be an immature specimen of the
Sumatran race of Horsfield’s Babbler Malacocincla sepiaria barussana (Hoogerwerf 1966, Mees, 1995).
The Amik Gölü population has not been seen since 1962 due to the drainage of its wetland habitat.
Considered valid by Roselaar (1995), but probably synonymous with nominate Bearded Tit P. b. biarmicus
(Kirwan 2006).
Known only from the type in the Liverpool Museum. Although there was little supporting evidence, the
specimen was originally and erroneously thought to have come from Rodrigues Island in the Mascarenes.
Now known to be an albinistic trembler of the genus Cinclocerthia from the West Indies (Olson et al. 2005).
The Rodrigues Starling Necropsar rodericanus was described from subfossil remains and from a description
by Tafforet (see p. 312), who mentioned the starling’s ability to predate or scavenge giant tortoises or turtles.
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Hachisuka (1953) compared it with the closely related, but generally frugivorous Réunion Crested Starling
Fregilupus varius (see p. 310), and suggested that as the Rodrigues Starling could not possibly prey on such
large prey items as giant tortoises and turtles, it was probably frugivorous as well. Therefore Hachisuka,
based entirely on Tafforet’s description, concluded that Tafforet had in fact observed a small crow or chough,
not a starling, naming it Testudophaga bicolor. The basis for this assignation is based entirely on the muddled
imagination of Hachisuka, and thus must be disregarded as pure fantasy.
The description is considered to be based on Coracias striata = Striated Starling Aplonis striata.
Known only from the type, a female, at Harvard, collected at Mengtse, Yunnan on May 3, 1921. Now placed
in the synonymy of the Black-throated Robin Luscinia obscura (Cheng 1987).
Known only from Latham’s description. Considered to be a possible immature Hawaii ’Elepaio Chasiempis
sandwichensis (Olson 1989).
Formerly occurred on the island of Yakushima, where the last known specimen was collected on September
14, 1911. Mayr & Paynter (1964) recognise no races of the Izu Islands Thrush, and this race is relegated to a
synonym of M. celaenops.
Considered to be an anomalous population of the nominate Canary Islands Chat, S. d. dacotiae (Collar &
Stuart 1985). The type specimens all fall within the range of individual variation of the extant taxon (Taylor
& Clement 2006).
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Known only from the type, now lost. Considered dubious (BirdLife International 2011).
Believed extinct since 1939, but the reason for its extinction is not known. Vaurie (1955a) disputed its
validity, as did Bannerman & Bannerman (1958); the Cyprus birds appear to be inseparable from Caucasus
populations.
Known only from the type, presumed to be a male collected between May 1925 and April 1926. Considered
an artefact, made up in part from an Olive-backed Sunbird C. jugularis (Ripley 1950).
Known only from the plate of Daubenton, based on a specimen no longer extant. Considered to represent
the Madagascar Fody F. madagascariensis (Cheke & Hume 2008).
Known only from the type, a male trade skin of ‘Rio’ (Brazilian) make, in Tring. Hellmayr (1936) considered
it to be a probable hybrid between the Chestnut-bellied Euphonia E. pectoralis and Orange-bellied Euphonia
E. xanthogaster.
Known only from Buffon’s description and the accompanying plate in Daubenton’s Planches Enluminées.
Sharpe (1888) thought that it represented a female and a young male of the Blue Grosbeak Passerina
caerulea.
Known only from the descriptions of Pennant and Latham, clearly from the same specimen. Unidentifiable,
and ‘not an American bird’ (McAtee 1963).
Known only from Buffon’s description and Daubenton’s accompanying plate. Gmelin’s name was formerly
used for the bird now known as Sporophila intermedia, the Grey Seedeater. Unidentifiable, and ‘not an
American bird’ (McAtee 1963).
Known only from the descriptions of Edwards and Linnaeus. Never identified.
Known only from the type, a male, collected on Hawaii on 25 September 1891 by Perkins, in the Kona
region of Hawaii. Considered to be a hybrid Common ’Amakihi Viridonia virens x Hawaiian Creeper
Paroreomyza mana (Bryan & Greenway 1944; Amadon 1950).
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Known only from the type, of unknown origin. Considered an artefact (Olson & Schifter 1989).
Known only from the type specimen, collected on November 1, 1870. Considered by the AOU (1957) to be a
hybrid Common Redpoll Carduelis flammea x Pine Siskin Spinus pinus.
Known only from the descriptions of Wilson (and later Audubon 1829). Chapman (1888) listed it as
hypothetical.
Known only from the descriptions of Pennant and others. Considered unidentifiable (McAtee 1963).
Known only from the descriptions of Pennant and others. Considered unidentifiable (McAtee 1963).
Known only from the descriptions of Pennant and others. Considered unidentifiable (McAtee 1963).
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Known only from the description of Catesby, from which are derived those of Pennant and others.
Considered unidentifiable (McAtee 1963).
Known only from the description of Edwards, and never satisfactorily identified.
Known only from Pennant’s description. Bechstein (1795) considered it to be an earlier name for Acrocephalus
palustris. Unidentifiable.
Known only from the type. Considered to represent a hybrid Blue-winged Warbler Vermivora pinus x
Kentucky Warbler Oporornis formosus (Ridgway 1880; Graves 1988).
Known only from Latham’s description, based on a specimen in the Leverian Museum. Not identifiable
with any known species, but the name was formerly used, incorrectly, for Masked Yellowthroat Geothlypis
aequinoctialis.
485
Known only from Audubon’s description, and plate of two specimens obtained in May 1811. Considered
synonymous with the Cape May Warbler Dendroica tigrina (Jardine 1876), or a hybrid between Cape May
Warbler and Blackpoll Warbler Dendroica striata (Coues 1884).
Known only from the plates of Wilson and Audubon. Equated with Black-throated Green Warbler Dendroica
virens (Mayr et al. 1986).
Known from a few specimens; generally regarded as a hybrid Yellow-throated Warbler Dendroica dominica x
Northern Parula Parula americana (Dunn & Garnett 1997).
Known only from the type. Meyer de Schauensee (1945) concluded that it is an artefact, being a typical
example of a Mountain Cacique Cacicus chrysonotus leucoramphus with yellow feathers glued below.
Known only from the type, collected sometime between 1862 and 1865 by Don Manuel Villavicencio.
Considered a possible hybrid, or immature Yellow-tailed Oriole I. mesomelas (Meyer de Schauensee 1966).
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Known from the descriptions of Brisson and Buffon, from which those of Pennant and Latham derive.
Unidentifiable, and ‘not an American bird’ (McAtee 1963).
Known only from the descriptions of Latham and Pennant. Unidentifiable (McAtee 1963).
Known only from the descriptions of Pennant and Latham. Unidentifiable (McAtee 1963).
Known only from Pennant’s description of a specimen, now lost, in the Leverian Museum. McAtee (1963)
considered it a partial albino, and unidentifiable.
Now considered to fall within the range of variation seen in A. henslowii (Browning 1990).
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Amak Island Song Sparrow Melospiza melodia amaka Gabrielson & Lincoln
Melospiza melodia amaka Gabrielson & Lincoln 1951, p. 253 (Amak Island, Aleutians)
Status Invalid taxon.
Range Amak Island, Aleutians, Alaska.
The island’s population of song sparrows disappeared at the start of the 1980s; the form amaka is now
considered synonymous with Aleutian Song Sparrow M. m. sanaka (Pruett et al. 2003).
The Hooded Seedeater was collected only once in October 1823, from a lake 15km north of Registro do
Araguaia, on the east bank of the Rio Araguaia in extreme west-central Goiás, Brazil (Ridgely & Tudor
1985). A recent genetic study has shown that it is probably a Dark-throated Seedeater Sporophila ruficollis,
showing melanism on the feathers of the cap (Areta et al. 2017).
Known only from the type, an adult male collected by E. W. Gifford on January 18, 1906. Considered an
aberrant Woodpecker Finch C. pallidus or hybrid C. pallidus x Warbler Finch Certhidea olivacea (Lack 1947;
Paynter 1970).
Known from two specimens, both adult males collected by Rollo Beck on February 28, 1906. Now consid-
ered to be a hybrid Small Tree Finch Camarhynchus parvulus x Warbler Finch Certhidea olivacea (McCarthy
2006).
Known only from the type, an adult male collected by E. W. Gifford on January 25, 1906. Now considered to
be a hybrid Small Tree Finch Camarhynchus parvulus x Warbler Finch Certhidea olivacea (McCarthy 2006).
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Known only from the type in the Zoological Museum, Rome, an aviary bird. Considered to be a melanistic
specimen of the Red-cowled Cardinal Paroaria dominicana (Hellmayr 1938).
Known only from the type in Tring, and a second specimen in the Rivoli (Massena) Collection in the
Academy of Natural Sciences of Philadelphia. Bond (1947) and Storer (1970) considered it to be a hybrid
Brassy-breasted Tanager Tangara desmaresti x Gilt-edged Tanager Tangara cyanoventris.
Known only from the type, in Paris, a cage bird of uncertain origin. Considered a hybrid Chestnut-backed
Tanager T. preciosa x Burnished-buff Tanager T. cayana (Storer 1970).
Known only from the type, an adult male in Tring of uncertain origin. Considered to be a hybrid R
ed-legged
Honeycreeper Cyanerpes cyaneus x Green Honeycreeper Chlorophanes spiza (Storer 1957).
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The rediscovery of a bird that was previously believed to have become extinct is a notable event, especially
if a long time has elapsed between the last confirmed record and the rediscovery. These time-spans can
range from years to decades to more than a century. However, the decision to declare a species ‘extinct’ or
simply ‘very rare’ is difficult and subjective. On rare occasions, the rediscovery of a species has shown that
the bird has adapted to modified conditions, or that previously unknown populations exist, and the outlook
for the species is more optimistic than previously predicted. However, the vast majority of species that are
rediscovered remain in deep peril, and the chance of them disappearing again – permanently this time – is
high. In Appendix 3, we have decided to include accounts of birds rediscovered since 1990; species redis-
covered before 1990 are included in Table 1 below. We have tried to make this list as complete as possible,
but it is certainly not an exhaustive one. It is hoped that many more supposedly extinct species will be redis-
covered in the future, but as habitat destruction continues the chances of the discovery of viable populations
of these birds becomes increasingly unlikely.
Table 1. Taxa rediscovered prior to 1990. After Donald et al. (2010); BirdLife International (2011); or published
sources.
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The Huila Black Tinamou occurred on the western slope of the east Andes in Huila, and was thought to be
possibly extinct because of severe deforestation due to the narcotics industry, agriculture, urban develop-
ment and over-hunting (Hume & Walters 2012). It was rediscovered in December 2008, when a Huila Black
Tinamou was photographed in the Cordillera de Huacamayos, Napo, Ecuador (Brinkhuizen & Córdova
Saeteros 2011), confirming that this subspecies had a much larger range than previously thought.
Bruijn’s Brush Turkey was known originally from 21 skins, the last collected in 1938 (Voisin et al. 2000). A
number of specific ornithological expeditions to locate the bird proved unsuccessful, until it was finally
rediscovered in 2002. It was found in hill ridgetop forest on Mount Nok near Majalibit Bay, where incuba-
tion mounds were discovered in a restricted area (Mauro 2005). The population appears to be declining due
to selective logging, cobalt and nickel mining, hunting and increased predation from introduced mammals,
especially dogs (Mauro 2006). Measures have been put in place to preserve the habitat and to prevent
Bruijn’s Brush Turkey from becoming extinct.
This elusive bird was considered common during the late 19th century (Finn 1911), but it declined rapidly
due to habitat destruction and over-hunting. It was last recorded with certainty in 1932 and feared extinct.
Remarkably, it was rediscovered after a period of 74 years in the Manas National Park, Assam in June 2006,
when a single individual was seen (BirdLife International 2011). The Manipur Bush-quail remains extremely
vulnerable as its damp grassland habitat continues to be destroyed.
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The Madagasar Pochard was considered common at Lake Alaotra on the central plateau of Madagascar in
the early 20th century, but severe habitat destruction, the introduction of exotic predatory fish and
over-hunting drastically reduced the population (Young & Kear 2006). A single male was captured in 1991
and kept in captivity, but it died the following year. The species was then thought to be extinct, but a group
of nine birds with chicks were sighted in November 2006 in another lake, 360km north of Lake Alaotra
(René de Roland et al. 2007). A captive breeding programme was initiated with some success, but the wild
population still numbers less than 30 birds and it is considered critically endangered.
Originally known only from two specimens, the female type collected by Rollo Beck on January 6, 1928
during the Whitney South Sea Expedition, and a male collected on May 18, 1928 by Beck’s successor,
Hannibal Hamlin, north-east of Rendora Island. At least twenty individuals were recently recorded from
the Bismarck Archipelago (Shirihai 2008), confirming the survival of the species. Its breeding grounds are
still unknown and the bird is considered critically endangered.
Salvin (1896) described the type of the Fiji Petrel as an adult male, collected at Ngau (Gau) in October 1855
by F. M. Rayner on the voyage of the Herald, one of a number of vessels sent to find the ill-fated ship and
crew of the Franklin Expedition to the Arctic. In 1984 Watling (1986) discovered a petrel on Gau Island,
which he believed to be Fiji Petrel, but the specimen was not collected. Shirihai et al. (2009) rediscovered the
species after an intensive survey in May 2009. Its rarity is blamed on predation by cats and rats.
The New Zealand Storm-petrel was known from three specimens taken in 1850. It was not seen again until
January 23, 2003, when Saville et al. (2003) recorded what they considered to be the lost storm-petrel, near
the Mercury Islands off the Coromandel Peninsula of New Zealand’s North Island. The seabird’s redis-
covery was officially confirmed by photographic evidence off Great Barrier and Little Barrier Islands in the
Hauraki Gulf on November 17 of the same year (Flood 2003).
The New Zealand S torm-petrel remains mysterious. The location of its distribution and breeding areas
are still unknown.
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The Príncipe Dwarf Olive Ibis was originally considered to be a race of the Olive Ibis Lampribris olivacea, but
recent work has shown it to be a subspecies of the São Tomé Dwarf Ibis Bostrychia bocagei. The ibis was
considered rare even when first collected by Leonardo Fea in 1901 (Bannerman 1930a). It disappeared
rapidly due to severe deforestation and hunting pressure (Snow 1950). The bird was long thought extinct,
but remarkably two ibises were seen in August 1991 (Sargeant 1994). The Príncipe Dwarf Olive Ibis remains
critically endangered (Jones & Tye 2006).
The Tasman Booby was described from subfossil remains from both Norfolk and Lord Howe Islands, and
disappeared from Lord Howe in 1788 due to over-hunting by visiting mariners (van Tets et al. 1988). It had
been extirpated from Norfolk Island some time before. Recent re-analysis of the skeletal material has shown
that the recently described and extant Lord Howe Booby S. d. fullagari is the same subspecies (Priddel et al.
2005), so the earlier name of S. d. tasmani should apply (Holdaway & Anderson 2001).
This large eagle was last collected in 1930, and was thought by some to be extinct, although this notion was
described in 1985 as ‘irresponsibly pessimistic’ (Collar & Stuart 1985). Madagascar has suffered some of the
worst deforestation in the world and this was considered the likely cause of the eagles’ extinction. Reliable
observations were finally made in 1988 and 1990 (Collar et al. 1994), but it was not until 1993 that a small
population was discovered by the Peregrine Fund (Thorstrom et al. 1995). The Madagascar Serpent-eagle
occurs in lowland rainforest in eastern Madagascar and is more widespread than previously thought. Its
habitat is under direct threat from human disturbance, and with its slow reproduction rate of just one young
every one to two years (Thorstrom & René de Roland 2000), the long-term future of the species is far from
secure.
The widespread Peregrine Falcon F. peregrinus is divided into at least 18 subspecies, of which some have
very limited distributions. The Volcano Islands Peregrine F. p. fruitii is known from very few specimens and
even fewer sightings. It was thought to have become extinct by 1945 (King 1981), possibly due to habitat
destruction and disturbance. It has certainly been extirpated from Iwo Jima and probably from Minami Iwo
Jima, but it was reported alive on Kita Iwo Jima in the early 1990s (White et al. 1994). The distribution of the
race is not known and it may also occur in the Ogasawara Islands, but the numbers must be extremely
small.
The little known Black Rail Laterallus jamaicensis occurs in scattered populations in mainland North America,
and on the coasts of Peru and Chile (Ripley 1977; Taylor & van Perlo 1998). The nominate subspecies from
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Jamaica and Puerto Rico was thought to have become extinct as a result of the introduction of the predatory
mongoose. However, the rail was rediscovered in the Black River Great Morass in southwest Jamaica in
2011 (Szabo et al. 2012).
This rail was common when first described in 1828, but it had become rare by the turn of the 20th century.
Only two specimens have been taken since 1901, the last of which was collected in 1959 (Taylor & van Perlo
1998). However, the rail was officially rediscovered when four birds were heard calling in January 1998
(Mazar Barnett et al. 1998). Reasons for the bird’s decline are uncertain, but habitat destruction seems to be
an important factor. This extremely elusive bird remains vulnerable today.
The Sakalava Rail appears to have always been scarce, with a rather disjunct distribution in the lowlands of
western Madagascar (Taylor & van Perlo 1998). Specimens were taken between 1930 and 1962, after which
the bird was feared extinct. However, a single bird was recorded in 1995, and there have been a number of
sightings since (Rabenandrasana & Sama 2006). It is now considered rare and localised.
The Utila Chachalaca is a subspecies of the w ide-ranging Plain Chachalaca O. vetula, restricted to Utila
Island off the Caribbean coast of Honduras. It was thought to have become extinct due to over-hunting, and
was last recorded in 1953 (Seutin 1998). A thorough survey of Utila between September 11 and November
15 2005 rediscovered a population of the birds, but the numbers were low and were restricted almost
entirely to mangrove and remnant patches of dry forest (Glowinski 2007). Hunting pressure is still a
concern, and habitat loss due to tourism is negatively impacting on the population. The Utila Chachalaca is
in grave danger of extinction.
The poorly-known Chinese Crested Tern was first described in 1863, but it was not recorded again until
1906. It was considered comparatively common in 1937, when 21 specimens were taken on islets off the
coast of Shandong, China, from what was believed at the time to be a breeding colony (the true location of
the breeding grounds was not found until 1951) (Gochfeld & Burger 1996). Since then, non-breeding birds
were reported in 1978 and 1991, but it was not until summer 2000 that four breeding pairs were located
(BirdLife International 2011). The Chinese Crested Tern is known from just two breeding sites on the east
coast of China, and the population may number less than 10 pairs. It is critically endangered, with the birds
still threatened by hunting, egg-harvesting and coastal development (Chan et al. 2010).
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This poorly differentiated Egyptian race of Chestnut-bellied Sandgrouse was thought to be extinct following
a record of 10 birds between Isna and Idfu in March 1979 (de Juana 1997). However, in March 2012, more
than 100 birds were recorded in the Nile Valley, and a number were photographed (Khil et al. 2012). Unlike
many rediscovered birds, the future of the Fayum Sandgrouse looks quite secure.
Cebu Amethyst Brown Dove Phapitreron amethystina frontalis (Bourns & Worcester)
Phabotreron frontalis Bourns & Worcester 1894, p. 10 (Cebu, Philippines)
Status Believed extinct since 1892, but rediscovered in November 2004.
Range Cebu, Philippines.
The Cebu Amethyst Brown Dove, like so many of Cebu’s forest birds, was thought to have become extinct
as a result of catastrophic deforestation (Brooks et al. 1995). It was last recorded in 1892 but was remarkably
relocated in November 2004, when two individuals were seen (Paguntalan & Jakosalem 2008). It has
managed to survive in the remnant Alcoy forest and possibly elsewhere, but its numbers must be extremely
low.
The Silvery Pigeon was originally a common species most often observed on small islands, but it disap-
peared due to over-hunting and deforestation, with the last official sighting reported in 1931 (Collar et al.
2001, Gibbs et al. 2001). It was feared to be extinct. However, Lee et al. (2009) confirmed its continued survival
with a photograph. The Silvery Pigeon is often confused with the similar Pied Imperial Pigeon Ducula
bicolor in the field, so it remained overlooked for nearly 80 years. The population appears to be small, but
this pigeon may be more widespread than previously thought.
Little was recorded about the Bonin Islands Wood Pigeon, which was thought to have disappeared due to
severe deforestation and perhaps over-hunting. It was thought to have become extinct some time in the
1980s (BirdLife International 2011), but the bird was rediscovered and photographed in 1998 (Ministry of
Environment 2002). The population numbers fewer than 40 birds (Brazil 2009) and remains in serious
danger of extinction.
The Solomon Islands White-bibbed Ground Dove is known from two adult and two juvenile skins, and was
last collected in the 1930s. A pair was reputedly seen by a Dr W. J. Marsteller feeding near the mouth of the
Malimbiu River, Guadalcanal, in the early 1940s (Donaghho 1950). It was thought to be extinct, but Dutson
(2011) considers it still extant.
The only specimen of this subspecies was taken in the Viga-Gigmoto watershed in 1971, and none have
been collected since. In 2008, Wamelink (in Szabo et al. 2012) reported recent sightings, but its current status
is not known.
Discovered in 1921, the Blue-throated Macaw was known only from captive birds from 1922, and its natural
distribution remained a mystery. It was not until 1992 that the first wild population was discovered near the
upper Río Mamoré, Beni, in northern Bolivia (Duffield & Hesse 1997). Like most macaws, this species has
suffered from severe deforestation and collecting for the bird trade; even as recently as the 1980s, more than
1,200 wild-caught birds were exported from Beni, Bolivia alone (Yamashita & Machado de Barros 1997). The
Blue-throated Macaw is considered critically endangered, but there are signs that numbers are slowly
increasing.
This large, distinctive ground cuckoo is known from a handful of specimens, all collected prior to 1916, after
which the bird was considered possibly extinct. However, it was rediscovered in November 1997, when a
single individual was caught and photographed in the Bukit Barisan Selatan National Park in the Barisan
Mountains (Zetra et al. 2002). Since then the cuckoo has been observed only occasionally and it is considered
to be critically endangered (BirdLife International 2011). Deforestation is the main factor contributing to its
rarity.
This little-known barn owl was known from a single female collected by J. J. Menden on October 20, 1938. It
was not recorded again and was thought to be possibly extinct. The owl was rediscovered on Taliabu, the
largest of the Sula Islands, in 1991, after seven weeks of surveys (Stones et al. 1997). The population appears
to be very small, and continuing habitat destruction threatens its l ong-term survival.
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The Anjouan Scops Owl had apparently not been seen since Humblot collected the type series in the mid-
1880s. As Anjouan has been subject to severe deforestation, it was assumed that the population had become
extinct. The owl was rediscovered in June 1992, in remnant upland forests on Anjouan (Safford 1992), and
the population appears to be stable. The continuing deforestation of the island must, however, remain a
concern.
This elusive owl was collected in the mountains of south-west Flores in 1896, and was not seen again until
1994 when two individuals were captured in the Ruteng Mountains (Hutchinson et al. 2007). The Flores
Scops Owl remains extremely difficult to locate, so its population density is unknown. It has suffered from
the severe deforestation that has taken place on Flores (BirdLife International 2011), and is presently consid-
ered endangered.
The Forest Owlet was first collected in 1873 and was last observed in 1884 (Rasmussen & Collar 1998). Only
seven specimens were known to exist. It occurred in two widely separated areas, northern Maharashtra and
south-east Madhya Pradesh/western Orissa in central India (Rasmussen & Collar 1998). The Forest Owlet
was considered long extinct when, after an intensive search of recorded localities, it was rediscovered in
1997 after 113 years (King & Rasmussen 1998). The Forest Owlet is now known to occur at a number of
localities, but remains critically endangered due to the increasing destruction of its habitat.
The elusive and cryptic Sulawesi Eared Nightjar was last recorded in 1931, but the bird was rediscovered in
1996 and found to be widespread in north and central Sulawesi (Riley & Wardill 2003). Because of the diffi-
culty in assessing numbers of such birds, the present status of this nightjar is hard to determine; however, it
appears to be thinly distributed and genuinely rare (BirdLife International 2011). Deforestation is the
primary cause of its rarity, and it is currently listed as vulnerable.
Status First collected in 1937. Long thought extinct but rediscovered in 2011.
Range Aneityum, Erromango and Tanna Islands, Vanuatu, South Pacific.
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The Vanuatu White-rumped Swiftlet inhabited the islands of Aneityum, Erromango and Tanna, in the south
of Vanuatu. It was thought to be possibly extinct as there were no recent reports. Dutson (in Szabo 2012)
reported seeing the bird in 2011, but its present status is not known.
The Mexican Woodnymph has sometimes been considered to be a race of the Purple-crowned Woodnymph
T. colombica. For many years it was only known from the type specimen, collected in the late 19th century. It
was not seen again until the 1950s (Collar et al. 1992). It is now known to survive with a patchy distribution
in south Nayarit, Jalisco and Colima states, western Mexico, where it is uncommon to locally common
(BirdLife International 2011). The population appears to be declining due to the loss of habitat for
agriculture.
The last known specimen of the nominate race of the Blue-banded Kingfisher, A. e. euryzona, was taken in
July 1937, in the tributaries of the Kali Sanen River in east Java (Kooiman 1940), and the last unconfirmed
records date from the 1950s. It was thought to be possibly extinct. However, it was mist-netted five times in
2000–2001 at Halimun National Park (Noske et al. 2011), and the area has changed little since (Szabo et al.
2012). Its present status is not known.
The White-mantled Barbet is restricted to the north-central Andes and the western slope of the Andes in
Colombia. It is divided into three subspecies. The nominate northern form was rather restricted in range
and had not been recorded since the late 1940s. It has recently been reported from the southern part of its
range (Cuervo et al. 2008; Fundación ProAves de Colombia 2011), but continued deforestation may pose a
threat in the future.
The Botero White-mantled Barbet had not been recorded since the 1950s, and was thought to be possibly
extinct. Its disappearance was attributed to the severe deforestation that has taken place in Colombia. It was
rediscovered near Pradera in the Central Andes, just 1.8km from the type locality, and is considered fairly
common (Cuervo et al. 2008).
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The Buff-rumped Woodpecker M. tristis is widely distributed across South-east Asia and Indonesia, in
subtropical dry forests including secondary growth. However, the Javan nominate race was thought to be
extinct (Winkler & Christie 2002) due to the lowland forests having been almost completely logged. The
woodpecker was a bird of lowland primary rainforest in the west of the island, and appears to have always
been scarce. The last confirmed record was in 1880, with one unconfirmed report in 1920, but birds were
seen at Halimun in 2010 (R. Hutchinson & B. van Balen, in litt. 2010), after a lapse of 130 years.
Kaempfer’s Woodpecker was known from just one individual, a female collected in Piauí, Brazil, although
it was only recognised as a distinct taxon (then thought to be a race) nearly 50 years later. The woodpecker
was thought to be possibly extinct until a male was mist-netted in 2006, in the state of Tocantins (Prado
2006).This was prior to the construction of a proposed Belém–Brasilia highway, which was going to dissect
the habitat in which the woodpecker was found. It has since been observed in neighbouring states and its
range, although rather patchy, is quite large (Pinheiro & Dornas 2008). The woodpecker appears to have an
association with the bamboo Gadua paniculata growing in cerrado and babassu palm forest (Pacheco et al.
2007), with the species’s range linked to the bamboo’s distribution. Therefore, increasing destruction of its
bamboo habitat is the main cause of its rarity, and the rediscovered population may already be declining.
This small, elusive manakin was first collected in 1957 when five specimens were taken (Sick 1959). It was
not seen again until 2002, when a male was mist-netted on the west bank of the Rio Jamanxim, which is situ-
ated about 200km from the type locality (Olmos & Pacheco 2003). Other small populations exist but
continued logging of its habitat has reduced its range in recent years (Kirwan & Green 2011). The Golden-
crowned Manakin is considered vulnerable, with a declining population (BirdLife International 2011).
The Cocha Antshrike was only known from the type specimen, described in 1937, until it was rediscovered
in 1991 near the type locality in Ecuador. The species is considered to be local and uncommon (Ridgely &
Tudor 1994). It appears to be under no immediate threat, but there are plans to develop parts of its habitat.
This is an example of an ‘expected’ rediscovery, when habitat is still extensive and the range is continental
but seldom visited (Donald et al. 2010).
The White-masked Antbird was known from one specimen taken at Andoas (probably the Río Tunigrama),
Peru, in September 1937. It was rediscovered in July 2001, on the Río Morona in western Loreto, northern
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Peru (Lane et al. 2006). This species requires low-elevation forest for foraging, and has suffered from severe
habitat loss. It is considered near-threatened and declining in numbers, but it seems to be relatively wide-
spread (van Osten et al. 2007).
This cryptic antpitta was known from 10 specimens collected between 1911 and 1942. It was not seen again
until it was rediscovered in Ucumari Regional Park, Risaralda, in 1994 (Kattan & Beltrán 1997). The bird has
subsequently been found in other localities, but it has a restricted range. Severe habitat destruction continues
to threaten the population. The bird is presently considered endangered and declining in numbers (BirdLife
International 2011).
The Northern Giant Antpitta Grallaria gigantea lehmanni occurred on both slopes of the Cordillera Central, at
the head of the Magdalena Valley in Colombia, where specimens were taken in the departments of Cauca
and Huila before the 1940s (Wetmore 1945). One was taken in 1941 in the area of what today is the Puracé
National Park (Wege and Long 1995), but the bird has not conclusively been seen since. Plenty of habitat
still remains, and birds of this race are said to have been recorded at La Planada Nature Reserve in Nariño in
1988 and 1989 (de Soye et al. 1997); Donegan (in Szabo et al. 2012) reports that it is likely to still exist.
The Ecuadorian Giant Antpitta is known from only a few localities on the western slope of the Andes in
Pichincha and Cotopaxi, Ecuador (Ridgely & Greenfield 2001). Severe deforestation has taken place over
much of its range, including conversion of habitat for agriculture and the narcotics industry, which had seri-
ously reduced the population. The bird has been recently observed on numerous occasions, and occurs in a
number of localities protected within the private ranges of landowners (Freile et al. 2010; Szabo et al. 2012).
This elusive antbird was comparatively common when first discovered in the latter half of the 19th century,
and specimens were taken up to 1950. One was seen at an army ant swarm in 1965 in Colombia, but this was
the last record until the bird was rediscovered in November 2004 (Colorado 2008). Since then, its distribu-
tion has been found to be much greater than previously expected. The lack of observations was primarily
due to its inconspicuous nature and security difficulties in the region preventing ornithologists from
accessing its habitat (Tobias et al. 2006). It is presently considered endangered.
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Stresemann’s Bristlefront M. stresemanni was thought to have become extinct since the mid-20th century in
its comparatively restricted range in eastern Brazil. It was rediscovered in 1995 in two localities (Collar et al.
1992; BirdLife International 2011), but the destruction of its favoured humid v alley-bottom forest for
logging, cattle ranching and agriculture continues to adversely affect the population. It is considered criti-
cally endangered, and the population is now tiny (BirdLife International 2011).
The Obscure Berrypecker Melanocharis arfakiana was originally known from two specimens, one from Moris
in the Arfak Mountains collected by Bruijn in 1867, the other from Matsika, south-eastern New Guinea,
collected by the Archbold expedition in 1933. The bird was not recorded again and was thought to be
possibly extinct. The Obscure Berrypecker was rediscovered in the 1990s; it is considered not uncommon
(Coates 1990), and has since been found in other areas. It appears to be tolerant of habitat disturbance.
The Cebu M ottled-breasted Bulbul was last collected by McGregor in 1906, and was long thought to be
extinct (Brooks et al. 1995). Its disappearance was blamed on the almost complete destruction of Cebu’s
forests. However it was rediscovered in the 1990s at Tabunan (BirdLife International 2011), where it is
uncommon. Like many Cebu birds, the bulbuls have managed to persist in small, isolated pockets of
secondary forest.
The Sooty Babbler was collected from just two localities in Laos (Baker 1920, 1930). It eluded ornithologists
for 74 years until its rediscovery in 1994, whereupon it was found to be locally common in central Vietnam
(BirdLife International 2011). Its conservation status is considered to be of least concern, but the population
is declining due to habitat destruction.
The Rusty-throated Wren-babbler was described from a single individual netted on January 5, 1947 in the
Mishmi Hills (Ripley 1948), and not seen again. It was thought to be possibly extinct until a population was
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rediscovered on November 18, 2004, through the use of song play-back (King & Donahue 2006). The bird
was found to be locally common but declining, and is now listed as near-threatened.
This babbler was last recorded in 1938–39, when five specimens were taken at two localities (Gyldenstolpe
1939). It was rediscovered in 1994, when it was found to have a restricted distribution but to be locally
common (BirdLife International 2011). The bird is listed as endangered, with the population decreasing due
to severe habitat destruction on the Da Lat plateau.
The enigmatic C herry-throated Tanager was known from a single specimen collected in 1870, and long
thought to be extinct. It was reported to still exist in the 1940s, but it was not until February 22, 1998 that the
bird was officially rediscovered (Pacheco 1998; Bauer et al. 2000). A second population was also found, but
its distribution is still very restricted. Reasons for its rarity are not clear but it is classed as critically endan-
gered, with a population of probably less than 250 birds (BirdLife International 2011).
This tanager was originally known from four specimens, all collected prior to 1946 (Stiles 1998). It was
rediscovered in 1992 at the Alto de los Galápagos, and has been observed at other localities, where it is
described as being common to abundant (Stiles 1998). Deforestation is the primary reason for its scarcity,
and it has now probably been extirpated at the type locality due to logging. The Gold-ringed Tanager is
listed as endangered and declining in numbers.
The Cone-billed Tanager was discovered in 1938 but then not seen for so long it was thought to be possibly
extinct. However, the bird was rediscovered in Emas National Park in 2003, and also along the Alto Rio
Juruena (Buzzetti & Carlos 2005). It is a rare species, threatened by the continuing deforestation of the Mato
Grosso and by future hydroelectrical dam projects.
The Daitoˉ Bush Warbler was last observed on Minami Daitoˉ -jima around 1942 and was thought to be
extinct due to the removal of vegetation for military airfields by the Japanese military during World War II.
However, analysis of the two extant colour forms known to occur on Okinawa have shown that one is a
winter visitor, C. d. riukiuensis, which breeds on Sakhalin to the north, and the resident population is C. d.
restricta (Manibu et al. 2002). The Daitoˉ Bush Warbler is not uncommon, and the population appears to be
stable.
This bird was known from a series of six individuals collected before 1928 on São Tomé. It was not seen
again until its rediscovery in 1990, after a lapse of 62 years (Atkinson et al. 1991). It is now known to occur in
almost every forested river basin in the Agua Ribeira, Formoso Grande, and around São Miguel (BirdLife
International 2011). The population size is unknown but it appears to be comparatively stable, though there
is evidence that populations near human-disturbed areas are declining.
The Large-billed Reed Warbler was long known from just one individual, collected around 1869 (Hume
1869). Its relationships were unknown, and it was variously considered to be related to the B lunt-winged
Warbler Acrocephalus concinens or the Paddyfield Warbler A. agricola (Vaurie 1955b). Ali & Ripley (1973)
suggested that it represents a moulting individual of some form of Clamorous Reed Warbler A. stentoreus.
It was finally confirmed as a valid – and rediscovered species – when a specimen was captured in
Thailand in 2006 and DNA feather sampling matched that of the type specimen (Round et al. 2007). In addi-
tion, the discovery of existing (mislabelled) specimens at Tring, New York and the Michigan Museum by
Lars Svensson and Pam Rasmussen revealed 10 new examples of the bird (Svensson et al. 2008). Significantly,
four of these were collected in July in north-east Afghanistan by Walter Koeltz (Lars Svensson, pers. comm.).
This hinted at a possible breeding locality, and following recording of a mystery warbler song in 2009,
Robert Timmins and Stephane Ostrovsky led an expedition to the region (Timmins et al. 2009) that resulted
in the successful discovery of a breeding area. The bird seems to be restricted to the Wakhan corridor,
between northwest Pakistan and south-east Tajikistan, while Ayé et al. (2010) report breeding birds further
north into Tajikistan. The birds migrate through northern India and Burma to Thailand.
The species evaded detection for some 140 years in part by breeding in remote valleys far from regular
birder traffic, but also by being cryptic – it is frustratingly similar to Blyth’s Reed Warbler A. dumetorum
(Lars Svensson, pers. comm.).
This thicketbird was described from three specimens taken in 1890 and bought by the Museum of Natural
History, Berlin, and from a male collected November 14, 1894 on Vitu Levu, Fiji, and not recorded again
(Kinsky 1975). The species was thought to be extinct. In 1974, there were some unconfirmed reports that the
Long-legged Thicketbird was still extant. Its survival was confirmed in 2002, when a number of birds were
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observed by BirdLife International researchers during ornithological surveys of the island (Dutson &
Masibalavu 2004). The L ong-legged Thicketbird appears to be comparatively widespread and the popula-
tion stable. Unless major changes occur, it is in no immediate threat of extinction.
The nominate race of Jerdon’s Babbler was thought to have disappeared due to habitat destriction, espe-
cially as it was last recorded in 1941 (Smythies 1953). Its habitat of tall grass had been almost entirely
destroyed by rice cultivation, so the chances of any birds surviving appeared bleak. Remarkably, the babbler
was rediscovered on 30 May 2014 in tall grassland on a flood plain (Rheindt et al. 2014); the bird had
managed to survive in an area of just 50 hectares of land near an abandoned agricultural research station.
This babbler is under severe risk from any development of its last remaining habitat, and there are presently
no protective measures in force. Unfortunately, the future of this incredible survivor is still far from secure.
The Lompobatang Flycatcher was considered common when it was first discovered, but numbers subse-
quently plummeted, primarily as a result of severe habitat destruction. The last sighting of this bird was
made in 1931, and it was thought to be possibly extinct. However, the flycatcher was rediscovered in 1995,
when two individuals were seen above Malino (BirdLife International 2011). There have been only sporadic
observations since then, and this bird is considered endangered with a decreasing population. With contin-
uing loss of habitat, the long term future of the Lompobatang Flycatcher appears to be bleak.
The Island of Cebu is one of the most devastated of the larger islands in the Philippines. Almost all of the
original dipterocarp forest has been cleared, of which only 0.03% of this forest type is left intact (Dutson et
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al. 1993). Even when Bourns & Worcester were collecting birds in the 1890s (Bourns & Worcester 1894) they
remarked on how little forest remained. The Four-coloured Flowerpecker was last collected in 1906
(McGregor 1909b), and as Rabor (1959) found no trace of it while collecting on Cebu from 1947 to 1956, it
was thought to be extinct.
In February 1992, in a remnant patch of forest at Tabunan, Central Cebu National Park, the Four-coloured
Flowerpecker was reported to still exist when a pair was seen (Timmins 1992). Further sightings by Dutson
et al. (1993), which included a juvenile, not only confirmed that the bird had survived undetected for 88
years, but also that the population was a breeding one. It was still resident in Tabunan in 2007 (BirdLife
International 2011). Other tiny populations were found in June 1998 at Nug-As and at Mount Lantoy in
early 1999, both on the south-eastern side of the island, and about 10–15 birds were discovered at Dalaguete,
also in the southeast.
The Four-coloured Flowerpecker remains highly endangered, and the total population probably
numbers less than 100 birds. The continuing deforestation of Cebu makes the future of the species far from
secure.
The São Tomé Fiscal Lanius newtoni was originally known from two records, one in 1888, when the type
specimen was collected, and one in 1928. It was not recorded again and thought to be possibly extinct.
However, it was rediscovered when a single bird was seen near the Rio Xufexufe, in the s outh-west of the
island (Atkinson et al. 1991). A number of sightings have been recorded from other areas on São Tomé, and
the bird appears be fairly widespread but very sparse in numbers. The São Tomé Fiscal is considered endan-
gered, and the population is declining (Dallimer et al. 2009).
This beautiful paradise-flycatcher was known from a single specimen, collected at Tabukan in 1873 (Meyer
1878). It then went undetected for 125 years, despite searches being made specifically for it. It was consid-
ered to be probably extinct (Whitten et al. 1987) until its rediscovery in October 1998 (Riley & Wardill 1998),
although some unconfirmed sightings had previously been made. Habitat destruction is the greatest threat
to the surviving population, which numbers between 20 and 135 individuals (BirdLife International 2011).
It is listed as critically endangered.
The Fiji Shrikebill Clytorhyncus vitiensis occurs in American Samoa, Fiji and Tonga, and it has been divided
into 12 subspecies. C. v. powelli occurred on three islands in American Samoa, Manu’a, Ofu and Olosega, but
it gradually declined as the native forests were cut. It was thought to be possibly extinct. However, Watling
(2004) recorded it on all three islands. It remains vulnerable to cyclones due to fragmentation of its forest
habitat.
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The Banggai Crow Corvus unicolor was last positively recorded in 1884–1885, and was long feared extinct. It
was not found in surveys made during 1991 and 1996, but was rediscovered on Peleng Island in 2007
(Indrawan et al. 2010). The Banggai Crow has subsequently been seen on a number of occasions in mountain
forests above 500m, and at lower altitudes in the centre of the island (Madge & Marzluff 2009). The popula-
tion is small, with estimates ranging from between 30 and 300 individuals. Deforestation is the primary
cause of its rarity, and the selective logging of Peleng is continuing to threaten the population. The species is
considered critically endangered.
Restricted to King Island in the Bass Strait, this bird’s decline was caused by intense habitat destruction. It
was recorded only infrequently in the 20th century, the last record being of two birds m
ist-netted at Loorana
in 1971 (McGarvie & Templeton 1974). The thornbill was thought to be possibly extinct until two individ-
uals were located in March 2002 (Donaghey & Lloyd 2003). It is considered critically endangered.
This bird – sometimes considered a race of Black-throated Shrikebill C. nigrigularis – was first collected by
the Whitney South Sea expedition in 1927, when two birds were collected after seven days of collecting on
the island of Nendö (Mayr 1933c). It was not recorded again and thought to be possibly extinct. However,
two pairs were located in 2004 in forest above Luselo (Dutson 2006). The population is small and declining;
it is subject to increasing habitat destruction and perhaps predation by introduced mammals, and the
species is considered to be critically endangered.
The original specimen of the W hite--tailed Tityra, an immature male, was collected by Natterer in October
1829; the bird was not seen again for nearly two centuries. There has been some doubt as to its taxonomic
status as Hellmayr (1929b), after initially confirming the validity of the species, later considered the bird to
possibly be an aberrant specimen of White-collared Tityra T. inquisitor albitorques. However, Whittaker
(2008) rediscovered a population in 2007, and in the process confirmed that the species was indeed a valid
taxon. Its scarcity is due to habitat destruction, and although the size of the population is unknown, it must
be extremely small.
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Salvin’s Purpletuft Iodopleura pipra leucopygia was known from two specimens of unknown origin, although
the labels on them stated ‘British Guiana’ (Guyana) (Salvin 1885). The Guyana locality is almost certainly
inaccurate, and it appears that the birds were actually collected from the east Brazilian coast, north of the
range of the nominate form (Snow 1982; Collar et al. 1992). A specimen was collected in July 1957 at
Pernambuco, and others were located at Alagoas in the early 1990s (Collar et al. 1992). The bird is still extant
but endangered (BirdLife International 2011); this is primarily due to the severe deforestation that has
occurred in the eastern Atlantic forests of Brazil.
The St Lucia Forest Thrush was considered common in the 19th century, but declined rapidly during the
20th century due to severe habitat degredation, overhunting for food, predation from introduced predators,
and increased brood parasitism by Shiny Cowbirds Molothrus bonariensis (Keith 1997). It was thought
possibly extinct by 1980, but a population was rediscovered in the Castries Waterworks Reserve and in the
Rainforest Sky Rides Park in July 2007, where it is considered rare.
The Large Wren Babbler Turdinus macrodactyla occurs in Malaysia and Thailand, where it is declining due to
loss of habitat (Collar & Robson 2007). The poorly known Javan race was not been recorded from the mid-
20th century and was thought to be possibly extinct due to the severe deforestation that has taken place
throughout Java. However, the bird was rediscovered in small numbers in the Alas Purwo National Park,
eastern Java, in 2000 (Grantham 2000).
Archer’s Lark was first collected on 23 September 1918 by G. F. Archer, after whom the species was named
(Clark 1920). It was originally thought to have been restricted to a small area of the Wajale plains, northwest
Somalia, with the last record within the type locality in 1922. However, a small population was discovered
in 1955, 100km to the northwest at Buramo (Collar & Stuart 1985), but it was not seen subsequently. An
unidentified Heteromirafra lark was photographed in 2011 in neighbouring eastern Ethiopia, near Jijiga
(Spottiswoode et al. 2013), which appeared to be closely related to the Liben (formerly Sidamo) Lark H. sida-
moensis, found in southern Ethiopia to the southwest. Using molecular, morphological and sonogram
analyses, Spottiswoode et al. showed that Archer’s, Liben and the unidentified population of larks all
belong to the same species, and represent three isolated populations. Liben Lark, as the species is now
known, is considered to be Critically Endangered, primarily due the spread of invasive weeds, especially
from the noxious American weed Parthenium hysterophorus.
The Timor Stubtail Urosphena subulata is a poorly known species, with the nominate occurring on the island
of Timor. A race, U. s. advena, occurs on Babar Island, to the east of Timor, which was thought to have
become possibly extinct due to extensive deforestation on the island. However, it was found to be common
on the island in 2009 and 2011 (Trainor, in Szabo 2012).
The Lord Howe Silver-eye Zosterops tephropleura was once considered to be a race of the wide-ranging
Silver-eye or Grey-backed White-eye, Z. lateralis. It is quite distinct, however, and it has now been given
specific status. It was sympatric with the extinct Robust Silvereye Z. strenua (see p. 303). Hindwood (1940)
recorded it as still extant at the time of writing, but it was thought to have subsequently become extinct
(Fuller 2000), possibly due to deforestation and predation from Black Rats Rattus rattus. This presumption
was premature, however, as although it is considered vulnerable, it is one of the commonest surviving
endemic birds on the island (Schodde & Mason 1999).
The Eastern race of the Thick-billed Grasswren A. t. modestus was originally widely distributed across South
Australia, but it declined across all of its range. It disappeared from New South Wales by 1912 (Higgins et al.
2001). This decline was primarily due to habitat destruction for livestock rearing and agriculture, droughts,
and the introduction of rabbits. This cryptic, elusive bird was feared extinct, but it was found to survive in
low numbers in parts of South Australia (Parker et al. 2010). The bird is considered to be critically
endangered.
Three subspecies of the Black-eared Thrush Zoothera cameronensis are generally recognised, cameronensis
from Cameroon and Gabon, graueri from DR Congo and kibalensis from south-western Uganda (Clement &
Hathaway 2000). The distinctive Z. c. kibalensis is sometimes regarded as a full species. It is known from two
adult males collected in 1966, at 1,525m on the edge of the Kibale Forest near Fort Portal (Prigogine 1978;
Fishpool & Evans 2001) and not recorded since. However, it is still considered extant as suitable habitat still
exists (Fishpool in Szabo et al. 2012).
The Lifou Thrush, a race of the widespread Island Thrush T. poliocephalus, was last collected in 1878 on
Lifou, when six birds were taken (Layard 1878). It must have disappeared between then and 1939, as
Macmillan (in Greenway 1967) searched for it in that year on Lifou without success. The reasons for its
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extinction are unclear, but Black Rats Rattus rattus are abundant on the island (Robinet et al. 1998), and no
doubt would have negatively impacted on the thrush population. Although it was reported on Tanna and a
skin exists, it was thought unlikely that the same race was involved (Hume & Walters 2012). Collar (2005),
however, has confirmed that the Lifou and Tanna populations are conspecific, and the thrush is still present
on Tanna.
The Black-hooded Thrush T. olivater occurs in Brazil, Colombia, Guyana, Suriname and Venezuela, and is
generally considered unthreatened over most of its range (Clement & Hathaway 2000). The Cauca subspe-
cies suffered from severe deforestation due to agriculture and the narcotics industry, and has not been seen
for some decades; it was thought to be possibly extinct. However, the thrush appears to tolerate habitat
degradation, so is likely to be extant (Donegan in Szabo et al. 2012). It was possibly seen in 2009.
The status of the Lufira Masked Weaver has been open to doubt, with some authorities considering it a
hybrid, a full species or a subspecies of Tanzanian Masked Weaver P. reichardi (Dowsett & Forbes-Watson
1993). It is now generally given specific status. The bird was common in 1960 when first described, but
appears to have declined drastically due to the construction of a hydroelectric dam, and was thought
possibly extinct. However, Louette & Hasson (2009) rediscovered the bird on an island in the centre of Lake
Lufira in February 2009, where it was found to be fairly common and the population relatively stable.
The Pale-headed Brush-finch was not seen from 1969 until its rediscovery in November 1998, when exten-
sive surveys found five pairs and two probable immatures in the Yunguilla valley, Girón, Azuay (Agreda et
al. 1999). It had declined due to habitat destruction and brood parasitism from the Shiny Cowbird Molothrus
bonariensis (Krabbe et al. 2010). Since then, numbers have increased following intensive conservation effort
(Krabbe et al. 2010), and the future of this bird appears to be comparatively secure.
Sillem’s Mountain Finch was described from two specimens collected on 7 September 1929 on a plateau at
5,125m at Kushku Maidan, in southern Xinjiang, China. It was feared extinct until it was rediscovered and
photographed in 2012, about 1,500km to the east in Qinghai (Hirschfeld et al. 2013). There are no presently
no details about population size or potential threats to it.
509
The São Tomé Grosbeak was first collected in the late 19th century and not seen again, despite several exten-
sive surveys in the first half of the 20th century (Collar & Stuart 1985). The island has been subject to severe
deforestation, especially for cocoa plantations, and the birds were thought to have been victims to habitat
loss. However, the grosbeak was rediscovered in 1991 (Peet & Atkinson 1994), and has been seen on a few
occasions since then. A number of individuals were photographed alive in 2009 (Ottaviani 2011). The popu-
lation is extremely small and is classed as critically endangered.
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Appendix 4
Museums and institutions
A list of museums and institutions around the world that hold extinct bird material discussed in this book.
BULGARIA DENMARK
Sofia Copenhagen
National Museum of Natural History Zoologisk Museum
Uppsala
CANADA Museum of Evolution
Kingston, Ontario
Biological Museum
511
FIJI Bremen
Suva Überseemuseum
Fiji Museum Dresden
Staatliches Museum für Tierkunde
FINLAND Frankfurt am Main
Helsinki Forschungsinstitut und Naturmuseum
Finnish Museum of Natural History Senckenberg
FRANCE Görlitz
Autun Staatliches Museum für Naturkunde Görlitz
Musée d’Histoire Naturelle de Ville d’Autun Göttingen
Besançon Zoologisches Museum
Musée d’Histoire Naturelle Halberstadt
Bordeaux Museum Heineanum
Musée d’Histoire Naturelle de Bordeaux Hamburg
Caen Zoologisches Institut und Zoologisches
Musée d’Histoire Naturelle de Caen Museum
Cassel Kassel
Musée d’Histoire Naturelle de Kassel Naturkunde Museum im Ottoneum
La Rochelle Kiel
Muséum d’Histoire Naturelle Zoologisches Museum der Christian-
Lyon Albrechts-Universität
Musée Guimet d’Histoire Naturelle
Leipzig
Metz
Musée Municipal Naturkundermuseum Leipzig
Mountauban Munich
Musée d’Histoire Naturelle de Mountauban Zoologische Staatssammlung
Nancy Stuttgart
Muséum–Aquarium de Nancy Staatliches Museum für Naturkunde
Nantes Wiesbaden
Muséum d’Histoire Naturelle de Nantes Landesmuseum Wiesbaden
Paris
Muséum National d’Histoire Naturelle GUADALOUPE
Poitiers Point a Pitre
Musée d’Histoire Naturelle Musée l’Herminier
Réunion
Muséum d’Histoire Naturelle ITALY
Rouen Bologna
Musée d’Histoire Naturelle de Rouen Museo dell’Istituto Nazionale per la Fauna
Strasbourg Selvatica
Musée Zoologique de l’Université Louis Florence
Pasteur Museo Zoologico ‘La Specola’, Sezione del
et de la Ville de Strasbourg Museo di Storia Naturale
Toulouse Genoa
Muséum d’Histoire Naturelle de Toulouse Museo Civico di Storia Naturale ‘Giacomo
Troyes Doria’
Musée des Sciences Naturelles
Milan
Vernon
Museo Civico di Storia Naturale de Milan
Musée de Vernon
Naples
GERMANY Centro ‘Museo delle Scienze Naturali’
Augsberg Pisa
Naturwissenschaftliches Museum Museo Storia Naturale e del Territoro
Berlin Rome
Museum für Naturkunde Museo Civico di Zoologia di Roma
Bonn Siena
Zoologisches Forschungsinstitut und Museum Museo Zoologico dell’ Accadamia dei
A. Koenig Fisiocritici
Braunschweig Turin
Staatliches Naturhistorisches Museum Museo Regionale di Scienze Naturali
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Appendix 4: Museums and institutions
INDONESIA NORWAY
Cibinong, Bogor Bergen
Museum Zoologicum Bogoriense Bergens Museum
IRELAND Oslo
Dublin Zoologisk Museum
National Museum of Ireland
PAPUA NEW GUINEA
JAMAICA Port Moresby
Kingston National Museum and Art Gallery
The Institute of Jamaica
THE PHILIPPINES
JAPAN Manila
Tokyo Philippines National Museum,
Imperial University UPLB Museum of Natural History,
Ministry of Agriculture and Forestry Los Baños, Luzon
Yamashina Institute for Ornithology
POLAND
MADAGASCAR Stettin
Antananarivo, Madagascar Museum des Staat Stettin
Museum of Ethnology and Palaeontology
PORTUGAL
THE NETHERLANDS Azores
Amsterdam Museu Carlos Machado, São Miguel, Azores
Zoölogisches Museum, University of Lisbon
Amsterdam Museo Bocage
Leiden Madeira
Naturalis, Nationaal Natuurhistorisch Museum of Natural History Funchal, Funchal,
Museum Madeira
Nijmegen
Natuurmuseum Nijmegen QATAR
Doha
NEW CALEDONIA Al-Wabra Wildlife Preservation Center
Nouméa Qatar National Museum
Départment des Sciences Humaines
RODRIGUES
NEW ZEALAND Rodrigues
Auckland François Leguat Museum
Auckland Institute and Museum
Christchurch RUSSIA
Canterbury Museum Moscow
Dunedin Darwin Museum
Otago Museum St Petersburg
Invercargill Zoologicheskii Institut, Rossiiskoi Akademii
Southland Museum and Art Gallery Nauk (Russian Academy of Sciences)
Manawatu
Te Manawa Museum of Art, Science and SINGAPORE
History Raffles Museum of Biodiversity Research
Nelson
Nelson Provincial Museum SOUTH AFRICA
Norfolk Island Durban, South Africa
Norfolk Island Museum Durban Natural Science Museum
Otorohanga
Waitomo Caves Museum SPAIN
Wanganui Tenerife
Whanganui Regional Museum Museo de Ciencias Naturales de Tenerife
Wellington Museo Canario Santa Cruz de Tenerife
Museum of New Zealand Te Papa Tongarewa Departamento de Zoologie de la Universidad
de la Laguna
513
SWEDEN Liverpool
Gothenburg World Museum
Göteborgs Naturhistoriska Museum London
Lund Grant Museum of Zoology and Comparative
Zoological Institute and Musuem of the State Anatomy
University Museum of the Royal College of Surgeons of
Malmö England
Malmö Natural History Museum The Natural History Museum
Stockholm Manchester
Naturhistoriska Riksmuseet The Manchester Museum
Uppsala Newcastle-upon-Tyne
Evolutionsmusset, Zoologi–aektionen, The Hancock Museum
Uppsala Universitet Norwich
Norwich Castle Museum
SWITZERLAND Nottingham
Basel Nottingham Natural History Museum
Naturhistoriches Museum Oxford
Geneva Oxford University Museum of Natural
Muséum d’Histoire Naturelle History
Lausanne Sheffield
Musée Zoologique de l’Université Weston Park Museum
Neuchâtel
Tring
Muséum d’Histoire Naturelle
The Natural History Museum
Wakefield
TAHITI
Wakefield Museum
Institut de Paléontologie, Muséum National
d’Histoire Naturelle
UNITED STATES
THAILAND Albany, New York
Bangkok New York State Museum
Thailand Institute for Scientific and Alfred, New York
Technological Research. Allen Steinheim Museum of Natural
History
UNITED KINGDOM Andover, Massachusetts
Bath Phillips Andover Academy
Bath Royal Literary and Scientific Museum Ann Arbor, Michigan
Birmingham Michigan Academy of Science, Arts and
Birmingham Museum and Art Gallery Letters
Bolton Museum of Zoology, University of Michigan
Bolton Museums, Art Gallery, and Aquarium Baton Rouge
Brighton Louisiana State University Museum
Booth Museum of Natural History Berkeley, California
Cambridge University of California Museum of Vertebrate
University Museum of Zoology Zoology
Cardiff Bismarck, North Dakota
National Museum and Galleries of Wales Museum of Natural History of the State
Edinburgh University and School of Mines of North
National Museums of Scotland Dakota
Royal Museum of Scotland Bloomfield Hills, Michigan
Exeter Cranbrook Institute of Science
Royal Albert Memorial Museum and Art Buffalo, New York
Gallery Buffalo Museum of Science
Glasgow Burlington, Vermont
Hunterian Museum and Art Gallery Robert Hull Fleming Museum (University of
Huddersfield Vermont)
Tolson Memorial Museum Cambridge, Massachusetts
Leeds Museum of Comparative Zoology, Harvard
Leeds Museum Resource Center University
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Appendix 4: Museums and institutions
515
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Alexander, W. B. 1921. Notes on the fauna of King Island from the logbooks of the ‘Lady Nelson’. Emu 21: 318–319.
Alexander, W. B. 1924. Sonnerat’s voyage to New Guinea. Emu 23: 299–305.
Alexander, W. B. 1954. Notes on Pterodroma aterrima Bonaparte. Ibis 96: 489–491.
Alexander, W. B. & Fitter, R. S. R. 1955. American land birds in Western Europe. British Birds 48(1): 1–14.
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Ali, S. 1960. The pink-headed duck Rhodonessa caryophyllacea (Latham). Wildfowl Trust 1lth Annual Report 54–58.
Ali, S. 1968. Dodo in Maalvi and Rahman A. (eds). Jahangir the Naturalist. New Delhi: National Institute of Sciences of India.
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