Extinct Birds 9781472597540

This is the first comprehensive review of the hundreds of bird species and subspecies that have become extinct over the

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Foreword by Errol Fuller

To many people, the word extinct conjures up visions of gigantic creatures living aeons 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 man. 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 mammals one might single out the Thylacine of Tasmania, or the Quagga of southern Africa. Among birds there are many species that have fascinating stories attached to their names, or are weird and wonderful in appearance. There is the Passenger Pigeon, for instance, or the Huia, or 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 startling. 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 book puts all the relevant information concerning recently extinct birds at their fingertips. 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 the study of a lifetime. In this particular case it represents the study of two lifetimes! Julian Hume and Michael Walters have brought together their great knowledge, experience, and just plain passion for the subject, into the production of this book. For many years Michael Walters held together the bird department at The Natural History Museum. From time to time others may have given themselves lofty titles, but it was he who knew exactly where everything was, and it was he who could answer the most obscure inquiry with both politeness and enthusiasm. He has brought these qualities and his vast knowledge to the present work. Julian Hume has contributed both his scholarship and his enormous practical experience in the field to this book. For years he has travelled the world seeking out snippets of information or gaining insights into the actual terrains that extinct birds once inhabited. Like me, Julian 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 to 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 7

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unturned in the authors’ pursuit of facts – some obscure, some easier to come by – and the result of their efforts is presented in a form that is crystal clear and easily accessible. Anyone coming to this work may rest assured: Julian Hume and Michael Walters have ruthlessly uncovered whatever information exists, and set it down in this truly encyclopaedic volume.

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Acknowledgements

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 inspirational, and our endless discussions have provided me with so many insights. I am forever grateful for that. I especially thank my partner, Lorna Steel, and Jim Martin from T & AD Poyser, who have both put up with my constant grumbling during the production of this book. Jim has worked closely and patiently with me on the project for its entirety; his inspired editorial skills and background knowledge kept things moving forwards, often when seemingly impassable obstacles loomed ahead. The book is much improved by Lorna and Jim’s efforts. 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 ­Pryfls-­Jones, who has been an inspiration throughout my later career. In the United Kingdom I thank 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 and Andrew Kitchener. Special thanks to Nigel Collar, who provided the ultimate proof-read; the book benefited greatly from his input. 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 and Dave Steadman. In France I thank Cécile ­Mourer-­Chauviré, Roger Bour, Christian Jouanin, Jean–François Voisin, Daniel Goujet, Clair 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, Jean-Michel Vincent, the late Alfred ­North-­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, Michele Sabelli, and the staff of the Mauritius Institute. In Rodrigues I thank Aurel Andre, Richard Payandee, 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, and all of the Dutch team who have been working on the Mare aux Songes project over the years. In Australia and New Zealand I thank Steve Bourne, 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. 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 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 offer thanks to my family. To my mother, brothers and sister, my late Nan, and to my father, who would have been so proud of this project, and to my daughters and partner, whose love and support helped see me through. Julian P. Hume

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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 extinction 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 our opinion) the best ink drawings 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 endangered 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. 11

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Extinct Birds

This book In keeping with tradition, this book is also entitled Extinct Birds, but it has a major difference from its predecessors; 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 2003), though we have amended to follow a ­common-­sense approach where appropriate, especially for common names of ­long-­extinct ­subfossil-­only species.

The main sections The book is divided into two main parts, followed by four appendices. 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, we 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. We have tried to be as comprehensive as possible but acquiring detailed information can be fraught with difficulties, especially when dealing with obscure scientific references, the location of museum specimens, and the sometimes spectacularly confused literature. We 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 virtually impossible to identify. Others represent plumage variations, aberrant specimens, hybrids, artefacts, or even ­dreamed-­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.

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 unkonwn), or are extremely skulking or cryptic. While not formally treated as extinct, these taxa may now be gone; 12

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Introduction

however, they may equally well survive (though doubtless in tiny numbers). We hope that this book will inspire the next generation of field ornithologists to attempt to establish whether populations of these birds cling on. Appendix 2: Doubtful and invalid taxa The second appendix includes taxa that have been described from poor, often hearsay evidence, by untrained observers, from inadequate illustrations and descriptions where the original specimen is now lost, or from single specimens without provenance data. Some can be considered to be ­dreamed-­up birds, others multiple descriptions of the same species. They are included here because their names are sometimes associated with extinct birds. We hope that this section helps clear up a number of the taxonomic muddles that pepper the world of palaeornithological nomenclature; as elsewhere, we have attempted to be as comprehensive as possible. A number of these taxa may be genuine, but the basis on which these forms have been described is at best inadequate, and too much credence has been placed upon the available evidence. Where specimens exist, genetic and morphological work has proved them to be aberrant individuals, hybrid forms, or even artefacts made from two or more specimens of different species. Appendix 3: Rediscovered taxa The third appendix covers birds once thought to have been extinct, but which have been rediscovered in recent years. Some have been relocated in areas away from their original range, others have survived in small populations in remote, undisturbed areas, or have adapted to disturbed habitats. Some of these rediscovered species have proved to be more numerous than once thought, whereas others are critically endangered and likely to disappear – permanently this time – within the next decade. Appendix 4: Museums and institutions The fourth appendix lists the museums around the world that hold specimens of the extinct birds – be they skins, pickled specimens or subfossil material – that feature in this book.

A note on the format Due to the wealth of material covered, we have chosen to follow the format of Greenway (1958, 1967), to make referencing to a particular species more convenient. Each species or subspecies has a synonymy list, with page number and type localities, followed by details on specimens held in museums and other institutions around the world – these are listed in full in Appendix 4. There then follows brief information on status and range, followed by a description of the bird, where specimens or good written accounts of the birds in life are available. The bulk of the species account summarises as much as is known on the bird’s biology, history and extinction, including sections on habitat and taxonomy where relevant. For many species a detailed account of the history of specimens is also presented here. All illustrations are by Julian P. Hume. These are based where possible on museum specimens.

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Glossary

The world of zoological nomenclature is riddled with often obscure terms and abbreviations. This is especially 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. From Latin affinis, meaning a species related to but not identical with the named species. 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. 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)’. 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. aff. alien

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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 specimen 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. 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 interbreed, 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 designated. 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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Extinct Birds The accounts

Tinamidae (Tinamous) Tinamous are one of the most ancient lineages of birds, closely related to the ratites, and occur in South and Central America, north to Mexico. There are nine recognised genera. They are predominantly terrestrial birds that fly only when pressed. Tinamous are generally considered not threatened, but some species are in danger of extinction due to ­over-­hunting and habitat destruction.

Huila Black Tinamou  Tinamus osgoodi hershkovitzi

Blake

Tinamus osgoodi hershkovitzi Blake 1953, p.199. (San Adolfo, Rio Aguas, Acevedo, Huila, 1500m, Colombia) Specimens  The type is in Chicago. Status  Possibly extinct. Not seen since 1976. Range ­South-­central Colombia. Description  40–46cm (16–18in). Adult almost uniform black; belly sooty; and undertail rufescent barred with black.

The Black Tinamou is divided into two subspecies, nominate T. o. osgoodi from ­south-­east Peru, and T. o. hershkovitzi from Huila, ­south-­central Colombia. The nominate subspecies is declining over its range due to habitat destruction and ­over-­hunting, whereas the Huila subspecies may now be extinct. The Huila Black Tinamou occurred on the west slope of the east Andes in Huila, but severe deforestation due to the narcotics industry, agriculture and urban development coupled with hunting may have exterminated the population. It was last certainly recorded in 1976 (Hilty & Brown 1986). There have been reports of its continued survival in Cueva de los Guácharos National Park, Huila, but deforestation and illegal hunting continue unabated even there. If the bird still survives, its future looks very bleak indeed. Habits Little is known about the Huila Black Tinamou. The birds appeared to prefer lower montane and valley cloud forests at an altitude between 600 and 1,400m.

Magdalena Tinamou  Crypturellus (erythropus) saltuarius

Wetmore

Crypturellus saltuarius Wetmore 1950, p. 171 (Ayacucho, Colombia, South America) Specimens  The type is in Washington, D. C Status  Probably extinct, and known only from the type, collected in 1943. Possible captive bird in the 1990s. Range  Ayacucho, in Cesar Department, and Chiminá Feligresia, near Mariquita in Tolima Department, both in the Magdalena Valley, Colombia. Description  32cm (13in). Upperparts dark ruddy brown with black barring on rump, wings brown with buff barring; underparts paler with dark barring on lower flanks and vent; chin white with greyish tinge to throat. It differed from nominate C. e. erythropus in being lighter on the belly, and in having a more greyish wash on the breast and a darker crown.

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; 17

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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 ­Red-­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).

Struthionidae (Ostriches) The Ostrich Struthio camelus is the world’s largest living bird and is presently found throughout ­sub-­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.

Syrian Ostrich  Struthio (camelus) syriacus

Rothschild

Struthio camelus syriacus Rothschild 1919, p. 81 (Syrian Desert) Specimens  Skins and eggs are in Tring. Status  Extinct. Last recorded sometime between 1940 and 1945, though there were later unconfirmed sightings. Range  Formerly inhabited the deserts of Syria, Jordan, Iraq and Saudi Arabia. Description  Male up to 210cm, female up to 175cm. The Syrian Ostrich was smaller than nominate S. c. camelus, with neck and legs red, and head with naked shield. Egg smaller.

Rothschild described the Syrian Ostrich at a meeting of the British Ornithologists’ Club on May 14, 1919, and his account reads as follows: The fact that Ostriches inhabited the Syrian Desert and Arabia has been known for a long time – in fact, there are several passages in the Bible relating to this bird. Some years ago, Mr. J. Aharoni received a number of Ostrich eggs from his Arabian hunters from the Syrian Desert, and sent them to Tring. These eggs agree with those of the ­North-­African Ostrich in being smooth and not pitted, but are much smaller and more highly polished than those of that bird. I at once urged Aharoni to procure for me some adult Syrian Ostriches. He managed to procure a pair of nearly ­full-­grown young ones alive, and as soon as they were fully adult he skinned them and sent them to me. These birds agree with the ­so-­called Struthio camelus Linn. in having the neck and legs light red, and they have the black and white (male) and brown (female) plumage peculiar to all Ostriches; but they are distinctly smaller. The bill, though not much shorter, is much narrower, while the tarsus and tail are distinctly shorter. They have the same naked shield on the head as in S. camelus, but not a horny shield as in S. molydophanes. Meinertzhagen (1954) said that the bird was common in the deserts until 1914, but precision rifles were brought to Arabia during the First World War, and later motor vehicles, so the birds, which must have been easy targets, were doomed. Previously, Arabs had hunted Syrian Ostriches on horseback, using falcons to corner them; this had a far smaller impact on numbers. Meinertzhagen 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 domestication. 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. 18

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Casuariidae

Habits Burckhart in Prater (1921) recorded the breeding habits: This bird breeds in the middle of winter and lays from twelve to one and twenty 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.

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.

King Island Emu  Dromaius ater

Vieillot

Casoar de la Nouvelle Hollande. Péron 1807, p. 467 Casuarius diemenianus Jennings 1828, p. 382 (Kangaroo Island; in error) Dromaeus parvulus ‘Gould’ (fide Broderip 1842, p. 145) Dromaeus minor Spencer 1906, p. 140 (King Island) Dromaeus bassi Legge 1907, p. 119 (King Island) (proposed in ignorance of Spencer 1906) Specimens  A skin is in Paris; skeletons in Paris and Florence, and subfossil remains are in Adelaide. Status  Extinct. Range  Formerly King Island, off the coast of Tasmania. Description  87cm (34in). A small, dark emu, about half the size of the Australian mainland Emu D. novae­hollandiae, with blackish feathers intermixed with brown on the body and extensive black feathering on the head and neck.

The nomenclature of the two extinct emus of King and Kangaroo Islands is exceedingly confusing. Both the Kangaroo Island and King Island birds were discovered by Captain Matthew Flinders in 1802, but he did not note them as different from those of the mainland. 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. The expedition collected at least three specimens of emu, of which two, a male and a female, were brought back alive to the Jardin des Plantes, Paris. The female died in April 1822 and is preserved as the Paris skin. The male 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. Latham’s (1823) account of Van Diemen’s Cassowary, based on the small size, may be referring to the King Island Birds, though it has never been definitely assigned: This bird has been thought by some to be the same with the New Holland [=Australian] Cassowary, or last described, but by no means answers to the description of that bird. To what size it arrives when full grown is not said, but certainly not so large as the New Holland species: we have met with two specimens alive in a London exhibition, which appeared to exceed the bulk of a large Bustard, though giving the idea of a still bigger bird, owing to the fullness of the plumage: the bill is broad at the gape, lessening by degrees to the point, where it is a trifle bent; the nostrils placed near the edge, about the middle; colour dusky blue; the space round the eyes and some part of the neck nearly bare, and of the same colour as the bill; irides brown; the top of the head, and hind part clothed with a few straggling crisped feathers . . . the thighs not at all visible, as in the ­New-­Holland species, and the toes are placed all three forwards, as in that bird . . . This species inhabits Van Diemen’s Land [= Tasmania] and we have very little reason to doubt its being a distinct species, as it differs in size and 19

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manners. – the author above quoted for a figure, calls it Casuarius Novae Hollandiae, supposing it to be the same bird . . . Mr. Jeffreys says that the Island of Van Diemen, in general, affords more amusement to the sportsman than all New Holland put together . . . There the Emeu is said to come down in vast numbers to feed on some chosen spot; and it is customary for a party to lie in wait for them, and often meet with flocks of 70 or 80 of each. The Tasmanian Emu (see p. 22) was not so obviously smaller than the mainland Australian Emu D. novaehollandiae, compared with the King Island bird, thus this reference is more likely to refer to the latter. To complicate the matter further, Brazil (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 not see any living emus. Furthermore, a coloured plate by Le Sueur accompanying Péron’s account (see ­Milne-­Edwards & Oustalet 1879) was erroneously assigned to King Island birds (Mathews 1910b) (see Kangaroo Island Emu, below). Subfossil King Island Emu Dromaius ater remains were subsequently discovered on both islands (Spencer 1906; Parker 1984), and Giglioli (1901) and Morgan & Sutton (1928) considered them undifferentiated. Thus, for many years it was believed that all the extant skin specimens came from Kangaroo Island, the King Island bird being known only from subfossil bones and not differing from those from Kangaroo Island. After review, however, Jouanin (1959) and Dottrens (1960) found than none of the skin specimens had come from Kangaroo Island after all. Baudin’s expedition consisted of two ships, Le Naturaliste and Le Géographe, and the former never visited Kangaroo Island. Le Naturaliste left King Island in December 1802 (this was evidently the visit referred to by Brazil), and in June 1803 arrived in France with a live adult emu and the skin of a chick, both collected at Sydney, New South Wales. However, Le Géographe did collect live emus on King Island and Kangaroo Island and left in June 1803, arriving in Paris in March 1804 with two live adult emus (the ­above-­mentioned Paris specimens) collected on King Island and five skins of young birds, some from each island. Of the five skins of young birds, there is one preserved in Paris and the other in Turin (the rest are lost); the provenance of these juvenile skin specimens in much less certain, however. The reason for the species’s extinction is not known. There were apparently plenty on the island at the time of Péron’s visit in 1803, but they had disappeared by 1836, when a few settlers arrived from England. Brazil (1914) suggested that dogs taken to the island by sealers and trained to catch kangaroos may have killed them off, and natural and ­human-­induced fires have also been cited as a cause. Habits Péron (in ­Milne-­Edwards & Oustalet 1879) gained a considerable amount of information on the habits of the King Island Emu from the inhabitants of the island, who were evidently careful observers: 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 20

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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.

Kangaroo Island Emu  Dromaius baudinianus

Parker

Dromaius ater Vieillot 1817 (a synonym of D. novaehollandiae) Casuarius diemenianus Jennings 1828, p. 382 (Kangaroo Island) Dromaeus parvulus Mathews 1901 Dromaius peroni Rothschild 1907a Dromaius baudinianus Parker 1984, p. 20 (Kangaroo Island) Peronista peroni Mathews 1913. Specimens  Known from Le Sueur’s plate. Subfossil bones are in Adelaide; a unique egg is in Tring. Status  Extinct. Range  Kangaroo Island, off the coast of Australia.

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 conflagrations, 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 1984). 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 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 1879), 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 21

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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.

Tasmanian Emu  Dromaius diemenensis

Le Souëf

Dromaeus diemenensis Le Souëf 1904a (based on an egg in his own collection) Dromaeus diemenensis Le Souëf 1904b, p. 114, p. 229 Dromaeus diemenensis Le Souëf 1907, p.13 (first description of bird) Specimens  Skins and eggs are in Tring; subfossil remains are in Augsberg; Frankfurt; Gothenburg; Launceston; London; and Nijmegen. Status  Extinct since the 19th century, but formerly common. Range  Tasmania. Description  150–190cm (59–75in). The Tasmanian Emu differed from the Australian mainland species by the lack of black feathers on the throat and foreneck, these parts being entirely white.

Although Tasmanian Emus had been recorded by Gunn (1852), 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. 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 made them very scarce by the 1830s (Gould 1846). By the early 1850s, Australian mainland birds had been imported from Victoria, Australia, and these may have hybridised with the remaining birds. The Tasmanian Emu must have become extinct in the wild around this time. A pair of emus were said to have been given to a Baron von Stieglitz, 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 may have been one of the mainland imports (as may von Stieglitz’s birds). Eggs and bones Of the eggs, one was in Le Souëf’s possession (Le Souëf 1904a), while the other egg was formerly in J. W. Mellor’s collection and was mentioned by Campbell (1900): . . . the granulations on a lighter ground appear finer than those from the mainland, and the egg is very dark green in colour. Bones have been unearthed at various times. Scott (1923) reported some collected by Tom Edwards at Mowbray Swamp, and some found by E. W. Clarke at Mole Creek. Scott drew attention to the robust pelvic elements, which he interpreted as an adaptation for powerful scratching and digging. The bones vary considerably in size, and Gunn (1852) believed that this represented sexual dimorphism, with males being slightly larger than the females. A tibiotarsus recovered in 1920 from Pleistocene swamp deposits at Irish Town, ­north-­west Tasmania, is referred to this species, but it is apparently as large as the tibiotarsus of the mainland emu. Scott (1923) insists: 22

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The point to be noted here is that the bone is beyond all question of Tasmanian origin, since its inclusion into the peaty matrix of the swamp was certainly at a much earlier date than that at which any mainland emus were imported into Tasmania, and therefore it stands as the earliest known specimen of a Tasmanian ­tibio-­tarsal shaft. Gunn (1852), who had been keeping several specimens of the ‘small Tasmanian species’ in captivity for some years, noted that one of them had just died, and continued: A leg of a Tasmanian Emu is now in my possession, and so far as I can judge from it, as a very imperfect specimen, 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 that fragments of emu eggshell had been found. No other evidence is known. The status of emus on Flinders Island remains unclear, but according to McCarthy (1965), they were extensively hunted by Native Australians there, and entered into their rituals and mythology. 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 ­father-­in-­law, Major W. Gray, owned an upland moor called Kearney’s Bogs, where emus occurred regularly 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.

Apteryigidae (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 ­over-­hunting, habitat loss and predation from introduced mammals, and some species are in critical danger of extinction.

North Island Little Spotted Kiwi  Apteryx owenii iredalei

(Mathews)

Apteryx owenii Gould 1847, p. 94 (New Zealand) Stictapteryx Iredale & Mathews 1926, p.76 Stictapteryx owenii iredalei Mathews 1935b, p. 180 Specimens  The type is in Tring. Status  Extinct. Last recorded in 1875. Considered a dubious subspecies by some authorities. Range  North Island, New Zealand. Description  35–45cm (14–18in). The Little Spotted Kiwi Apteryx oweni is a uniform pale ­buffish-­grey, irregularly finely mottled with white and brownish black; bill ivory or pinkish; iris blackish; legs and feet pale ­greyish-­white. If there are any plumage differences between A. o. iredalei and other races they are not clearly defined.

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 23

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the North Island form is known from just one specimen, taken in 1875 from the Tararua Range. The South Island Little Spotted Kiwi was considered common in the 1870s (Sales 2005), but numbers plummeted, and demand for specimens from European museums increased. The birds were regularly hunted by prospectors and farmers for food, and the flightless, terrestrial kiwis were extremely vulnerable to predation by feral dogs and cats. 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 ­offshore 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 ­over-­hunting extirpated the population before protective measures could be put in place. The race is now considered to be extinct. Habits Nothing 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 acute sense of smell to locate prey items. The call is somewhat similar to the Brown Kiwi Apteryx australis, but is more tremulous.

Aepyornithidae (Elephant birds) The Aepyornithidae or elephant birds were confined to the island of Madagascar. One species, Aepyornis maximus, was easily the heaviest bird known to science, and laid the largest egg of any vertebrate, living or extinct; even larger than that of any dinosaur. The entire elephant bird family is extinct due to ­over-­hunting, destruction of habitat and possibly natural climate change.

Giant Elephant Bird  Aepyornis maximus

Geoffroy Saint-Hilaire

Aepyornis maximus Geoffroy ­Saint-­Hilaire 1851, p. 104 (cf. Geoffroy ­Saint-­Hilaire 1850, p. 209) (Masikoro, Madagascar) Aepyornis modestus ­Milne-­Edwards & Grandidier 1869 (Ambolisatra, Madagascar) Aepyornis ingens ­Milne-­Edwards & Grandidier 1894, p. 124 (west coast between Bélo and Mouroundava, Madagascar) Aepyornis titan Andrews 1894, p. 18 (Itampulu Vé, Madagascar) Specimens Subfossil remains and eggs are in many museums around the world, with the largest collections in London, Madagascar, Paris and Tring; almost complete skeletons in Madagascar and Paris. Status Extinct. Known only from subfossil remains. Last recorded around the end of the 17th century. Range Madagascar.

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 ­well-­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). Elephant bird taxonomy The taxonomy of the elephant birds is extremely confused, and there has been much debate as to the actual number of species involved. The present consensus is that seven or eight species once occurred on 24

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Giant Elephant Bird Aepyornis maximus

Madagascar, four in the genus Aepyornis and three or four in Mullerornis (Lamberton 1934, Brodkorb 1963). However, some or all may represent just one variable or sexually dimorphic species within each genus (see also the Dinornithidae, the Moas, p. 28). The term elephant bird was probably derived from the explorer Marco Polo and his account about the legend of the rukh (or Roc), a giant ­eagle-­like bird that supposedly could seize an elephant in its talons. Goodman (1994) suggested that this legend arose from an erroneous attribution to the Malagasy Crowned Eagle Stephanoaetus mahery and its perceived association with Aepyornis eggs (see p. 81). The elephant birds were once widespread over Madagascar, from the northern to southern tip, including the arid regions of the ­south-­west and the central highlands (Hawkins & Goodman 2003). They appear to have been numerous until the arrival of human beings in Madagascar 2,000 years ago, but they declined rapidly afterwards. Extinction A number of hypotheses have been proposed for the extinction of the elephant birds. It is possible that introduced domestic fowl brought diseases, against which the elephant birds were completely defenceless (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). 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 25

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Australia and New Guinea. Elephant birds may have been important ­seed-­dispersers of forest trees and shrubs (see Dransfield & Beentje 1995). 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 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. 81), whereas eggs and chicks may have provided food for the largest Madagascan carnivore, the ­now-­extinct Giant Fossa Cryptoprocta spelea. Coevolution with native plants An endemic genus of Madagascar plants, Uncarina (Pedaliaceae), may have required elephant birds to disperse its fruit. The genus has characteristic large, prickly fruits that attach themselves to large vertebrates. Two types of burr are produced. Some are sticky (known as adhesive burrs), and others have grappling ­hook-­like structures (trampling burrs) (Midgley & Illing 2009). Midgley & Illing suggest that the trampling burrs evolved to attach to hooves or feet, and would have been perfectly adapted to adhere to the feet of elephant birds. Uncarina generally occur in the dry deciduous and scrubby arid regions of Madagascar, essentially areas favoured by Aepyornis, so it is likely that Uncarina coevolved with these birds and relied on them for ­fruit-­dispersal. In the absence of elephant birds, introduced hoofed mammals have now become the main dispersal agent of Uncarina fruit.

Hildebrandt’s Elephant Bird  Aepyornis hildebrandti

Burckhardt

Aepyornis hildebrandti Burckhardt 1893, p. 127, pl. 13–16 (Madagascar) Aepyornis mulleri ­Milne-­Edwards & Grandidier 1894, p. 124 (Antsirabé, Madagascar) Specimens  Subfossil remains are in Berlin; Paris; Washington, D. C.; and Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Range  Madagascar.

Hildebrandt’s Elephant Bird was described from a tarsometatarsus, but a nearly complete skeleton was subsequently found, which is now in Paris. 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.

Gracile Elephant Bird  Aepyornis gracilis

Monnier

Aeopyornis gracilis Monnier 1913, p. 15, pl. 8, fig. 10 (Madagascar) Specimens  The type is in Paris. Status  Extinct. Known only from subfossil remains. Range  Madagascar.

This dubious species is known from a single femur, and differed only in its less robust size. If more specimens become available, it is likely to fit well within the individual variation of Hildebrandt’s Elephant Bird A. hildebrandti.

Greater Elephant Bird  Aepyornis medius

­Milne-­Edwards & Grandidier

Aepyornis medius ­Milne-­Edwards & Grandidier 1866, p. 97, note 2 Aepyornis grandidieri Rowley 1867, p. 892 (in text) (Cape Sainte Marie, Madagascar) Aepyornis cursor ­Milne-­Edwards & Grandidier 1894, p. 124 (No locality) Aepyornis lentus ­Milne-­Edwards & Grandidier 1894, p. 124. (No locality) Specimens  The types are in Paris. Status  Extinct. Known only from subfossil remains. Range  Madagascar.

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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.

Robust Elephant Bird  Mullerornis rudis

Milne-­Edwards & Grandidier

Mullerornis rudis ­Milne-­Edwards and Grandidier 1894, p. 126, pl. 26 (west coast between Bélo and Mouroundava, Madagascar) Flacourtia rudis Andrews 1895, p. 25 Specimens  The types are in Paris. Status  Extinct. Known only from subfossil remains. Range  Madagascar.

The genus Mullerornis comprised three or four species, all of which were smaller than their closest relatives, 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 ­post-­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.

Coastal Elephant Bird  Mullerornis agilis

­Milne-­Edwards & Grandidier

Mullerornis agilis ­Milne-­Edwards & Grandidier 1894, p. 125 ­(south-­west coast near Mouroundava, Madagascar) Specimens  Subfossil remains are in Paris and Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Range ­South-­west and ­south-­east Madagascar.

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 genus and Aepyornis, but the differences are less striking between the various species of Mullerornis. If 27

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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. ­Eggshells 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.

Dinornithidae (Giant moas) The moas were a group of large, flightless birds endemic to New Zealand. They are usually considered to belong to the order Struthioniformes (ostriches, cassowaries, emus and rheas), but because of their distinctness they are sometimes placed in their own order, the Dinornithiformes. Unlike Struthioniformes, they completely lacked even vestigial wings, and include among their number the tallest known bird. Recent analysis has shown that the moas may be more closely related to tinamous than to the kiwis, as previously thought. The moas all became extinct prior to the arrival of Europeans in New Zealand. The taxonomy of the moas is extremely complex and it is not within the scope of this book to cover the subject in its entirety. For comprehensive overviews see Worthy & Holdaway (2002) and Worthy (2005b).

North Island Giant Moa  Dinornis novaezealandiae

Owen

Dinornis novaezealandiae Owen 1843, p. 8, 10 (Poverty Bay, North Island, New Zealand) Dinornis ingens Owen 1844a, p. 247 (in text) (North Island, New Zealand) Dinornis struthoides Owen 1844a, p. 244 (in text) (North Island, New Zealand) Dinornis giganteus Owen 1844b, p. 144 (No locality) Dinornis gracilis Owen 1855, p. 246 (North Island, New Zealand) Dinornis maximus Haast 1869, p. 87 Dinornis maximus Owen 1869, p. 498 (in text) (Middle Island = South Island, New Zealand) Dinornis altus Owen 1879, pp. 253, 361, pl. 79, fig. 4 (South Island) Dinornis torosus Hutton 1891, p. 247 (Takaka) Dinornis excelsus Hutton 1891, p. 247 (Te Aute) Dinornis validus Hutton 1891, p. 247 (Glenmark Swamp) Dinornis firmus Hutton 1891, p. 247 (Poverty Bay) Palapteryx plenus Hutton 1891, p. 248 (South Island) Dinornis strenuus Hutton 1891, p. 8 (Enfield) Dinornis potens Hutton 1892, p. 115 (New Zealand) Dinornis gazelle Oliver 1949, p. 166 (Te Aute) Dinornis hercules Oliver 1949 p. 174 (Coonoor) Specimens  Many museums around the world hold specimens, including subfossil bones, eggs and eggshell fragments, feathers and footprints attributed to Dinornis, with the most important in Auckland, Canterbury, Dunedin and Wellington, New Zealand; London; and Tring. Status  Extinct. Known only from subfossil remains. Date of extinction c.1350–1450. Range  North Island, New Zealand. Description  The North Island Giant Moa was a ­long-­necked, ­small-­headed bird with a long body, and massive robust legs. The bill was broad and flattened with ­well-­developed nasal bones, implying a good sense of smell. The eyes were disproportionately small. Estimates of size suggest that the females weighed between 76kg and 242kg, and stood 1.2 to 1.9m tall to the middle of the backs, whereas the much smaller males weighed between 34kg and 85kg, and stood from 0.9 to 1.2m tall (Bunce et al. 2003). The posture of the bird is thought to be like a cassowary Casuarius sp., with the head and neck held forward (Worthy & Holdaway 2002). The plumage comprised long and ‘decomposed’ feathers on the main body and shorter, downier feathers on the head and neck. There may have been a small comb on top of the head.

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 28

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North Island Giant Moa Dinornis novaezelandiae nesting

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 ­(Vickers-­Rich et al. 1995). Holdaway & Jacomb (2000) have postulated that the disappearance of all moas may have occurred in less than a century. 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 fecundity (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 ­egg-­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 flattened bill. The gizzard contained large numbers of stones to aid in the 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 & 29

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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. All moa species were ­K-­strategists – species with low fecundity and slow maturation rates (Turvey et al. 2005).

South Island Giant Moa  Dinornis robustus

Owen

Dinornis ingens var. robustus Owen 1846, p. 48 (Middle Island = South Island, New Zealand) Specimens  Many museums around the world hold subfossil specimens including bones, feathers, footprints, eggs and eggshell fragments attributed to Dinornis, with the most important in Auckland, Canterbury, Dunedin and Wellington, New Zealand; London; and Tring. Mummified remains are in Otago and York. Status  Extinct. Known only from subfossil remains. Date of extinction c.1350–1450. Range  South Island, New Zealand.

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. 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). 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 ­well-­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. 78). 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. Coevolution with plants As in the elephant birds, it is likely that many of New Zealand’s endemic plants coevolved with moas. It appears that these plants evolved defensive strategies to prevent browsing by a suite of moa species. These adaptations included divaricating foliage (the wide spreading of foliage from a point of axis), in which the edible material is interlaced and more difficult to browse, and also heterophylly, where juvenile plants have leaves of a different shape to the adults (see Atkinson & Greenwood 1989; Worthy & Holdaway 2002). Heterophyllic plants are often spiny or have unpalatable-looking leaves, and often produce toxins. In New Zealand, plants with such defences generally lose them from a height of around 3m upwards, high enough to be out of reach of the tallest browsing moa (Greenwood & Atkinson 1977). A similar scenario occurred on the Mascarene Islands, where plants were under extreme browsing pressure from the endemic Cylindraspis giant tortoises, and heterophylly is pronounced (Eskildsen et al. 2004), although divaricating growth may also have a climatic basis (McGlone & Webb 1981). 30

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Emeidae (Lesser moas) Little Bush Moa  Anomalopteryx didiformis

(Owen)

Dinornis didiformis Owen 1844a, p. 242, pl. 27, figs 3–6 (Poverty Bay, North Island, New Zealand) Dinornis dromaeoides Owen 1844a, p. 253, pl. 22, figs 1–2 (Poverty Bay, North Island, New Zealand) Dinornis dromioides Owen 1846, p. 46–47 (emendation) Dinornis parvus Owen 1883, p. 233, pl. 51–58 (Pokororo, South Island) Dinornis oweni Haast 1886a, p. 482, nomen nudum Anomalopteryx antiquus Hutton 1892, p. 124 Anomalopteryx fortis Hutton 1893, p. 9 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 and South 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.

Broad-­billed Moa  Euryapteryx curtus

(Owen)

Dinornis curtus Owen 1846, p. 48 (East Coast district, North Island) Eurapteryx exilis Hutton 1897, p. 552, pl. 48, fig. C (Wangaehu, North Island) Euryapteryx tane Oliver 1949, p. 105 (Doubtless Bay, North Island) Specimens  Subfossil remains are in Auckland, Canterbury, Wellington, and Whanganui, New Zealand; and London. Status  Extinct. Known only from subfossil remains. Date of extinction c.1350–1450. Range  North and South Islands, New Zealand.

The ­Broad-­billed 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 ­rain-­shadow forest, coastal dunelands, open grasslands, and shrublands. Unlike all other moas, the ­Broad-­billed 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.

Stout-­legged Moa  Pachyornis geranoides

(Owen)

Palapteryx geranoides Owen 1848a, pp. 1, 7; Owen 1848b, p. 361, pl. 54, figs 1–5 (Te Rangatapu, South Island) Euryapteryx gravis Owen 1870, p. 141, pl. 14 (Kakanui, South Island) Emeus gravipes Lydekker 1891, p. 297 (Kakanui, South Island) Euryapteryx pygmaeus Hutton 1891, p. 249 (Takaka, South Island)

31

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Extinct Birds Emeus boothi Rothschild 1907a, p. 210 (Shuy River, South Island) Emeus haasti Rothschild 1907a, p. 210 (Glenmark, South Island) Emeus parkeri Rothschild 1907a, p. 211 (Shag Point, South Island) Euryapteryx kuranui Oliver 1930, p. 52 (Castle Point, South Island) Dinornis expunctus Archey 1927, p. 152 Pachyornis mappini Archey 1941, p. 41 Pachyornis septentrionalis Oliver 1949, p. 61 (Pohue, North Island) Specimens  Subfossil remains are in Canterbury, Nelson and Wellington, New Zealand; and London. Status  Extinct. Known only from subfossil remains. Date of extinction c.1350–1450. Range  South Island, New Zealand.

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. The Stout-legged 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).

Crested Moa  Pachyornis australis

Oliver

Pachyornis australis Oliver 1949, p. 70 (Salisbury tableland, South Island) Specimens  Subfossil remains are in Canterbury and Wellington, New Zealand. 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 ­Heavy-­footed Moa P. elephantopus and ­ tout-legged Moa P. geranoides. It stood around 1.2m high and weighed around 75kg. The frontal region of S 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 subalpine areas (Worthy & Holdaway 1993).

Heavy-­footed Moa  Pachyornis elephantopus

(Owen)

Dinornis elephantopus Owen 1856, p. 54 (Awamoa, South Island) Dinornis crassus var. minor Hutton 1874b, p. 276 (Hamilton Swamp, South Island) Pachyornis immanis Lydekker 1891, p. 343, fig. 66B (South Island) Euryapteryx ponderosus Hutton 1891, p. 249 (Hamilton Swamp, New Zealand) Pachyornis rothschildi Lydekker 1892, p. 479 (unknown locality) Pachyornis inhabilis Hutton 1893, p. 11 (Canterbury?, New Zealand) Pachyornis valgus Hutton 1893, p. 12 (Enfield, South Island) Pachyornis murihiku Oliver 1949, p. 67 (Greenhills near Bluff, South Island) Specimens  Subfossil remains are in Canterbury and Dunedin, New Zealand; London; and Tring. Mummified remains are in Cambridge, England; the remarkable egg with embryo is in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction c.1350–1450. Range  Eastern South Island, New Zealand.

The ­Heavy-­footed Moa was one the most bizarre of all of the moas. It was a massive, ­thick-­set species with disproportionately large legs and feet. The females stood about 1.2m tall and may have weighed as much as 163kg (Bunce et al. 2009). The bill, which was large and robust, was capable of applying great force, and was probably used to clip the fibrous leaves of New Zealand Flax Phormium tenax and twigs up to at least 8mm in diameter (Wood 2007). The olfactory system was ­well-­developed so this species had a keen sense of 32

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smell. The reason for the hyperdevelopment of the hind limbs is not clear (see Andrews 1983). It has been speculated that moas may have evolved long intestines to ferment their herbivorous diet, with corresponding increase in body size. In Pachyornis, whose diet included extremely tough fibrous leaves, the increase in bulk resulted in massive supporting limbs. The ­Heavy-­footed Moa was widespread in the lowlands east of the Alps in the South Island and occupied 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 coloration at least, white eggs were laid by this genus, whereas ­greenish-­blue ones were laid by Megalapteryx didinus (see below).

Eastern Moa  Emeus crassus

  Heavy- footed Moa Pachyornis elephantopus

(Owen)

Dinornis crassus Owen 1846, p. 46 (Waikouaiti, South Island) Dinornis casuarinus Owen 1846, p. 47 (Waikouaiti, South Island) Dinornis rheides Owen 1851, p. 8 (indeterminate) Dinornis huttoni Owen 1879, p. 430 (Hamilton Swamp, South Island) Euryapteryx compacta Hutton 1893, p. 11 (Enfield, South Island) Emeus crassus Reichenbach 1850 Specimens  Subfossil remains are in Auckland, Canterbury and Dunedin, New Zealand; and London. Mummified remains are in Dunedin. Status  Extinct. Known only from subfossil remains. Date of extinction c.1350–1450. Range  South Island, New Zealand.

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 mummified 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. 33

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Megalapterygidae (Upland Moa) Upland Moa  Megalapteryx didinus

(Owen)

Dinornis didinus Owen 1883, p. 257, pl. 59–61 (Queenstown, South Island, New Zealand) Megalapteryx hectori Haast 1886b, p. 161, pl. 30 (Takaka, South Island) Megalapteryx tenuipes Lydekker 1891, p. 251, fig. 69a (Lake Wakatipu, South Island) Palaeocasuarius haasti Forbes 1893b, p. 451, nomen nudum Palaeocasuarius velox Forbes 1893b, p. 451, nomen nudum Megalapteryx hamiltoni Rothschild 1907a, p. 197 (Waingongoro, North Island = a doubtful locality) Megalapteryx benhami Archey 1941, pp. 35, 138 (Mount Arthur, South Island) Specimens  Subfossil remains are in Auckland, Canterbury, Dunedin and Wellington, New Zealand; London; and Tring. Status  Extinct. Known only from subfossil remains. Date of extinction c.1350–1450. Range  South Island, New Zealand.

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 (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 mummified leg shows that the tarsi were feathered, which was another adaptation 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 ­greenish-­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 hochstetteri, 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. 31) was the smallest of all moa species. There were also size differences between ­high-­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.

Megapodidae (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.

Consumed Megapode  Megapodius alimentum

Steadman

Megapodius alimentum Steadman 1989a, p. 538 (Lifuka, Ha’apai [Hapace] Islands, Tonga) Specimens  Subfossil remains are in Fiji. Status  Known only from subfossil bones. Disappeared soon after the arrival of Polynesians. Range  Fiji and Tonga.

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, 34

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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.

Pile-­builder Megapode  Megapodius molistructor

Balouet & Olson

Megapodius molistructor Balouet & Olson 1989, pp. 9–11 (Pindai Cave, Nepoui Peninsula, New Caledonia) Specimens  Subfossil remains are in Paris. Status  Known from bones collected in Pindai Cave. Date of extinction not known. Range  New Caledonia, possibly Vanuatu and Fiji.

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. 327), may represent 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.

Viti Levu Scrubfowl  Megapodius amissus

Worthy

Megapodius amissus Worthy 2000, p. 342 (Udit Cave, Wainibuku, Viti Levu, Fiji) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil bones. Date of extinction unknown. Range  Viti Levu, Fiji.

This megapode was the same size or slightly larger than the extant ­Orange-­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.

Noble Megapode  Megavitiornis altirostris

Worthy

Megapodius altirostris Worthy 2000, p. 351 (Udit Tomo, Wainibuku, Viti Levu, Fiji) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Range  Viti Levu, Fiji.

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 ­age-­related characters. It had a markedly 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.

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Sylviornithidae (New Caledonian Giant Scrubfowl) The relationships of Sylviornis are still debated, but in general this bird is 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.

New Caledonian Giant Scrubfowl  Sylviornis neocaledoniae

Poplin

Sylviornis neocaledoniae Poplin 1980, p. 691 (L’Île de Pins, ­Nouvelle-­Calédonie) Specimens  Subfossil remains are in Paris; and Wellington, New Zealand. Status  Extinct. Known only from bones, some of which are from the Holocene. Range  New Caledonia and Isle of Pines.

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. Like other New Caledonian Giant Scrubfowl megapodes, Sylviornis was a Sylviornis neocaledoniae ­mound-­builder, and giant mound structures called tumuli, at first thought to be human graves, are now considered to be the nesting mounds (Green & Mitchell 1983). In life, Sylviornis may have been herbivorous, feeding on low vegetation and possibly digging for roots and tubers with its specialised bill and feet, but ­Mourer-­Chauviré & Balouet (2005) argue that the jaws are indicative of another feeding strategy, perhaps an invertebrate predator. 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 ­Mourer-­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.

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. 36

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Aptornithidae

North Island Adzebill  Aptornis otidiformis

(Owen)

Dinornis otidiformis Owen 1844a, p. 247 (East Cape, New Zealand). Aptornis otidiformis Owen 1848b, p. 347 (description of genus) Specimens  Subfossil remains are in Auckland, Canterbury, Otago and Wellington, New Zealand; and London. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  North Island, New Zealand.

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, flightless 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 North Island Adzebill Aptornis otidiformis with a petrel chick 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 invertebrates, or preyed on burrowing birds and reptiles by digging them out of their tunnels. Equally they may have been opportunistic omnivores, 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 shrublands and grasslands rather than subalpine shrubland or dense forests (Worthy & Holdaway 2002). Subfossil remains have been found 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 ­over-­hunting was combined with the introduction of dogs and the Pacific Rat Rattus exulans, which were probable predators of eggs and chicks.

South Island Adzebill  Aptornis defossor

Owen

Aptornis defossor Owen 1870b, p. 354, pls. 40, 41, 43, 44 (Oamaru) Specimens  Subfossil remains are in Canterbury, New Zealand; and London. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  South Island, New Zealand.

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. 37

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It was probably a ­slow-­moving, cumbersome species, whose only natural enemy as an adult was the giant Haast’s Eagle Harpagornis moorei (see p. 78). 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.

Numididae (Guineafowl) Guineafowl are ­medium-­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 transported around the world. The guineafowl are generally not threatened, but continued habitat destruction may impact on populations in the future.

Moroccan Guineafowl  Numida meleagris sabyi

Hartert

Numida sabyi Hartert 1919a, p.69 (District of Zemmour, Western Morocco) Specimens  The type is in Tring. Status  Possibly extinct, last recorded with certainty in the 1950s, but possible captive population survived until the 1980s. Range  Between River Oum er Rbia and River Sebou, Morocco. Description  58cm (23in). It differs from nominate N. m. meleagris in being larger and completely lacking the wide pale vinous ­bluish-­grey collar, the feathers being black or ­brown-­black spotted and barred with white from the chest to middle of neck; all ­wing-­quills black with white crossbars and spots, not vermiculated with light grey, and spots and bars not bordered with black; ­ground-­colour of the ­tail-­feathers and ­tail-­coverts almost pure black, except outer webs being mottled with grey, and white spots on coverts smaller; back much darker; entire under surface darker; white spots being smaller and on many feathers less numerous; head unfeathered; hind head and neck are differently feathered from any known Numida; feathering reaches to middle of neck, a few bristly feathers reach chin, but on occiput a bunch of apparently forward and upward directed feathers cover the horn from behind; bony knob and wattles as in old males of N. m. meleagris.

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 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.

Phasianidae (Grouse and quails) The Phasianidae are a diverse group of birds divided into 38 genera and widely distributed in the Old World. They are terrestrial birds and generally exhibit high sexual dichromatism, with the males being spectacularly coloured during the breeding season. Some species like the Red Junglefowl Gallus gallus have had a long association with humans. All species are considered game birds, and hunted indiscriminately. Many are endangered, some critically.

Heath Hen  Tympanuchus cupido cupido

(Linnaeus)

Tetrao cupido Linnaeus 1758, p. 160. (Virginia, based on Catesby 1743, appendix p. 1) Specimens  Specimens are in Ann Arbor, Michigan; Berkeley, California; Basle; Brussels; Cambridge, Pittsfield and Springfield, Massachusetts; Charleston, Southern Carolina; Chicago; Edinburgh; Exeter and London, England; Geneva;

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Phasianidae Gothenburg; Los Angeles; New York; Nijmegen; Oslo; Paris; Philadelphia; Pemberton and Trenton, New Jersey; Providence, Rhode Island; Rouen; San Francisco; Springfield, Illinois; Vienna; and Winnipeg, Canada. Status  The last Heath Hen, a male, died shortly after March 11, 1932, on Martha’s Vineyard Island. It had been alone since December 1928, and none has been seen since. Range  Formerly occurred widely in eastern North America. Description  43cm (17in). Not ­well-­differentiated from other subspecies of prairie chickens, but genetic analysis shows that the population was clearly distinct (Palcovacs et al. 2004). Very similar to the Greater Prairie Chicken T. c. pinnatus, but differed in being slightly smaller; plumage with more reddish hue; barring thicker on breast and sides; pinnae (horns) fewer in number, more pointed; tail ­greyish-­brown.

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. attwateri; 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 distribution 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 erroneous, 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 introduced 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 eighteenth-century record for ‘Grouse’ (Belknap 1792) may refer to the Heath Hen, but Greenway (1967) considered the record to be unconvincing. 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 mainland 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 1916). 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 unenforced. 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. 39

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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. Decline and extinction on Martha’s Vineyard By the end of the nineteenth century, the Heath Hen was confined to Martha’s Vineyard. In 1890, a total population of 120–200 was recorded, and from 1906 a population census was taken each year for 20 years. In 1906 there were only 77 after fire hit the island, and in 1908 this had fallen to 60. By 1910 there were 300, which increased to 1,000 in 1914 and almost 2,000 in 1916. But a devastating fire during the breeding season in that year, followed by heavy raptor predation in 1917, reduced the population to 150 by 1918 (Gross 1928). Poultry disease killed off many birds, and cats and poaching also took their toll. Despite these setbacks, by 1920 the population had risen to 600, but it never completely recovered from the crash, and only three birds were counted in 1925, though 25 were estimated (Phillips 1926, Gross 1928). From 1925 onwards the Federation of New England Bird Clubs employed special wardens to protect the birds, who trapped and killed 120 cats on the reservation, and uncovered indirect evidence of poachers. But their efforts proved to be in vain. Excessive inbreeding and disease had reduced the population so that it ceased to be viable. DNA analysis has shown that genetic variability was much reduced at least 30 years prior to extinction (Johnson & Dunn 2006), making the population extremely vulnerable to diseases. On 9 February 1932, after an interval of nine months during which he had not been seen, the last Heath Hen, a male affectionately nicknamed ‘Booming Ben’ reappeared, ­out-­surviving the rest of his kind by nearly four years (Cokinos 2000). He was observed at regular intervals until 11 March. 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. 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 ­Sharp-­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

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Phasianidae 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 ­over-­hunting (Silvy & Hagen 2004). The New Mexico population was restricted to the high mesas in Colfax and Union counties, New Mexico, and probably 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 ­south-­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 grasslands 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.

Amik Gölü Black Francolin  Francolinus francolinus billypayni

Meinertzhagen

Francolinus francolinus billypayni Meinertzhagen 1933, p. 21 (Lake Antioch, Syria) Specimens  The type is in Tring. Status  Possibly extinct. Last recorded in the late 20th century. Range  Amik Gölü, ­south-­central Turkey. Description  34cm (13.5in). Male nominate predominantly black with white patch on cheek; presence of a chestnut collar; belly rufous; white spots on flanks; back and wings scalloped with shades of golden brown with ­sub­terminal ­tawny-­buff bands and pale edges; tail black with narrow white or greyish bars; iris brown; legs and feet ­reddish-­brown to red. Female similar to male, but mainly brown with wider brown bars on lower back and tail; head and underparts buff; rump and ­uppertail-­coverts light brown; white ­cheek-­patch absent; and chestnut collar replaced by a nuchal patch.

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 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.

New Zealand Quail  Coturnix ­novae­zelandiae ­novaezelandiae

Quoy & Gaimard

Coturnix novaezelandiae Quoy & Gaimard 1830, p. 242 (Baie Chouraki = Hauraki Gulf, North Island, New Zealand) Specimens  Specimens are in Auckland; Christchurch; Wellington, New Zealand; Cambridge, England; Cambridge, Massachusetts; Dresden; Edinburgh; Liverpool; New York; Paris; Philadelphia; Pittsburgh; Toronto; Tring; Vienna; and Washington State. Status  Extinct since at least 1870. Range  Formerly occurred on North, South, and Great Barrier Islands, New Zealand.

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Extinct Birds Description  19cm (7.5in). Adult male: top of head, back, primaries and outer secondaries, tail coverts dark brown, upper face rufous brown variably striped with black and white, sides of face, chin and throat rufous with narrow white shaft lines, dark band through the eye to the side of throat, lower neck mottled black and white, flanks rufous brown, abdomen fulvous white, bill black, iris light hazel, feet pale flesh brown. Adult female similar to male but lighter, no rufous colour on face or throat.

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 1870 (Buller 1887–88). 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 ­north-­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. 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).

Canary Islands Quail  Coturnix gomerae

Jaume et al.

Coturnix gomerae Jaume et al. 1993, p. 154–155 (Bujera del Silo, La Gomera) Specimens  Subfossil remains are in Tenerife. Status  Extinct. Known only from subfossil remains. Possibly survived into historic times. Range  El Hierro, La Palma, Tenerife and Fuerteventura, Canary Islands; probably also Gran Canaria and Lanzarote.

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 disappeared 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 a more terrestrial species.

Canary Islands Quail Coturnix gomerae

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Anatidae

Himalayan Mountain Quail  Ophrysia superciliosa

(Gray)

Rollulus superciliosus J. E. Gray 1846, p. 8, pl. 16 (‘India’ = Mussoorie Hills, Himalayan foothills, eastern Punjab) Specimens  Specimens are in Liverpool; New York; and Tring. Status  Last seen in June 1868, when a specimen was shot by Captain J. Hutton. May still survive in low numbers. Range  Formerly occurred in the foothills of the Himalayas from near Mussoorie in east Punjab and Naini Tal in Kumaon. Unconfirmed reports have come from eastern Kumaon (Ripley 1952). Description  25cm (10in). Male greyish with distinctive ­black-­and-­white head and neck; forehead white merging into grey; superciliary stripe and eye-stripe white; head and neck mottled black and white, throat black; back dark brown, below greyer, feathers edged with black giving a mottled appearance; wings browner; ­undertail-­coverts black spotted with white; female cinnamon brown, undersides paler spotted with black; top of head and throat pale brown, white feathers restricted around eye. Bill bright red, legs and feet pale red. Immature male, duller, with buff mottling on wing.

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. 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 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 specimen was shot. They have not been seen since, and I never saw them before.

Anatidae (Ducks, geese and swans) The Anatidae comprise a group of diverse, generally aquatic birds that are found throughout the world except Antarctica. The family contains 40 genera. A number of species were flightless island endemics that had become entirely terrestrial. Most species are regularly hunted, so ­over-­hunting and habitat destruction has impacted on populations. As a result, extinctions within this group are high.

South Island Goose Cnemiornis calcitrans

South Island Goose Cnemiornis calcitrans

Owen

Cnemiornis calcitrans Owen 1865, p. 396 (Tunaru) Cnemiornis minor Forbes 1892d, p. 185 (Omaru) Specimens  Subfossil remains are in Canterbury and Wellington, New Zealand; and London. Status  Known only from subfossil bones. Date of extinction unknown, but probably 16th century. Range  South Island, New Zealand.

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According to most authors, the genus Cnemiornis most closely resembles the extant Cape Barren Goose Cereopsis novaehollandiae, but Cnemiornis was larger and totally flightless. These geese were massive (15–18kg), being about three times heavier than the Canada Goose Branta canadensis and Cape Barren Goose (Worthy & Holdaway 2002), but with tiny wings. The bill was deep and ­squared-­off, which was presumably an adaptation for grazing grasses. It appears that the South Island Goose was exterminated soon after the Polynesian arrival to the islands, perhaps exacerbated by the loss of suitable grassland habitats due to climate change.

North Island Goose  Cnemiornis gracilis

Forbes

Cnemiornis gracilis Forbes 1892d, p. 187 (Kaiiwi, New Zealand) nomen nudum at date of publication, ibid Dawson 1958, p. 235 Cnemiornis septentrionalis Oliver 1955, p. 602. Specimens  Subfossil remains are in London; and Wellington, New Zealand. Status  Known only from subfossil bones and perhaps one account. Date of extinction not known, but perhaps as recent as the 1870s. Range  North Island, New Zealand.

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 subsequently 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 historical account (see Worthy & Holdaway 2002), quoted from ­Cockburn-­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.

Bering Cackling Goose  Branta hutchinsii asiatica

Aldrich

Branta hutchinsii asiatica Aldrich 1946b, p. 95 (Bering Island, Siberia) Specimens  Specimens are in Washington, D. C. Status  Extinct, last recorded in the early 20th century. Considered conspecific with Aleutian Cackling Goose B. h. leucopareia by many authorities. Range  Komandorskie and Kurile Islands, north Pacific. Probably migrated southward to Japan. Description  55cm (22in). A small, dark form of the Cackling Goose with short neck and bill. Generally ­greyish-­brown on breast, back and wings, belly and ­undertail-­coverts white; head, neck and tail black; white ­cheek-­patches separated by black line under throat; rump white; conspicuous wide white neck-ring subtended by a ring of dark feathers; bill, legs and feet blackish. Differed from B. h. leucopareia in having a wider white ­neck-­ring.

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 population was exterminated due to ­over-­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.

Giant Hawaiian Goose  Branta sp. Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Disappeared c.1500. Range  Hawaii, Hawaiian Islands.

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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 determined. 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 ­moa-­nalos (see p. 56) (Olson & James 1992). 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 Harpagornis moorei (see p. 78). 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).

Greater Hawaiian Goose ­(Nene-­Nui)  Branta hylobadistes

Olson & James

Branta hylobadistes Olson & James 1991, p. 45 (Auwahi Cave, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct, only known from subfossil bones. Date of extinction unknown. Range  Found in Holocene deposits on Maui, Hawaiian Islands.

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, ­as-­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.

Chatham Islands Black Swan  Cygnus sp.

(Forbes)

Chenopsis sumnerensis Forbes 1892d, p. 188 (Monck’s Cave, Sumner, New Zealand) Cygnus chathamicus Oliver 1955, p. 605 (Chatham Islands) Specimens  Subfossil remains are in London and Wellington, New Zealand. Status  Extinct. Date of extinction not known. Range  Known only from subfossil bones collected in New Zealand and the Chatham Islands.

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. 45

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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 probably represent yet another ­now-­extinct species from these islands.

Mauritius Sheldgoose  Alopochen mauritiana

(Newton & Gadow)

Goose. Marshall 1668 (in Khan 1927) Sarkidiornis mauritiana Newton & Gadow 1893, p. 290–91, pl. 34, figs. 9–10 (Mauritius) Specimens  Subfossil remains are in Cambridge and Tring. Status  Extinct since c.1700. Known only from bones and vague travellers’ accounts. Range  Endemic to Mauritius.

Geese were mentioned by the Dutch on Mauritius during the 17th century (Cheke & Hume 2008), but without providing any details. The English mariner Marshall (1668) 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.

Mauritius Sheldgoose Alopochen mauritiana

The Mauritius Sheldgoose probably disappeared due to ­over-­hunting and perhaps predation of eggs and chicks by introduced predators.

Réunion Sheldgoose  Alopochen kervazoi

(Cowles)

Mascarenachen kervazoi Cowles 1994, p. 88 (Grotte des Premiers Français, near ­Saint-­Paul, Réunion) Alopochen kervazoi ­Mourer-­Chauviré et al. 1999, p. 11. Specimens  Subfossil remains are in St Denis, Réunion. Status  Extinct. Known from subfossil remains and accounts only. Not seen since c.1700. Range  Endemic to Réunion, Mascarene Islands.

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: 46

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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.

Crested Shelduck  Tadorna cristata

(Kuroda)

Pseudotadorna cristata Kuroda 1917, p. 1, fig. 1 (Naktung River, near Fusan, Korea) Specimens  Specimens of a male and female are in Tokyo; a female is in Copenhagan. Status  Probably extinct. Possible but unconfirmed sightings as recently as 1984. The last specimen taken was in September 1916 along the Naktung River, near Fusan, Korea. Range  Probably bred in ­south-­east Russia, North Korea and northern China; possibly migrated south to Korea and Japan. Description  60–63cm (23.5–24.5in). A distinctive parti-coloured and crested duck. Male: Top of head dark metallic green, black line, which separates grey face, extending from forehead and below eye to middle of hind neck; nuchal feathers long, greyish at base tipped greenish black; pale spot beneath eye, chin with dark green patch; face and neck grey with indistinct brown barring; upper breast with wide metallic green band narrowing onto hind neck; back, lower breast and abdomen dark grey with narrow, white vermiculations; greater ­wing-­coverts white, ­median-­coverts grey with white vermiculations, shading to brown; inner secondaries chestnut, outer secondaries green, greater ­wing-­coverts white; primaries and tail greenish black; undertail-coverts brown shading to white; bill probably pinkish. Female: Top of head and nape black extending like spectacles around eye; forehead and eye surround, throat and neck white; back, ­median-­coverts, flanks and underparts ­greyish-­brown with thin white vermiculations.

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).

New Zealand ­Pink-­eared Duck (Scarlett’s Duck) Malacorhynchus scarletti

Olson

Malacorhynchus scarletti Olson 1977a, pp. 132–35 (Pyramid Valley, South Island, New Zealand) Specimens  Subfossil remains are in Canterbury, New Zealand. Status  Known only from subfossil bones found in marshes and middens. Date of extinction unknown, but certainly caught by Polynesian colonists. Range  North and South Islands, New Zealand, and the Chatham Islands.

This duck was related to, but was considerably larger than, the Pink-eared Duck Malacorhynchus membranaceus of Australia and Tasmania. From a study of its anatomy (Worthy & Holdaway 2002), the New Zealand 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.

Finsch’s Duck  Chenonetta finschi

(van Beneden)

Anas finschi van Beneden 1875, p. 123, pl. 3 (Earnscleugh Cave, New Zealand) Chenonetta finschi Worthy & Olson 2002, p. 1–17 Specimens  Subfossil remains are in Bremen; Berlin; Canterbury and Wellington, New Zealand. Status  Known only from subfossil bones; date of extinction unknown. Range  North and South Islands, New Zealand.

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Finsch’s Duck is closely related to the Australian Wood Duck Chenonetta jubata, and probably derived from it (Olson & Worthy 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 eagle Harpagornis: 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 ­wing-­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. 45). 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.

Coues’s Gadwall (Washington Island Gadwall) Anas (strepera) couesi

(Streets)

Chaulelasmus couesi Streets 1876, Bulletin of the Nuttall Ornithological Club, vol. 1, p. 46 (Washington Island, Line Islands) Specimens  Specimens are in Washington, D. C. Status  Extinct, not seen since 1874. Doubtfully distinct. Range  Washington Island (Teraina), Line Islands, Kiribati. Description  Approximately the size of a teal, but otherwise resembled young Common Gadwall, except that the bill and feet were black and the number of lamellae in the bill was greater.

Only two specimens of this duck are known, the co-types collected in 1874 (Streets 1876). 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 differentiated 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). Nothing was ever recorded of the bird’s ecology.

Amsterdam Island Duck  Anas marecula

Olson & Jouventin

‘animals’ W. de Vlaming 1696, in Anon. 1854, p. 75–81 and 262–265 (Amsterdam Island); also Schilder 1976 ‘small brown duck’. Barrow 1806, p. 148. (Amsterdam Island) Anas marecula Olson & Jouventin 1996, pp. 1–9 Specimens  Subfossil remains are in Washington, D. C.

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Amsterdam Island Duck Anas marecula

Status  Known only from vague early accounts and subfossil bones (Amsterdam Island only). Probably exterminated by sealers and rats. Range  Amsterdam Island and St Paul Island, southern Indian Ocean. Description  Differs markedly from any other duck due to its small size and reduction of the wings. The only similar species is the flightless Auckland Islands Teal Anas aucklandica, which is a larger bird. The bill of Amsterdam Island Duck is very short and rounded at the tip, and more closely resembles the wigeons than any other group.

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 was by William de Vlaming, who in 1696 thought he saw two ­four-­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. The island was again 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 not much larger than a thrush, which 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 dress, so that the description of the Amsterdam bird as ‘small and brown’ suggests a native population in degenerate plumage. 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. Bones from cavities Olson & Jouventin (1996) confirmed the presence of an endemic duck by examining bones unearthed by Martinez (1987), who collected thousands of specimens from cavities in basaltic rocks on Amsterdam, most of which were seabirds. Bones of a small duck were also common and found throughout most of the island habitats from sea level to 500m. All the main bones of the skeleton were present and Martinez concluded that they represented a small duck of the genus Anas, with robust legs and weak wings suggesting poor flight or even flightlessness. Salt glands were also poorly developed, inferring that the bird was not a coastal species. Olson & Jouventin (1996), in describing the species, confirmed the generic proposal of Martinez as Anas is the only genus which has successfully adapted to terrestrial habitats on oceanic islands. 49

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Nicéforo’s Pintail  Anas georgica niceforoi

Wetmore & Borrero

Anas niceforoi Wetmore & Borrero 1946, p. 68 (Laguna de Tota, Boyac, Colombia) Specimens  Specimens are in Washington, D. C. Status  Extinct since 1952. Reasons for extinction are unclear. Range  Formerly occurred in Central Colombia between 1,000 and 3,000m in the upper Cauca Valley, central Cordillera Oriental, the Bogota Savanna and Cundinamarca. Description  43–55cm (17–21.5in). Nicéforo’s Pintail was most similar to Anas georgica spinicauda but generally darker and richer in colour; top of the head dark brown with little or no rufescent suffusion; head and necked more streaked; black speculum lacked green reflections; tail less pointed; longer line of culmen straighter, nail smaller.

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. Nothing was recorded about its habits.

Tristram’s Pintail  Anas acuta modesta

(Tristram)

Dafila modesta Tristram 1886, p. 79, pl. 7 (Sidney Island, Phoenix Group, central Pacific) Specimens  Specimens are in Liverpool. Status  Known only from the three co-types, formerly in Tristram’s collection, collected by J. V. Arundel on Sidney Island. Never seen since. Of doubtful validity. Range  Sidney Island (Manra), Phoenix Group, Kiribati. Description  The male was similar to a nominate Northern Pintail Anas acuta acuta but smaller and dull-coloured, and in the process of moulting into breeding dress. Salvadori (1895) pointed out that the male type had been kept in confinement as its right wing had been clipped. He was unable to distinguish the birds from ‘some’ specimens of Northern Pintail (presumably the smaller and duller), but when considering the location, he felt it unsafe to synonymise the two.

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 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.

Rennell Island Teal  Anas gibberifrons remissa

Ripley

Anas castanea remissa Ripley 1942, p. 94 (Rennell Island, Solomon Islands) Specimens  The type is in New York. Status  Extinct since 1959. Reasons for extinction are unclear. Range  Formerly occurred on Rennell Island, Solomon Islands. Description  Differed from A. g. gracilis (New Guinea, Australia, New Zealand and adjacent islands) in being smaller and darker, and from A. g. gibberifrons (islands from Java to Timor) in having a smaller bill and streaking on the underside of the neck.

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 & Wolff (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.

Mascarene Teal  Anas theodori

Newton & Gadow

Anas theodori Newton & Gadow 1893, pp. 282, 291–92, pl. 34 (Mauritius) Specimens  Subfossil remains are in Cambridge, England; London and Paris. Status  Known from inadequate accounts and subfossil bones. Range  Mauritius and Réunion, Mascarene Islands.

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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.

Réunion Pochard  Aythya sp. Aythya sp. ­Mourer-­Chauviré et al. 1999, p. 20 (Réunion) Specimens  Subfossil remains are in St Denis, Réunion. Status  Extinct. Known from possibly one inadequate account and subfossil remains. Range  Réunion, Mascarene Islands.

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.

Chatham Duck  Pachyanas chathamica

Oliver

Diaphorapteryx sp. Milne-Edwards 1896, p. 126 Pachyanas chathamica Oliver 1955, p. 599 (Chatham Islands) Specimens  Subfossil remains are in Canterbury, New Zealand. Status  Known only from subfossil bones. Probably became extinct in the 16th century. Range  Chatham Islands, New Zealand.

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’. It was larger than the Mallard Anas platyrhynchos, with a heavy cranium and ­well-­developed salt glands, suggesting it was an inhabitant of the Chatham Islands’ lagoons and coastal regions (Worthy & Holdaway 2002). It had comparatively reduced wings so had reduced powers of flight, which would have made it vulnerable to ­over-­hunting by humans and predation from introduced mammals, especially the Pacific Rat Rattus exulans. This duck appears to have become extinct during the 16th century.

Pink-headed Duck  Rhodonessa caryophyllacea

(Latham)

Anas caryophyllacea Latham 1790, p. 866 (India) Specimens ­Pink-­headed Ducks are represented in many museums, including Brussels; Leiden; Liverpool; Mumbai, India; New York; Paris; Stockholm; Tring; Washington, D.C.; and Yale. Status  Probably extinct. The last wild specimen was said to have been collected in 1922, but the species survived until 1932 in captivity at Foxwarren Park, England (Ezra 1942), and probably at Clères, France and in Calcutta, until about 1940–44. The last accepted record from the wild was from Bihar in 1948–9. Range  Formerly occurred in scattered localities from Bhutan south and west to central Bihar, along the Ganges and south along the coast of India to Madras. Description  60cm (24in). Particularly distinctive in coloration. Generally brownish, paler beneath, back, sides and belly speckled with pinkish-white; entire head and neck bright pink. Wings buff, speculum dull salmon-pink. Tail dark brown, bill reddish-pink, eyes ­orange-­red, feet dark with reddish tinge. Female paler than male, throat pink, not brown; immature bird with white (not pink) head.

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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 Pink-headed Duck long, it is a great pity that so few studies were ever Rhodonessa caryophyllacea carried out, particularly as the bird appears to have been unusual in many ways. The great ­19th-­century Indian ornithologist, A. O. 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, Gujurat, Kutch, Kathiawar, Sind, Rajputana, Central Indian Agency or the Central Provinces (Hume & Marshall 1879–81). There were very few records from the North West Provinces of India. Hume saw or heard of it only once in 20 years, but one was shot near Fatehpur, while others were apparently obtained in the Dun and at Bareilly. It was ‘extremely rare’ in Burma, and found occasionally in Nepal, Bhutan and Bangladesh (complete review of records in Collar et al. 2001). It most often occurred in the area of the Himalayan foothills west of Brahmaputra and north of the Ganges, which appears to have been the centre of distribution. Simson (1884) says that it was of very local and scattered distribution and was little known to most ornithologists in Bengal, although it was ‘far from uncommon’ in certain parts of the province. 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. 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, crocodileinfested rivers, heavily inundated by floods and inhabited sparsely by humans, but commonly by tigers. This area has become more populated in the 20th century and has been drained and cultivated. The last generally accepted record was from Manroopa Lake, Bihar in 1948–49 (Collar et al. 2001). However, in 2004, a team from BirdLife International made a possible sighting in Burma. While considered extinct, this duck may just cling on. 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. They never formed large flocks, nor associated with other duck species. They were not particularly good to eat, but were often shot nevertheless; however, Jerdon (1864) claimed that they were good eating. Blanford (1898) said that the duck was found throughout the year in small flocks of 4–10, or occasionally up to 20 or 40 in the cool season, and in pairs from April to September. F. A. Shillingford 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 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 for ducks, being quite round and china white (interestingly, Baker 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. Birds never bred in captivity, despite being kept under seemingly ideal conditions. Bucknill (1924) reported that the species appeared to have virtually disappeared by the time of writing, 52

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and inferred that it may have been partly migratory, for at Lucknow it was only a very rare winter visitor in cold weather, and that its distribution may have been to some extent dependent on the rains. Bucknill spoke of a number of known ‘sportsmen’ in the province of Bihar and Orissa who had been attending big duck shoots in Bengal for over 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 it, 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 halfcentury . . . 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. J. C. Parker reported to Hume that the flight of the duck was very powerful and rapid. Shillingford commented that the call was like that of the Mallard drake, ‘with a slight musical ring about it.’ Hume expressed the opinion that Rhodonessa was very closely related to Anas, and that he would have merged the two, but for the very extraordinary eggs. However, Johnsgard (1961) believed the species to be closely related to the pochards, a fact supported by examination of museum skin osteology and feather protein analysis (Woolfenden 1961; Brush 1976).

Labrador Duck  Camptorhynchus labradorius

(Gmelin)

Labrador Duck. Pennant 1785, p. 559 Anas labradoria Gmelin 1789, p. 537 (Arctic America, Connecticut and Labrador). Fulica grisea Leib 1839, p. 170 (described as new a bird found in Philadelphia market on December 15, 1839. The specimen, now in the Academy of Natural Sciences of Philadelphia, is generally considered to be a young bird) Specimens  Labrador Ducks can be found in many major museums around the world. The number of surviving specimens is given by Hahn (1963) as 54. Status  Extinct. The last authentic specimen was shot in the autumn of 1875 on Long Island, New York. Range  Occurred sparsely on the Atlantic coast of northern North America. Description  54cm (22in). Male black and white, somewhat reminiscent of a Common Eider Somateria mollissima. Head and neck white with black line on top of head, and black collar; back, rump, upper tail coverts and tail black; ­wing-­coverts white, primaries black; underparts black. Legs and feet whitish ash, spattered blackish on hind part, webs black. Bill blackish-brown, base orange or yellow; iris reddish hazel to yellow. Female more ­brownish-­grey, ­whitish-­grey line behind eye, throat and speculum white. Immature like female, except darker.

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 disappeared 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.

Labrador Duck

Camptorhynchus labradorius

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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 ­Red-­breasted Merganser Mergus serrator and Mallard Anas platyrhynchos (Chilton & Sorenson 2007). Audubon’s son John was shown empty nests on the tops of bushes in Labrador, which were thought possibly to be nests of the Labrador Duck (Audubon 1838), but this seems doubtful. In Rothschild (1907), Alfred Newton remarked that the bird bred on rocky islets, but Rothschild suggests that this can be no more than conjecture. Nevertheless it is a reasonable conjecture, for Zusi & Bentz (1978) examined the duck’s osteology and musculature, found skeletal characters shared only with the eiders, and concluded that the Labrador Duck must have been most closely related to them. Its general appearance concurs with this, and as eiders often nest on rocky islets, it is probable that the Labrador Duck did so too. The Labrador Duck had a peculiar bill, with swollen cere, an unusually large number of lamellae ­(tooth-­like projections on the jaws) and prominent lateral ­bill-­flaps, suggesting that it had a specialised feeding habit and diet. Giraud (1844) said that it fed on small shellfish and other fish, obtained by diving; Leib (1839) recorded that the stomach of his specimen contained broken shells.

Auckland Island Merganser  Mergus australis

Hombron & Jacquinot

Mergus australis Hombron & Jacquinot 1841, p. 320 (Auckland Islands) Specimens  Specimens are in Christchurch, Otago and Wellington, New Zealand; Dresden; New York; Tring; and Vienna; subfossil remains are in Wellington. Status  Not seen since 1901, and considered extinct. Range  Confined to the Auckland Islands, New Zealand. Bones discovered in kitchen middens at several sites in New Zealand may be referable to this species, and if so, will extend the previously known range. Description  58cm (23in). Head with crest brownish red, rusty on throat and foreneck; back ­blackish-­brown, feathers edged with grey; sides of body grey, underparts barred irregularly with grey and white; a double white patch on grey wing. Sexes similar, but males larger, with longer bill and crest. Both sexes resembled the female of northern species of merganser, but there was no male nuptial plumage. It was the smallest species of merganser, and had a more slender bill and shorter toes than other species. It had reduced wings but could still fly well; reports that it was flightless are erroneous.

The Auckland Islands were discovered on 18 August 1806 by Captain Abraham Bristow and named after the English politician Lord Auckland. The Auckland Island Merganser was first recorded by Jacquinot in 1840, when it was confined to the Auckland Islands, but in 1945 a subfossil mandible was found in a midden at a Polynesian site at Wairau Bar, Marlborough, South Island, New Zealand. Seven years later a premaxilla and cranium were found at Lake Grassmere and various other subfossil bones from the mainland of New Zealand have been discovered. These certainly refer to some form of Mergus, but although probably M. australis, the material is insufficient to establish the identity beyond question. 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), who had previously seen the merganser at the Auckland Islands in 1840 (McCormick 1884), but the reference is vague and he might have confused it with Auckland Islands Teal Anas aucklandica (Williams & Weller 1974). 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. 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 54

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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 destruction. Waite (1909) emphasised that there was little doubt that the pigs had caused havoc among ­ground-­nesting birds. Habits The merganser inhabited fresh and brackish waters of creeks and upper estuaries, occasionally wandering into salt-water harbours. It possibly fed on freshwater shrimps, but this is unconfirmed. Hutton (1902) claimed to have taken a large fish from the bill of a merganser, while the stomach of the only known preserved, complete specimen, at Tring, contained macerated fish bones, the mandibles of a polychaete worm and an unidentified gastropod (P. J. K. Burton, pers. comm. to Kear). Little was ever recorded of its breeding. Kear & Scarlett (1970) believed that the pair-bond may have been a long one. Four 7 to 10 day old ducklings were found in the month of December, but the nest site, the eggs and the clutch size were never described. Adult birds in pairs were recorded in October, November, January, May and July. Nothing else is known of its habits.

New Zealand Musk Duck  Biziura delautouri

Forbes

Biziura delautouri Forbes 1892a, p. 417 (Omaru, Otago, New Zealand) Biziura lautouri Forbes 1892d, p. 188, nomen nudum Specimens  Subfossil remains are in Auckland, Canterbury, and Wellington, New Zealand; and London. Status  Extinct since the 16th century. Known only from subfossil bones. Range  North and South Islands, New Zealand.

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. The New Zealand Musk Duck may have preferred larger prey than other duck species, including crayfish, large insects, molluscs and fish (Tennyson & Martinson 2006). Like all extinct New Zealand anatids, this species probably succumbed to ­over-­hunting.

New Zealand ­Stiff-­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 ­Blue-­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. 55

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Maui Nui ­Moa-­nalo  Thambetochen chauliodous

Olson & Wetmore

Thambetochen chauliodous Olson & Wetmore, 1976, p. 252 (Moomomi dunes, Molokai, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D.C. Status  Extinct. Only known from subfossil bones. Date of extinction unknown. Range  Molokai and Maui, Hawaiian Islands.

The Hawaiian Islands, due to their isolation and absence of terrestrial mammals, witnessed an extraordinary radiation of ducks and geese, which often reached great size and evolved specialised adaptations. Most remarkable were the ­moa-­nalos, which superficially resembled geese, but actually evolved from dabbling ducks (Sorenson et al. 1999). So different were these ­goose-­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 disproportionately 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 Rattus exulans. The ­moa-­nalo Thambetochen chauliodous from Molokai and Maui was the first species discovered, and appears to have been a lowland species.

Oahu ­Moa-­nalo  Thambetochen xanion

Olson & James

Thambetochen xanion Olson & James 1991, p. 35 (Barbers Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D.C. Status  Extinct. Only known from subfossil bones. Date of extinction unknown. Range  Oahu, Hawaiian Islands.

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 landbridge. 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.

Stumbling ­Moa-­nalo  Ptaiochen pau

Olson & James

Thambetochen pau Olson & James 1991, p. 39 (Auwahi Cave, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D.C. Status  Extinct. Only known from subfossil bones. Date of extinction unknown. Range  Maui, Hawaiian Islands.

The Stumbling ­Moa-­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 chauliodous on Maui. The mountain ranges of the Hawaiian Islands provided the last refuge for many ­now-­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 of ­over-­hunting with predation or competition by introduced exotic animals.

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Kaua’i ­Turtle-­jawed Goose  Chelychelynechen quassus

Olson & James

Chelychelynechen quassus Olson & James 1991, p. 33 (Makawehi dunes, Kaua’i, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D.C. Status  Extinct. Only known from subfossil bones. Date of extinction unknown. Range  Kaua’i, Hawaiian Islands.

Kauai Turtle-jawed Goose Chelychelynochen quassus

Kaua’i, the oldest of the main Hawaiian Islands, harboured the most distinctive species of ­moa-­nalo. The ­Turtle-­jawed Goose was ­so-­named because of the extraordinary deep, short jaws with bony ­tooth-­like projections, reminiscent of a turtle. It was large, ­robust-­legged and completely 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).

Kaua’i Mole Duck Talpanas lippa

Olson & James

Talpanas lippa Olson & James, 2009. In Iwaniuk et al. 2009, p. 59 (Makauwahi Cave, Maha’ulepu/Pa’a district, Kaua’i, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct, known only from subfossil remains. Date of extinction unknown. Range  Kaua’i, Hawaiian Islands.

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 presumably 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.

Hunter Island Penguin  Tasidyptes hunteri

van Tets & O’Connor

Tasidyptes hunteri van Tets & O’Connor 1983, pp. 1–13 (Hunter Island, Tasmania) Specimens  Subfossil remains are in Launceston. Status  Extinct. Known only from incomplete skeletons of at least two individuals found in 13th century aboriginal middens. The date of extinction is not known, but believed to be between 760 years ago to possibly less than 200. Range  Hunter Island, Tasmania.

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Kauai Mole Duck Talpanas lippa

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 ­as-­yet unknown breeding locality (van Tets & O’Connor 1983). The Hunter Island Penguin may have disappeared as recently as c.1800 due to ­over-­hunting by Tasmanian Aborigines, who themselves were wiped out by European settlers just 75 years later.

Chatham Islands Penguin  Eudyptes sp. Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known from subfossil remains only. Last recorded around 1872. Range  Chatham Islands, New Zealand.

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. pachyrhynchus (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. 58

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Waitaha Penguin  Megadyptes waitaha

Boessenkool et al.

Megadyptes waitaha Boessenkool et al. 2009, p. 817 (Lake Grassmere, Marfells Beach, Marlborough, South Island) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Disappeared around c.1500–1550. Range  Southern South Island, New Zealand.

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, pollution, and predation from introduced mammals. Some species are in critical danger of extinction.

Waitaha Penguin Megadyptes waitaha

Jamaican Petrel  Pterodroma caribbaea

Carte

?The larger dark Petterill or Shearwater. Browne 1756, p. 482. ? Proc[ellaria] Jamaicensis Bancroft 1832–34, p. 81, nomen nudum (Summit of the Blue Mountain Peak, Jamaica) Pterodroma caribbaea Carte 1866, p. 93, pl. 10 (Blue Mountains, Jamaica) Specimens  Specimens are in Cambridge, England; Liverpool; and Tring. Status  Probably extinct since about 1880. However, Bond (1936) recorded seeing one at sea west of the Bimini Group, Bahamas, in 1936. Range  Formerly bred in the Blue Mountains of Jamaica, but may have also occurred on Guadeloupe and Dominica. Description  40cm (16in). General coloration dark sooty brown, paler on the abdomen and ­undertail-­coverts. ­Uppertail-­coverts and bases of all the tail feathers dirty white. Pale rump patch conspicuous when wings expanded, concealed when closed. Lacks the white spots on the neck and rump of P. hasitata.

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 59

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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 domicile, 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 sometimes 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. 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 Attwood (1791) on Dominica, but the latter is much more likely to represent P. hasitata. 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.

Mount Pitt Petrel (Norfolk Island Petrel)  Pterodroma (solandri?)

(Gray)

Norfolk Island Petrel. Phillip 1789, p. 161 and plate Procellaria phillipii G. R. Gray 1862b, p. 246 (Norfolk Island); based on the above Status  Extinct. Last recorded in 1790. Known from illustrations and accounts only. Range  Norfolk Island.

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 subspecies related to P. solandri. 60

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Slaughter of the petrels Clark was an Englishman who had been sent to Sydney with the Royal Marines in 1788 and was drafted to Norfolk Island in March 1790, as Quartermaster General under one Major Ross. His copious notes earned him the nickname of ‘the Pepys of early settlement’, and he penned horrific accounts of the mass killing of the Mount Pitt birds ­(so-­called because they nested on the slopes of Mount Pitt). A penal colony was established in 1788 and for some periods, attempts were made to subsist off the land (Greenway 1967). Clark indicated that a total of 170,000 birds were killed in the three months from 10 April to 10 July 1790, and occasionally 5,000 were killed in a single night’s hunting (Whitley 1934). Iredale (1930) records a Captain Hunter’s account of killing the birds: In the month of April we found that Mount Pitt, which is the highest ground on the island, was during the night crowded with birds. This hill is as full of holes as any rabbit warren; in these holes at this season these birds burrow and make their nests, and as they are an aquatic bird they are, during the daytime, frequently seen at sea in search of food. As soon as it is dark they hover in vast flocks over the ground where their nests are. Our people (I mean seamen, marines and convicts) who are sent out in parties to provide birds for the general benefit, arrive upon the ground soon after dusk, where they light small fires which attract the attention of the birds, and they 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 plentiful 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. Understandably, the Norfolk Island Petrel was unable to sustain such huge losses, and became extinct less than 13 years after its discovery. The introduction of predatory 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.

Lesser Hawaiian Petrel  Pterodroma jugabilis

Olson & James

Pterodroma jugabilis Olson & James 1991, p. 17 (Barbers Point, Oahu) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from Holocene subfossil remains. May have survived into recent times. Range  Oahu and Hawaii, Hawaiian Islands.

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 ­over-­hunting by humans and predation by the Polynesian Rat on the main islands, which no doubt led to their extermination. 61

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Greater St Helena Petrel  Pseudobulweria rupinarum Pterodroma sp. Ashmole 1963b, p. 398 (St Helena) Pterodroma rupinarum Olson 1975b, p. 14 (Prosperous Bay, St Helena) Pseudobulweria rupinarum Imber 1985 Specimens  Subfossil remains are in London and Washington, D. C.

(Olson)

Greater St Helena Petrel Pseudobulweria rupinarum

Status  Extinct. Disappeared sometime after 1502. Range  St Helena. Known only from Pleistocene and Holocene subfossil deposits.

The Greater St Helena Petrel was a ­medium-­sized petrel belonging to the rostrata group of Pseudobulweria petrels, but smaller than Tahiti Petrel P. rostrata, and larger and with longer wings than either Mascarene P. aterrima or Beck’s P. becki Petrels (Olson 1975b). The heavy, down-angled bill is characteristic of the rostrata group, and distinguishes them from other members of the genus. Tahiti Petrel P. rostrata is distinguished in flight by high arching glides and slow determined beats of wings, which are not held straight as in many petrels but are bent at the wrist. P. rostrata breeds in the Marquesas, New Caledonia and the Society Islands, P. aterrima is known only from a handful of specimens taken on Réunion and Mauritius (see p. 442), and P. becki was, until 2007, known from only two specimens taken at sea near the Solomon Islands (Shirihai 2008; see p. 443). Species of the rostrata group, so far as is known, breed on high mountainous islands. Olson (1975b) comments: Although St Helena does not afford the great altitudes of the known breeding sites of the P. rostrata group, its relief and climate come closer to approximating those conditions than do any of the other South Atlantic islands. The reason for the bird’s extinction is not certainly known, but it is probably a result of severe deforestation during the 16th century. The population appeared to be declining as a result of natural environmental pressures before the arrival of humans.

Lesser St Helena Petrel  Bulweria bifax Olson Pterodroma sp. Ashmole 1963b, p. 400 (St Helena) Bulweria bifax Olson 1975b, Smithsonian Contributions to Palaeontology, no. 23, p. 17 (near Sugarloaf Hill, St. Helena) Specimens  Subfossil remains are in London and Washington, D. C. Status  Extinct. Probably disappeared sometime after 1502. Range  St Helena. Known from subfossil bones occurring in Pleistocene and recent deposits.

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 appearance, 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 influence. Any surviving population was probably extirpated after the arrival of Europeans and their commensals in 1502. 62

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Scarlett’s Shearwater  Puffinus spelaeus

Holdaway & Worthy

Puffinus spelaeus Holdaway & Worthy 1994, p. 203 (Te Ana Titi Cave, Fox River, Westland, New Zealand) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known from subfossil bones. Date of extinction unknown but may have survived until at least the 15th century or more recently. Range  South Island, New Zealand.

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.

Lava Shearwater  Puffinus olsoni

McMinn et al.

Puffinus olsoni McMinn et al. 1990, p. 65 (Cueva de las Palomas, Montaña de la Arena La Oliva, Fuerteventura) Specimens  Subfossil remains are in Lanzarote and Washington, D. C. Status  Extinct. Known only from subfossil remains. Disappeared at some point in the 15th century. Range  Lanzarote and Fuerteventura, Canary Islands.

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).

Lava Shearwater Puffinus olsoni

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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 ­over-­hunting and predation from introduced predators, especially rats.

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 ­over-­hunting, habitat loss and introduced predators.

Guadalupe Petrel Oceanodroma macrodactyla Bryant Oceanodroma leucorhoa macrodactyla W. E. Bryant 1887, p. 450 (Guadalupe Island) Specimens  Specimens are in Berkeley, California; Cambridge, Massachusetts; Chicago; Los Angeles; New York; Philadelphia; Pittsburgh; Providence, Rhode Island; San Francisco; Tring; and Washington, D. C. Eggs are in Tring. Status  Last seen in 1911 and presumed extinct. Range  Formerly bred on Guadalupe Island, west of Baja California, Mexico. Description  23cm (9in). General coloration sooty black above glossed with plumbeous grey, chocolate brown below with a moderately forked tail; ­uppertail-­coverts white with darker median stripe extending to lateral coverts; bill and feet black. Indistinguishable in the field from Leach’s Petrel and could only be identified by larger size and paler underside to the wings.

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 ­over-­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 64

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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.

Alaotra Grebe  Tachybaptus rufolavatus

Delacour

Podiceps rufolavatus Delacour 1932, p. 2 (Lake Aloatra) Specimens  Specimens can be found in many major museums, including Antananarivo, Madagascar. Status  Extinct. Last seen in December 1982 when twelve were located on Lake Alaotra, with a further two seen near Andreba on Lake Alaotra in September 1985. Range  Madagascar; generally confined to Lake Alaotra and surrounding lakes. Description  25cm (10in). A rather small grebe with characteristic long, stout bill and short wings. Breeding plumage: crown and hind part of neck dark blackish with greenish gloss, throat and ­ear-­coverts pale cinnamon to white (lacking in African Little Grebe T. ruficollis capensis), with mottled dusky grey and rufous underparts; breast, flanks and abdomen darkest grey; back dark brown. Distinct pale yellow iris. Bill black with white tip.

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. They were usually found in pairs, sometimes in association with Little Grebe, a species that colonised Madagascar as recently as 1945 (Milon 1946). It 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 lakes.

Atitlán Grebe Podilymbus gigas Griscom Podilymbus gigas Griscom 1929, p. 5 (Panajachel, Lake Atitlán, Guatemala) Specimens  Specimens can be found in most major museums. Status  Extinct. Last seen in 1986. Range  Confined to Lake Atitlán, Guatemala. Description  36–40cm (14–16in). The Atitlán Grebe was similar to the Pied-billed Grebe Podilymbus podiceps in colour and form, differing only in having a darker head, neck and belly, but was almost twice the weight, and as the wing hardly differed from that of the smaller species, it was probably flightless.

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, who remarked (1932): 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 65

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Atitlán Grebe Podilymbus gigas

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 enormously 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 population 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 hybridised 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. 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 Atitlán Grebe is believed to have fed mainly on crabs, populations of which were disturbed by the introduction of Largemouth Bass Micropterus salmoides to the lake in 1960.

Colombian Grebe (Tota Grebe)  Podiceps andinus

(Meyer de Schauensee)

Colymbus caspicus andinus Meyer de Schauensee 1959, p. 55 (Lake Tota, Boyacá, Colombia) Podiceps nigricollis andinus Storer 1979, p. 153 Specimens  At least 18 skins have been preserved, most of which are held at the University of Bogotá, Colombia. Status  Extinct. Last recorded February 1977. Range  Formerly occurred in the temperate zone lakes of the Eastern Andes in Boyacá and Cundinamarca, Colombia. Subsequently confined to Lake Tota, Boyacá. Description  30cm (12in). Breeding: generally dark brownish-black above with fore-neck and sides of body chestnut; centre of underparts white; crown, nape and throat blackish-brown; tufts of plumes on head-sides golden-buff to chestnut; bill black; iris red; legs and feet dark. Non-breeding plumes absent; overall duller, more brownish-grey above, underparts entirely white.

When the Tota 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 66

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Threskiornithidae

seen in 1972, and the last definite sighting was of three birds in February 1977. An International Council for Bird Preservation expedition to Lake Tota in 1982 failed to find it (Varty et al. 1986). 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. Habits The Tota 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.

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.

Réunion Ibis (Réunion Solitaire)  Threskiornis solitarius

(Sélys-­Longchamps)

Apterornis solitarius Sélys-Longchamps, 1848, p. 293 Threskiornis solitarius ­Mourer-­Chauviré et al. 1999, pp. 1–38 Specimens  Subfossil remains are in Réunion. Status  Extinct. Last recorded c.1708. Range  Réunion, Mascarene Islands.

Until the discovery of subfossil remains, it was generally believed that a species of white dodo (see p. 137), 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 2003). 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 visted Réunion in 1672–73, 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. 137) 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). The Réunion Ibis appears to have been a montane species, perhaps restricted in its distribution. It gradually retreated toward the remote mountaintops as Réunion became more populated, and the last definite account refers to them in 1708 (Cheke & Hume 2008). ­Over-­hunting and the introduction of predators such as rats and cats are probable reasons for its extinction.

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Réunion Ibis Threskiornis solitarius

Extinction The introduction of predators such as rats and 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 mountains. Their food is only worms and filth, taken on or in the soil. Réunion’s endemic ibis became extinct soon after.

Molokai Flightless Ibis  Apteribis glenos

Olson & Wetmore

Apteribis glenos Olson & Wetmore 1976, p. 252 (Moomomi dunes, Molokai, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Range  Molokai, Hawaiian Islands.

The discovery of a flightless ibis on the island of Molokai represented another remarkable addition to the Hawaiian Islands avifauna. It was first described in 1976 from subfossil remains discovered on the 68

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Moomomi dunes by natural historian Joan Aidem, and from fragmentary material found on Maui (Olson & Wetmore 1976). Since then, better material has been discovered on both islands, enabling Olson & James (1991) to separate the two island populations into discrete species. Undescribed subfossil remains have also been found on Lanai, but not on the islands of Oahu and Kaua’i to the west, or on Hawaii to the east. Therefore, Olson & James suggest that the ibises were Maui Nui endemics (Maui Nui being the name used collectively for the central Hawaiian islands of Maui, Molokai, Lanai and Kaho’olawe); these islands were connected until at least the ­mid-­Pleistocene (Price & ­Elliott-­Fisk 2004). Molokai Flightless Ibis Remarkably, the Lanai subfossil specApteribis glenos imen still retained feathers; Carla Dove and Storrs Olson have compared these with modern ibis genera (Dove & Olson 2011), concluding that the Lanai bird (and all Hawaiian ibises) was closely related to (and probably derived from) the South American genus Eudocimus. This concurs with earlier DNA analysis (Fleischer & Mcintosh 2001). Furthermore, Dove & Olson were able to establish plumage coloration; the Lanai ibis at least was a mixture of brown (light and dark brown shades) and ­ivory-­beige or very light brown – most similar to a juvenile American White Ibis Eudocimus albus. The Molokai Flightless Ibis was a small bird resembling the New Zealand kiwi Apteryx sp., with extremely reduced wings and large, robust legs. The bill was long and strongly decurved, with a narrow, pointed tip. Olson & James (1991) suggest that the ibis may have been the main predator of a diverse terrestrial snail fauna, which once occurred throughout Maui Nui. Being flightless and probably easy to catch, the ibis disappeared shortly after the arrival of Polynesians and their commensal animals. The ibis would have also been susceptible to introduced pigs and dogs, and the Pacific Rat Rattus exulans was probably a dangerous predator of eggs and chicks.

Maui Flightless Ibis  Apteribis brevis

Olson & James

Apteribis brevis Olson & James 1991, p. 23 (Auwahi Cave, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Range  Maui, Hawaiian Islands.

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 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. 69

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Ardeidae (Herons, egrets and bitterns) The Ardeidae are a diverse group of generally aquatic birds, with long necks and legs. They have a cosmopolitan distribution, occupying all continents except for Antarctica. Their taxonomy is unresolved and the family has been divided into at least 20 genera. Some species have a restricted distribution, but in general are not considered threatened. A number of island endemics became more terrestrial, with corresponding reduction in ­wing-­size and more robust legs.

New Zealand Little Bittern  Ixobrychus minutus novaezelandiae

(Potts)

Ardeola Novae Zelandiae Potts 1871, p. 99 (Westland, South Island, New Zealand) Specimens  Specimens are in Canterbury, New Zealand and Tring. Status  Probably extinct. Not collected since the 19th century. Range  Known only from a single specimen from North Island, and fewer than 10 specimens from South Island. Description  36cm (14in). General coloration bright rufous on face, neck and upper breast; below white; superciliary stripe and throat white; crown and hindneck black; back, rump and tail black; ­wing-­coverts buff; secondaries white; primaries black; bill ­orange-­yellow; legs yellowish.

The New Zealand Little Bittern was first obtained at Touranga 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. Colensoe (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 ultimately 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 ­salt-­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, studiously 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 water . . . It is stated that the Little Bittern is so quiet in his habits that it will remain still when approached, and almost suffer itself to be taken by the hand. Buller continued: I had an opportunity of observing one of these birds in a state of captivity at Hokitika, in May 1871. It had been taken only a few days before, and was already comparatively tame. Its usual posture was one of repose, with the head drawn in and resting on the shoulders; but when alarmed or excited it assumed a very different attitude, 70

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standing bolt upright, with the body resting, as it were, on the tarsal joints, these being brought close together, the neck stretched upwards to its full extent and perfectly rigid, the beak elevated, and the eyes directed outwards and downwards in such a way as to command a full view in front without having to move the head. On being turned out in the verandah it ran quickly and spread its wings, but did not make any attempt to fly, and after a short interval endeavoured to ­re-­enter its cage. It evinced great alarm on the appearance of a cat, stretching up its neck and emitting a peculiar snapping cry. At other times when molested it uttered a cry not unlike that of the Kingfisher, although not so loud. Mr. McNee, to whom the bird belonged, informed me that he could not get it to eat anything till he produced a dish of water containing some ‘mudfish’, which it instantly seized and devoured. This singular fish . . . is very common in the Hokitika district, being found in all the creeks and ­surface-­ponds in the woods which here cover the whole face of the country. The remarkable part of their history is that on the pools becoming dry these mudfish burrow into the moist soil or clay, often to the depth of two feet, remaining there for an indefinite time, or till the return of rainy weather . . . I think it highly probable that the mudfish constitutes the chief food of the Little Bittern; for as many of the surface pools are never dry, there would be no difficulty in finding a supply all the year round. I may mention also that Mr. Clapcott’s bird, while alive in his possession, was fed on worms, and that it would only take them when placed in a saucer or other vessel containing water. Oliver (1955) believed the bird to be a distinct form deserving specific status, it being much larger than the Australian Little Bittern Ixobrychus minutus. It apparently occurred mainly on the wooded margins of salt water lagoons on the sea shore – which would be most unusual for a little bittern. Some authorities (e.g. Hancock & Elliot 1978) state that there was never a resident New Zealand population, and that the known specimens were wandering individuals from the Australian population. However, after examination of subfossil remains, Worthy & Holdaway (2002) affirm that the leg bones were more robust than the Australian Little Bittern; thus its status as a probably extinct New Zealand endemic can be maintained.

Mauritius Night Heron  Nycticorax mauritianus

(Newton & Gadow)

Butorides mauritianus Newton & Gadow 1893, pp. 289–90 (Mauritius) Specimens  Subfossil remains are in Cambridge, England. Status  Extinct since c.1700. Known only from subfossil remains. Range  Mauritius, Mascarene Islands.

The Mauritius Night Heron was placed in Butorides by Newton & Gadow, but Rothschild (1907), 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. 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 predators such as cats and rats were primarily responsible.

Mauritius Night Heron Nycticorax mauritianus

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Rodrigues Night Heron  Nycticorax megacephalus

(Milne-Edwards)

Ardea megacephala Milne-Edwards 1874a, p. 10 (Rodrigues) Nycticorax megacephala Günther & Newton 1879, pp. 434–7, pl. 41, fig. G Specimens  Specimens are in Cambridge, England; London; Paris; and Tring. Status  Extinct. Known only from subfossil bones and travellers’ accounts. Range  Rodrigues, Mascarene Islands.

Both Leguat (1708) and Tafforet (1726) 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 1726 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 would follow his party around to take advantage of disturbed lizards. Nothing more is known about it. 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.

Réunion Night Heron  Nycticorax duboisi

(Rothschild)

Megaphoyx Hachisuka 1937c, p. 148 Ardea duboisi Rothschild 1907a, p. 114 Nycticorax duboisi ­Mourer-­Chauviré et al. 1999, p.5 Specimens  Subfossil remains are in Paris and Réunion. Status  Extinct. Known only from Dubois’s description and subfossil remains. Extinct c.1700. Range  Réunion, Mascarene Islands.

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 any extant species. Hachisuka (1937) said little more except to place the bird into his own (now invalid) genus, Megaphoyx.

Bonin Night Heron  Nycticorax caledonicus crassirostris

Vigors

Nycticorax crassirostris Vigors 1839, p. 27 (Peel Island or Chichijima, Bonin Islands) Specimens  Specimens are in Tring. Status  Last seen in 1889 and presumed extinct. Range  Formerly occurred on Chichijima (Peel Island) and ­Nakōdo-­jima in the Bonin Islands. Description  61cm (24 inches). Crown black, ornamented with two or three long white plumes extending over hind neck; back cinnamon-brown, underparts white. Bill black, legs and feet orange. Differed from nominate N. c. caledonicus by a straighter, more robust bill.

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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 in 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.

Niue Night Heron  Nycticorax kalavikai

Steadman et al.

Nycticorax kalavikai Steadman et al. 2000, p.169 (Anakuli, Niue) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Niue, ­south-­west Pacific.

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. 74) did on Bermuda (Olson & Wingate 2006). The reasons for its extinction are probably ­over-­hunting by humans and predation by introduced mammals.

Ascension Night Heron  Nycticorax olsoni

Bourne et al.

Nycticorax olsoni Bourne et al. 2003, p. 46 (Chapel Grotto, Ascension) Specimens  Subfossil remains are in Tring and Washington, D. C. Status  Extinct, known only from subfossil remains. May have survived until the 16th century. Range  Ascension Island, South Atlantic.

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. 100) 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. 73

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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).

Bermuda Night Heron  Nyctanassa carcinocatactes

Olson & Wingate

Nyctanassa carcinocatactes Olson & Wingate 2006, p. 329 (Devil’s Sinkhole, Hamilton Parish, Bermuda) Specimens  Subfossil remains are in New York and Washington, D. C. Status  Extinct. Known only from subfossil remains. Probably became extinct in the early 16th century. Range  Bermuda.

The Bermuda Night Heron was almost certainly a derivative of the ­Yellow-­crowned Night Heron N. violacea, 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 terrestrial, probably succumbed to introduced predators and perhaps a shortage of prey.

Sulidae (Boobies) The boobies generally occurs 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). 74

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Ua Huka Booby  Papasula abbotti costelloi

Steadman et al.

Papasula abbotti costelloi Steadman et al. 1988, p. 490 (Hanamiai Site, Tahuata, Marquesas) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known only from subfossil remains. Date of extinction around c.1200. Range  Ua Huka and Tahuata, Marquesas Islands, South Pacific.

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 was taken on Assumption Island, in the ­south-­west Indian Ocean (Ridgway 1893), subfossil remains have been found on the Mascarene Islands of Mauritius and Rodrigues, and Steadman et al. (1988) have collected subfossil remains from Tikopia, an island in the Solomon Islands in the Pacific. The Assumption population disappeared as a direct result of habitat destruction during guano excavation (Stoddart 1981), and it was probably hunted to extinction on the Mascarenes (Cheke & Hume 2008). The relict Christmas Island birds are endangered. The Ua Huka subspecies of Abbott’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). The subfossil remains of the Ua Huka Booby are from archaeological midden sites, which indicate that it was regularly hunted by Polynesians. Abbott’s Booby today nests only in tall trees, so the Ua Huka Booby would have also been badly affected by habitat destruction. It probably disappeared rapidly after human colonisation of the Marquesas Islands.

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.

Mascarene Reed Cormorant  Phalacrocorax (africanus) nanus

(Newton & Gadow)

Plotus nanus Newton & Gadow 1893, pp. 288–9, pl. 34, figs. 1–5 (Mare aux Songes, Mauritius) Specimens  Subfossil remains are in Cambridge, England and London. Status  Extinct. Last recorded in the early 17th century. Range  Mauritius and Réunion, Mascarene Islands.

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. Habits The Mascarene Reed Cormorant was never recorded alive on Mauritius, but on Réunion it was mentioned by two competent observers; Dubois in 1672–73 (Dubois 1675) 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.

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Spectacled Cormorant (Pallas’s Cormorant)  Phalacrocorax perspicillatus  Pallas Phalacrocorax perspicillatus Pallas 1811, p. 305 (based on Steller’s MS) (Bering Island) Specimens  Specimens are in Dresden; Helsinki; Leiden; St Petersburg; and Tring. Subfossil remains are in St. Petersberg and Washington, D. C. Status  Extinct since about 1852. Range  Formerly occurred on Bering Island (Ostrov Beringa), Commander Island (Komandorskiye Ostrova) and probably on other islands in the region. The islet of Riy Kamen (Aji Kamen) may have been its last refuge. Reports of midden material from Aimchitka, Aleutians, are erroneous (see Olson 2005). Description  97cm (38in). A very large, bronze-green, ­double-­crested cormorant weighing 6.0–6.8kg (12–14 pounds), with small wings and steel-blue reflections on the neck. Head and upper neck with long hair-like, pale yellowish feathers. Tail black. Forehead and skin around bill naked, variably coloured white, vermillion and blue. Bare skin spectacles surrounding eyes white. Sexual dimorphism present, female smaller and lacking the crests and spectacles.

The explorer Georg Steller, who visited Bering Island on Bering’s ­ill-­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. Steller Georg Wilhelm Steller was born in Windsheim, near Nuremberg, on 10 March 1709, and studied at Wittenberg, Leipzig, Jena and Halle Universities. 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 expedition 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

Spectacled Cormorant Phalacrocorax perspicillatus

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considered 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 progressively 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, 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 trappers 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, ­long-­necked birds with a long, sharp bill and a wide distribution, occurring 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. 77

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Levant Darter  Anhinga rufa chantrei

(Oustalet)

Plotus chandrei Oustalet 1882, p. 7 (Lake of Antioch) Specimens  Specimens are in Leiden and Paris. Status  Probably extinct. Last recorded in the 1990s. Range  Amik Gölü, ­south-­central Turkey, and marshes of the Lower Tigris and Euphrates, southern Iraq and Iran. Description  80cm (31.5in). Male mainly glossy black with white streaking; thin white lateral neck stripe; chin, throat, underside of neck, and undersides rufous; maxilla ­bluish-­grey, mandible pinkish; legs and feet blackish. Females and immature browner.

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 ­north-­east Israel, with lesser numbers reaching the coastal plain of ­north-­west Israel. The population 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 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.

Accipitridae (Eagles, kites and hawks) The Accipitridae comprises a diverse range of birds of prey, including eagles, buzzards, sparrowhawks and harriers. They have colonised a number of remote island archipelagos. All have suffered from persecution and habitat destruction, and many species are on the verge of extinction.

Haast’s Eagle  Harpagornis moorei

Haast

Harpagornis moorei Haast 1872, p. 192 (Glenmark, New Zealand) Harpagornis assimilis Haast 1873, p. 62 (Glenmark, New Zealand) Specimens  Subfossil remains are in Canterbury and Wellington, New Zealand. Status  Known only from bones, date of extinction not known. Range  South Island, and doubtfully from North Island, New Zealand.

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 the 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 supposed 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 measurements of bones of moorei and assimilis (the two species originally described by Haast), which differed only in size, and concluded that Haast’s Eagle was sexually dimorphic. The two sexes, therefore, were different in their locomotion and predatory potential, and may have taken different prey; such sexual dimorphism (and partitioning of prey resources) is relatively common in raptors. Holdaway suggested that Harpagornis was closely related to Aquila, which has now proved to be the case (Lerner et al. 2005). DNA analysis (Bunce et al. 2005) has shown that Haast’s Eagle formed a clade with two small forest eagles, Aquila 78

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Haast’s Eagle Harpagornis moorei with Euryapteryx moa.

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) H. moorei (Braithwaite 1992) make Haast’s Eagle 30% heavier than the world’s largest living eagle, the Harpy Eagle Harpia harpyja from South America. Furthermore, it evolved to become the world’s largest eagle in less than 0.7–1.8 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 of the known 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 as 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. 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 1873), may well be referable to Harpagornis, indicating 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 eylesi (see p. 80). 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.

Hawaiian Harrier  Circus dossenus

Olson & James

Circus dossenus Olson & James, 1991, p. 65 (Molokai, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from bones. Date of extinction unknown. Range  Molokai and Oahu, Hawaiian Islands.

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 deforestation by the Polynesians. The Hawaiian Harrier was probably a ­bird-­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. 79

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Eyles’s Harrier  Circus eylesi

(Forbes)

Circus teauteensis Forbes 1892d, p. 186 (Te Aute swamp, North Island, New Zealand) Circus hamiltoni Forbes 1892d, p. 186. (North Island, New Zealand) Circus eylesi Scarlett 1953, p. 245 (Pyramid Valley swamp, South Island, New Zealand) Specimens  Specimens are in Canterbury and Wellington, New Zealand; and London. Status  Known only from bones. Date of extinction not known, but possibly 1800s. Range  North and South Islands, New Zealand.

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 ­re-­examined to confirm this, while Tennyson & Martinson (2006) use teauteensis. 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. Regardless of it specific status, 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 (see p. 79).

Powerful Goshawk  Accipiter efficax

Balouet & Olson

Accipiter efficax Balouet & Olson 1989, p.6 (Pindai Cave, Nepoui Peninsula, west coast of New Caledonia) Specimens  Subfossil remains are in Paris. Status  Extinct. Known only from subfossil material. Reasons for extinction unclear. Range  New Caledonia, Melanesia.

Accipter hawks are generally ­forest-­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.

Gracile Goshawk  Accipiter quartus

Balouet & Olson

Accipiter quartus Balouet & Olson 1989, p.7 (Pindai Cave, Nepoui Peninsula, west coast of New Caledonia) Specimens  Subfossil remains are in Paris. Status  Extinct. Known only from subfossil material. Reasons for extinction unclear. Range  New Caledonia, Melanesia.

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 comparatively recently (Balouet & Olson 1989).

Car Nicobar Sparrowhawk  Accipiter butleri butleri

(Gurney)

Astur butleri Gurney 1898, p. 27 (Car Nicobar Island) Specimens  Specimens are in Tring. Status  Possibly extinct, but data deficient. Last recorded with certainty in 1901; unconfirmed report in 1977. Range  Car Nicobar, northern Nicobar Islands, India.

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Accipitridae Description  29.5cm (11.7in). Upperparts pale ­blue-­grey, palest on head, primaries and secondaries darker; throat and chin whitish; breast pink finely barred pale rufous and white; thighs white; tail grey with indistinct subterminal blackish bar; wing quills grey, primaries darker mottled white basally on inner webs; iris bright orange; cere, legs and feet yellow. Juvenile extremely distinctive; upperparts dark chestnut, darker on nape, feathers with dark centres producing a mottled appearance; primaries and secondaries indistinctly barred; breast and sides buff, heavily streaked dark ­reddish-­brown; belly whitish with rufous blotching; tail ­cinnamon-­red with a terminal and central dark brown bar; bill black, base bluish; cere pale green; iris ­greyish-­white; legs and feet pale yellow.

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 ­rufous-­crowned young birds probably bred in March or April, and at the same time both adult cocks killed were in a state of breeding.

Bermuda Hawk  Bermuteo avivorus

Olson

Bermuteo avivorus Olson 2008b, p. 132 (Bermuda) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil bones. May have survived to historic times. Range  Bermuda, western Atlantic.

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.

Malagasy Crowned Eagle  Stephanoaetus mahery

Goodman

Stephanoaetus mahery Goodman 1994a, pp. 421–428 (Madagascar) Specimens  Subfossil remains are in London; Paris; and Antanarivo. Status  Extinct. Known only from subfossil remains. Possibly survived into comparatively recent times. Range  Madagascar.

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. 81

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Malagasy Crowned Eagle Stephanoaetus mahery catching a Ruffed Lemur Varecia variegata

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 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 questioned by Csermely 1996). The Malagasy Crowned Eagle was probably a forest inhabitant, and the top predator in Madagascar. The destruction of forests and hunting of prey species by the Malagasy would have played a major role in the eagle’s disappearance, and it appears to have become extinct perhaps as recently as the 16th century.

Falconidae (Falcons and caracaras) The falcons are streamlined birds with long, pointed wings and slender bodies. Some species are ­high-­speed aerial hunters. Unlike the falcons, the caracaras are generally ­slow-­flying terrestrial birds that scavenge for food. The falcons and caracaras have colonised remote island archipelagos, where a number of species have evolved. They have been persecuted throughout their ranges, and island forms in particular have suffered from anthropogenic changes. 82

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Guadalupe Caracara  Polyborus lutosus

Ridgway

Polyborus lutosus Ridgway 1876a, p. 459 (Guadalupe Island) Specimens  About forty skins have been preserved, together with a few chicks, skeletons, and a few doubtful eggs. Specimens are in Cambridge and Lancaster, Massachusetts; Chicago; New York; Pittsburgh; Providence, Rhode Island; San Diego, California; and Washington, D. C. in the USA: Tring and Norwich in the UK. Status  Extinct. Last seen December 1, 1900, when specimens were collected by Rollo Beck. Range  Guadalupe Island, off Baja California, Mexico. Description  54cm (22in). Closely related to the Crested Caracara Polyborus cheriway, differing in having less white on upper breast, belly barred like the breast, not black. Adult P. lutosus brownish above, becoming blacker on crown, primaries, shoulders and tip of tail. Most of the feathers irregularly barred brown and white. Young birds dark brown above, paler brown, with buff or whitish streaks through middle of feathers; wings and tail more brown, less grey.

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 caracaras, 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 (1876), 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. Salvation denied by greed The Guadalupe Caracara could have been saved by captive breeding. In 1897, Daniel Cleveland, a member of the San Diego Natural History Society, recalled a fisherman who had brought six live birds from Guadalupe to San Diego, and others seem to have arrived there as well. That they could survive in captivity was less important than the rarity of the bird and its associated monetary value (Cleveland, in Abbott 1933): About 1897, a fisherman captured six of the living eagles on the island, and brought them to San Diego, where he kept them in a large cage, and where, as I remember, all of them died within about a month. The man professed to want to sell these birds, but demanded $150 each for them, and refused all offers for less. I was very anxious to purchase a pair of them, but could not afford to pay the price demanded. So the man’s greed resulted in our failure to rear some of these birds in captivity and in his own loss from his failure to sell the birds. 83

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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.

Puerto Rican Caracara  Polyborus latebrosus

Wetmore

Polyborus latebrosus Wetmore 1920, p. 77 (Cueva Toraño, Puerto Rico) Specimens  The type is in New York. Status  Extinct; known only from subfossil remains. Date of extinction unknown. Range  Puerto Rico, West Indies.

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.

Bahamas Caracara  Polyborus creightoni

Brodkorb

Caracara creightoni Brodkorb, 1959, p. 353 (New Providence) Polyborus plancus Olson 1976a, p. 363 Specimens  Subfossil remains are in Cuba and Florida. Status  Extinct. Known only from bones. Date of extinction unknown. Range  Bahamas and Cuba.

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.

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Jamaican Caracara Caracara tellustris

Jamaican Caracara  Caracara tellustris

Olson

Caracara tellustris Olson 2008a, p. 267 (Skeleton Cave, Jackson’s Bay cave system, south coast of Portland Ridge, Clerndon Parish, Jamaica) Specimens  Subfossil remains are in New York. Status  Extinct. Known only from subfossil bones. May have survived to comparatively recent times. Range  Jamaica.

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 terrestrial 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.

Réunion Kestrel  Falco duboisi

Cowles

Falco duboisi Cowles 1994, p. 89 (Réunion) Specimens  Subfossil remains are in Lyon; Paris; and Réunion. Status  Extinct. Known only from subfossil remains. Last recorded in 1672–73. Range  Réunion, Mascarene Islands.

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. 85

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Cuban Kestrel Falco kurochkini

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.

Cuban Kestrel  Falco kurochkini

Suárez & Olson

Falco kurochkini Suárez & Olson 2001a, p. 35 (Cuba, West Indies) Specimens  Subfossil remains are in Cuba and Washington, D. C. Status  Extinct. Known only from subfossil remains. May have survived into the historic period. Range  Cuba.

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 intermediate 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 potential 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.

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.

Moroccan Bustard  Ardeotis arabs lynesi

(Bannerman)

Choriotis arabs lynesi Bannerman 1930b, p.432 (Camp Boulhaut [Sidi Ben Sliman], ­south-­east of Rabat, Morocco) Specimens  Specimens are in Tring. Status  Probably extinct. Last recorded in 1993.

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Otididae Range  Western Morocco. Description  Male 100cm (39.5in); female 75cm (29.5in). Back, wings and central tail ­sandy-­brown with fine black markings; some broad black and white subterminal bands on outer rectrices; head, neck, and breast finely barred ­grey-­brown on whitish background; crown black with grey centre with slight crest; superciliary stripe white; belly white; bill light greyish horn, darker at tip; iris, legs and feet yellowish. The Moroccan form differs from the nominate in being darker with more coarse vermiculation above, and darker barring on neck.

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 ­north-­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 ­semi-­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 relationships 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. 36), 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 endangered due to predation from introduced mammals and deforestation.

Lowland Kagu  Rhynochetos orarius

Balouet & Olson

Rhynochetos orarius Balouet & Olson 1989, p. 28 (Pindai Cave, Nepoui Peninsula, western New Caledonia) Specimens  Subfossil remains are in Paris. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  New Caledonia, Melanesia.

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 extinct lowland form has only been found in areas at or near the coast, the two 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.

Lowland Kagu Rhynochetos orarius

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Rallidae (Rails and coots) The Rallidae comprises a diverse range of birds that have colonised every continent except Antarctica, and almost all islands around the world. They occupy a range of habitats, with a preference for marshland and dense forests. Terrestrial species are generally cryptic and extremely difficult to observe, and a large number of island species have become flightless. The Rallidae have suffered more recent extinctions than any other bird group. The rail radiation probably incorporated almost all habitable islands across the Pacific (see Steadman 2006a). Each population gradually evolved unique characters, and over millennia many became flightless endemics, preventing contact with rails from the other island groups. Steadman estimated the number of rails that might have once occurred in the Pacific, and suggested a very conservative 442 to nearly 1,600, which may still not reflect the true diversity of the region’s rails before the arrival of humans.

Goldman’s Yellow Rail Coturnicops noveboracensis goldmani

(Nelson)

Porzana goldmani Nelson 1904, p. 151 (Lerma, Mexico) Specimens  Specimens are in Ithaca and Washington, D. C. Status  Possibly extinct. Last recorded July 22, 1964. Range  Marshes of the Rio Lerma Valley, central Mexico. Description  19cm (7.5in). The Yellow Rail is a generally ­tawny-­yellow bird below, upperparts blackish with ­tawny-­buff streaks and narrow white bars; broad ­blackish-­brown eye stripe and crown; superciliary stripe ­tawny-­buff; flanks and ­undertail-­coverts ­blackish-­brown, irregularly barred buff and white; bill yellowish; iris red or yellowish; legs and feet brown or greenish. Goldman’s Yellow Rail was larger, darker on the upperparts, with blacker crown and with black ­mid-­feather stripes on nape; underparts more cinnamon. Juvenile lacked white spotting on cheeks and malar areas and upper flanks.

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 vegetation 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.

Jamaican Black Rail Laterallus jamaicensis jamaicensis

(Gmelin)

Rallus jamaicensis Gmelin 1789, p. 718 (Jamaica) Specimens  Specimens of the Jamaican subspecies are in New York; Philadelphia; and Washington D. C. Status  Now extinct, disappearing from Jamaica and Puerto Rico in the 1870s. Jamaican birds known from three skins. Considered by some to be conspecific with the mainland United States population of L. j. jamaicensis. Range  Jamaica, Puerto Rico and a few reports from Cuba.

The little known Black Rail Laterallus jamaicensis occurs in scattered populations in eastern and western North America, and on the coasts of Peru and Chile (Ripley 1977; Taylor & van Perlo 1998). Birds from eastern North America winter in the Caribbean today, but a resident population living ­year-­round in Jamaica and Puerto Rico has now died out. Gosse (1847) stated that the bird was formerly common and a resident species on Jamaica, but by the turn of the century, it had become very rare, possibly extinct, as a result of the introduction of the predatory mongoose. 88

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Doubt has been expressed as to whether these populations were actually subspecifically distinct from those of North America (Taylor & van Perlo 1998). Coale (1923) examined the three known examples collected on Jamaica and concluded that they differed from the mainland form in having a longer and heavier bill, longer middle toe and in being overall duller with the white markings on the upperparts smaller.

Fiji ­Bar-­winged Rail Nesoclopeus poeciloptera

Hartlaub

Rallina poeciloptera Hartlaub 1866b, p. 171 (Viti Levu, Fiji) Nesoclopeus poeciloptera Peters 1932, p. 348 Specimens  Specimens are in Cambridge, Massachusetts; Berlin; Boston; Leiden; Liverpool; Tring; and New York. Status  Last seen with certainty in 1890 and now probably extinct. Range  Occurred on Viti Levu and Ovalau in the Fijian Islands. Description  35cm (14in). A large, probably flightless rail with a powerful bill. Upperparts brown; underparts dark grey; whitish patch on the middle of throat; wings with bright chestnut and black bars; bill orange and yellow; iris light brown; legs and feet yellow. Male smaller than female.

Many naturalists, including members of the Whitney South Seas Expedition (an extensive series of explorations 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 in 1890 (Reichenow 1891a), but Holyoak (1979) claimed to have seen a bird in 1973. Introduced mongooses were 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.

Wake Island Rail Gallirallus wakensis

(Rothschild)

Hypotaenidia wakensis Rothschild 1903, p. 78 (Wake Island) Specimens  Specimens are in New York; Tring; and Washington D. C. Status  Last seen in 1945, now extinct. Range  Wake and Wilkes Islands, north Pacific. Description  22cm (9in). In seasonally worn birds dark ashy brown above, fading to earthy brown, ­ear-­coverts and lores dark brown; pale superciliary line, chin and upper throat whitish; neck grey, rest of underside ­ash-­brown, narrow white bar on breast, two or three more on the flanks and abdomen; tail uniform brown. Wings and tail soft, ends not clearly defined, pattern indistinct. In fresh plumage overall brighter, upperparts brighter chestnut; barring and white bar on breast more distinctive; wings and tail sharp and clearly defined barring. Juvenile dark ­blackish-­brown, paler below. Bill, legs and feet brown. Iris red.

Wake, Wilkes and Peele Islands are a group of low, scrub-covered atolls in the North Pacific. The Wake Island Rail was flightless with robust legs, but was a derivative of the volant Gallirallus philippensis group (Taylor & van Perlo 1998; Olson 1999b). 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). Japanese troops were stationed on the islands during World War II, and were forced to survive on a starvation diet, so any likely food source would have been consumed. The bird had evidently gone by 1945. Bailey (in Greenway 1967) reported the bird extinct on Wake, but thought it might survive on Wilkes and Peele, which was unlikely. 89

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Peele is separated from Wake by a 100m channel, which appears to have restricted the rail to Wake and Wilkes. Habits Little was recorded of the rail’s habits, but most of what we know was recorded by Wetmore in 1923 (in Ripley 1977). It was omnivorous, feeding on molluscs, insects and worms and exposing them in loose soil with sideward thrusts of its bill. Courtship and copulation took place during late July, with nesting occurring in ­mid-­August. The bird made a low chattering sound, and at close quarters a gentle cluck. It was timid, and would scamper into cover at the least disturbance.

Macquarie Island Banded Rail  Gallirallus philippensis macquariensis

Hutton

Rallus macquariensis Hutton 1879, p. 455 (Macquarie Island) Specimens  Specimens are in Dunedin, New Zealand and Tring. Status  Extinct since about 1880. Range  Formerly resident on Macquarie Island, south-­west Pacific. Description  30–33cm (12–13in). The Macquarie Island population was a dark form of the Buff-banded Rail Gallirallus philippensis, somewhat resembling immatures of the New Zealand race G. p. assimilis. Head chestnut on top with fine blackish streaks; eyebrow white with chestnut ­eye-­stripe extending onto sides of neck; cheeks, chin and throat greyish; upperparts ­olive-­brown, patterned with blackish patches and white spots. Underparts greyish, breast and flanks with fine black and white barring, a broad rufous band across breast; belly white. Bill dark pink; legs and feet greyish; iris red.

The remote and inhospitable subantarctic island of Macquarie was once home to a rail and a parakeet (see p. 170). 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 ­over-­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).

Dieffenbach’s Rail (Moeriki)  Gallirallus dieffenbachii

(Gray)

Rallus dieffenbachii Gray 1843, p. 197 (Chatham Islands) Gallirallus dieffenbachii Olson 1973b, p. 396 Specimens  Specimens are in Tring and Bremen. Status  Date of extinction not known. Range  Formerly inhabited Chatham Island and perhaps Pitt Island, New Zealand. Description  28cm (11in). A ­medium-­sized rail, resembling ­Buff-­banded Rail Gallirallus philippensis, except throat banded black and white; breast barred irregularly brown and black; bill stouter and more decurved at tip. Crown and hind neck ­reddish-­brown with chestnut stripe extending from bill through eye to hind neck; grey line above eye; chin and throat pale grey; band of black feathers, tipped with white, separated throat and breast; belly barred black and white; upper back barred black and brown, lower back indistinctly barred darker brown; bill ­yellowish-­brown with darker tip; legs and feet pale brown; iris ­reddish-­brown.

The type of this species was collected in 1840. Ernst Dieffenbach (who first discovered it) stated that it was: . . . 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. 90

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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). Confusion with Chatham Islands Rail For a number of years Dieffenbach’s Rail was confused with the other extinct Chatham rail G. modestus (see below), which was considered to be the immature form of G. dieffenbachii (Buller 1873). Hutton (1874) recognised them as different and created a new genus, Cabalus, for G. modestus, based on modifications associated with flightlessness. Andrews (1896b) confirmed that they differed in a number of characters: plumage of G. modestus much looser than in G. dieffenbachii owing to the almost complete absence of barbules; beak relatively much longer and more slender in G. modestus, and less sharply decurved at the extremity; and sternum in G. modestus more reduced, keel almost obsolete. DNA analysis has shown that Dieffenbach’s Rail is quite distinct from its probable ancestor, the ­Buff-­banded Rail G. philippensis, despite morphological similarities (Trewick 1997b), and it is not a subspecies of it as was once thought (e.g. Ripley 1977; Fuller 2000).

Chatham Islands Rail  Gallirallus modestus

(Hutton)

Rallus (Cabalus) modestus Hutton 1872, p. 247 (Chatham Islands) Ocydromus pygmaeus Forbes 1892b, p. 252, nomen nudum. Specimens  Known from about 26 skins, one egg and a number of subfossil bones. Skins are in Cambridge, England; Cambridge, Massachusetts; Auckland, Christchurch and Wellington, New Zealand; Liverpool; New York, Paris; Philadelphia; Pittsburgh; and Tring. Subfossil remains are in London. The unique egg is in Tring. Status  Now extinct. The last known specimen was collected between 1895 and 1900. Range  Known from Mangare Island, Chatham Island and Pitt Islands, Chatham Group, New Zealand. Description  22cm (8.75in). Male olivaceous brown, bases of feathers plumbeous; feathers of breast slightly tipped with pale fulvous; those on abdomen and flanks with two narrow bars of same colour; throat dark grey, each feather tipped with brown; quills soft brown, first three faintly barred with reddish fulvous, fourth and fifth longest; tail soft, short and brown; iris light brown; bill and legs light brown (Hutton 1872). Female smaller, more faintly barred. Juvenile uniformly brownish black. Egg creamy white with faint pale reddish and purplish spots (Oliver 1930), but the only known specimen is virtually white with the faint suggestion of a few, very tiny dots (see Walters 1994).

The first specimens of this rail were collected by H. H. Travers on Mangare Island (Hutton 1872, 1874). They consisted of an adult and an immature skin, plus an individual preserved whole in spirits. Mangare 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 earlier date, so the Chatham Island 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 extermination, 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 Mangare, 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 G. modestus may have been derived from a more recent colonisation, evolving into a nocturnal, almost ­kiwi-­like niche, thus remaining in sympatry with its more generalist congener G. dieffenbachii. However, Holdaway et al. (2001) question this scenario. Olson (1973b) suggested that the smaller G. modestus disappeared from 91

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Chatham Island and Pitt Island due to competition with the larger G. dieffenbachii, but it appears that they might only have been sympatric on Mangare Island (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 Mangare, as there is no sand on Mangare 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.

Tahiti ­Red-­billed Rail  Gallirallus pacificus

Gmelin

Pacific Rail. Latham 1785, p. 235 (Otaheite) Rallus pacificus Gmelin 1789, p. 717 (Tahiti). Based on Latham. Status  Extinct. No specimens survive; known only from the description of J. R. Forster, and Georg Forster’s plate. Range  Tahiti and possibly (see below) nearby Mehetia, South Pacific. 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 ­blood-­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 ­Red-­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 Red-billed Rail Gallirallus pacificus

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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 Gallirallus philippensis ecaudatus (this race sometimes occurs in lists of extinct birds, but it is actually extant). The matter was clarified by Lysaght (1953); Plate 128 of Forster depicts Gallirallus pacificus. In Rothschild (1907a), there is a figure of the Tahiti ­Red-­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 bird is faithfully reproduced, except the legs are coloured red instead of being ­flesh-­coloured (see Fuller 2000).

Rota Rail  Gallirallus temptatus

Kirchman & Steadman

Gallirallus temptatus Kirchman & Steadman 2006a, p. 8 (Route 100 archaeological site, Rota, Northern Mariana Islands) Specimens  Subfossil remains are in Florida. Status  Extinct. Known from archaeological subfossil remains only. Date of extinction unknown. Range  Rota, Mariana Islands, ­south-­west Pacific.

The Rota Rail was a ­medium-­sized species, overall smaller than the ­Buff-­banded Rail G. philippensis and Barred Rail G. torquatus, from which many of the Pacific island populations were almost certainly derived (Kirchman & Steadman 2006a). It was flightless, but the legs did not exhibit the usual robustness, a character normally associated with flightlessness.

Aguiguan Rail  Gallirallus pisonii

Kirchman & Steadman

Gallirallus pisonii Kirchman & Steadman 2006a, p. 10 (Pisonia Rockshelter archaeological site, Aguigan. Northern Mariana Islands) Specimens  Subfossil remains are in Florida. Status  Extinct. Known from archaeological subfossil remains only. Date of extinction unknown. Range  Aguiguan, Mariana Islands, ­south-­west Pacific.

The Aguiguan Rail was a small to ­medium-­sized species, smaller than the Rota Rail (above), but similar to the ­Buff-­banded Rail G. 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.

Tinian Rail  Gallirallus pendiculentus

Kirchman & Steadman

Gallirallus pendiculentus Kirchman & Steadman 2006a, p. 18 (Unai Chulu archaeological site, Tinian, Northern Mariana Islands) Specimens  Subfossil remains are in Florida. Status  Extinct. Known from archaeological subfossil remains only. Date of extinction unknown. Range  Tinian, Mariana Islands, ­south-­west Pacific.

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The Tinian Rail was a ­medium-­sized species, smaller than ­Buff-­banded Rail G. philippensis, with disproportionately shorter wing elements. It was therefore likely to have been flightless, but less so than other flightless species derived from G. philippensis, such as Wake Island Rail G. wakensis (see p. 89) (Kirchman & Steadman 2006a).

New Ireland Rail  Gallirallus ernstmayri

Kirchman & Steadman

Gallirallus ernstmayri Kirchman & Steadman 2006a, p. 24 (Balof archaeological site, New Ireland) Specimens  Subfossil remains are in Florida. Status  Extinct. Known from archaeological subfossil remains only. Date of extinction unknown. Range  New Ireland, Bismarck Archipelago, ­south-­west Pacific.

This was a large rail, equal in size to the New Britain Rail Gallirallus insignis and Solomon Rail G. woodfordi (Kirchman & Steadman 2006a), with a total length of approximately 33cm. Subfossil remains are fragmentary but the bird appears to have been flightless. Along with G. insignis, the New Ireland Rail is only the second known from the Bismarck Islands, but future palaeontological work will doubtless reveal more species.

Tahuata Rail  Gallirallus roletti

Kirchman & Steadman

Gallirallus roletti Kirchman & Steadman 2007, p. 149 (Hanamiai archaeological site, Tahuata, Marquesas Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known from subfossil remains only. Date of extinction unknown. Range  Tahuata, Marquesas Islands, South Pacific.

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 ­medium-­sized, probably flightless species, equal in size to ­Buff-­banded Rail G. 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.

Ua Huka Rail  Gallirallus gracilitibia

Kirchman & Steadman

Gallirallus gracilitibia Kirchman & Steadman 2007, p. 156 (Hane Dune archaeological site, Ua Huka, Marquesas Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known from subfossil remains only. Date of extinction unknown. Range  Ua Huka, Marquesas Islands, South Pacific.

This was a small to ­medium-­sized flightless rail, equal in size to ­Buff-­banded Rail G. 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.

Nuku Hiva Rail  Gallirallus epulare

Kirchman & Steadman

Gallirallus epulare Kirchman & Steadman 2007, p. 158 (Ha’atuatua, archaeological site, Nuku Hiva, Marquesas Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known from subfossil remains only. Date of extinction unknown. Range  Nuku Hiva, Marquesas Islands, South Pacific.

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. 94

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Huahine Rail  Gallirallus storrsolsoni

Kirchman & Steadman

Gallirallus storrsolsoni Kirchman & Steadman 2006b, p. 285 (Fa’ahia archaeological site, Huahine, Society Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known from very well preserved skeletons. Date of extinction perhaps 700 years ago, soon after the arrival of Polynesians. Range  Huahine, Society Islands, South Pacific.

The Huahine Rail was a ­medium-­sized flightless rail, about the size of the volant ­Buff-­banded Rail G. 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 Gallirallus rails known to occur in the Society Islands (the other was the extinct Tahiti ­Red-­billed Rail G. pacificus, which is only known from a painting; see p. 92). 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 introduced animals.

Niue Rail  Gallirallus huiatua

Steadman et al.

Gallirallus huiatua Steadman et al. 2000, p. 175 (Anakuli, Niue) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Range  Niue, South Pacific.

The Niue Rail was a ­medium-­sized flightless rail, about the size of volant ­Buff-­banded Rail G. philippensis, but it differed in a number of characters which suggests that it may not have been part of the same radiation as other Gallirallus 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 colonists, it would have succumbed quickly to ­over-­hunting and anthropogenic changes to the environment.

Mangaia Rail (Ripley’s Rail)  Gallirallus ripleyi

Steadman

Gallirallus ripleyi Steadman 1987, p. 42 (Te Rua Rere Cave, Tava’enga District, Mangaia, Cook Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Range  Mangaia, Cook Islands, South Pacific.

The Mangaia Rail was a small species, about the size of Wake Island Rail G. wakensis (see p. 89) at 22–25cm in total length. It had a slender bill and strong, robust tarsometatarsi (Steadman 1987). It occurred sympatrically on Mangaia with another extinct, flightless rail, the Mangaia Crake Porzana rua (see p. 103), 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). ­Over-­hunting, habitat alteration and introduction of mammalian predators appear to be the causes of its extinction.

Eua Rail  Gallirallus vekamatolu

Kirchman & Steadman

Gallirallus vekamatolu Kirchman & Steadman 2005, p, 468 (Eua, Tonga) Specimens  Subfossil remains are in Florida. Status  Extinct. Known only from subfossil remains. Range  Eua, Tonga.

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 ­Buff-­banded Rail G. philippensis (Kirchman & Steadman 2006). The wing elements do not superficially indicate that this species was flightless, being 95

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approximately the same size as the ­Buff-­banded Rail, but more robust. However, correlations between other ­similar-­sized but extant flightless species (e.g. Guam Rail Gallirallus 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 Rail  Gallirallus steadmani

Worthy & Bollt

Gallirallus steadmani Worthy & Bollt 2011, p. 77 (Atiahara archaeological site, Tubuai, Austral Islands) Specimens  Subfossil remains are in Sydney. Status  Extinct. Known from extremely well preserved skeletons and numerous subfossil remains. Date of extinction perhaps c.700 years ago, soon after the arrival of Polynesians. Range  Tabuai, Austral Islands, South Pacific.

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 ­Buff-­banded Rail G. 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 G. 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.

Norfolk Island Rail  Gallirallus sp. Specimens  Subfossil remains are in Norfolk Island. Status  Extinct. Known from subfossil remains and perhaps a contemporary painting. Range  Norfolk Island.

Meredith (1991), followed by Holdaway & Anderson (2001), reported the presence on Norfolk Island of a flightless Gallirallus rail, closely related to but larger than ­Buff-­banded Rail G. 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 Porzana tabuensis (the latter, or a race of it, may well be the Dark Rail Porzana 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 Gallirallus species. It appears that the rails were victims of ­over-­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 Gallirallus 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  Tricholimnas lafresnayanus

(Verreaux & Des Murs)

Gallirallus lafresnayanus Verreaux & Des Murs 1860, p. 437. (New Caledonia) 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.

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The few known specimens of New 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 New Caledonian Wood Rail Tricholimnas lafresnayanus (1967) believed originated with the hydrographer Monsieur Bouquet de la Gry. 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 inhabited the same type of woodland as the Kagu Rhynochetos jubatus. 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 motionless, 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.

Cleland’s Rail  Lewinia pectoralis clelandii

(Mathews)

Rallus pectoralis clelandii Mathews 1911, p. 189 (Oceania = New South Wales) Specimens  Specimens are in New York and Tring. Status  Extinct. Not seen since 1932. Range  Extreme ­south-­west Australia, from the Margaret River to King George’s Sound. Description  23–27cm (9–10.5in). A ­medium-­sized, stocky rail with a comparatively small head, a moderately long bill, and a short tail. Larger than nominate L. p. pectoralis, bill longer and deeper; breast brighter grey, with reduced ­olive-­buff feather tips. Top of head and neck chestnut or rufous with heavy black streaking; cheeks and throat grey; rest of upperparts streaked black and ­olive-­brown; throat and breast grey; belly, flanks and ­undertail-­coverts black with white or buff barring; wings dark brown with narrow white and ­olive-­brown barring above, ­brownish-­grey below; bill pinkish red with dusky tip; legs and feet grey with pinkish tinge; iris brown, ­reddish-­brown or red.

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 97

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1932 (Whittell 1933). 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.

Flores Rail  Lewinia pectoralis exsul

(Hartert)

Hypotaenidia brachypus exsul Hartert 1898, p. 50 (Mangari District, Flores, Lesser Sunda Islands, Indonesia) Specimens  The type is in New York. Status  Possibly extinct. Last recorded in 1958–59. Range  South and west Flores, Lesser Sunda Islands, Indonesia. Description  23–27cm (9–10.5in). Differed from nominate L. p. pectoralis in having a dark ­rufous-­chestnut crown with indistinct black streaking towards nape, lacking black central stripe; back more olive, less ­rufous-­tinged; feathers of back and rump with 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 is known from only four specimens, and was last seen in 1958–59 (Paynter 1963). Recent searches for it have been unsuccessful and it might now be extinct (Taylor & van Perlo 1998). It appears to have disappeared as a result of habitat destruction. Nothing is known of its habits.

Assumption ­White-­throated Rail  Dryolimnas cuvieri abbotti

(Ridgway)

Rougetius abbotti Ridgway 1894, p. 74 (Assumption Island) Specimens  Specimens are in Tring and New York; a skeleton is in Tring. Status  Extinct. Disappeared between 1908 and 1937. Range  Assumption Island, ­south-­west Indian Ocean. Description  The nominate is greenish olive above; chestnut red on the head and breast; throat white; flanks dusky with pale barring. Assumption race is smaller, paler and more greyish olive on upperparts; black streaks on the back narrower, white on the chin and throat more extensive; white barring on the lower abdomen broader and coarser.

The ­White-­throated Rail Dryolimnas cuvieri is an Indian Ocean island endemic; the nominate, volant subspecies 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 ­semi-­flightless species once occurred on Mauritius (JPH, in prep.). 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 (1909), 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 98

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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 developed 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 (1909) 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 ­Vesey-­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. Habits Other than Nicholl’s (1909) 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.

Réunion Rail  Dryolimnas augusti 

Mourer-­Chauviré et al.

Dryolimnas augusti ­Mourer-­Chauviré et al. 1999, p. 21 (Réunion) Specimens  Subfossil remains are in Réunion. Status  Extinct since c.1670. Known from subfossil remains from one individual and perhaps one contemporary account. Range  Réunion, Mascarene Islands.

The Réunion Rail was a large, robust bird, much larger than ­White-­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 ­over-­hunting and predation by introduced mammals, particularly rats and cats.

St Helena Rail  Atlantisia podarces

(Wetmore)

Aphanocrex podarces Wetmore 1963, pp. 379–81, pl. 9 (St Helena) Atlantisia podarces Olson 1973a, pp. 13–17 Specimens  Subfossil remains are in Washington D.C. Status  Known only from bones. Disappeared during the early 16th century. Range  St Helena.

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 99

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forests and grasslands in the lowlands, but systematic destruction of the habitat began soon after human colonisation (Ashmole & Ashmole 2000). This resulted in the loss of almost all of the forest, leaving the majority of the island barren and arid. Forest clearance with fire, ­over-­grazing by introduced goats, and predation by pigs, rats, cats and mice was responsible for the extinction of almost all of the native birds. The only surviving species is an endemic plover, the Wirebird Charadrius sanctaehelenae, which has actually benefited from the removal of the forests, although it remains endangered. The island was once a major breeding site for thousands of seabirds, nearly all of which have now disappeared. The St Helena Rail was one of the largest rails and was comparable in size to a New Zealand Weka Gallirallus australis, only more slender (Olson 1973a). It was closely related to the Inaccessible Island Rail Atlantisia rogersi, a small black, flightless rail which survives on Inaccessible Island in the Tristan da Cunha group. The St Helena Rail occurred sympatrically with the much smaller St Helena Crake Porzana astrictocarpus (see p. 100). Ashmole & Ashmole (2000) suggest that of the three islands inhabited by Atlantisia rails (Tristan, Ascension and St Helena), St Helena is the largest and most ecologically diverse, this allowing two species of rail to coexist. Despite being flightless, the St Helena Rail had comparatively long wings and long claws, which Olson (1977b) suggests were adaptations for clambering and fluttering up the steep valley walls on St Helena. It probably foraged among the island’s abundant seabird colonies for food, feeding on their eggs and chicks, but it may also have predated the island’s abundant snail and invertebrate fauna (Olson 1973a). The St Helena Rail, being flightless and presumably easy to catch, probably disappeared within a few decades of humans arriving on the island. Its eggs and young would have been extremely vulnerable to introduced mammalian predators.

Ascension Rail  Atlantisia elpenor

Olson

Fowle. Peter Mundy in Temple & Anstey 1936, p. 83 Crecopsis sp. Fisher & Peterson 1964, pp. 241, 273 Atlantisia elpenor Olson 1973a, pp. 3–13 (Ascension Island) Specimens  Subfossil remains are in London and Washington D.C. Status  Known from subfossil bones and a traveller’s account. Became extinct sometime after 1656. Range  Ascension Island, South Atlantic.

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

Ascension Rail Atlantisia elpenor

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bird species could survive on such a place, but an endemic rail and a night heron (see p. 73) certainly did. Mundy landed on Ascension on June 7, 1656 (Temple & Anstey 1936), and made the only known description of the Ascension Rail from life: . . . much bigger than our sterlings [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 themselves a little with their wings . . . shortt billed, cloven footed, thatt can neither fly nor swymme. It was more than ordinary dainety meatt, relishing like a roasting pigge. Subfossil remains of the rail have been found in the 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 ­medium-­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 exterminated virtually every bird on the island. However, the rail had probably long disappeared before their arrival.

Jamaican Wood Rail (Uniform Crake) Amaurolimnas concolor concolor

(Gosse)

Rallus concolor Gosse 1847, p. 369 (Jamaica) Specimens  Specimens are in Cambridge, Massachusetts; Chicago; New York; Tring; and Washington, D. C. Status  Extinct since about 1890. Other races range widely but irregularly over Central America and northern South America. Range  Formerly inhabited Jamaica. Description  25cm (10in). A ­rufous-­chestnut rail, paler than the mainland races A. c. guatemalensis and A. c. castaneus. Rufous above, more chestnut on scapulars and ­wing-­coverts; primaries sepia above, edged lighter, dusky brown below; inner secondaries chestnut like back; tail chestnut, crown and nape dusky brown; slightly rufescent on forehead; lores and face dull rufous, browner on the ­ear-­coverts; under surface ­vinous-­chestnut, paler on chin and centre of breast and abdomen; ­underwing-­coverts and axillaries dusky brown; bill yellowish green; feet purplish; iris vermilion.

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 predator, 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, beautifully 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.

Kosrae Island Crake  Porzana monasa

(Kittlitz)

Rallus monasa Kittlitz 1858, p. 30 (Kosrae Island, Carolines) Ortygometra tabuensis Finsch 1880, p. 297, 307 Kittlitzia monasa Hartlaub 1893, p. 391

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Extinct Birds Aphanolimnas monasa Sharpe 1892, p. 20 Porzana monasa Ripley 1977, p. 231 Specimens  Specimens are in St Petersburg. Status  Extinct. Not recorded since December 1827–January 1828. Range  Formerly confined to Kosrae (formerly known as Kusaie) Island in the Caroline Islands, Western Pacific. Description   18cm (7in). A small, flightless black rail. General coloration black with bluish-grey reflections; chin whitish fading to brown on middle of throat, primaries and tail browner; white spots on the ­undertail-­coverts; inner ­wing-­coverts brownish, spotted with white; iris and feet red; bill black at tip, lighter at base.

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). 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 examination of the specimens, concluded it was another dark endemic Porzana, derived from Spotless Crake P. tabuensis (Olson 1973b) or Henderson Crake P. 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.

Iwo Jima Crake (Mamijiro Kuina)  Porzana cinerea brevipes

Ingram

Porzana cinerea brevipes Ingram 1911, p. 21 (Sulphur Island, Bonins = Iwo Jima) Specimens  Specimens are in Amsterdam; Tokyo; and Tring: eggs are in Tring. Status  Last collected in 1911, but possibly survived until 1925. Range  Formerly occurred on Naka Iwo Jima (Sulphur Island), and doubtfully Minami Iwo Jima (San Augustino), Volcano group, Bonin Islands. Description  17–21cm (7–8in). Similar in appearance to the nominate Ashy Crake Porzana cinerea cinerea, which occurs on Indonesia and the Malay Peninsula (Taylor & van Pedro 1998), but with the lower flanks and ­undertail-­coverts more rufous, the rump and lower back ruddier, the bill deeper at the base, and the tarsus shorter. Head ­blackish-­brown, neck and upper back brown, rump and lower back feathers with dark centres and ­brownish-­olive edges; underparts white, washed with pale brown; bill yellowish olive with red base; iris red; feet and legs olive.

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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.

Mangaia Crake  Porzana rua

Steadman

Porzana rua Steadman 1987, p. 28 (Te Rua Rere Cave, Tava’enga District, Mangaia, Cook Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Range  Mangaia, Cook Islands, South Pacific.

This was a ­medium-­sized, ­robust-­limbed crake, larger in size than the Spotless Crake P. tabuensis, but with more reduced wing elements than any other Porzana, except the extinct St Helena Crake P. astrictocarpus (Steadman 1987). It was most similar in size and skeletal morphology to the extant Henderson Island Crake P. atra, which was probably derived from Baillon’s Crake P. pusilla (Olson 1973a); it is likely that the Mangaia Crake had the same ancestral origins (Steadman 1987). The Mangaia Crake probably disappeared shortly after the arrival of Polynesians, but a Porzana species might have survived on Mangaia into the historic period (see p. 331).

Western Hawaiian Crake  Porzana sandwichensis sandwichensis

(Gmelin)

Sandwich Rail. Latham 1785, p. 236 Rallus sandwichensis Gmelin 1789, p. 717 (Hawaiian Islands = Island of Hawaii, on the authority of Stresemann 1950) Pennula wilsoni Finsch 1898, p. 77 (Hawaiian Islands) Porzana sandwichensis sandwichensis Olson 1999b, p. 5 Specimens  Specimens are in New York and Leiden; subfossil remains are in Washington, D. C. Status  Extinct, the last specimen taken in 1864. Range  Leeward side of Hawaii. Description  14cm (5.5in). Top of head and ­ear-­coverts brown with faint greyish tinge; back, wing and ­tail-­coverts brown; feathers of mantle and back with dark brown centres and light edges giving a mottled appearance; throat whitish, shading to deep ­reddish-­brown on breast, to greyish chocolate on belly flanks and ­undertail-­coverts. Bill dusky ash or black, edges of mandible yellowish; legs ­dusky-­flesh or possibly orange.

Considerable confusion existed about the number of species of Hawaiian crakes and their nomenclature (see Olson 1999b). Two geographical subspecies are now recognised, P. s. sandwichensis for the lighter leeward population, and P. 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) 103

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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 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’.

Eastern Hawaiian Crake  Porzana sandwichensis millsi

Dole

? Dusky Rail. Latham 1785, p. 237 ? Rallus obscurus Gmelin 1789, p. 718. Based on Latham ? Moho. Pease 1862, p. 145. Pennula millei Dole 1878, p. 14. (Hawaii) (usually amended to millsi, following Newton 1889, p. 5) Pennula ecaudata Wilson & Evans 1894, p. 171 Porzana sandwichensis millsi Olson 1999b, p. 5 Specimens  Specimens are in Cambridge, England; Honolulu; Tring; and Vienna. Status  Extinct. Last recorded in the 1860s. Range  Windward side of Hawaii. Description  14cm (5.5in). Differed from P. s. sandwichensis in having back dark reddish chocolate, lower back with darker centres to the feathers, and lacking the mottled appearance of the back feathers.

Five skins of P. 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. 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 ornithologist, 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 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!). He never found one, but described the bird as follows: PENNULA MILLEI [A printer’s error for millsi] Moho. Not previously described. 6 and a half in. long. Bill three quarters of an in. long, black, straight, sides compressed, curved at tip. Tail not visible. Wings rudimentary, hidden in the long, loose, hairy feathers. Plumage dark, dull brown, ashy under the throat; feathers loose, hairy, long. Lower part of tibia naked. Legs long, set far back. Toes 3 front, 1 back. Habitat, uplands of Hawaii. Nearly extinct. Specimen in Mills’ Coll. I feel confident that this remarkable bird belongs to the Rallidae, but am unable to fix its place more definitely. It is the only bird which the natives call Moho, which word is nearly synonymous with the New Zealand word Moa, which is their name for a gigantic wingless bird of that country. Regarding it as a new genus I have taken the liberty of naming it as above, gladly thereby recognising Mr. Mills’ valuable services in preserving specimens of this bird, and giving others opportunities of studying it. Habits It 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. 104

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Ziegler’s Crake  Porzana ziegleri

Olson & James

Porzana ziegleri Olson & James 1991, p. 51 (Barbers Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known from subfossil remains only. Date of extinction unknown. Range  Oahu, Hawaiian Islands.

Only two Porzana crake species are known historically from the Hawaiian Islands, the Hawaiian Crake P. sandwichensis (see above) and Laysan Crake P. palmeri (see p. 106). The discovery of subfossil remains has now shown that each of the main Hawaiian Islands had at least one species of rail, or even up to three on the larger islands (Olson & James 1991), and more await description. All 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 Porzana 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 P. keplerorum (see p. 106) on Maui.

Molokai Crake  Porzana menehune

Olson & James

Porzana menehune Olson & James 1991, p. 53 (Ilio Point, Molokai, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known from subfossil remains only. Date of extinction unknown, but this or another ­as-­yet undiscovered species may have survived until recent times. Range  Molokai, Hawaiian Islands.

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  Porzana keplerorum

Olson & James

Porzana keplerorum Olson & James 1991, p. 55 (Auwahi Cave, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known from subfossil remains only. Date of extinction unknown. Range  Maui, Hawaiian Islands.

Keplers’ Crake appears to be descended from the same ancestral species as Molokai P. menehune and Ziegler’s P. ziegleri crakes (Olson & James 1991). It was a small crake, larger than P. menehune but smaller than P. ziegleri. It also differed from the latter in having a shorter, wider bill and more reduced wing elements.

Ralphs’ Crake  Porzana ralphorum

Olson & James

Porzana ralphorum Olson & James 1991, p. 59 (Barbers Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known from subfossil remains only. Date of extinction unknown. Range  Oahu, Hawaiian Islands.

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 P. 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.

Severns’s Crake  Porzana severnsi

Olson & James

Porzana severnsi Olson & James 1991, p. 60 (Auwahi Cave, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known from subfossil remains only. Date of extinction unknown. Range  Maui, Hawaiian Islands.

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).

Laysan Crake (Laysan Rail)  Porzana palmeri

(Frohawk)

Porzanula palmeri Frohawk 1892, p. 247 (Laysan Island) Specimens  Specimens are in Berlin; Cambridge, England; Honolulu; Massachusetts; Chicago; Tring; New York; Philadelphia; and Washington, D. C. Status  Extinct. Last recorded in June 1944, or possibly June 1945. Range  Endemic to Laysan Island but subsequently introduced to other islands in the Hawaiian chain. Description  15cm (5.9in). In general coloration it was a paler version of Baillon’s Crake Porzana pusilla, a closely related species from which it was almost certainly derived (Olson 1973b). Forehead, sides of face, throat, breast and supercilium ash grey; back, wings and top of the head ­sandy-­brown with centres of feathers ­blackish-­brown; scapulars and flanks ­sandy-­brown; pale white barring on lower belly and ­undertail-­coverts; bill, feet and legs yellowish ­olive-­green; iris ruby red; sexes alike; immature birds with underside pale brown; downy young all black; legs and feet black; bill yellow.

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. 106

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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 by 1936. 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, a US landing craft accidentally introduced rats to Midway which 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; and see Olson 1999b 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. 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, fearless 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 clambered 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. 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’.

St Helena Crake  Porzana astrictocarpus

Olson

Porzana astrictocarpus Olson 1973a, p. 17 (St Helena) Specimens  Subfossil remains are in London and Washington, D. C. Status  Became extinct after 1502. Known only from subfossil bones. Range  St Helena.

This flightless crake, along with the Laysan Crake Porzana palmeri (see p. 106), was considered by Olson (1973a) to be derived from Baillon’s Crake P. pusilla. It had evolved for a long period in isolation, so had characters not present in Baillon’s. Both P. palmeri and P. astrictocarpus had equally reduced wing elements, while the latter had less robust pelvis and leg elements. More importantly, P. 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. 99), and may have similarly utilised seabird colonies for food (Ashmole & Ashmole 2000), perhaps feeding on regurgitated 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). 107

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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.

Mauritius Red Rail  Aphanapteryx bonasia

Sélys-Longchamps

Aphanapteryx bonasia Sélys-Longchamps 1848, p. 292 Didus broeckii Schlegel 1854, p. 256 (based on Van den Broeck’s drawing) Pezophaps broeckii Schlegel 1873a, p. 4 Aphanapteryx imperialis Frauenfeld 1868, p. 6 tab. 2 (based on a vellum painting by George Hoefnagel of c.1610 in the library founded by Emperor Francis I of Austria). Specimens  Almost complete associated skeleton in Mauritius; various subfossil bones are in London; Paris; and Cambridge, England. Status  Extinct. Known from bones, and from contemporary paintings and drawings of varying quality. Range  Mauritius, Mascarene Islands. Description  We can be fairly certain about the morphology of the Mauritius Red Rail. Based on the available evidence, it was ­chicken-­sized, with a long decurved bill, had reddish ­hair-­like plumage with a nape crest; the wings, being tiny, were hidden under the plumage; and it had extremely robust legs and feet. There was also sexual dimorphism in body size, a common phenomenon in Rallidae (Olson 1977b).

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 (1648), 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 (1638) 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.

Mauritius Red Rail Aphanapteryx bonasia

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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 (1650) in 1638 described ‘red hens’ that could be caught with red cloth, while Mundy (in Temple 1919) in 1638 also described this phenomenon, but refers to the bird as a ‘Mauritius Henne, of a yellowish Wheaten Coullour’. Marshall (in Khan, 1927), 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 rendition, 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 individuals; the red rail painting remains the only known colour depiction. A Red Rail is also supposedly illustrated 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 associated 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 introduction 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 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 1937). 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. 109

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Rodrigues Rail (Grey Rail, Leguat’s Rail) Erythromachus leguati

Milne-Edwards

Gelinotes. Leguat 1708, p. 71 Erythromachus leguati Milne-Edwards 1874a, p. 6, pls. 11–12 (based on Leguat) Aphanapteryx leguati Günther & Newton, 1879, p. 431 Specimens  Subfossil remains are in Cambridge, England; Paris; London; and Rodrigues. Status  Extinct. Known from bones and from contemporary accounts. Range  Rodrigues, Mascarene Islands

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 1626, who gave more details (from A. Newton’s 1875 translation): 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 excellent 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 severe deforestation by tortoise hunters from 1735, who burned off the forest to collect giant tortoises Cylindraspis sp. for their meat and oil (Cheke & Hume 2008), may have also contributed significantly to its demise. Subfossil remains Milne-­Edwards (1874b; 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  Nesotrochis debooyi

Wetmore

Nesotrochis debooyi Wetmore 1918, p. 516, pl. 82 (St. Thomas) Specimens  Subfossil remains are in Washington, D. C. Status  Known only from bones. Date of extinction unknown. Range  Formerly occurred on Puerto Rico, St Thomas, St John, St Croix and possibly Virgin Gorda in the Virgin Islands.

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 110

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been found in archaeological middens, which suggest that it must have been an important food item for the island’s ­pre-­Columbian inhabitants. ­Over-­hunting was probably the main reason for its extinction.

Haiti Rail  Nesotrochis steganinos

Olson

Nesotrochis steganinos Olson 1974, p. 439–450 (Hispaniola) Specimens  Subfossil remains are in Washington, D. C. Status  Known only from bones. Date of extinction unknown. Range  Haiti.

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 considerable size variation in the specimens.

Picapica Rail  Nesotrochis picapicensis

(Fischer & Stephan)

Fulica picapicensis Fischer & Stephan 1971, p. 593–607 (Cuba) Specimens  Subfossil remains are in Cuba. Status  Known only from bones. Date of extinction unknown. Range  Cuba.

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.

New Zealand ­Snipe-­rail  Capellirallus karamu

Falla

Capellirallus karamu Falla 1954, p. 241 (Karamu, Coonoor, Waitanguru, New Zealand) Specimens  Subfossil remains are in Auckland, Canterbury, and Otorohanga, New Zealand. Status  Known from subfossil remains including complete skeletons. Date of extinction unknown. Range  North Island, New Zealand.

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 adaptations for probing (Olson 1975c). The legs were particularly robust, and the ­snipe-­rail may have filled a ­kiwi-­like role, probing the ground for invertebrates.

Hawkins’s Rail (Mehonui)  Diaphorapteryx hawkinsi

(Forbes)

Aphanapteryx hawkinsi Forbes 1892b, p. 252 (Chatham Islands) Diaphorapteryx hawkinsi Forbes 1893a p. 253 Specimens  Subfossil remains are in London; Tring; and Wellington, New Zealand. Status  Known only from complete skeletons and individual bones, but may have survived into historic times. Range  Chatham Islands.

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 111

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Hawkins’s Rail in life. Both Dannefaerd and 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):

Hawkins’s Rail Diaphorapteryx hawkinsi

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. Shand (see 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.

Viti Levu Rail  Vittirallus watlingi

Worthy

Vittirallus watlingi Worthy 2004, p. 303 ­(Qara-­ni-­vokai, Voli Voli, Viti Levu, Fiji) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Date of extinction unknown. Range  Viti Levu, Fiji.

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 sympatric 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 ­over-­hunting, habitat destruction 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 particularly susceptible to fire.

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Hova Gallinule  Hovacrex roberti

(Andrews)

Tribonyx roberti Andrews 1897, p. 356, pl. 9, figs 4–7 (Sirabé, central Madagascar) Hovacrex Brodkorb 1965, p. 197 Specimens  Subfossil remains are in London. Status  Known only from subfossil remains. Date of extinction unknown. Range  Madagascar.

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.

Réunion Blue Gallinule (Oiseaux Bleu) Porphyrio caerulescens

(Sélys-Longchamps)

Oiseau bleu. Dubois 1671–72 Gros oiseau. Brown 1724, p. 313 Apterornis caerulescens Sélys-Longchamps 1848, p. 294 Porphyrio caerulescens Sharpe 1894 Status  Now extinct. Known only from travellers’ accounts. Range  Formerly occurred on Réunion, Mascarene Islands.

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 characters – 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. 113

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The bird had probably disappeared by the end of the 17th 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).

North Island Takahe (Moho)  Porphyrio mantelli

(Owen)

Notornis mantelli Owen 1848b, pp. 347, 366 (Waingongoro, New Zealand) Notornis parkeri Forbes 1892d, p. 185. Porphyrio mantelli mantelli Greenway 1967, p. 249. Porphyrio mantelli Trewick 1997b, p. 439. Specimens  Subfossil remains are in London; and Otorohanga and Wellington, New Zealand. Status  Extinct. Probably survived until the late 19th century. Known from subfossil remains and one account. Range  North Island, New Zealand.

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 1998). 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).

Lord Howe Gallinule  Porphyrio albus

(White)

? White Gallinule. Callam 1783 (fide Gray 1862b, p. 240) White Gallinule. Phillip 1789, p. 273 and plate Fulica alba White 1790, pp. 134–35, 238 and coloured plate. (Lord Howe Island) Gallinula alba Latham 1790, p. 768. (Norfolk Island). Based on Phillip Porphyrio stanleyi Rowley 1875, p. 37, pl. 9. Porphyrio raperi Mathews 1928, p. 6. Based on an unpublished drawing by George Raper. Specimens  Specimens in Vienna and Liverpool. Status  Extinct, last seen in c.1844. Known from two skins, subfossil remains and contemporary illustrations. Range  Formerly endemic to Lord Howe Island (Ball’s Pyramid off Lord Howe in error); occurrence on Norfolk Island is unsubstantiated. Description  36cm (14in). Predominantly white in the skins, but evidence from contemporary illustrations show that some of the adults were tinged with blue, especially on the wings; feathers of neck and breast with yellowish reflections, rest of body with blue; legs yellow; bill, shield and iris red. Gray (1862b) stated that young were entirely black, becoming ­bluish-­grey and finally pure white with maturity.

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The Lord Howe Gallinule is known from two skins, subfossil remains, and a number of illustrations and accounts (Forbes 1901; Iredale 1910; Holdaway & Anderson 2001). Although there was some variation in coloration, it was a large, predominantly white, flightless bird that was tame and easily killed. It must have been a spectacular bird. These colour differences were almost certainly a result of individual variation, with some birds being blue, some white, and some a mixture of the two, perhaps related to age and sex. 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).

Lord Howe Gallinule Porphyrio albus

Literature muddles There is no doubt that there was a population of largely white gallinules on Lord Howe Island that disappeared 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 (see Iredale 1910; Hindwood 1932). It is also uncertain whether the Lord Howe Gallinule was a distinct species. Purple Gallinule Porphyrio porphyrio of the Australasian race melanotus still occur on Lord Howe Island (Ripley 1977), so Hindwood’s (1965) assertion that the island population was white but that occasional blue birds arrived from Australia and interbred is indeed possible. However, the original resident population was flightless, or at least behaviourally so, and the two surviving skins exhibit soft rectrices, a character associated with flightlessness. Of the Porphryio species still to be found on Lord Howe Island, some show partial albinism in their plumage (Mayr 1941a), and Mayr thought it likely that the ­blue-­phase individuals might have survived after the white ones were exterminated, since the blue birds were less conspicuous. Early encounters The Lord Howe Gallinule was first described in 1789 by Arthur Phillip (1789), Governor of New South Wales: This beautiful bird greatly resembles the purple Gallinule [Porphyrio porphyrio] in shape and make, but is much superior in size, being as large as a dunghil[l] fowl. The from end of bill to that of the claws is two feet three inches. The bill is very stout, and the colour of it, the whole of the top of the head and the irides red; the sides of the head round the eyes are reddish, very thinly sprinkled with white feathers; the whole of the plumage is, without exception, white. The legs the colour of the bill. The species is pretty common on Lord Howe Island, Norfolk Island, and other places and is a very tame species. The other sex, supposed to be the male, is said to have some blue on the wings. White (1790) described the ease at which the gallinules could be caught: 115

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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 in the late 18th century, the Lord Howe Gallinule disappeared comparatively quickly, possibly before 1834, the year when the island was first settled (Hindwood 1940). Prior to this date, whalers and sealers used the island for supplies, and they quite possibly hunted the bird to extinction. Rats and cats did not arrive until later (Hindwood 1932). Foulis (in Hindwood 1940) was on the island from 1844 until 1847, and undertook an ornithological survey. He made no mention of a white gallinule, which must have been extinct by that time.

New Caledonia Gallinule  Porphyrio kukwiedei

Balouet & Olson

Porphyrio kukwiedei Balouet & Olson 1989, p. 23 (Pindai Cave, Nepoui Peninsula, west coast of New Caledonia) Specimens  Subfossil remains are in Paris. Status  Extinct. Known from subfossil remains only. Date of extinction unknown, but may have survived until the 1860s. Range  New Caledonia, Melanesia.

This was a very large, flightless gallinule, almost equalling the extant South Island Takahe Porphyrio hochstetteri 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 dimorphism, 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.

Huahine Gallinule  Porphyrio mcnabi

Kirchman & Steadman

Porphyrio mcnabi Steadman & Steadman 2006b, p. 290 (Fa’ahia archaeological site, Huahine, Society Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known from subfossil remains only. Date of extinction unknown, but soon after the arrival of Polynesians. Range  Huahine, Society Islands, South Pacific.

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 Gallinula storrsolsoni (see p. 95), 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.

Marquesas Islands Gallinule  Porphyrio paepae

Steadman

Porphyrio paepae Steadman 1988, p. 163 (Hanatekua Shelter, Hiva Oa, Marquesas Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known from subfossil remains and one possible contemporary painting. Date of extinction unknown, but may have survived until the early 20th century. Range  Hiva Oa and Tahuata, Marquesas Islands, South Pacific.

Until the discovery of the Huahine Gallinule (see above), the Marquesas species was the only known gallinule in eastern Polynesia (Steadman 1988); however, other species are waiting to be described. This 116

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suggests that like Gallirallus 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 proportions 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.

New Zealand Flightless Gallinule  Gallinula hodgeni

(Scarlett)

Rallus hodgeni Scarlett 1955b, p. 265 (Pyramid Valley, New Zealand) Pyramidia [sic] hodgeni Oliver 1955, p. 596 [Oliver proposed the new genus as Pyramida, but captioned the species Pyramidia. The earlier name has priority, and is clearly what he intended] ? Gallirallus hartreei Scarlett 1970, pp. 68–71 (New Zealand near Napier) Tribonyx hodgeni Olson 1975c, p 68 Gallinula hodgeni Olson 1975c, p 68 Specimens  Subfossil remains are in Canterbury and Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Range  North and South Islands, New Zealand.

The New Zealand Flightless Gallinule was originally known only from incomplete postcranial material, which made determination of its relationship problematic. The discovery of skull and jaws showed that the bird was closely related to the Australian gallinule group Tribonyx, which is now considered to be a subgenus of Gallinula (Olson 1975c, 1977b; Taylor & van Perlo 1998; Worthy & Holdaway 2002). The New Zealand Flightless Gallinule had reduced wings with short, heavy tarsi. The skull was most similar to the extant but equally flightless Tasmanian Native Hen Gallinula mortieri. The wings were even more reduced than in G. mortieri. Olson (1977b) hypothesised that the New Zealand species, along with the Australian flighted and flightless forms, were all descended from a volant ancestor. Being flightless and easy to catch, this species probably died out shortly after the arrival of humans on New Zealand.

Tristan Island Moorhen (Tristan Island Gallinule)  Gallinula nesiotis

Sclater

Gallinula nesiotis P. L. Sclater 1861c, p. 261, pl. 30 (Tristan da Cunha) Specimens  Only the type skin and two incomplete skeletons of Gallinula nesiotis are known with certainty to have been collected on Tristan da Cunha (Beintema 1972). All are in Tring. Status  Extinct. Last collected in 1861, and not seen since 1873. Range  Tristan da Cunha, South Atlantic. Description  25cm (10in). Virtually flightless, but otherwise rather like a small Common Moorhen Gallinula chloropus, with a black (not grey) neck and underparts, and more robust in the legs. G. nesiotis differed from Gough Moorhen G. comeri in being smaller and less robust in measurements of the skeleton. In G. nesiotis, head and neck black; back and ­wing-­coverts washed dark ­olive-­brown; wings, tail and underparts black with a few pale streaks on sides; ­undertail-­coverts white, with black feathers in middle; edge of outer primary white; bill and shield bright red with yellow tip; iris ­reddish-­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 ­north-­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 undisturbed 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 117

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(Taylor & van Perlo 1998). Olson (1973a) suggests that the islands were populated by two different colonisation 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 ­moor-­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.

Samoan Gallinule  Pareudiastes pacifica

Hartlaub & Finsch

Pareudiastes pacificus Hartlaub & Finsch 1871, p. 25, pl. 2.(Savaii, Samoan Islands) Specimens  Specimens are in Bremen; Hamburg; Liverpool; New York; and Tring; eggs in Tring. Status  Discovered in 1869, last officially recorded in 1873. Reports of its existence in August 1984 lack confirmation. Range  Formerly occurred on Savai’i Island in Samoa, and perhaps also on Upolu Island. 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 ­undertail-­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 ­reddish-­brown, tip darker brown; shield dirty yellowish; legs and feet reddish ­horn-­brown; claws ­horn-­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 bird may have survived until the end of the century. 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). Reports of sightings in 1984 (see Taylor & van Perlo 1998) hint that it may still survive, although there has been no further confirmation. 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). 118

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Rallidae

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 recovered 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.

Makira Gallinule (San Cristobal Gallinule)  Gallinula silvestris

(Mayr)

Edithornis silvestris Mayr 1933a, p.1–2 (San Cristobal, Solomon Islands) Pareudiastes silvestris Olson 1973b, p. 410 Gallinula sylvestris Taylor & van Perlo 1998, p. 489 Specimens  The type specimen is in New York. Status  Probably extinct. Last collected on December 4, 1929. Unconfirmed reports from the 1950s and 1970s. Range  Makira (formerly San Cristobal), Solomon Islands. Description  26cm (10in) Male dark bluish slate on breast, throat, sides of the head, crown and hind neck, becoming blackish on chin and cheeks; forehead covered by dark ­grey-­blue shield; scapulars, ­wing-­coverts and secondaries brownish black tinged with olive; lower flanks, ­underwing-­coverts, ­undertail-­coverts, thighs, lower back and rump dull brownish black; iris chocolate brown; bill and feet bright red.

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 apparently 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 disappearance, 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 119

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forest, native plantations and secondary growth, and had many brooks and creeks which cut into the mountainsides, 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.

Mascarene Coot  Fulica newtoni

Milne-Edwards

Waterhen. Dubois 1674 (Réunion) Fulica newtoni Milne-Edwards 1867, pp. 194–220, pls. 10–13 (Mauritius) Specimens  Subfossil remains are in Cambridge, England; London; and Réunion. Status  Extinct since c.1700. Known from subfossil remains and contemporary accounts. Range  Mauritius and Réunion, Mascarene Islands.

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. newtoni 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 ­Mourer-­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 ­water-­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 ­water-­hens on the Etang de St. Paul which was formerly covered in them. Both populations were presumably exterminated through ­over-­hunting and introduced predators.

Chatham Island Coot  Fulica chathamensis

Forbes

Fulica chathamensis Forbes 1892b, p. 252 (Chatham Islands) Palaeolimnas newtoni; error for chathamensis Forbes 1893c, p. 544 Palaeolimnas chathamensis ­Milne-­Edwards 1896, p. 130 Nesophaleris chathamensis Brodkorb & Dawson 1962, p. 268 Fulica chathamensis chathamensis Olson 1975c, pp. 63–79 Specimens  Subfossil remains are in London; and Canterbury and Wellington, New Zealand. Status  Extinct. Known only from subfossil bones. Date of extinction unknown. Range  Chatham Island, Chatham Group, New Zealand.

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 (1955) 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 120

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characters of the humerus and skull from those of New Zealand. Worthy & Holdaway (2002) ­re-­examined the subfossil remains, however, and concluded that the differences are in fact enough to warrant full species status. The Chatham Island Coot was extremely large and heavy, with reduced wings and robust legs. It differed from the New Zealand species 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 flightless 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.

New Zealand Coot  Fulica prisca

Hamilton

Fulica prisca Hamilton 1893, p. 98 (Castle Rocks) Palaeolimnas chathamensis Scarlett 1955a, p. 262 Nesophaleris prisca Brodkorb & Dawson 1962, p. 268 Fulica chathamensis prisca Olson 1975c, pp. 63–79 Specimens  Subfossil remains are in London; and Canterbury and Wellington, New Zealand. Status  Extinct. Known only from subfossil bones. Date of extinction unknown. Range  North and South Islands, New Zealand.

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 (1907) claimed that the New Zealand species was smaller than the Chatham bird, with less reduced wings, but both were in fact large and flightless. Olson (1977b) suggests that the coot was derived from Eurasian Coot Fulica atra, first in New Zealand whereupon it achieved great size, then subsequently colonising the Chathams; thus the two populations must have evolved flightlessness independently. The New Zealand Coot would have made easy prey for the Polynesian colonists, and their eggs and young would have been susceptible to introduced predators. It probably disappeared soon after permanent settlement of the islands.

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 ­over-­hunting and deforestation.

New Caledonia Buttonquail  Turnix varius novaecaledoniae

Ogilvie-­Grant

Turnix novaecaledoniae ­Ogilvie-­Grant 1889, p. 599 (New Caledonia) Turnix varius novaecaledoniae ­Ogilvie-­Grant 1893, p.552 (note) Specimens  The type specimen is in Tring; subfossil remains are in Paris. Status  Extinct. Known only from the type collected before 1889, and numerous subfossil remains. Range  New Caledonia, Melanesia. Description  19cm (7.5in). The Painted Buttonquail is ­greyish-­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 unthreatened throughout its range. A distinctive subspecies known only from the type specimen once occurred on 121

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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 introduced 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.

Tawi-­tawi Small Buttonquail  Turnix sylvatica suluensis

Mearns

Turnix suluensis Mearns 1905, p. 83 (Jolo, Sulu Island, Sulu Archipelago) Specimens  The type specimen is in Washington, D. C. Status  Possibly extinct. Not reliably recorded since the mid-20th century. Range  Jolo and ­Tawi-­tawi, Sulu Archipelago, Philippine Islands. Description  12cm (4.75in). Upperparts ­walnut-­brown, finely vermiculated with grey and black; sides of head and neck white speckled with clove brown; nape and mantle ­walnut-­brown, vermiculated with black, grey and pale cinnamon; lower back, rump, and ­uppertail-­coverts ­clove-­brown edged with grey and cinnamon; tail greyish drab, cross banded dusky; ­wings-­coverts rufous; primaries greyish; chin, upper throat, and malar region whitish, malar region speckled with ­blackish-­brown; upper neck and lower breast clay coloured, bordered by chain of black spots; lower breast and middle of belly whitish; sides of neck, chest and breast chestnut mixed with black and clay. Juvenile more spotted on underparts.

The Small Buttonquail Turnix sylvatica has an extremely wide distribution, occurring from ­north-­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 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 introduced predators. Habits Little is known about its habits. Other Small Buttonquail races occur in dense thickets, scrub, open grassland 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 tropical 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.

Canary Island ­(Meade-­Waldo’s) Oystercatcher Haematopus meadewaldoi

(Bannerman)

Haematopus niger meade-waldoi Bannerman 1913a, p. 33 (Jandia, Fuerteventura, eastern Canary Islands) Haematopus meadewaldoi Hockey 1982, p. 81 Specimens  Specimens are in Liverpool; Manchester; and Tring. Status  Not reliably recorded since June 1913, when the last specimen was taken, and now presumed extinct. Range  Eastern Canary Islands.

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Canary Island Oystercatcher Haematopus meadewaldoi

Description  40–45cm (16–17.5in). Very similar to the African Black Oystercatcher Haematopus moquini, but with smaller wings, more slender tarsi and a longer bill. Overall glossy black with white patch at base of inner webs of primaries; iris red, eye ring reddish orange; bill vermilion with lighter tip; legs and feet dark pink; nails ­ivory-­coloured.

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 lighthouse 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. ­Meade-­Waldo (1893) described the bird as uncommon, always seen in pairs and very tame – this habit may 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. 123

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Charadriidae (Plovers) Plovers are ­medium-­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.

Javan Lapwing  Vanellus macropterus

(Wagler)

Vanellus tricolor Horsfield’s 1821, p. 186 (Java) (not Charadrius tricolor of Vieillot 1818) Rogibyx tricolor Mathews 1913, p. 41 Charadrius macropterus Wagler 1827, p. 77 (with the merging of Rogibyx and Zonifer in Vanellus, Horsfield’s name becomes preoccupied by Vieillot’s; the next available name is that of Wagler) Specimens  Specimens are in Amsterdam; Leiden; Paris; Liverpool; New York; Washington D.C..; and Tring. Status  Probably extinct. Not reliably recorded since 1940. An unconfirmed report is given for 2002 (BirdLife International 2011), but no further information is available. Range  Formerly occurred in Java, and perhaps also on Sumatra and Timor. Description  27–29cm (10.5–11.5in). A generally dark bird and the only Vanellus to have black thighs. Head, belly patch and flight feathers deep black; upperparts, throat and upper breast grey; lower breast and upper belly dark brown; ­uppertail-­coverts, vent and basal half of tail white; rest of tail black with narrow white terminal band; bill black with ­flesh-­coloured base; wattles yellow; iris ­brownish-­black; legs and feet orange merging to yellow on tarsus.

Very little is known of this bird. Nearly all of the specimens 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. The 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, but considered local and uncommon, in the 1920s, and the last record was in Meleman on the south coast of east Java in 1939 (Bock 1958). This lapwing occurred in open areas near fresh water ponds, but probably also agricultural areas (King 1981). The reasons for its disappearance are unclear, but ­over-­hunting and habitat modification may be the primary causes. This latter explanation is favoured by Collar et al. (2001).

Javan Lapwing Vanellus macropterus

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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. Nothing else is known about this species.

Scolopacidae (Snipes and sandpipers) The Scolopacidae form a diverse group of wading birds. They exhibit a wide range of bill shapes and lengths, each designed for a specific feeding niche. These waders are sought after as game birds, and as a result some species have seriously declined in number. Island populations have also been severely impacted by introduced predators.

North Island Snipe (Little Barrier Island Snipe) Coenocorypha barrierensis

Oliver

Coenocorypha aucklandica barrierensis Oliver 1955, p. 275 (Little Barrier Island, New Zealand) Specimens  The unique type skin and subfossil remains are in Wellington, New Zealand. Status  Known only from the type, collected in 1870. Range  North Island and Little Barrier Island, New Zealand. Description  19–24cm (7.5–9.5in). Differs mainly from the South Island Snipe C. iredalei in having a greater area of buffy white on chin and throat; absence of barring on lower abdomen; crescent-shaped marks on upper abdomen; and in being less rufous overall.

The Coenocorypha snipe are now confined to the New Zealand area, and are distinct from the more widespread 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, particuNorth Island Snipe larly with specimens from the Coenocorypha barrierensis Auckland Islands. 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. 127); 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 125

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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) have referred subfossil remains discovered on North Island to this species, the Little Barrier Island population outliving the mainland population by centuries. 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).

South Island Snipe (Stewart Island Snipe)  Coenocorypha iredalei

Rothschild

Coenocorypha aucklandica iredalei Rothschild 1921b, p. 63 (Jack Lees Island, New Zealand) Specimens  Skins are in Canterbury, New Zealand and Tring; subfossil remains are in Tring; and Canterbury and Wellington, New Zealand. Status  Became extinct around 1964 due to depredations of rats. Range  Formerly occurred on South Island, Big South Cape Island, Jacky Lee Island and probably other outliers of Stewart Island, New Zealand. Description  19–24cm (7.5–9.5in). General coloration brown, with dark ­eye-­stripe; scapulars on wings mottled. Differs from extant Snares Island Snipe Coenocorypha huegeli by being more rufous above and lighter below, with the chin buffy white.

Big South Cape Island seems to have been the last island on which this snipe occurred, though it had previously 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 1921). 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 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 existence 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 ­mutton-­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 ­Short-­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 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, sometimes 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 126

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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 ­mouse-­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.

Forbes’s Snipe  Coenocorypha chathamica

Forbes

Gallinago chathamica Forbes 1893c, p. 545 (Chatham Islands) Specimens  Subfossil remains are in London and Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. May have survived into historic times. Range  Chatham Islands, New Zealand.

The Chatham Islands are the only island group on which two species of Coenocorypha snipe lived sympatrically. 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 mammalian 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.

Viti Levu Snipe  Coenocorypha miratropica

Worthy

Coenocorypha miratropica Worthy 2003, p. 95 (Vatuma Cave, Nadi, Vitilevu, Fiji) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Range  Viti Levu, Fiji.

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 Zealand species and an ­as-­yet undescribed bird of this genus from Norfolk Island. As the Pacific oceanic island fossil record becomes more complete, it will probably reveal new species of Coenocorypha on other island archipelagos, indicating that these seemingly ­reluctant-­to-­fly birds are very capable island colonists. The Viti Levu Snipe disappeared almost certainly as a result of human colonisation of the island, and the establishment of pigs, dogs and Pacific Rat Rattus exulans. The date of extinction is not known.

Puerto Rico Woodcock  Scolopax anthonyi

(Wetmore)

Gallinago anthonyi Wetmore 1920, p. 78 (Cueva Catedral, near Morovis, Puerto Rico) Scolopax anthonyi Olson 1976b, p. 266. Specimens  Subfossil remains are in New York and Washington, D. C. Status  Extinct, known only from subfossil remains, but may have survived into comparatively recent times. Range  Puerto Rico.

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 127

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discovered were originally forested, which is a habitat preferred by the genus. Nothing else is known of it, or why it became extinct.

Eskimo Curlew  Numenius borealis

(Latham)

Scolopax borealis J. R. Forster 1772, pp. 411, 431 (Fort Albany, Hudson Bay) Specimens  Many museums hold skins of the Eskimo Curlew. For a comprehensive listing see Hahn (1963). Status  Probably extinct. Not definitely recorded since 1963. Probable sighting in 1981, unconfirmed reports of its existence up to 2006. 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 ­cinnamon-­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 ­rusty-­yellow; ­wing-­linings cinnamon in flight; bill blackish, with base ­fleshy-­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 ­non-­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 ­mid-­western states (Environment Canada 2007). As befits a ­long-­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. 144) 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

Eskimo Curlew Numenius borealis

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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. 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 associated with ­mixed-­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.

White-winged Sandpiper  Prosobonia leucoptera

(Gmelin)

White-winged Sandpiper. Latham 1785, p. 172, pl. 82 (Tahiti and Eimeo = Moorea) Tringa leucoptera Gmelin 1789, p. 678. Based on Latham. Specimens  The type specimen is in Leiden. Status  Extinct. Last seen in the 18th century, and known from one specimen and at least one contemporary illustration. Range  Tahiti, Society Islands, South Pacific. Description  18cm (6.5in). Wings and back sooty-brown, paler on crown and sides of head and neck; chin and gular region buffish-white; underparts and rump russet; narrow pale russet ring round eye, supercilium pale russet anteriorly, changing to white above auricular region; crescent-shaped patch of white on lesser ­wing-­coverts near bend of wing; central rectrices sooty-brown tipped with russet, outer ones with prominent russet tips, becoming progressively more heavily barred with russet towards lateral pair (part of tail is missing); bill dark above, lower bill slightly paler; legs straw-coloured, slightly tinged green.

White-winged Sandpiper Prosobonia leucoptera

The Polynesian Sandpipers of the genus Prosobonia are poorly known. Only one species now survives, the Tuamotu 129

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Sandpiper P. parvirostris, which is considered to be endangered. The genus was originally widespread in the South Pacific, with ­as-­yet undescribed birds on Mangaia in the Cook Islands, Ua Huka in the Marquesas, and Henderson Island (Wragg 1995), and possibly on Kiribati (see p. 332). The ­White-­winged Sandpiper P. leucoptera was observed by J. R. Forster during the 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 p. 332). Habits Virtually nothing was recorded about the ­White-­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.

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.

Huahine Gull (Society Islands Gull)  Larus utunui

Steadman

Larus utunui Steadman 2002b, pp. 3–4 (Fa’ahia, Huahine, Society Islands) Specimens  Subfossil remains are in Honolulu and Tahiti. Status  Extinct. Known only from archaeological subfossil remains. Range  Huahine, Society Islands, South Pacific.

The Huahine Gull is known from the subfossil remains of two individuals, collected with a host of other bird species from a Polynesian midden site on Huahine, Society Islands. Gulls were probably once resident throughout French Polynesia, but the Huahine bird is the only species described from physical evidence. No Larus species occurs in Polynesia today. The closest populations are the Silver Gull L. novaehollandiae, ­Black-­billed Gull L. bulleri and Kelp Gull L. dominicanus in New Zealand, 3,800km to the ­south-­west of Huahine (Steadman 2002b). Morphological analysis of the postcranial elements suggests that the Huahine population was closely related to the Silver Gull. All birds were utilised by the Polynesians for their meat, bones, feathers and eggs, so the Huahine Gull was probably hunted to extinction along with many other seabirds. Steadman suggests that habitat loss, disease and predation from introduced mammals may have also impacted on the population.

Huahine Gull Larus utunui

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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.

Great Auk  Pinguinus impennis

(Linnaeus)

Alca impennis Linnaeus 1758, p. 130 (‘in Europa arctica’ = Norwegian seas) Specimens  Many institutions hold specimens of Great Auks, which comprise 81 mounted skins, around 75 eggs and 24 complete skeletons (see Fuller 1999). Status  Extinct. Its range had been contracting since prehistoric times. The last pair was taken in June 1844, but there was a sighting of an individual on the Grand Banks of Newfoundland in 1852. Range  Formerly occurred on the shores and islands of the North Atlantic, with archaeological remains found as far south as Florida and Gibraltar. It also once occurred in the Mediterranean. Description  75–85cm (30–33in). Head, neck and back overall glossy black; underside white; feet and claws black, webbing ­brownish-­black; bill with white grooves; inside of the mouth yellow; iris hazel or chestnut. Three distinct plumage moults, one while breeding, one during the ­non-­breeding season and one in the winter. Breeding plumage wide white patch over eye; ­non-­breeding white eye patch replaced with white band, and grey line from eye to ear. Summer moult chin and throat ­blackish-­brown; bill with up to seven white grooves on rostrum, 12 on mandible. Winter moult; throat white, bill with less white grooves. Juvenile neck mottled black and white; white eye spot absent; grey line through eye to below ear; beak grooves less prominent. Egg ground colour yellowish white to light ochre, with a varying pattern of black, brown or greyish spots and lines, often concentrated on the large end.

The Great Auk was the largest and most penguin-like of all the auks; indeed, the name ‘penguin’ was originally 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

Great Auk Pinguinus impennis

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­ orth-­eastern United States, Scandinavia, the west coast of Greenland, Iceland, the British Isles, France and n 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 ­south-­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 relatively 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 disappearing 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 ­egg-­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). Last stronghold The Geirfuglasker, a volcanic island off the ­south-­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. 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, 132

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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 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 ­Gare-­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 ­Razor-­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.

Pteroclididae (Sandgrouse) The sandgrouse are restricted to the Old World and divided into two genera. They prefer open country, especially plains and ­semi-­deserts, and are cryptically coloured with feathered legs and toes. They frequently fly long distances in search of water. They are hunted over much of their range, and some species are considered threatened.

Fayum Sandgrouse  Pterocles exustus floweri

Nicoll

Pterocles exustus floweri Nicoll 1921, p. 128 (Fayum, Egypt) Specimens  The type specimen is in Tring. Status  Probably extinct. Last recorded March, 1979. Range  Nile Valley, central Egypt. Description  31–33cm (12.25–13in). Differed from the nominate in being darker, with more grey on head; female less brownish, with the ­breast-­band paler. Male general coloration ­sandy-­beige with pale brown back; rufous face, and throat; upper belly chestnut; lower belly blackish; narrow black chestband; black tip to ­wing-­coverts and secondaries; tail with long black central feathers; bill ­slate-­blue with dark tip. Female brighter, back mottled tan, brown and dark brown.

The ­Chestnut-­bellied Sandgrouse Pterocles exustus is widely distributed across Africa and the Middle East to India. It is not considered threatened anywhere in its range, but it appears that the population that once inhabited the Nile Valley in Egypt has disappeared. This poorly differentiated subspecies was last recorded when 10 birds were seen between Isna and Idfu in March 1979 (de Juana 1997). The reasons for the population’s disappearance are unclear. Habits Little is known about the Nile Valley birds, but the nominate P. e. exustus inhabits sparse scrub in arid environments, and can travel long distances on a daily basis for water. The birds are cryptic and remain motionless when threatened. They generally live in small scattered groups, but concentrate in larger numbers around watering holes (Maclean 1996). They feed generally on leguminous plant seeds and beans, but will take plant shoots and some insect food.

Columbidae (Pigeons and doves) The pigeons and doves are have a ­world-­wide distribution, and have been divided into at least 50 genera. They are a diverse group with considerable variation in size and shape. They occur in a range of habits, 133

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from arid regions to dense forest. Some species are sedentary and terrestrial, whereas other genera are adept at crossing large expanses of water, and have colonised most island archipelagos. One continental species, the Passenger Pigeon Ectopistes migratorius, was once considered the most numerous species on earth. Once isolated on islands, three species have evolved into giant, flightless birds, most notably the Dodo Raphus cucullatus and Solitaire Pezophaps solitaria in the Indian Ocean, but also on Fiji in the Pacific Ocean and perhaps St Helena, in the South Atlantic. Pigeons and doves are hunted indiscriminately for their meat, and combined with habitat destruction and predation from introduced mammals, a large number of species have become extinct.

Dodo  Raphus cucullatus

(Linnaeus)

Cygnus cucullatus. Nieremberg 1635, p. 231 (based on Clusius’s description and figure). Struthio cucullatus Linnaeus 1758, p.155 (‘India’) Didus ineptus Linnaeus 1766, p. 267 ? Didus nazarenus Bartlett 1851, p. 284 Specimens  Subfossil remains of the Dodo are held in many museums including Berlin; Brighton, England; Cambridge, England; Cambridge, Massachusetts; Copenhagen; Durban; Frankfurt; Leeds; Leiden; Liverpool; Mauritius; London and Tring; New York; Oxford; Paris; Qatar; Réunion; Royal College of Surgeons, London; Stuttgart; Tokyo; Vienna; Washington, D. C.; and York. Status  Extinct. Date of extinction not certain, given variously as 1662 to 1693. Range  Mauritius, Mascarene Islands. George Pine’s account of Dodos on the Isle of Pines (Neville 1668) is fictitious. Description  Approximately 70–75cm (28–29in). Much confusion has arisen over the external morphology of the Dodo, exacerbated by ridiculous assumptions based on inadequate evidence (e.g. Oudemans 1917; Hachisuka 1953). It appears to have been variable in colour, some light or dark grey, brownish, yellowish-brown, black, or even white. Face devoid of feathers, ­ash-­coloured or bluish; bill light ash, blackish or yellow with blue spot on the mandible; iris white; legs yellowish or blackish; claws black. Some of these descriptions may be a result of sexual dimorphism, individual variation, or ageing, of which we know nothing. Analysis of subfossil remains confirms size dimorphism, the males being larger than the females (Livezey 1993).

More has been written about the Dodo Raphus cucullatus of Mauritius than any other extinct bird species (see Fuller 2002). But in reality, virtually nothing is known about it in life. Early Arab traders probably 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 ­Vice-­Admiral Wybrandt Warwijck that in September 1598 claimed Mauritius for the Netherlands (Barnwell 1948; Moree 1998), at which point the documented history of the 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

Dodo Raphus cucullatus (top); Rodrigues Solitaire Pezophaps solitaria (bottom)

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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 (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, translated in Wissen 1995) 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 ­well-­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 starvation diet until rescued. One party led by Vilkert 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. 108). Whoever actually 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 ­17th-­century human population on Mauritius was small, never more than 50 people, occupying an island 61km long by 47km wide, and covering an area of 1,860km2. 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 mummified 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 individual exhibited in London, of which the head and foot still reside in Oxford. Various drawings and paintings exist, some of which are direct copies of earlier works, others based on indeterminate material, with pitifully few considered authentic. Of these, two deserve additional comment. A Dodo attributed to court artist Ustad Mansur of c.1625 (Ali 1968), who painted zoological specimens for the above-mentioned 135

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Jahangir, is easily the best coloured rendition, and the 1601 ink sketch of a recently killed Dodo by Joris Joonstenzoon Laerle (see Hume 2003) unequivocally reproduces the bird as seen in life. All traditions of the Dodo on the island had died out by the 1800s, and doubts had begun to arise as to whether the bird had ever really existed. Duncan (1828) and Broderip (1837) accumulated all of the available information to confirm the validity of the bird, and the publication of the unparalleled Dodo monograph by Strickland & Melville (1848) initiated a rush to obtain fossil Dodo material. George Clark, after a ­tip-­off from a railway engineer, Harry Higginson, discovered a lowland, coastal marsh in ­south-­west Mauritius called the Mare aux Songes in 1865, and from this one place almost all Dodo subfossil remains were subsequently recovered. Bitter rivalry and academic jealousy ensued over claims to the material (Hume et al. 2008), but ultimately Richard Owen, superintendent and comparative anatomist at the then British Museum (Natural History) formally described the Dodo’s skeletal anatomy to science (Owen 1866a, 1872a). An almost complete, associated specimen was discovered by the collector Etienne Thirioux in the early 1900s, and a second associated individual (named ‘Dodo Fred’) was recovered from a mountain cave in Bois Cheri in 2007 (see Cheke & Hume 2008). Thirioux’s Dodo was probably collected in a valley cave on the slopes of Le Pouce Mountain. The discovery of these highland specimens has shown that the Dodo also once occurred in the central mountains as well as in the lowlands and coastal regions. Affinities The affinities of the Dodo have been investigated by a number of authors. The bird has been considered a member of a large array of bird orders; it has been thought to be a diminutive ostrich, a rail, and even a vulture. After examining a skull in Copenhagen, Professor J. T. Rheinhardt proposed that the Dodo was related to pigeons and doves (Rheinhardt 1842). This claim was initially met with ridicule, but after Strickland & Melville (1848) confirmed his theory by examination of the Oxford Dodo head, the idea became universally accepted (see Owen 1866a). DNA studies have now concluded that the Dodo and closely related Rodrigues Solitaire Pezophaps solitaria (see p. 137) are a sister clade, nested within the Columbidae and derived from the same common ancestor as the ­south-­east Asian Nicobar Pigeon Caleonas nicobarica (Shapiro et al. 2002), though this is open to debate (Johnson & Clayton 2000). 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. 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 (Rijkdijk et al. 2009) 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 & ­Mourer-­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.

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Rodrigues Solitaire  Pezophaps solitaria

(Gmelin)

Solitary Dodo. Latham 1785, pp. 3–4 Didus solitarius Gmelin 1788, p. 728. Based on Latham Pezophaps minor Strickland 1852 ? Didus nazarenus Bartlett 1851, p. 284 Specimens  Many institutions hold subfossil remains. Complete skeletons, probably associated individuals rather than composites, are in Cambridge, England; Natural History Museum, London and Royal College of Surgeons, London; also individual bones in Cambridge, Massachusetts; Glasgow; Liverpool; Mauritius; Paris; Stuttgart; and Washington, D. C. Status  Extinct. Known only from bones and from contemporary accounts. Probably became extinct around the 1760s. Range  Endemic to Rodrigues, Mascarene Islands. Description  Approximately 70cm (27.5in) in female, 90cm (35.5in) in male. Male general colour light greyish with blackish back, frontlet of black feathers; female light buff or brown; feathers of thighs distinctly rounded; frontlet of tan feathers; bare skin parts undescribed; beak short (2.5cm); iris black.

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, but see p. 136). 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 biodiversity 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 1874a), and his description of the Solitaire adds more to our knowledge 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 quickness, 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 137

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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 sometime after 1726, and the tortoise hunters indiscriminately burned off the vegetation; 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 1789 in a cave deposit by Labistour, a resident on the island. 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 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. 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. 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.

Madeira Wood Pigeon  Columba palumbus maderensis

Tschusi

Columba palumbus maderensis Tschusi 1904, p. 227 (Boaventura, Madeira) Specimens  Specimens are in Madeira. Bannerman & Bannerman (1965) believed that Tschusi’s type specimen was probably destroyed during one of the World Wars. Status  Extinct. Not recorded since 1904. Range  Formerly occurred on the island of Madeira. Description  42cm (16.5in). This race was larger (Gibbs et al. 2001 state that is was smaller) and darker than nominate palumbus, and the greyish wine-red of the crop area was more pronounced.

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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 misidentification of the rare Trocaz Pigeon Columba trocaz, a closely related species. The reason for its extinction is unknown.

Lord Howe Pigeon  Columba vitiensis godmanae

(Mathews)

Raperia godmanae Mathews 1915, p. 24 (Lord Howe Island) Specimens  Three paintings in The Natural History Museum, London (George Raper); Alexander Turnbull Library, Wellington, New Zealand (Anonymous); and the National Library, Canberra, Australia (John Hunter). Status  Now extinct, last recorded in 1853. Known only from paintings. Range  Formerly occurred on Lord Howe Island. Description  40cm (15.5in). Head and breast purple with white throat; mantle metallic green; back and ­wing-­coverts brown. In C. v. halmaheira, the closest race, the bird is entirely glossed purple when looked at from one angle, and entirely green when viewed from another.

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) 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. 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 specimens (Hindwood 1938).

Ryukyu Pigeon  Columba jouyi

(Stejneger)

Janthoenas jouyi Stejneger 1887a, p. 583 (Northern part of Liu Kiu = Ryukyu Islands) Specimens  Specimens are in Liverpool; Tokyo; and Tring. Status  Extinct. Last seen on Okinawa in 1906, and on the Daito Islands in 1936. Range  Ryukyu Islands (Okinawa, Iheya, Izena, Yagahi and Zamami) and Daito Islands ­(Kita-­daita-­jima and ­Minami-­daito-­jima), Japan (Gibbs et al. 2001).

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Ryukyu Pigeon Columba jouyi Description  45cm (17.5in). A large black pigeon, glossed with rose-purple on head; green on lower back; distinctive pale grey stripe across shoulders; bill ­greenish-­blue with yellow tip; iris brown; feet purplish red.

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.

Bonin Islands Pigeon  Columba versicolor

Kittlitz

Columba versicolor Kittlitz 1832, p. 5, pl. 5, fig. 2 (Bonin Islands: Peel Island or ­Chichi-­jima) Columba kitlizii Temminck 1835, livr. 98, no. 578. (Virtually unpaginated = vol. 4, p. 1076 of Tring bound copy) Columba metallica Vigors 1839, p. 25 Columba iris Kittlitz 1858, p. 175 Specimens  Specimens are in Frankfurt; St. Petersburg; and Tring. Type formerly in the collection of the Zoological Society, London, but now lost. Status  Extinct. Last recorded in 1889. Range  Formerly known to have occurred on ­Nakōdo-­jima and Chichijima (Peel Island) in the Ogasawara (Bonin) Islands, Japan. Description  45cm (17.5in). Differs from Japanese Wood Pigeon Columba janthina and Ryukyu Pigeon C. jouyi in being paler with a golden-purple (not green) back. Head metallic green, shading to green and purple on the hind neck; back golden purple; rump green; ­wing-­coverts with paler, more golden reflections; underparts light grey; bill ­greenish-­yellow; iris blue; feet dark red.

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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) 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).

Mauritius Wood Pigeon  Columba sp. Specimens  Subfossil remains are in Paris. Status  Extinct. Known from subfossil remains and contemporary accounts. Range  Mauritius, Mascarene Islands.

Subfossil remains have shown that a small species of pigeon, probably referable to the genus Columba, once existed on Mauritius (JPH, in press). The Mauritius Wood Pigeon was briefly mentioned in some early accounts, which generally only describe the ease with which it could be caught. It was an arboreal species, and probably disappeared around1730 as a result of ­over-­hunting, predation from Black Rats Rattus rattus and severe deforestation. Nothing else is known about this bird.

Réunion Pink Pigeon  Nesoenas duboisi

Rothschild

Nesoenas duboisi Rothschild 1907a, p. 166 (Réunion) Specimens  Subfossil remains are in Réunion. Status  Extinct since the end of the 17th century. Known from subfossil remains and the account of Dubois. Range  Réunion, Mascarene Islands.

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 1672–73, 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 1704), 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 (Cheke & Hume 2008), appears to have been a major factor in their rapid disappearance, as deforestation was negligible at this time.

Rodrigues Turtle Dove  ? Nesoenas rodericana

(Milne-Edwards)

Columba rodericana Milne-Edwards 1874a, p. 16, pl. 12, ff. 1, 1a, 1b, 1c (sternum) ? Turtur picturatus Milne-Edwards 1874a, p. 14 Specimens  Subfossil remains are in Cambridge, England; London; Rodrigues; and Tring. Status  Extinct. Known only from subfossil remains and contemporary accounts. Range  Rodrigues, Mascarene Islands.

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Rodrigues Turtle Dove ?Nesoenas rodericana

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 referable 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. 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. 161). 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.

Mauritius Turtle Dove  Nesoenas sp. Specimens  Subfossil remains are in Cambridge, England; and Paris. Status  Extinct. Known from subfossil remains and contemporary accounts. Range  Mauritius and possibly Réunion, Mascarene Islands.

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 142

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from Madagascar from at least 1770 (Cheke & Hume 2008). Subfossil remains have shown that Mauritius once harboured a distinct species (JPH, in press), 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.

Seychelles Turtle Dove Nesoenas picturata rostrata

(Bonaparte)

Turtur rostratus Bonaparte 1855a, p.16 (Mariannes = Seychelles) Specimens  Specimens are in Liverpool and Tring. Status  Probably extinct. No pure-bred specimens have been seen in recent years. Range  Seychelles, ­south-­west Indian Ocean. Description  28cm (11in). A small, heavy built terrestrial pigeon with long wings and legs. Seychelles form smaller than nominate; wings shorter, more rounded; no grey on head, overall darker and browner; lower flanks and lower breast dark grey; ­undertail-­coverts white; bill ­purplish-­red at base, pale ­blue-­grey tip; bare orbital skin ­purplish-­red; iris brown; legs and feet ­blood-­red.

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.

Amirante Turtle Dove  Nesoenas picturata aldabrana

(Sclater)

Turtur aldabranus Sclater 1872a, p. 623. (Aldabra, in error) Specimens  Specimens are in Tring. Status  Probably extinct. No ­pure-­bred specimens have been seen in recent years. Range  Amirante Islands, ­south-­west Indian Ocean. Description  28cm (11in). Amirante birds differed from nominate N. p. picturata in being generally overall more vinaceous, except for ­wing-­coverts; also ­undertail-­coverts grey.

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 seems to have disappeared due to hybridisation with the nominate race N. p. picturata, introduced from Madagascar. The race was originally described from captive-bred birds held by Edward Newton while he was colonial 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 be established. Pure-bred Amirante birds have not been reported for some time, and the subspecies may well now be extinct.

Marquesas Cuckoo Dove  Macropygia heana

Steadman

Macropygia heana Steadman 1992, p. 333 (Hane Site, Ua Huka, Marquesas Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Ua Huka, Marquesas Islands, South Pacific.

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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.

Society Islands Cuckoo Dove  Macropygia arevarevauupa

Steadman

Macropygia arevarevauupa Steadman 1992, p. 338 (Fa’ahia Site, Huahine, Society Islands) Specimens  The type is in Tahiti. Status  Extinct. Known only from the type, a subfossil tibiotarsus. Date of extinction unknown. Range  Huahine, Society Islands, South Pacific.

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 capability 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.

Passenger Pigeon  Ectopistes migratorius

(Linnaeus)

Columba migratoria Linnaeus 1766, p. 285 Specimens  Specimens exist in almost all major institutions. For full listings see Greenway (1967) and Hahn (1963). Status  Extinct. The last individual died in Cincinnati Zoo in 1914. Range  Formerly abundant in North America. 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 ­slate-­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 ­greyish-­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 ­buff-­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 and Washington, ­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 144

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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. 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 development 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. 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. 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 later, as there were a number of sight records between 1900 and 1907, but those cases that were investigated 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. By 1909, only the Cincinnati Zoological Gardens birds were still alive; two females and a single male. On September 1, 1914, just before one in the afternoon, the last individual, Martha, died (Fuller 2000). Causes for extinction The extinction of the Passenger Pigeon is unique in the annals of bird extinction, not only for its very rapid decline from one of the world’s most abundant species, but for the enormous volume of documentation that exists (see Blockstein 2002). Hunting alone cannot explain the bird’s extinction. The Passenger Pigeon appears to have been inextricably linked to the eastern deciduous forest area of the US and Canada, and although it was able to disperse far from its normal range, it never colonised any other forest type. The western forest, which is in many ways ecologically similar, was already occupied by the closely related ­Band-­tailed Pigeon Patagioenas fasciata, which may have resulted in competitive exclusion. The breeding area was limited by the occurrence of oak and beech trees; a combination of intense hunting pressure and the huge increase in deforestation that took place in the eastern United States during the latter half of the 19th century were the major factors in the pigeon’s extinction. Furthermore, the social organisation of the species was very important for finding food (Bucher 1992). Solitary birds would have found it much more difficult to detect areas of high mast than large flocks. As the birds declined and forests were destroyed, the remaining populations became thinned and fragmented, and perhaps were unable to locate crucial feeding grounds. It appears that this communally breeding species required large densities of birds for breeding success. Blockstein & Tordoff (1985) suggest that disturbance of the nesting colonies, which resulted in the adults abandoning their nests, was a deciding factor in the Passenger Pigeon’s downfall, and that as a result of human activities many potential nesting sites were abandoned during the latter half of the 19th century (Blockstein 2002). This concurs with Forbush (1927), who suggested that the continuous destruction of young birds, and failure to rear young by the adults, ultimately finished off the species. 145

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It is not inconceivable that the enormous periodic concentrations of birds comprised most of the entire population, if not all of it, which dispersed into smaller groups at other times. This would have made the bird particularly susceptible to extinction. Once the Passenger Pigeon population had reached a critical low point, the species was doomed, even though thousands of individuals may still have remained. Bucher (1992) suggested that the Passenger Pigeon was descended from Zenaida stock which had occupied the central plains of North America and become secondarily adapted to woodland habitats. However, recent DNA analysis has shown, surprisingly, that the Passenger Pigeon was not related to the Zenaida doves, but was in fact closest to the aforementioned ­Band-­tailed Pigeon, and is part of a radiation that includes the Old World Columba pigeons, Eurasian turtle doves Streptopelia sp., and Macropygia cuckoo doves (Johnston et al. 2010). In captivity, the Passenger Pigeon hybridised with domesticated Eurasian Collared Dove Streptopelia decaocto, although the hybrids were infertile (Deane 1908). 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 ­pre-­nesting behaviour (in captivity at least), without the bowing, strutting or ­bill-­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).

St Helena Pigeon  Dysmoropelia dekarchiskos

Olson

Dysmoropelia dekarchiskos Olson 1975b, p. 29 (Sugarloaf Hill, St Helena) Specimens  Subfossil remains are in London and Washington, D. C. Status  Extinct. Known only from subfossil remains from deposits of mid-Pleistocene age, but may have survived until the 16th century. Range  St Helena.

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. 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. It is difficult to determine whether 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 1963) was probably the introduced Zebra Dove Geopelia striata and not an endemic form, and that introductions 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 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. 146

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Puerto Rican Quail Dove  Geotrygon larva

(Wetmore)

Oreopelia larva Wetmore 1920, p. 79 (Cueva Clara, Puerto Rico) Specimens  Subfossil remains are in New York and Washington, D. C. Status  Extinct, known only from subfossil bones. Date of extinction unknown. Range  Puerto Rico.

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 ­Grey-­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.

Kanaka Pigeon  Caloenas canacorum

Balouet & Olson

Caloenas canacorum Balouet & Olson 1989, p. 14 (Pindai Cave, Nepoui Peninsula, New Caledonia) Specimens  Subfossil remains are in Paris. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  New Caledonia and Tonga.

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 enigmatic 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 terrestrial 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.

Spotted Green Pigeon (Liverpool Pigeon)  ? Caloenas maculata

(Gmelin)

Spotted Green Pigeon. Latham 1783, p. 642 Columba maculata Gmelin 1788, p. 780 Specimens  The unique type specimen is in Liverpool. Status  Extinct. Known only from descriptions, and the type specimen. Possibly survived until the 1820s. Range  Unknown, but may have been Tahiti, Pacific Ocean. Description  30cm (12in). Dark glossy green above; head and neck darker; feathers of the neck longer; ­wing-­coverts and scapulars tipped with triangular cinereous white spots; primaries black; tail black, tipped with cinereous white; belly, vent, and thighs dusky black; bill black, tip pale yellow; legs and feet ­reddish-­brown. Has been sometimes depicted with a knob at the base of the beak; this is not apparent in the specimen.

The Spotted Green Pigeon is an enigma. It bears no close relationship with any extant genera save a rather superficial resemblance to the Nicobar Pigeon Caloenas nicobarica. Wagler (1827) thought that it might have been the young of the Nicobar Pigeon Caloenas nicobarica, of which Rothschild & Hartert (1901) commented: 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. 147

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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. We agree with these authors, and note that this pigeon is like no other species, with only superficial resemblance to other genera. The wings were short and rounded, the tail long (unlike in Caloenas nicobarica) and the tarsi short and stout, which indicates that it was probably an arboreal species, not a terrestrial one, with a poor ability to migrate between islands. The bill was typical of a frugivorous pigeon. The Spotted Green Pigeon may be the same bird called the ‘titi’ by the Tahitians, who described a speckled green and white bird that had a loud call, titi (Henry 1928). No such bird exists there now, so it probably disappeared due to ­over-­hunting and predation by introduced animals.

Norfolk Island Ground Dove  Gallicolumba norfolciensis

(Latham)

Columba norfolciensis Latham 1801b, p. 60 (Norfolk Island) Specimens  John Hunter’s plate is in the National Library, Canberra, Australia; subfossil remains are in Norfolk Island. Status  Extinct. Known from Latham’s description, contemporary illustrations and subfossil remains. Range  Norfolk Island. Description  36cm (14in). Head and neck white; back and wings deep dusky purple, with a few darker markings; breast, belly and vent black; tail dull purple, with inner webs of feathers dusky; bill black, legs and feet red.

Latham (1801b) described under Columba norfolciensis three quite distinct birds, two now identified as the Common Emerald Dove Chalcophaps indica and the White-headed Pigeon Columba leucomela. The Norfolk Island Ground Dove was first illustrated by John Hunter (1793); this was reproduced by Hindwood (1965), who 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. 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), once occurred on Norfolk Island during the early years of settlement. Furthermore, subfossil material, almost certainly referable to this bird, has now been found (Higgins & Davies 1996). Doubtless it was soon exterminated by ­over-­hunting and predation from cats and rats. 148

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Taxonomy Goodwin (1983) believed that the bird was probably a Gallicolumba, very similar in appearance to the Whitethroated Ground Dove G. xanthonura of Yap and the Marianas Islands, and showing similar sexual dimorphism. He also pointed out, however, that Hunter’s plate (Hunter 1793) is closer in appearance to the Shy Ground Dove G. stairi of the Fijian, Tongan and Samoan Islands than is Latham’s description. This, he suggests, is due to individual variation, or differing plumages between the sexes.

Society Islands Ground Dove  Gallicolumba erythroptera ssp. ? Columba pectoralis W. W. Ellis 1777, BMNH no. 71 (York Island, or Eimeo or Imaio = Moorea). Not Peristera pectoralis Peale; it is probable that the name pectoralis was not assigned to the bird by Ellis, but was added to his picture later, after the publication of Peale’s description. Garnet-­winged Pigeon. Latham 1783, p. 624 (Eimeo = Moorea) Garnet-­winged Pigeon. var. from Otaheite. Latham 1783,p. 624 (Otaheite = Tahiti) Columba erythroptera Gmelin 1789, p. 775 (based on Latham) Columbe erythroptere. Temminck & Knip 1808–1811, pl. 55 Pigeon à ailes rouges. Sonnini 1799–1805, p. 223 Columba erythroptera (part) Gray 1856b, p. 6 (York Island or Eimeo = Moorea) Phlogoenas erythroptera Sharpe 1906a, p. 205 ? Gallicolumba erythroptera Lysaght 1959, p. 334 Specimens  A specimen is in Leiden. Status  Extinct. Not recorded since the birds were first collected in the late 18th century. Range  Moorea and Tahiti, Society Islands. Description  24cm (9.5in); Forehead white, continuing over eye as a streak almost to hindneck; nape and hindneck black; lower neck, shoulders, and wing-coverts deep garnet red; back, primaries, lower breast, belly, and vent black; tail basal part ­greyish-­ash, terminal part black; bill variably ­dusky-­yellow to black; legs and feet brown.

The Polynesian Ground Dove Gallicolumba erythroptera 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 G. e. erythroptera is considered critically endangered, whereas the ­White-­headed Polynesian Ground Dove G. e. albicollis is probably extinct (see below). 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 ­over-­hunting, 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 1906).

White-­headed Polynesian Ground Dove Gallicolumba erythroptera albicollis

Salvadori

Gallicolumba erythroptera albicollis Salvadori 1892 (volume dated 1893), p. x (Bow Island) Specimens  A specimen is in Tring. Status  Possibly extinct; not seen since the 1950s. Range  Hao, Hiti and probably Tahanea, Tuamotu Archipelago. Description  25cm (10in). Differed from nominate G. e. erythroptera by head, neck and breast being entirely white, with some variable dusky maroon mottling on centre of crown. Male G. e. erythroptera mostly dark grey; throat, chest, forehead, and eyebrow white; upper back, scapulars and shoulders reddish purple. Female dark grey with paler grey head; forehead, eyebrow and face ­off-­white; chest rufous with paler feather edges; bill blackish; iris dark brown; legs and feet blackish.

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Tanna Ground Dove Gallicolumba ferruginea

Two forms of Polynesian Ground Dove G. erythroptera 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 ­white-­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 disappeared, 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 G. 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.

Tanna Ground Dove (Dove of Tanna)  Gallicolumba ferruginea

(Wagler)

Garnet-winged Pigeon. var from Tanna. Latham 1783, p. 625 Columba erythroptera var gamma. J. F. Gmelin 1789, p, 755 Garnet-winged Pigeon. var B. Latham 1823, p. 72 (Island of Tanna) Columba ferruginea Wagler 1829, col. 738 (Island of Tanna) Specimens  The only known illustration is Georg Forster’s drawing no. 142, in the Natural History Museum, London. Status  Extinct. Known only from J. R. Forster’s (1844) description and Georg Forster’s drawing. A specimen, formerly in the Banksian collection, is now lost. Range  Tanna Island, southern Vanuatu. Description  27cm (10.5in). Head rusty brown; back ­reddish-­purple, wings dark green and ­brownish-­grey primaries with very narrow pale edges; breast rusty brown; belly grey; bill black; iris yellowish; feet red.

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): 150

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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. 149] 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 forehead, 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, G. sanctaecrucis, has disappeared for the same reason over much of its range (Gibbs et al. 2001). Nothing more is known about it.

Henderson Island Ground Dove  Gallicolumba leonpascoi

Worthy & Wragg

Gallicolumba leonpascoi Worthy & Wragg 2003, p. 773 (Henderson Island) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Henderson Island, South Pacific.

Doves of the genus Gallicolumba 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 G. 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 ­over-­hunting soon after humans arrived on the island.

Great Ground Dove  Gallicolumba nui

Steadman

Steadman, D. W. 1992, p. 330 (Hane Site, Ua Huka, Marquesas Islands) Specimens  Subfossil remains are in Honolulu; Washington State; and Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown, but before European contact. Range  Marquesas Islands and Cook Islands.

This ground dove, based on the tarsometatarsus, was the largest Gallicolumba 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 Gallicolumba 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.

New Caledonian Ground Dove  Gallicolumba longitarsus

Balouet & Olson

Gallicolumba longitarsus Balouet & Olson. 1989, p. 16 (Pindai Cave, Nepoui Peninsula, New Caledonia) Specimens  Subfossil material is in Paris. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  New Caledonia and Île des Pins, Melanesia.

This was one of the largest Gallicolumba 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 151

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establish. 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.

Thick-­billed Ground Dove  Gallicolumba salamonis

Ramsey

Gallicolumba salamonis Ramsay 1882, p. 299 (San Cristobal, Solomon Islands) Specimens  Specimens are in New York and Sydney. Status  Probably extinct. Not seen since September 1927. Range  Ramos and Makira (San Cristobal), Solomon Islands. Description  23cm (9in). Male forehead dark rufous shading to rich chestnut on crown; nape, upper back, scapulars and ­wing-­coverts, with glossed violet–purple on upper back, ­wing-­coverts, and side of breast; primaries and secondaries rich dark chestnut; lower back, ­uppertail-­coverts and tail dark chestnut, tail tip pale rufous; chin, throat pale buff shading into darker, browner ­ear-­coverts and cinnamon buff breast shield; abdomen rich chocolate; bill brown; feet carmine.

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 ­over-­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).

Solomons ­White-­bibbed Ground Dove Gallicolumba jobiensis chalconota

Mayr

Gallicolumba jobiensis chalconota Mayr 1935, p.2.(Vella Lavella Island, Solomon Islands) Specimens  The type is in New York. Status  Possibly extinct. Last collected in the 1930s; a possible sighting in the 1940s. Range  Guadalcanal, Vella Lavella and Makira (San Cristobal), Solomon Islands. Description  25cm (10in). Differs from nominate G. j. jobiensis in white of lores and supercilium reduced, and restricted purple on upperback. Male head and neck black; lores, supraloral area and supercilium to rear of eye white; upper back, scapulars, side of breast, and ­wing-­coverts blackish, with ­purple-­edged iridescence; tertials, primaries and ­uppertail-­coverts dark brown; back and rump ­dark ­olive-brown; tail ­blackish-­brown; iris dark brown; bill black; legs and feet ­orange-­red to pale maroon. Female head ­blackish-­brown with indistinct grey supercilium; upperparts dark brown, upper back to ­uppertail-­coverts, ­wing-­coverts, tertials and secondaries glossed dark green; chin buff; throat and breast dull cinnamon; belly ­earth-­brown.

The Solomons ­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), but no recent records exist. If it does still survive, which seems doubtful, it is excessively rare. Nothing was recorded about the Solomons form, but the nominate prefers primary forest with dense undergrowth. It feeds on seeds, fruit and invertebrates.

Sulu ­Bleeding-­heart  Gallicolumba menagei

Bourns & Worcester

Gallicolumba menagei Bourns & Worcester 1894, p. 10 (Tataan, ­Tawi-­tawi, Sulu Archipelago) Specimens  Specimens are in Washington, D. C. Status  Possibly extinct, no specimens taken since the types were collected in 1891. Range  ­Tawi-­tawi, Sulu Archipelago, Philippines.

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Columbidae Description  30cm (12in). A ­medium-­sized, rather ­short-­tailed pigeon: forehead dark grey shading into iridescent green nape and upper back, extending to lower breast, sharply demarcated from white throat and sides of breast; upperparts dark chestnut, glossed deep reddish purple; wings uniform brownish chestnut; belly ­ash-­grey, shading into ­creamy-­white vent and ­undertail-­coverts; bill black with ­slate-­grey tip; iris light ­silver-­grey; legs and feet light red; claws light brown.

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 ­bleeding-­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. ­Tawi-­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, preferring primary forest with dense undergrowth, and fed on fruit, seeds and invertebrates.

Catanduanes ­Bleeding-­heart  Gallicolumba luzonica rubiventris

Gonzales

Gallicolumba luzonica rubiventris Gonzales 1979 (Catanduanes, Luzon) Specimens  The type is in Manila, Philippines. Status  Possibly extinct, known only from the type specimen, which was collected in 1971. Not recorded since. Range  Catanduanes Island, off Luzon, Philippines. Description  28cm (11in). Differed from nominate G. l. luzonica in having smaller wings; breast patch more extensive and darker red; belly pale ­pinkish-­brown, washed strongly pink; flanks pale chestnut; ­undertail-­coverts pale brown; bill black; iris ­greyish-­violet; legs and feet dull crimson with grey claws.

The only specimen of this subspecies was taken on the ­Viga-­Gigmoto watershed in 1971, and none has been collected since. The nominate is listed as near threatened, but the Catanduanes population, if it still survives, must be critically endangered (Gibbs et al. 2001). The Luzon ­Bleeding-­heart G. luzonica is hunted everywhere, both for meat and the bird trade, and deforestation is also a major concern. Nothing was recorded about the habits of the Catanduanes ­Bleeding-­heart, but nominate birds are mainly terrestrial, foraging for fruit, seeds and invertebrates in primary and secondary forest, and mostly associated with the lowlands.

Henderson Island Ground Pigeon  Bountyphaps obsoleta

Worthy & Wragg

Bountyphaps obsoleta Worthy & Wragg 2008, p. 505 (Henderson Island) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction uncertain. Range  Henderson Island, South Pacific.

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 Gallicolumba 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 ­Tooth-­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 (JPH, in prep.). 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 153

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discovered in Polynesian middens, so these birds probably disappeared due to ­over-­hunting shortly after the human occupation of the island.

Choiseul Ground Pigeon  Microgoura meeki

Rothschild

Microgoura Meeki Rothschild 1904, p. 78 (Choiseul, Solomon Islands) Specimens  Skins are in New York and Tring; a unique egg is in Tring. Status  Extinct. Last seen in January 1904. Range  Choiseul Island, Solomon Islands. Description  31cm (12in). A large ground pigeon with a bluish-grey crest of ­hair-­like feathers: forehead and face black; top of head bluish-grey like crest; back darker with a grey wash; lower back and rump brownish; upper throat almost bare; belly bright cinnamon-brown; wings brown; upper mandible blue with black tip, lower mandible reddish; iris dark lemon; legs 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 expedition 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 (1967) searched for it in the 1960s, but failed to find it. Parker (1967a, b; 1972) 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 Choiseul Ground Pigeon (probably modern Lengatura), two Microgoura meeki 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 description is called ‘kukuru-nilova’, 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 154

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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 villages east of Sasamunga. At 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: 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 (1972) 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 presence 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 monotypic ­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 155

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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 (1972) 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.

Tongan ­Tooth-­billed Pigeon  Didunculus placopedetes

Steadman

Didunculus placopedetes Steadman 2006b, p. 235 (’Eua, Tonga, South Pacific) Specimens  Subfossil remains are in Florida. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Tongatapu, Lifuka, Ha’ano, ‘Uiha and Ha’afeva, Tonga, South Pacific.

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 ­Tooth-­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.

Negros Fruit Dove  Ptilinopus arcanus

Ripley & Rabor

Ptilinopus arcanus Ripley & Rabor 1955, p.1 (Negros, Philippines) Specimens  The type specimen is in Yale. Status  Possibly extinct. Only known from the type, collected in 1953. Range  Negros Island, Philippines. Description  16.5cm (6.5in). A very small, vivid ­dark-­green fruit dove: type female forehead ­ash-­grey; yellow ­eye-­ring broad and extensive; throat ­greyish-­white fading to green underparts; mantle, lower back, ­uppertail-­coverts green; lower belly whitish with green subterminal bands, shading to yellow vent; bright yellow ­undertail-­coverts; yellow fringes to tertials and greater coverts form narrow bar on folded wing; bill black; legs and feet probably dull purplish red; iris unrecorded.

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 ­pinkish-­purple, bordered in front of eye and rear of crown with yellow; back ­greyish-­green; wings dark glossy green; reduced emarginated tip to first primary; rump and ­uppertail-­coverts

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Columbidae 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 ­tail-­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 Greygreen 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 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. chrysogaster, P. hernsheimi or even P. coralensis [westen Society Islands race of ­Grey-­green Fruit Dove, Kosrae race of ­Purple-­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 ­undertail-­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 ­bread-­fruit tree. A Cuculus [doubtless ­Long-­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. 157

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Nuka Hiva ­Red-­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.

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 themselves 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.

Hiva Oa ­Red-­moustached Fruit Dove  Ptilinopus mercierii tristrami

Salvadori

Ptilinopus tristrami Salvadori 1892, p. 1 (Hivaoa Island, Marquesas group) Specimens  Specimens are in New York and Tring. Status  Extinct. Not seen since 1922. Range  Hiva Oa Island, Marquesas Islands, South Pacific. Description  22cm (8.5in). Differed from the Nuka Hiva nominate race in having a yellow band encircling the red cap; being brighter in colour; less generally tinged with yellowish green. Immature less yellowish on crown and hind neck; light ­feather-­edgings narrower on tertials and tail feathers; breast tinged with green with a few light yellowish ­feather-­edgings; abdomen paler yellow; and ­undertail-­coverts ­yellowish-­white.

Like the nominate ­Red-­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 nineteenth 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 ­Red-­moustached Fruit Doves occurred in mountain forests, generally higher up than the ­White-­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.

Mauke Fruit Dove  Ptilinopus rarotongensis ‘byronensis’ Columba Byronensis. Bloxam’s unpublished manuscript (see Olson 1986c, p. 200) (Mauke, Cook Islands) Ptilinopus rarotongensis cf. goodwini Olson 1986c, p. 202 Status  Extinct. Known only from Bloxam’s account of the lost type. Range  Formerly occurred on Mauke, Cook Islands, South Pacific.

Olson (1986) 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 (see p. 276). 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: 158

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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 ­belly-­patch (Gibbs et al. 2001), being replaced by a few orange feather tips (Holyoak 1974a). Olson (1986) noted that Bloxam’s description of the Mauke bird as 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.

Mauritius Blue Pigeon (Pigeon Hollandais)  Alectroenas nitidissima (Scopoli) Pigeon hollandais. Sonnerat 1782, p. 175, pl. 101 Hackled Pigeon. Latham 1783, p. 641 Columba nitidissima Scopoli 1786, p. 93, (Île de France = Mauritius) Columba franciae Gmelin 1789, p. 779 Columba batavica Bonnaterre 1790, p. 233 Ramier périssé. Levaillant 1808, pl. 267, p. 74 Columba jubata Wagler 1827, Columba, species 22 Specimens  Skins are in Edinburgh; Mauritius; and Paris; subfossil remains are in Cambridge, England; and Paris. Status  Extinct. Last specimen taken in 1826; perhaps survived until 1837. Range  Mauritius, Mascarene Islands. Description  30cm (12in). A relatively large pigeon, the largest of its genus, with a distinct red tail, white head and indigo blue body: head, neck and upper breast silvery-white, feathers long and stiffened, particularly on neck; general colour dull indigo blue; upper back, scapulars and wings metallic blue; tail and uppertail-coverts maroon red; base of outerwebs of outermost rectrices blackish blue; orbital skin bright red; iris reddish orange with yellow inner ring; bill greenish with darker tip; legs and feet dark ­slate-­grey. The legs and feet have often been described as red, but this is incorrect; also, there is no evidence for carunculations on the head, contra Gibbs et al. (2001).

Mauritius Blue Pigeon Alectroenas nitidissima

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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 ­Cloven-­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 (JPH in prep.). 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 ­Sonnerat-­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 & Cheke 2008). 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 (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. 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 predominantly on fruit, never alighting on the ground to feed.

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Réunion Blue Pigeon  Alectroenas sp. Status  Extinct. Known from vague 17th century accounts only. Last reported c.1672–73. Range  Réunion, Mascarene Islands.

Evidence for the existence of an Alectroenas blue pigeon on Réunion rests entirely on a few rather vague 17th century accounts. The first was a report by the Dutch navigator, Willem Bontekoe (Bontekoe 1646; translated by JPH), who in 1619 reported: 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 ­blue-­winged pigeon genus, and as two ­now-­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 1672–73 may have also observed them (see p. 141), and this is the last time that they were reported. The Réunion Blue Pigeon had probably disappeared by around 1700 due to ­over-­hunting and predation by introduced rats, as in 1704 the competent 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 generally all blue (Hume et al. 2006; Cheke & Hume 2008).

Rodrigues Blue Pigeon  Alectroenas sp. Specimens  A tarsometatarsus is in Tring. Status  Extinct. Known only from a single tarsometatarsus. Range  Rodrigues Island, Mascarene Islands.

A single tarsometatarsus, referable to the genus Alectroenas (JPH, in press), has shown that a species of blue pigeon once occurred on Rodrigues Island. 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 (1726) 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.

Providence Blue Pigeon  Alectroenas sp. Status  Extinct. Range  St Pierre and Providence Islands, Seychelles, ­south-­west Indian Ocean.

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 abundance, 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. 161

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Viti Levu Giant Pigeon Natunaornis gigoura

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.

Viti Levu Giant Pigeon  Natunaornis gigoura

Worthy

Natunaornis gigoura Worthy 2001, p. 767 (Udit Cave, Wainibuku, Viti Levu, Fiji) Specimens  Subfossil remains are in Wellington, New Zealand. 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 ­over-­hunting, and as for the Dodo and Solitaire, introduced mammalian predators would have been serious predators of eggs and chicks.

Lakeba Imperial Pigeon  Ducula lakeba

Worthy

Ducula lakeba Worthy 2001, p. 784 ­(Qara-­ni-­puqa Rockshelter, Lakeba Island, Fiji Group) Specimens  Subfossil remains are in Fiji and New Zealand. Status  Extinct. Known from Holocene subfossil remains. Date of extinction unknown. Range  Lakeba, Lau Group, Fiji.

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Lakeba Imperial Pigeon Ducula lakeba

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. 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 (see p. 154) 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).

‘Uvea Imperial Pigeon  Ducula david

Balouet & Olson

Ducula david Balouet & Olson 1987, p. 769 (Utuleve, Uvea, Wallis Group, South Pacific) Specimens  Subfossil remains are in New Caledonia. Status  Extinct. Known from late Holocene subfossil remains found in archeological middens. Date of extinction unknown. Range  ‘Uvea Island (Wallis Island), Wallis Group, South Pacific.

The Wallis Islands are situated 385km west of Samoa and 385km ­north-­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.

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Henderson Island Imperial Pigeon  Ducula harrisoni

Wragg & Worthy

Ducula harrisoni Wragg & Worthy 2006, p. 134 (Henderson Island) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Henderson Island, South Pacific.

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 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 ­as-­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.

Negros Spotted Imperial Pigeon  Ducula carola nigrorum

(Whitehead)

Ptilocolpa nigrorum Whitehead 1897, p. 34 (Negros, Philippines) Specimens  Specimens are in Chicago and Delaware. Status  Possibly extinct. Last recorded in the 1950s. Range  Negros and Siquijor, Philippines. Description  33cm (13in). Male head, neck and breast ­ash-­grey; white, ­crescent-­shaped band across centre of breast; blackish bar across lower breast; rest of underparts dark chestnut; upperparts grey tinged mauve, with dark markings and green gloss giving a spotted appearance; flight feathers and tail greenish black; bill base ­coral-­red tipped dull white; orbital skin ­blue-­grey; iris pale straw yellow; legs and feet ­coral-­pink. Female head and underparts darker; lacks white ­breast-­band; browner upperparts with spotting less discernible; and paler brown abdomen. Immature duller than female.

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.

Norfolk Island Pigeon  Hemiphaga novaeseelandiae spadicea

(Latham)

Chestnut Shouldered Pigeon. Latham 1801a, supplement p. 375 (Norfolk Island) Columba spadicea Latham 1801b, supplement, p. lx (no locality). Columba princeps Vigors 1833, p. 78 Specimens  Specimens are in Bremen; Cambridge, Massachusetts; Frankfurt; Leiden; Lisbon; Liverpool; Milan; Naples; New York; Philadelphia; Rouen; Tring; Vienna; Wiesbaden; and Lyons. Status  Extinct since the 1850s. Range  Formerly occurred on Norfolk Island. Description  50cm (19in). Head, foreneck and breast glossy deep green, paler on breast; shoulders deep ­chocolate-­red, or chestnut; hindneck glossed with copper; lower part of back, wings and tail pale blue; primaries dusky; breast, belly and vent white; bill deep red, tip yellow; iris crimson; legs and feet red.

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The New Zealand Pigeon or Kerer is known from three subspecies, nominate H. n. novaeseelandiae on North and South Islands and smaller offshore islands on New Zealand, H. n. chathamensis on the Chatham Islands, and H. n. spadicea on Norfolk Island (Gibbs et al. 2001). There may once have been a population on Lord Howe Island, and another on Raoul Island (see below). Norfolk Island was first discovered by Captain Cook in October 1774, after which Phillip Gidley King 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 The early settlers called the Norfolk Island Pigeon the ‘Wood Quest’ (Moore 1985), but no observations seem to have been made of the bird, and there is no record of when it was last seen in the wild. It was still common in 1838, as Ensign Best (in Moore 1985) mentioned that he and one of his companions killed at least 72 birds during 17 hunting expeditions, with the largest bag of 25 taken by two men on September 18, 1838. It must have disappeared extremely rapidly after this date as Peale (1848) failed to find it when he visited the island in 1839 with the United States Exploring Expedition. The last record of living birds was by Vigors (1833), when he received a pair at the Zoological Society, London. The birds he saw in New Zealand, and which he referred to as spadicea, were actually the nominate novaeseelandiae. ­Over-­hunting and predation by introduced mammals are the likely causes for this bird’s extinction.

Raoul Island Pigeon  Hemiphaga novaeseelandiae ssp. Specimens  The single humerus is in Auckland, New Zealand. Status  Known from a single humerus and a brief report. Disappeared sometime during the early 19th century. Range  Raoul (Sunday) Island, Kermadec Islands, South Pacific.

The New Zealand Pigeon or Kereru 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.

Lord Howe Island Pigeon  ? Hemiphaga sp. Very Large Pigeon from Lord Howe Island. Phillip 1789, p. 225 Status  Extinct. Known only from contemporary accounts. Date of extinction unknown. Range  Formerly occurred on Lord Howe Island.

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. 165

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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.

Strigopidae (Kakas, keas and kakapos) The New Zealand Kaka Nestor meridionalis, Kea N. notabilis, and Kakapo Strigops habroptila, are an ancient group of parrots that are considered distinct from the Psittacidae (Wright et al. 2008). They were once widespread, but all have declined due to habitat destruction and competition and predation from introduced animals. The Kakapo is now extirpated from mainland New Zealand and survives only on ­off-­shore island sanctuaries.

Norfolk Island Kaka  Nestor productus

(Gould)

Wilson’s Parrakeet Latham 1822, p. 170 (New South Wales; error) Plyctolophus productus Gould 1836a, p. 19 (no locality = Philip Island) Nestor productus Gould 1837a, pl. 25, fig. 1 (New South Wales or Norfolk Island) Nestor norfolcensis Pelzeln 1860a, p. 323 (based on an aberrant specimen) Specimens  Specimens are in Amsterdam; Göttingen; Leiden; Liverpool; Melbourne; New York; Philadelphia; Tring; and Washington, D. C. Status  Extinct. Last recorded in early 1850s. Range  Formerly occurred on Philip Island, and perhaps also on nearby Norfolk Island. Description  33cm (13in). Overall coloration ­greenish-­ash to brown, intermixed with orange; upper breast ­greyish-­brown; underparts variably mixed with yellow and dull orange; rump dull red; ­underwing-­coverts dull yellow; primaries brown marked on inner webs with whitish bars; thighs brown or ­reddish-­orange; tail brown; bill ­brownish-­grey; iris dark brown; legs and feet ­olive-­brown.

There is some confusion about the distribution and taxonomy of the Norfolk Island Kaka. All of the specimens 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. 383). This led some ornithologists, most notably Rothschild (1907a), to speculate that N. norfolcensis might have been an inhabitant of Lord Howe Island (see p. 333). All of these wild assumptions have fortunately now been quelled, and as the plumage of the Norfolk

Norfolk Island Kaka Nestor productus

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Island Kaka is extremely variable and bill deformities were apparently not uncommon in these parrots, N. norfolcensis 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 no information is available. 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 extermination 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: 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 ­white-­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 ­honey-­feeding Parrakeets . . . with a ­brush-­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.

Chatham Islands Kaka  Nestor sp. Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Chatham Islands.

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 possibly Lord Howe Island (see p. 333). Millener (1999) ­re-­examined the material, and has shown that the Chatham Island Kaka was indeed an endemic, undescribed species, and that it had reduced flying capabilities, hitherto only known among parrots in the Kakapo Strigops habroptilus and the ­Broad-­billed Parrot Lophopsittacus mauritianus of Mauritius (Hume 2007). No doubt a poorly volant, large terrestrial parrot would have been quickly exterminated once humans arrived on the islands.

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Psittacidae (Parrots and macaws) The Parrots and macaws are widely distributed, occurring in the Americas, Africa, Australia and eastwards through Polynesia in the Pacific. The main centre of distribution is the tropics, but the Carolina Parakeet 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.

Cebu Hanging Parrot (Colasisi)  Loriculus philippensis chrysonotus

Sclater

Loriculus chrysonotus Sclater 1872b, p. 324, pl. 11 (Cebu, Philippines) Specimens  Specimens are in Tring and Washington D. C. Status  Extinct since 1943, but doubtful, unconfirmed sightings in 2004. Range  Formerly inhabited the island of Cebu in the Philippines. Description  14cm (5.5in). The Philippine hanging parrots are overall green with red foreheads, chest patches and rumps, and varying amounts of yellow on crown and nape; bill ­coral-­red; iris dark brown; legs and feet orange. Cebu Hanging Parrot distinct in having frontal area red; rest of crown, nape and upper back rich golden yellow; fine reddish ­hind-­collar; more orange on lower part of ­throat-­patch; female similar but less distinct.

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 ­re-­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.

Siquijor Hanging Parrot  Loriculus philippensis siquijorensis

Steere

Loriculus Siquijorensis Steere 1890, p. 6 (Siquijor, Philippines) Specimens  Specimens are in Tring and Washington D. C. Status  Extinct since 1908. Range  Formerly occurred on Siquijor Island, Philippines. Description  14cm (5.5in). Similar to nominate Loriculus p. philippensis except that it lacks ­gold-­yellow to back of crown and head and gold-orange band to nape; smaller red bib; larger red ­crown-­patch. Females like males, except red throat and breast patch replaced with yellowish tinge; lores, sides of head and throat tinged pale blue.

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 168

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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 probably of escaped subspecies from elsewhere.

Sangihe ­Red-­and-­blue Lory  Eos histrio histrio

Müller

Psittacus histrio P. L. S. Müller 1776a, p. 76 (Sangir Island) Specimens  Specimens are in Leiden. Status  Possibly extinct. Last records of pure bred specimens occurred in the mid-1990s. Range  Sangihe, in the Sangihe Archipelago, Indonesia. Description  31cm (12in). Generally bright red and purplish blue, deeper red on rump. Forecrown red; broad purplish blue band on hindcrown; darker blue line from eyes to upper back; nape red; scapulars, flight feathers and thighs bluish black; ­wing-­coverts tipped with black; underparts red; broad blue band across breast; tail dark reddish purple; bill orange; bare skin around the eye black; iris red; legs and feet grey.

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 plantations, 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).

Challenger ­Red-­and-­blue Lory  Eos histrio challengeri

Salvadori

Eos indica Sclater 1878, p. 578 (nec. Gmelin) Eos histrio challengeri Salvadori 1891a, p. 22. Specimens  Specimens are in Tring. Status  Possibly extinct. No recent reports and last recorded in the late 19th century. Range  Uncertain; thought to be Miangas Island, Talaud Archipelago, between Mindanao and Sulawesi. Description  25cm (9.8in). Similar to nominate Eos histrio histrio, but smaller; blue colour on breast less extensive, and more mingled with red; also, blue line from eye does not reach blue mantle.

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).

Sinoto’s Lorikeet  Vini sinotoi

Steadman & Zarriello

Vini sinotoi Steadman & Zarriello 1987, p. 521 (Hane Site, Ua Huka, Marquesas Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known only from subfossil remains found in archaeological sites. Date of extinction unknown, but prior to the arrival of Europeans. Range  Marquesas Islands, South Pacific.

Sinoto’s Lorikeet was larger than any known Vini species. There appears to be size variation between specimens from different islands, with birds being larger on Hiva Oa and Tahuata than on Ua Huka (Steadman & Zarriello 1987). 169

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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.

Conquered Lorikeet  Vini vidivici

Steadman & Zarriello

Vini vidivici Steadman & Zarriello 1987, p. 523 (Hanatekua Shelter, Hiva Oa, Marquesas Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known only from subfossil remains found in archaeological sites. Date of extinction unknown, but prior to arrival of Europeans. Range  Marquesas Islands, South Pacific.

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).

New Caledonian Lorikeet  Charmosyna diadema

(Verreaux & Des Murs)

Psitteuteles diadema Verreaux & Des Murs 1860, p. 390 (New Caledonia) Specimens  The type specimen is in Paris. Status  Probably extinct. Known only from the type, a female, and a second female collected at the same time but now believed to be lost. Range  New Caledonia. Description  19cm (7.5in). Type female a small green parrot; paler on forehead, lores, ­ear-­coverts and sides of neck; cheeks and throat yellowish; crown blue; underparts pale green; vent red; tail green with the four outer feathers red at base and with a black band, yellowish on end of outer webs.

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.

Macquarie Island Parakeet  Cyanoramphus novaezelandiae erythrotis

(Wagler)

Psittacus erythrotis Wagler 1832, p. 426 (based on Vigors 1825) (Macquarie Island) Specimens  Specimens are in Tring. Status  Extinct. Last recorded in 1890. Range  Macquarie Island, ­south-­west Pacific. Description  27cm (10.5in). Generally green, darker above, paler and more yellowish below; forehead, crown, a band behind eyes and patch on either side of rump red; outer webs of the flight feathers ­violet-­blue; bill pale ­bluish-­grey, becoming dark grey towards tip, iris red; legs and feet ­greyish-­brown. Larger and paler than nominate C. n. novaezelandiae.

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 170

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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 abundant 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 Gallirallus philippensis macquariensis (see p. 90) – 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.

Lord Howe Parakeet  Cyanoramphus subflavescens

Salvadori

Parrot from Lord Howe Island. Phillip 1789, p. 225 Cyanoramphus subflavescens Salvadori 1891b, (Lord Howe Island) Specimens  Specimens are in Tring. Status  Extinct. Last reported in 1869. Range  Lord Howe Island. Description  27cm (10.5in). Uniform bright ­mid-­green washed with yellow, especially on cheeks and underparts; frontal region, lores and line thorough eye variably marked with small amounts of red; red patches on each side of rump; outer edges of ­flight-­feathers ­violet-­blue; uppertail green washed with brown; bill ­bluish-­grey, tipped ­blackish-­grey; iris dark red; legs and feet dark grey. Similar to the New Zealand Kakariki C. novaezelandiae but larger. Differed from C. cookii in being generally yellower in colour, with much less red on the forehead.

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. Hill (1870) observed a solitary pair in 1869, which proved to be the last record of their occurrence. 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).

Society Parrot  Cyanoramphus ulietanus

(Gmelin)

Society Parrot. Latham 1781, pp. 250–51, no. 52. Based on the type in the Leverian Museum, now in the Vienna Museum Psittacus ulietanus Gmelin 1788, p. 328 (Ulietea = Raiatea, Society Islands) Psittacus pacificus J. R. Forster 1844, p. 238 Platycercus tannaensis Finsch 1868, pp. 271–72 Psittacus fuscatus Pelzeln 1873, p. 30 Specimens  Specimens are in Tring and Vienna. Status  Extinct. Not recorded since the early 1770s. Range  Raiatea, Society Islands, South Pacific. Description  27cm (10.5in). Head ­blackish-­brown; upperparts deep ­olive-­green, each feather edged ­greyish-­brown giving a mottled effect; rump dull crimson; underparts olive yellow, each feather edged pale ­greyish-­brown; primaries and tail dusky, becoming ­cinereous-­brown towards tip; bill deep blue, with black tip; legs and feet black.

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). 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 171

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Society Parrot Cyanoramphus ulietanus

(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.

Black-fronted Parrot (Society Islands Parrot) Cyanoramphus zealandicus

(Latham)

Red-rumped Parrakeet. Latham 1781, p. 249. no. 50 Psittacus zealandicus Latham 1790, p. 102 (New Zealand; error = Tahiti) Platycercus pacificus var. 3. Vieillot 1823, p. 1387 Psittacus pacificus Wagler 1832, p. 524 (based on Forster; includes Kuhl’s erythronotus and novae-seelandiae as synonyms) Conurus phaëton Des Murs 1845, p. 449 Platycercus phaëton Des Murs 1849, pl. 16 (both Des Murs’s names appear to be based on the Paris specimen) Platycercus forsteri Finsch 1868, pp. 268–70 Specimens  Specimens are in Liverpool; Paris; and Tring. Status  Now extinct. Last collected in 1844. Range  Tahiti, Society Islands, South Pacific. Description  25cm (10in). Forehead dark purple; crown ­greenish-­chestnut, sides of head pale green; crimson streak from base of bill passing through eye; hindneck, neck, upper parts of body and ­wing-­coverts, dark green: middle of back pale ferruginous brown; rump crimson, with tinge of chestnut; underparts blackish green; primaries brown with bluish edges; secondaries dusky with green edges and pale ­rusty-­brown tips; tail bluish, the two middle feathers with green margins; bill deep blue, upper bill tipped black; legs and feet black.

This species was probably first obtained on Cook’s first voyage, as there is an unpublished painting in London based on type specimen by Sydney Parkinson, who had accompanied Cook on the voyage. It is plate 8, entitled ‘no. 5, Green Peroquit Otahite, Aá’ (Green Parakeet Tahiti, Parrot). Possibly at least two more were collected by Forster and Ellis on Cook’s second voyage. There are two specimens in Liverpool Museum; one of these, obtained from the collection of the Earl of Derby, is believed to have originally been in the possession of Sir Joseph Banks (subsequently in the Bullock Collection), and is therefore probably Latham’s type specimen (Latham 1781). It is a male, said to have come from Tahiti. The second specimen is a female, also from Tahiti, also from the Earl of Derby’s collection and apparently obtained by him from J. Leadbeater, a dealer whose specimens often lacked history. The last specimen was collected in 1844 by Lt M. J. de Marolles, and is now in Paris (Greenway 1967). 172

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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 mountains 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.

Paradise Parrot  Psephotus pulcherrimus

(Gould)

Platycercus pulcherrimus Gould 1845, p. 115 (Darling Downs, Queensland) Specimens  Specimens are in Brisbane; Brussels; Cambridge, Massachusetts; Dresden; Edinburgh; Exeter; Genoa; Glasgow; Helsinki; Leiden; Liverpool; Manchester; New Zealand; New York; Norwich; Oxford; Oslo; Paris; Philadelphia; Qatar; Queensland; Rouen; Stockholm; Sydney; Toronto; Tring; Victoria; Vienna; and Washington, D. C. Status  Now considered extinct. Last reliably recorded in November 1927. Range  Central and southern (and possibly northern) Queensland and northern New South Wales, Australia. Description  27–30cm (10.5–12in). Male forehead red; centre of crown black; area around eye yellow; cheeks, throat and breast ­emerald-­green to turquoise on sides of body; hindneck brownish black; back, lower back and flight feathers ­greyish-­brown edged blue; rest of wing, centre of abdomen, and thighs red; central ­tail-­feathers ­bronze-­green with ­blackish-­blue tips; lateral ­tail-­feathers ­greenish-­blue with ­blue-­white tips; bill grey; bare skin around eye grey; iris brown; legs and feet ­greyish-­brown. Female generally duller and lacking pronounced reds of male.

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 disappeared. 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. Habits The Paradise Parrot inhabited open savanna woodland and shrubby grassland, and was particularly associated with river valleys. 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. 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 considered rapid and undulating, but it spent almost all of its time on the ground.

Pacific Islands Eclectus Parrot  Eclectus infectus

Steadman

Eclectus infectus Steadman 2006c, p. 139 (Anatu, ‘Eua, Tonga) Specimens  Subfossil remains are in Florida.

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Extinct Birds Status  Extinct. Known only from subfossil remains. Date of extinction unknown, but may have been as recently as the 18th century. Range  ‘Eua, Lifuka, Uiha and Vava’u, Tonga; possibly also Vanuatu and Fiji.

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. 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  Psittacula bensoni

(Holyoak)

Lophopsittacus bensoni Holyoak 1973, pp. 417–18, pl. 8 (Mauritius) Psittacula bensoni Hume 2007, p. 17 Specimens  Subfossil material of the Mauritian species is in Cambridge, England; Mauritius and Paris. Status  Extinct. Known from subfossil remains and contemporary accounts. Last recorded in 1732 on Réunion and 1759 on Mauritius. Range  Mauritius and Réunion, Mascarene Islands.

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. 180). ­Re-­examination of the material, however, has indicated that it was in fact a grey, ­long-­tailed ­Psittacula-­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) and it appears that no live or dead bird was ever taken out of Réunion.

Seychelles Parakeet  Psittacula wardi

(Newton)

Palaeornis wardi E. Newton 1867a, p. 346 (Seychelles) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Liverpool; New York; Paris; and Tring. Status  Extinct. Disappeared some time between 1881 and 1906. Range  Formerly occurred on the islands of Mahé, Silhouette and possibly Praslin, Seychelles. Description  41cm (16in). Male generally green; back of head, nape and narrow stripes to cheek washed with blue; broad ­cheek-­stripe and obscure narrow line from cere to eye black; abdomen yellowish green; ­purple-­red patch on ­wing-­coverts; upperside of central ­tail-­feathers blue with yellow tips, lateral feathers green; underside of tail yellow; bill red with yellow tip; iris yellowish; feet grey. Female lacked black ­cheek-­stripe; immature as female, but with shorter ­tail-­feathers.

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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 is often regarded as conspecific. However, the osteology of the Seychelles bird suggests that is quite distinct (Hume 2007). 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 only 155km2 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, but 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) 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 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.

Rodrigues ­Ring-­necked Parakeet (Newton’s Parakeet) Psittacula exsul

(Newton)

Perroquets verds et bleus. Leguat 1708, pp. 67, 107 Palaeornis exsul A. Newton 1872, p. 33 (Rodrigues) Specimens  Specimens are in Cambridge, England. Status  Extinct. Last seen in 1875. Range  Formerly occurred on Rodrigues, Mascarene Islands. Description  40cm (16in). Male overall colour ­slate-­blue, not green; underside lighter ­grey-­blue; dark line from cere to eye; black collar from base of mandible to sides of neck; rostrum ­reddish-­brown; mandible black; iris yellow; feet and legs grey. Female differs from male in head greyer head; black collar less prominent and did not extend to back of neck; bill entirely black.

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: 175

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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 (1726) stated that there were three kinds of parrot on the island (see p. 178). The first was the Rodrigues Parrot Necropsittacus rodericanus. The second was almost certainly 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 the female P. exsul. This suggests that there were originally blue and green colour morphs, of which only the blue morph has been preserved (Hume 2007). 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 ­south-­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 (1726) account (above) describing a little red above the wings ­(shoulder-­patch) is characteristic of P. eupatria, and not P. krameri.

Réunion ­Ring-­necked Parakeet  Psittacula eques

(Boddaert)

Psittaca borbonica torquata Brisson 1760, p. 328, pl. 27, fig. 1. Psittacus alexandri var [gamma] Linnaeus 1766, p. 142 Perruche à double collier. Buffon 1770–83, p. 143 Perruche à collier, de l’Isle de Bourbon. Daubenton 1765–1780, pl. 215 Alexandrine Parrakeet, var C. Double-ringed Parrakeet. Latham 1781, p. 326 Psittacus eques Boddaert 1783, p. 13 Psittica torquata Var. B. Latham 1822, p. 161 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 ­blackish-­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 176

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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 type 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 supposed skin in the Royal Museum, Edinburgh (Jones 1987), might be the specimen used for Martinet’s illustration, but its provenance remains in doubt (Cheke & Hume 2008). 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).

Mascarene Parrot  Mascarinus mascarinus

(Linnaeus)

Psittacus mascarinus Brisson 1760, p. 315 Psittacus mascarin Linnaeus 1767–71, p. 524 (‘Mascarina’ = Réunion) Mascarinus obscurus Bonaparte 1854b, p. 154, no. 207 Mascarinus duboisi Forbes 1879, p. 304–06 Specimens  Specimens are in Paris and Vienna; subfossil remains are in Réunion. Status  Extinct. Last recorded at the end of the 19th century. Range  Formerly occurred on Réunion, and perhaps also on Mauritius. Description  35cm (14in). Head and neck ­ash-­grey; back, wings, rump and underparts very dark ­ash-­grey; tail dark ­ash-­grey, with lateral feathers white at base; bare skin around eye region, bill, bare skin around upper bill and iris bright red; legs and feet pale flesh; claws ­grey-­brown.

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. 174], and yet others, smaller, are green [Réunion ­Ring-­necked Parakeet; see p. 176]. 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: 177

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Mascarene Parrot Mascarinus mascarinus

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. The ­often-­quoted last living specimen was depicted by Hahn (1834) in the menagerie 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): bill red; base of bill black; head and throat ­bluish-­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. Taxonomy The relationships of this aberrant parrot are difficult to determine, and it has been duly placed in a number of genera, most recently with the Coracopsis black parrots of Madagascar, the Comoros and the Seychelles ,based on molecular evidence (Kundu et al. 2011). This seems extraordinary; the Mascarene Parrot differs drastically in morphology. It would also have had to evolve these differences within a short time-frame, as between 300,000 and 180,000 years ago the island’s original fauna was obliterated by explosive volcanic events (see Mourer-Chauviré et al. 1999). The bird may have recolonised Réunion from elsewhere after these events, as Kundu et al. (2011) suggest that the divergence time from Coracopsis was 4.6 million years, long before Réunion’s emergence from the sea, but no physical evidence has been found on any other island to substantiate this. Forbes (1879) considered it unlike any other parrot, and based on the distinctive large red bill, Garrod (1874), Forbes (1879) and Oustalet (1897) suggested a close relationship between Mascarinus and the psittaculine genera Tanygnathus and Psittacula. Anatomically, the Mascarene Parrot appears to belong in Psittaculini, differing in proportions of the cranium and jaws compared with Coracopsis (Hume 2007). Peter Mundy (Temple 1914) spoke of ‘russet parrots’ on Mauritius, which may refer to this species, but again no confirmatory fossil remains have been found.

Rodrigues Parrot  Necropsittacus rodericanus

(Milne-Edwards)

Psittacus rodericanus Milne-Edwards 1867, p. 151 (Rodrigues) Necropsittacus rodericanus ­Milne-­Edwards 1874a, p. 18 Specimens  Subfossil remains are in Cambridge, England; London; Rodrigues; and Tring. Status  Became extinct sometime after 1761. Known from subfossil bones and from contemporary accounts. Range  Rodrigues, Mascarene Islands.

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Rodrigues Parrot Necropsittacus rodericanus

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 1726: 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. Milne-Edwards (1867) described this species from a portion of maxilla (upper bill). Later, Günther & Newton (1879) described a ­near-­complete set of skull and postcranial bones. These subfossil remains were assumed to refer to the parrot described by Tafforet (1726). It was considered to be a large cockatoo-sized 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 179

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confused the issue, and described another species, Necropsittacus francicus (see p. 398), 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 introduced by human agency.

Broad-billed Parrot (Raven Parrot)  Lophopsittacus mauritianus

(Owen)

Psittacus mauritianus Owen 1866b, pp. 168–171 (Mauritius) Specimens  Subfossil material is in Cambridge, England; Mauritius; and Paris. Status  Known from bones, traveller’s accounts and illustrations. Became extinct sometime after 1674. Range  Mauritius, Mascarene Islands.

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 ­cockatoo-­sized parrot had once been a contemporary of the Dodo, but it was now extinct. Subsequently, Newton (1875b, 1876) reported the discovery of an illustration of a large parrot in the unpublished 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 might suggest dark coloration and 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

Broad-billed Parrot Lophopsittacus mauritianus

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species was flightless (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 but Hoffman in 1673–75 (1680, in Cassel 1931) left the best description: There are also geese, flamingos, three species of pigeon of varied colours, mottled and green perroquets [parakeets], red crows with recurved beaks and with blue heads, which fly with difficulty and have received from the Dutch the name of ‘Indian crow’. This was the last time it was mentioned. Being large and presumably easy to catch, it probably disappeared as a result of hunting, deforestation and nest predation by introduced monkeys and rats. 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 ­weak-­jawed or frugivorous as postulated by Holyoak (1971). Hachisuka’s (1953) statement 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. 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 1874a; 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.

Glaucous Macaw  Anodorhynchus glaucus

(Vieillot)

Guacamayo azul. Azara 1802, p. 402, no. 273 Macrocercus glaucus Vieillot 1816, p. 259 Specimens  Specimens are in Leiden; Liverpool: Tring; and Washington, D. C.; there is a skeleton in Leiden. Status  Not definitely seen since 1951 and now believed to be extinct. Range  Formerly occurred in a limited area in Paraguay, northern Argentina and ­south-­eastern Brazil. Description  68–72cm (26.5–28in). A large bird, though smaller than the familiar Hyacinth Macaw Anodorhynchus hyacinthinus; generally ­greenish-­blue in colour with a glaucous ‘bloom’; head and underparts with grey tinge; uppertail ­greenish-­blue; bill dark ­blackish-­grey, bare lappets at base of mandible yellow; bare orbital skin yellow; iris dark brown; legs and feet dark grey.

The Glaucous Macaw is a ­little-­known species; formerly in demand as a cage bird, 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 1951, and there are reports from Paraná in the early 1960s, but inhabitants in areas where it was known to occur had no knowledge of it. Very few specimens are known to survive. It formerly inhabited Paraguay, the provinces of Misiones and Corrientes in northern Argentina, ­south-­west Brazil and possibly also the Artigas area of ­north-­western Uruguay. The reasons for its disappearance 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

Glaucous Macaw Anodorhynchus glaucus

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tracts remain in other parts of the species’ former range, e.g. in ­south-­east Paraguay and ­north-­east Argentina. Almost nothing was known of its habits. In 1805, Azara (1802–1805) reported that it nested in holes in trees and cliffs, and laid two eggs; two other nineteenth-century explorers also saw them nesting on cliffs in different places (Collar et al. 1992). Apparently no further observations on its breeding were ever made. It was probably dependent on the Yatay Palm Butia yatay, the only palm in its range with the appropriate size of nut (Yamashita & Valle 1993).

Cuban Macaw  Ara tricolor

Bechstein

Le petit Ara. Daubenton 1770–86, Planches Enluminées, no. 641 L’Ara tricolor. Levaillant 1801, p. 13, pl. 5 Ara tricolor Bechstein 1811, p. 64, pl. 1 (South America, based on Levaillant; = Cuba) Sittace? lichtensteini Wagler (in Bonaparte 1856b) Ara cubensis Wetherbee 1985, p. 169 Specimens  Specimens are in Berlin; Cambridge, Massachusetts; Dresden; Görlitz; Havana; Leiden; Liverpool; Tring; New York; Paris; Stockholm; and Washington, D. C.; subfossil remains are in Cuba. Status  Extinct since about 1855. Known from skins and subfossil remains. Range  Formerly occurred on Cuba and the Isle of Pines, West Indies. Description  50cm (20in). Forehead red; crown yellowish; hindneck yellow; upper back red with green edges to feathers; ­wing-­coverts darker red; rump and ­uppertail-­coverts pale blue; wings ­purplish-­blue; dorsal surface of tail red below, blue above with red bases to the feathers; underparts red; ­undertail-­coverts blue; bill dark.

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. Only two species, the Cuban Macaw Ara tricolor and Puerto Rican Macaw A. autochthones (see p. 185) are known from skins and subfossil bones (Wetmore 1937a; Olson & Maíz 2008). There is illustrative and comparatively ­well-­documented evidence for a species of macaw on Jamaica and another, perhaps supported by subfossil remains, on Guadeloupe (see p. 183) (Williams & Steadman 2001, but see Olson & Maíz 2001); 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 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. 183) and Amazona martinicana (see p. 339), both of which he justified as being valid. All other supposed macaw species known only from accounts are open to alternative explanation. The Cuban Macaw The Cuban Macaw is the only extinct West Indian macaw known from skin specimens, and to have survived into historic times. But scarcely any information is known even about this species. Cuban natives called it ‘Guacamáyo’, simply meaning ‘macaw’, and they apparently not only killed the birds for food, but also took them from the nest to rear as pets. Gundlach (1856, 1876), who was an authority on the species, reported the bird only from the Zapata swamp and the Ensenada de Cochinos on the south coast, but it supposedly occurred in many parts of Cuba and on the Isle of Pines. Its range probably extended into Pinar del Río 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. 182

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The last known specimen was supposedly shot (but not collected) at La Vega, near the Ciénaga de Zapata on the south coast, in 1864 (Zappey, in Bangs & Zappey 1905), but this hearsay information was given to Zappey by the plantation owner where the bird was 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. Although Gundlach found the flesh ­evil-­smelling and ­foul-­tasting, ­over-­hunting for food by the locals 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. It fed on fruit, seeds, sprouts and buds. Small populations apparently 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 inextricably linked with this food source. Interestingly, on examination of a museum skin of the Cuban Macaw, a new species of chewing louse, Psittacobrosus bechsteini, was discovered. This insect was almost certainly unique to the macaw, and disappeared along with its host (Mey 2005). Thus the extinction of a single species can also spell doom for smaller organisms such as parasites that depend on it to survive.

Guadeloupe Macaw  Ara guadeloupensis

Clark

Les Arras. Du Tertre 1654, p. 294 Ara guadeloupensis Clark 1905, p. 272 Status  Presumed to have become extinct in the early 18th century. Known only from accounts, an illustration in Du Tertre (1667), and perhaps subfossil remains. Range  Guadeloupe, West Indies.

The Guadeloupe Macaw is as ­well-­documented as any bird can be in the absence of extant specimens. Several authors describe it in considerable detail, and all are agreed as to its appearance. 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 1905) 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 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 183

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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. 338) and Guadeloupe Parakeet (see p. 336). 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 ­tropical-­island birds. Labat Labat (1742, quoted by Clark 1905) 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 & Maíz (2001) 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. 185) 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 relatively 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. Paintings Daubenton’s (1779) 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.

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Puerto Rican Macaw  Ara autochthones

Wetmore

Ara autochthones Wetmore 1937a, p. 12 (St. Croix) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Puerto Rico and St Croix.

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 ­pre-­Columbian archaeological material on Puerto Rico (Olson & Maíz 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.

Mona Island Conure  Aratinga chloroptera maugei

(Souancé)

Psittacara Maugei Souancé 1856, p. 59 (no locality = Puerto Rico) Conurus gundlachi Cabanis 1881, p. 5 (Mona Island) Specimens  Specimens are in Chicago; Leiden; and Paris. Status  Last seen in 1892, and now extinct. Range  Formerly occurred on Mona Island and probably Puerto Rico. Description  32cm (12.5in). Generally green with forehead sometimes narrowly marked with red; bend of wing and ­underwing-­coverts red; red feathers often scattered through body plumage; bill ­pinkish-­yellow; bare skin around eye ­greyish-­white; iris ­yellowish-­orange; legs and feet grey. Compared to nominate A. c. chloroptera slightly smaller; bill darker; red lighter on under primary coverts; and lesser primary coverts entirely green without any red feathering.

The extinction of this subspecies is believed to have been due to forest destruction, but the date is uncertain. The last specimen was taken on Mona Island in 1892 by the collector W. W. Brown. The bird, if it had occurred on Puerto Rico, had already disappeared by that time. Greenway (1967) suggested that this occurred as early as 1860, and cited Stahl (1887) as saying that in 1883 it was known only from memories of the parents of the oldest inhabitants. However Gundlach (1874b) believed that it was still to be found in the eastern forests. Peters (1937) believed it still survived on Mona Island, but Bond (1956) pronounced it extinct. Greenway (1967) doubted that the species differed in any significant way from the nominate Hispaniolan Conure A. c. chloroptera.

Sinú Painted Parakeet  Pyrrhura picta subandina

Todd

Pyrrhura subandina Todd 1917, p. 6 (Jaraquiel, Bolivar, Colombia) Specimens  Specimens are in Pittsburgh and Washington, D. C. Status  Possibly extinct, last reliably recorded in 1949. Range  Sinú valley, Córdoba, ­north-­east Colombia. Description  22cm (8.5in). Generally green; frontal band and lores ­reddish-­brown; cheeks ­bluish-­green; ­ear-­coverts ­yellowish-­brown; feathers on side of neck, throat and upper breast edged with ­greyish-­buff; rump, central belly and tip to the tail maroon; ­primary-­coverts and outer webs of primaries blue; leading edge of wing red; bill ­brownish-­grey; iris ­yellowish-­brown; legs and feet grey.

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). 185

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Culebra Island Amazon  Amazona vittata gracilipes

Ridgway

Amazona vittata gracilipes Ridgway 1915, p. 106 (Culebra Island) Specimens  Specimens are in Washington D. C. Status  Extinct since 1899. Range  Culebra Island, off the eastern coast of Puerto Rico. Description  28cm (11in). Smaller than the Puerto Rico Amazon A. vittata, and had relatively smaller and more slender feet.

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; 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.

Aruba Amazon  Amazona barbadensis canifrons

(Lawrence)

Chrysotis canifrons Lawrence 1883, p. 381 (Aruba Island) Status  Known only from the type collected in 1882, now lost. The last reports date from the mid-1940s. Range  Formerly occurred on Aruba Island, Barbados. 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) 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.

Eastern Carolina Parakeet  Conuropsis carolinensis carolinensis

(Linnaeus)

Psittacus carolinensis Linnaeus 1758, p. 97 (Carolina and Virginia) Range  Florida to Virginia, eastern United States. Status  Extinct. Last definite record was February 21, 1918.

Western Carolina Parakeet (Louisiana Carolina Parakeet) Conuropsis carolinensis ludovicianus

(Gmelin)

Psittacus ludovicianus Gmelin 1788, p. 347 (Louisiana) Specimens  Luther (1996) lists about 720 skins and 16 skeletons in museums around the world. Of the 49 reported eggs, McKinley (1977) only assigns about 20 with certainty to the Carolina Parakeet.

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Psittacidae Range  Mississippi–Missouri basin, eastern United States. Status  Extinct since c.1910. Description  30cm (12in). The Carolina Parakeet was a small green parrot with a long tail, a yellow and orange head, yellow flanks, and some yellow on the wings. Male and female alike. Overall coloration green, paler on under parts; forehead, lores, bare skin around eye and upper cheeks orange, rest of the head and upper part of neck, bend of wing, carpal edge and thighs yellow; outer webs of primaries yellow at base; tail green; bill yellowish horn; iris light brown; legs and feet ­pinkish-­brown. Juvenile forehead, lores and bare skin around eye brownish orange; rest of head and body green; no yellow on bend of wing and thighs. The western race differed from the eastern by being greener, with less bluish tint; having less yellow on wings; and being generally paler.

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. 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) maintained 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. Reasons for extinction There seems to be little doubt that humans were directly responsible for the extinction of the Carolina Parakeet, but the exact reasons are unclear. They were shot in great numbers because of the damage done to fruiting crops, and it was easy to kill entire flocks, as when one was shot, the others would return to hover and scream over the dead. Furthermore, as they became rarer, the capturing of live birds for profit intensified. Their preferred habitat remained comparatively intact, so habitat destruction is probably not to blame. Nor is disease likely to have been an important factor, as this would have caused a much more rapid decline than in fact occurred. McKinley (1960) has suggested competition over nest sites with introduced bees as a possible cause, but the true reasons will now never be known; the United States’ only endemic parrot was lost before conservation actions could be put into place. The Carolina Parakeet represented a monotypic genus, not obviously related to any other group, and otherwise known only from a fossil ancestor, Conuropsis fraterculus (Wetmore 1926b), described from a single bone from Miocene deposits in Nebraska (though Olson 1985a casts doubt on this attribution). Captive birds Carolina Parakeets were often kept in captivity where they bred regularly, but mortality of adults and juveniles 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 187

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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). Habits 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 paroquets 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. Vocalisations comprised loud screams that could be heard over long distances, and quieter chattering (Snyder & Russell 2002).

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 threatened, but some island subspecies have declined due to deforestation.

St Helena Dwarf Cuckoo  Nannococcyx psix

Olson

Nannococcyx psix Olson 1975, p. 31 (in diagnosis of genus), p. 32 (Prosperous Bay Valley, St Helena) Specimens  The type is in Cambridge, Massachusetts. Status  Extinct since the early 16th century. Known only from a subfossil humerus. Range  St Helena, South Atlantic.

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St Helena Dwarf Cuckoo Nannococcyx psix

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 1975). 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 beginning of the 16th century, when most of the forests were destroyed.

Delalande’s Coua ­(Snail-­eating Coua)  Coua delalandei

(Temminck)

Coccycus Delalandei Temminck 1827, livr. 74, pl. 440 (Madagascar) Specimens  Specimens are in Brussels; Cambridge, Massachusetts; Leiden; Liverpool; Antananarivo, Madagascar; New York; Paris; Philadelphia; Stuttgart; Tring; and Vienna. Status  Extinct. Not reliably recorded since 1834. Range  Humid forests of the ­north-­east coast of Madagascar. Description  56cm (22in). The largest of the couas, being predominantly dark glossy blue above and chestnut and white below: head blue with an area of blue skin around eye, surrounded by black feathers; back blue; tail with greenish tinge, outer feathers with white tips; throat and upper breast white; belly chestnut; bill, legs and feet black; iris brown.

Delalande’s Coua was known principally from Île ­Sainte-­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). Peters (1940) described its range as: Formerly occurred on the island of Sainte Marie and opposite mainland of eastern Madagascar, probably from the head of Antongil Bay southward to Tamatave. Now extirpated on the former island and said to be confined only to a few patches of forest between Fito and Maroantsetra. However, there appears to be no direct evidence that it occurred on the mainland. All the known specimens (about thirteen) came from Île ­Sainte-­Marie, so there is a possibility that it was confined to the island, at least in recent times. Lavauden (1932) said that birds were occasionally trapped by natives near Fito and Maroantsetra, apparently during the 1920s, and that the natives prized the feathers. Rand (1936) was of the 189

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Delalande’s Coua Coua delelandei

opinion that it was extinct since none had been seen for some years, and Delacour (1932) noted that all the forests in the area had been destroyed. Large prizes were offered in 1932 to dealers in Antananarivo (Collar & Stuart 1985), but no specimens were forthcoming. Its extinction was probably due to deforestation, and perhaps ­over-­hunting, but introduced rats may have depleted snails, a major food source (Collar & Stuart 1985). 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 impatience, 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 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.

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Ancient Coua  Coua primaeva

­Milne-­Edwards & Grandidier 

Coua primaeva ­Milne-­Edwards & Grandidier 1895, p. 11 (Ankevo, Madagascar) Specimens  The type specimen is in Paris. Status  Extinct. Known only from Holocene subfossil remains. Range  Madagascar.

Among a collection of subfossil remains of birds from Madagascar, ­Milne-­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.

Bertha’s Coua  Coua berthae

Goodman & Ravoavy

Coua berthae Goodman & Ravoavy 1993, p. 26 (Grotte d’Anjohibe, Grotte Principale, Province of Mahajanga, Madagascar) Specimens  Subfossil remains are in Antananarivo, Madagascar and Paris. Status  Extinct. Known only from Holocene subfossil remains. Range  Madagascar.

This is the largest member of the genus Coua, living or extinct. Characteristics of the available skeletal material show that Bertha’s Coua was a large, ­swift-­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.

Assumption Island Coucal  Centropus toulou assumptionis

Nicoll

Centropus assumptionis Nicoll 1906b, p.105 (Assumption Island, Seychelles) Specimens  Specimens are in Tring and Washington D. C. Status  Extinct. Not recorded since the 1920s. Range  Assumption Island, Seychelles. Description  40cm (16in). Similar to C. t. insularis of Aldabra Island, but considerably smaller; wings darker maroon, especially underside of quills.

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 extinction 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 (Barn owls) Barn owls are nocturnal predatory birds with disproportionately large heads and ­heart-­shaped faces. They have a wide distribution, occurring in varied habitats on continental land masses and on a number of oceanic islands, and are divided into two genera. Many barn owls are poorly known, and some have only been recorded on a few occasions since their discovery. Deforestation is the main threat to island species and subspecies.

Puerto Rican Barn Owl  Tyto cavatica

Wetmore

Tyto cavatica Wetmore 1920, p. 80 (Cueva Torano, Puerto Rico) Specimens  Subfossil remains are in New York. Status  Extinct. Known only from subfossil remains. May have survived until 1912. Range  Puerto Rico.

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 ­cave-­roosting owls from as recently as 1912, which may have been references to this bird.

New Caledonian Barn Owl  ?Tyto letocarti

Balouet & Olson

Tyto? letocarti Balouet & Olson 1989, p. 18 (Gilles Cave, 5km WSW of Boulouparis, New Caledonia) Specimens  Subfossil remains are in Paris. Status  Extinct. Known from subfossil material only. Range  New Caledonia, Melanesia.

This owl, although clearly a member of Tytonidae, is very distinct, and pending the discovery of more material, 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. 36), the New Caledonian Wood Rail Tricholimnas lafresnayanus (see p. 96) and the New Caledonian ­Owlet-­nightjar Aegotheles savesi (see p. 352). 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.

Hispaniolan Barn Owl  Tyto ostologa

Wetmore

Tyto ostologa Wetmore 1922a, p. 3 (St. Michel de l’Atalye, Hispaniola) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Range  Hispaniola, West Indies.

This was an extremely large barn owl, with a tarsus as robust as that of a Snowy Owl Bubo scandiacus, but much longer (Wetmore 1922a). It was discovered in cave deposits along with a host of now-extinct West 192

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Indian mammals. Compared to Tyto neddi (below) it had long toes, which may have enabled it to prey on arboreal primates and thick-skinned edentates (Steadman & Hilgartner 1999). In particular, it may have been the chief predator of the recently discovered (but equally extinct) Hispaniolan monkey Antillothrix bernensis. The owl’s extinction can be blamed on deforestation and loss of prey species.

Barbudan Barn Owl  Tyto neddi

Steadman & Hilgartner

Tyto neddi Steadman & Hilgartner 1999, p. 76 (Pocket, Gun Shop Cliff, Two Foot Bay, Barbuda) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Range  Barbuda, West Indies.

The Barbudan Barn Owl T. neddi was one of the larger West Indian Tyto species, and the first to be described from the Lesser Antilles. As Barbuda was once connected to Antigua, it is likely that this species also occurred there (Steadman & Hilgartner 1999). Barbuda once harboured three species of owl – the Burrowing Owl Athene cunicularia, an undescribed Tyto sp., probably American Barn Owl T. alba pratincola, and T. neddi, 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 Barn Owl Tyto sp. took reptiles and small–medium mammals, while T. neddi 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, including hutias (Capromyidae) and rice rats Oryzomys sp., which resulted in the loss of the Barbudan Barn Owl’s preferred large prey.

Bahaman Barn Owl  Tyto pollens

Wetmore

Tyto pollens Wetmore 1937b, p. 436.(Great Exuma Island, Bahamas) Specimens  Subfossil remains are in Cambridge, Massachusetts. Status  Extinct. May have survived to the 16th century. Range  Great Exuma Island and possibly Andros Island, Bahamas.

Bahaman Barn Owl Tyto pollens with Bahaman Hutia Geocapromys ingrahami

This was another extremely large barn owl, larger than Tyto ostologa of Hispaniola (Wetmore 1937b). The tarsometatarsus was extremely robust, almost twice the width as in the extant American Barn Owl Tyto alba pratincola, but it was comparatively short, being about 15% longer (Brodkorb 1959). 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 sp., 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 stands 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 a bird called the ‘chickcharnie’, 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, chickcharnies would unleash terrible misfortune. 193

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Strigidae (True owls) The Strigidae are cosmopolitan in distribution and found on all continents except Antarctica. Their affinities are still unresolved and there is some doubt that true owls should be separated from the barn owls. The Strigidae have been divided into 25 genera; they are cryptic, almost exclusively nocturnal birds, that can be difficult to observe. In general, they are not considered threatened, but island populations have been severely affected by deforestation. Some species are known only from very few specimens.

Mauritian Lizard Owl (Sauzier’s Owl) Mascarenotus sauzieri

(Newton & Gadow)

Strix sauzieri Newton & Gadow 1893, pp. 286–88, pl. 33, figs. 11–18 (Mauritius) Strix sp. Newton & Gadow 1893, pp. 287–88 Scops commersoni Oustalet 1896, p. 35, fig. 3 (Mauritius) Strix newtoni Rothschild 1907a, p.79 Mascarenotus sauzieri ­Mourer-­Chauviré et al. 1994, p. 1699 Specimens  Subfossil remains are in Cambridge, England; Paris; and Tring; Jossigny’s illustration is in Paris. Status  Extinct. Known only from subfossil remains, contemporary accounts and illustrations. Range  Mauritius, Mascarene Islands. Description  Length c.42cm. A large, eared owl with bare tarsi. Upperparts dark brown, with feathers of head, neck and back edged with rufous; tail ­rufous-­brown marbled with light rufous; wings brown, banded with irregular bands of white, pale buff and brown; throat and sides of body whitish, rest of undersides dark buff with whitish spots.

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 evolution 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), presumably 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 deforestation 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, ­white-­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.

Réunion Lizard Owl  Mascarenotus grucheti

Mourer-­Chauviré et al.

Mascarenotus grucheti ­Mourer-­Chauviré et al. 1994, p. 1704 (Réunion) Specimens  Subfossil remains are in Lyon; Paris; and Réunion. Status  Extinct. Range  Réunion, Mascarene Islands.

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 probably a secretive inhabitant of dense and remote forests. It was approximately the same size as M. sauzieri but 194

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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).

Rodrigues Lizard Owl  Mascarenotus murivorus

(Milne-Edwards)

Strix (Athene) murivora Milne-Edwards 1874a, p. 13 (Rodrigues) Strix sp. Milne-Edwards 1874a. Bubo leguati Rothschild 1907a, p. 71. Based on Strix sp. of ­Milne-­Edwards. Specimens  Subfossil remains are in Cambridge, England; Paris; Rodrigues; and Tring. Status  Extinct. Disappeared between 1726 and 1761. Known from subfossil remains and contemporary accounts. Range  Rodrigues, Mascarene Islands.

The Rodrigues Lizard Owl was briefly mentioned by Leguat (1708) in 1691–93, and described by Tafforet (1726) in 1726, 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. In 1761 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. This was probably a result of severe deforestation caused by tortoise hunters burning off the vegetation. All of the Mascarene ­lizard-­owls probably nested in ­tree-­holes, so deforestation would have been particularly devastating for these birds.

Kaua’i Stilt Owl  Grallistrix auceps

Olson & James

Grallistrix auceps Olson & James 1991, p. 72 (Makawehi dunes, Kaua’i) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Kaua’i, Hawaiian Islands.

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. 194) and some other owl 195

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Kauai Stilt Owl Grallistrix auceps pursuing Kauai ’O’o Moho braccatus

genera, parallel the skeletal proportions of ­forest-­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.

Maui Stilt Owl  Grallistrix erdmani

Olson & James

Grallistrix erdmani Olson & James 1991, p. 80 (Owl cave near Puu Makua, Maui) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Maui, Hawaiian Islands.

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 altitudes 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 Sceloglaux albifacies did on New Zealand (Worthy & Holdaway 2002).

Oahu Stilt Owl  Grallistrix orion

Olson & James

Grallistrix orion Olson & James 1991, p. 74 (Barbers Point, Oahu) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Oahu, Hawaiian Islands.

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. 196

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Molokai Stilt Owl  Grallistrix geleches

Olson & James

Grallistrix geleches Olson & James 1991, p. 76 (Moomomi dunes, Molokai) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Molokai, Hawaiian Islands.

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.

Socorro Elf Owl  Micrathene whitneyi graysoni

Ridgway

Micrathene graysoni Ridgway 1886, p. 333 (Socorro Island) Specimens  Specimens are in Washington, D. C. Status  Probably extinct. Last recorded around 1970. Range  Socorro Island, Revillagigedo Islands, off Baja California. Description  12.5–14.5cm (5–5.75in). Generally brownish above with irregular rusty to orange spotting, becoming lighter on crown and head; underside brown with orange streaks; tip of scapulars, primaries and secondaries white; tail with light rusty bands with brown interspaces; eyebrows white; facial disc white to orange; bill greyish horn with pale yellow tip; iris yellow. Browner than nominate, which is more ­rusty-­brown on upperparts, with grey wash.

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 introduced 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.

Marie-­Galante Burrowing Owl  Athene cunicularia guadeloupensis

(Ridgway)

Speotyto guadeloupensis Ridgway 1874, p. 90 footnote. (‘Guadeloupe’; error = ­Marie-­Galante) Specimens  Specimens are in Cambridge, Massachusetts; Guadeloupe; New York; Tring; and Washington, D. C. Status  Extinct since the end of the 19th century, but probably never common. Range  Formerly occurred on ­Marie-­Galante Island, West Indies. Description  22cm (8.5in). Darker and more reddish in colour than other West Indian races of Burrowing Owl; spots smaller and less numerous; eyebrow brown rather than whitish; underparts darker but with narrower bars.

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 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 197

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(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 ­Marie-­Galante.

Antigua Burrowing Owl  Athene cunicularia amaura

(Lawrence)

Speotyto amaura Lawrence 1878b, p. 234 (Antigua, Nevis and St Kitts) Specimens  Specimens are in Chicago; New York; Tring; and Washington, D. C. Status  Extinct since the end of the 19th century. Range  Formerly occurred on Antigua, Nevis and St Kitts, West Indies. Description  20–22cm (8–8.5in). Upperparts deep brown marked with light fulvous spots; primaries ­blackish-­brown; tail dark brown barred ­reddish-­fulvous; superciliary streak pale fulvous; cheeks dark brown, feathers tipped fulvous; throat pale ­whitish-­buff, broad dark brown band across throat; underside fulvous, barred dark brown; bill light yellow, sides of upper bill black; feet dull ­yellowish-­brown; iris bright yellow. Female more strongly barred on abdomen.

The Antigua Burrowing Owl appears to have possessed demonic status among the inhabitants of the islands on which it occurred, and was much feared (Lawrence 1878b). Like the ­Marie-­Galante population (see p. 197), it was probably never common. Only five specimens are known. Lawrence could not find a single bird when on Antigua, so offered a reward, and within three hours of doing so obtained three living birds which had been dug out of a cliff in the chalk-hills. There are sight records from St Kitts, but no specimens were collected there. Bond (1950) cited the introduction of the mongoose as the cause of its extinction. Habits All observers agreed that the bird was nocturnal, and lived in old drains, holes in cliffs, and ruined walls. It was locally called ­Coo-­coo from its hoot at night. Captive birds frequently uttered a chattering cry, but they did not hoot. Lawrence (1878b) was told the owls fed on lizards and mice.

Lord Howe Boobook (Lord Howe Morepork) Ninox novaeseelandiae albaria

Ramsay

Ninox albaria Ramsay 1888, p. 36 (Lord Howe Island) Specimens  Specimens are in Sydney and Tring. Status  Extinct since about 1940. Range  Formerly occurred on Lord Howe Island. Description  30cm (12in). Generally ­rich-­fawn to ­rusty-­brown, richer on sides of chest; face, lores, forehead, chin, and upper throat buff or ­snow-­white; breast irregularly barred white; wings and tail rufous, barred dull ash; bill black; feet dull ­yellowish-­brown; iris golden yellow. Paler than nominate N. n. novaeseelandiae, and larger than Norfolk Island race N. n. undulata.

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 that it inhabited the native forests as well as occurring around the settlements on the island. 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 unsuccessful 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.

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Norfolk Island Boobook  Ninox novaeseelandiae undulata

(Latham)

Strix undulata Latham 1801a, supplement, p. 17 (Norfolk Island) Specimens  Specimens are in Liverpool; New York; and Tring. Status  Extinct. The last individual, a female, died in 1996. Range  Norfolk Island. Description  30cm (12in). Generally ­rusty-­brown above barred white, rufous below; wings barred buff and white; head and throat ­rusty-­brown; facial disc white to buff; bill ­bluish-­black; feet dull ­yellowish-­brown; iris golden yellow. Nominate N. n. novaeseelandiae less barred above, less ­rusty-­brown below.

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 ­pure-­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). This tragic scenario epitomises the finality of extinction, when a ­last-­ditch effort to save a species can only result in hybrid birds. 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).

South Island Laughing Owl  Sceloglaux albifacies albifacies

(Gray)

Athene albifacies G. R. Gray 1844b, p. 2. pl. 1 (Waikouaiti, South Island, New Zealand) Specimens  Specimens are in Bremen; Cambridge, England; Cambridge, Massachusetts; Christchurch, Dunedin, and Wellington, New Zealand; Edinburgh; Liverpool; New York; Norwich, England; Tring; and Washington, D. C. For a comprehensive listing see Worthy (1997). Status  Now almost certainly extinct. Last collected in July 1914, but existed until at least the 1920s. Range  Formerly widespread in South Island and Stewart Island, New Zealand. Description  32cm (13in). A large owl with ­yellowish-­brown plumage striped with brown; white stripes on scapulars (sometimes also feathers edged with white rather than ­yellowish-­brown on hind-neck and mantle); face white behind and below eyes, greyish towards the centre, feathers with brown shaft-lines; wings and tail brown with ­brownish-­white bars; bill black, ­horn-­coloured towards tip; iris ­yellowish-­brown; tarsus (feathered) yellowish to reddish-buff; toes ­flesh-­brown or pale yellow; claws black.

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 in 1927 at Lake Wairaumoana (Blackburn 1982), 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 199

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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), 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. Laughing Owls formerly also inhabited the Chatham Islands. These birds may have constituted an undescribed and ­now-­extinct subspecies. Laughing Owls in captivity Buller (1906) found that that Laughing Owls were extremely tame and docile birds. He kept some in captivity, and found that they allowed themselves to be handled freely without any attempt to bite or use their claws. He described the experience of a man who caught one: Mr. Jacobs, the taxidermist at Masterton, has given me a characteristic account of a very fine example which he obtained alive at Nelson, and afterwards mounted for the local Museum. A man, so he informed me, was travelling from Nelson to the West Coast, when he observed a large Owl squatting on the ground near the roadside. He dismounted from his horse and caught the bird. Then, selecting a retired nook in the adjoining woods, he drove a thick pole into the ground and secured his captive to it by the leg, allowing a sufficient of flax to permit of the Owl moving freely about over the ground. On his return by the same road two days later he found that the bird had snapped, or in some way had got disengaged from, the flax string, and was perched on the top of the pole, permitting itself to be recaptured without the slightest resistance. He took it on with him to Nelson, and, not knowing its value, sold it to the narrator for a few shillings. It now graces the collection in the Nelson Museum. 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, particularly worms. The call was reported to be a loud ­coo-­e-­e, while captive birds variously uttered a low chuckle like a turkey, a c­ at-­like mew, a p ­ uppy-­like yelp, and a tuneless whistle (Gurney 1896; Greenway 1967). Their diet varied depending on geographical location, feeding predominantly on petrels and parakeets in areas where they were abundant, or frogs, lizards and bats in others (Worthy & Holdaway 2002). They also preyed on rails, pigeons, small birds, lizards (including Tuatara Sphenodon punctatus), and even the extant New Zealand Morepork Ninox novaeseelandiae and extinct New Zealand ­Owlet-­nightjar Aegotheles novae­ zealandiae (see p. 203).

North Island Laughing Owl  Sceloglaux albifacies rufifacies

Buller

Sceloglaux rufifacies Buller 1904, p. 639 (Wairarapa, North Island, New Zealand) Specimens  The type specimen appears to be lost (Worthy 1997). Status  Known only from the type (now lost) and one other specimen. Last reliably reported in the 1889, but may have survived into the early 20th century. Range  North Island, New Zealand. Description  32–33cm (13–13.5in). Face dull ­rufous-­brown; crown and nape ­blackish-­brown, strongly marked rufous; primaries barred and edged rufous; tail uniform ­yellowish-­brown, faintly barred pale brown; bill lemon yellow; feet dull yellow; iris probably ­yellowish-­brown. Nominate S. a. albifacies was less rufous, larger, and had a white face (not ­rufous-­brown).

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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). A very doubtful sighting was made in the Kaimanawa Range around 1930 (Williams & Harrison 1972). Oliver (1930) stated that only two specimens were ever collected in North Island, the first by a taxidermist 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. The bird was apparently ­well-­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. Buller and the specimens Buller (1906) discussed the two specimens in some detail. Of the first specimen, he said: On my last visit to Nelson I was informed by Mr. Martin, the well-known taxidermist, and a most observant man, that when he came out to the Colony in 1856, in the ship ‘Creswell’, he was acting as servant to Captain King, a gentleman much devoted to natural history. Martin, who had been a pupil of Gardner, of Oxford Street, was employed by his master in making a collection of native birds; and almost immediately after landing at Wellington, he proceeded up the West Coast, shooting and skinning birds as he went, till he had collected enough to fill several large cases. When camped in the woods under Mount Egmont, in the Taranaki district, one evening at dusk a large bird flew over the party, making a loud laughing noise – ha, ha, ha! Martin shot the bird, and it proved to be an Owl. It was nearly as large as the specimen of Sceloglaux albifacies in the Nelson Museum, but much darker in plumage, and without any appearance of white on the face. The eyes, as he perfectly remembers, were of a dark-brown colour. This was impressed on his mind at the time, because he had shot and skinned many specimens of the Morepork [the other New Zealand owl, which still survives], of much smaller size, and with vivid yellow eyes. What ultimately became of Captain King’s collection nobody knows. On learning this story, Buller more critically examined the type specimen as it was the only one at that time known to have come from North Island, and commented: When I was living in Wellington, in the summer of 1868–69, the [type] was sent in to the Colonial Museum, freshly killed, by a settler in the Wairarapa district, about fifty miles from the city. It was carefully skinned and the sex determined by the Museum taxidermist, and the specimen placed in the type collection. I took an early opportunity of comparing it with the only specimen of Sceloglaux albifacies in the Museum collection – one received some time before from Otago – and although appreciably smaller in size, I took it to be a dark phase of that species. Two years later, I came to England to publish the first edition of my “Birds of New Zealand”, and . . . brought with me this owl, with other skins of New Zealand birds, for the purpose of having them mounted for exhibition in the Museum. This task was entrusted to the late Mr. Burton of Wardour Street. During this operation the tail of the Owl, probably on account of some defect in it, was replaced by one from another species; but it was so well matched that, although the bird remained in the Colonial Museum for five and twenty years, the counterfeit tail was never detected by Sir James Hector or myself. Rothschild (1907), after examining the type, found it to be a ­subadult, 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. 201

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New Caledonian Nightjar  Eurostopodus exsul

Mayr

Eurostopodus mystacalis exsul Mayr 1941b, p.6 (Tao, ­north-­western New Caledonia) Specimens  The type specimen is in New York. Status  Possibly extinct. Known from one specimen, taken in 1939. Range  Tao, ­north-­western New Caledonia. Description  26cm (10in). Differed from Eurostopodus mystacalis in being smaller, lighter above with a solid black crown, and in wing coloration. This was a ­medium-­sized ­silvery-­grey nightjar, lacking a collar around the ­hind-­neck and white markings in tail; throat patches white; small white bar in middle of outer wing.

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 ­White-­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.

Jamaican Pauraque (Jamaican Poorwill)  Siphonorhis americana

(Linnaeus)

Caprimulgus americanus Linnaeus 1758, p. 193 (‘America calidore’ = Jamaica) Specimens  Specimens are in New York and Tring; subfossil remains are in Florida. Status  Probably extinct. Known from only four specimens. Range  Jamaica. Description  23–25cm (9–10in). Male ­rufous-­brown, mottled and streaked with greyish and dark brown: scapulars heavily blotched with black, laterally streaked with whitish; ­wing-­coverts with buff terminal spots; primaries deep brown, irregularly banded with rufous, lighter on the outer webs; tail feathers dull rufous, barred with dark brown, a dark brown subterminal bar and a white terminal bar; throat and breast dark brown, with a white band across throat; lower breast tipped and barred with dull white; abdomen fawn, barred with brown. Female (juvenile?) slightly paler; white patch on throat duller; tail tipped buff instead of white.

All four specimens of this nightjar were taken in the western part of the island, in Westmoreland and Freeman’s Hall, in Trelawney. 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 Trelawney; 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 1998), but if it does still exist it must be extremely rare. 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, ­semi-­arid woodland or in more open country.

Cuban Pauraque (Cuban Poorwill)  Siphonorhis daiquiri

Olson

Siphonorhis daiquiri Olson 1985b, p. 528 (Cueva de Los Indios, near Daiquirí, Santiago de Cuba) Specimens  Subfossil remains are in Moscow and Washington, D. C. Status  Probably extinct. Known only from subfossil remains, but may have persisted into recent times. Range  Cuba.

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 202

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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 ­hot-­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.

New Zealand ­Owlet-­nightjar  Aegotheles novaezealandiae

(Scarlett)

Megaegotheles novaezealandiae Scarlett 1968, p. 254 Specimens  Subfossil remains are in Auckland and Canterbury, New Zealand. Status  Extinct. Disappeared after the arrival of Polynesians. Range  North and South Islands, New Zealand.

The New Zealand Owlet-nightjar was the largest known species in the family, living or extinct, and weighed an estimated New Zealand Owlet-nightjar 150–200 g, making it also the heaviest in Aegotheles novaezealandiae 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 Sceloglaux albifacies, which may have been its chief predator. It disappeared before European settlement 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 ­isotope­derived ages from ­owlet-­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. 203

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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.

Mangaia Swiftlet  Aerodramus manuoi

(Steadman)

Collocalia manuoi Steadman 2002a, p. 327 (Ana Manuku, Keia District, Mangaia, Cook Islands) Specimens  Subfossil remains are in Florida. Status  Extinct. Known only from subfossil remains. Range  Mangaia, Cook Islands, South Pacific.

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 Swiftlet Aerodramus manuoi

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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 disappearance. 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).

Southern Vanuatu ­White-­rumped Swiftlet Aerodramus spodiopygius ingens

(Salomonsen)

Collocalia spodiopygius ingens Salomonsen 1983, p. 80 (Eromanga Island = Erromango Island, New Hebrides = Vanuatu) Specimens  The type specimen is in New York. Status  Probably extinct. Not recorded since 1971. Range  Aneityum, Erromango and Tanna Islands, Vanuatu, South Pacific. Description  10cm (4in). Generally sooty black above with diagnostic dusky grey underparts; face dark brown, contrasting with paler underparts; rump conspicuously marked white; tail black, slightly forked; tarsi heavily feathered. Greater face contrast than in nominate; also darker underparts, and more heavily feathered tarsi.

The ­White-­rumped Swiftlet Aerodramus spodiopygius is divided into eleven subspecies. It occurs on many islands throughout Melanesia including Fiji, New Caledonia, Samoa, the Solomon Islands, Tonga, Vanuatu, and the Wallis group (Chantler 1999; Chantler & Driessens 2000). The Vanuatu race was first collected, but not described, on March 27, 1937 by L. Macmillan on Erromango Island in the south of Vanuatu (Salomonsen 1983). It also occurred on Aneityum and Tanna. It must have disappeared between 1937 and 1971, as Lord Medway, an authority on Collocalia swiftlets, failed to locate any birds during an intensive survey by the Royal Society Percy Sladen Expedition to Vanuatu in 1971 (Medway & Marshall 1975). The reasons for the bird’s disappearance are unknown. A closely related subspecies, A. s. epiensis, still occurs on islands in north and central Vanuatu. Habits Nothing is known about the Vanuatu birds, but the nominate A. s. spodiopygius feed above the forest canopy or under the trees near the ground. They are ­cave-­breeding birds, using echolocation when underground, and cementing their nests to the cave walls with fine plant fibres mixed with saliva.

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, 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 individuals. 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 extinction due to habitat destruction, the island endemics particularly so.

Coppery Thorntail  Discosura (Popelairia) letitiae

(Bourcier & Mulsant)

Trochilus letitiae Bourcier & Mulsant 1852, p. 143 (Bolivia) Specimens  The type specimen is in Tring. Status  Probably extinct. Known only from two males of uncertain locality. None has been seen for more than a century. Range  Bolivia. Description  c.9cm. (3.5in). Male crown and all underside glittering light green; back of neck and back golden or ­orange-­green (depending on light); ­uppertail-­coverts purplish red; tail purplish violet with green reflections on tips of two central tail feathers; wings ­purplish-­brown; tarsi white; upper bill ­flesh-­coloured at base, black for the remainder; lower bill flesh-coloured, tip black.

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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 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.

Elegant Hummingbird  Chlorostilbon elegans

(Gould)

Erythronota? elegans Gould 1860, p. 307 (no locality) Specimens  The type specimen is in Tring. Status  Known only from the unique type, an adult male. Previously believed to be an aberration, but now considered to be a valid but extinct species. Range  Possibly Jamaica, or the northern Bahamas. Description  10cm (4in). Upperparts golden glittering green; underparts darker golden glittering green; tail ­dark-­purple bronze; ­undertail-­coverts ­bronze-­green edged dusky white; bill blackish, pinkish at base of lower bill.

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 predators caused its Brace’s Emerald extinction. Chlorostilbon bracei

Brace’s Emerald

Chlorostilbon bracei

(Lawrence)

Sporadinus bracei Lawrence 1877, p. 50–52 (New Providence, Bahamas) Specimens  The type specimen is in Washington D. C. Status  Extinct. Known from the type, a male collected in 1877, and from subfossil remains. Range  New Providence, Bahamas. Description  9.5cm (3.75in). Male head and back bronze green, back with golden hue; white spot behind eye; throat ­blue-­green; abdomen green, feathers with ­ash-­grey tips; wings purplish; rectrices greenish; ­undertail-­coverts around cloacal opening (crissum) grey with indistinct cinnamon hue at edges; bill, legs and feet black. Female unknown.

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The type specimen of Brace’s Emerald was collected by Lewis J. K. Brace on July 13, 1877 near Nassau, New Providence (Graves & Olson 1987). Initially it was considered an aberrant Cuban Emerald Chlorostilbon ricordii, and synonymised with it. Bond (1936) recognised the distinctiveness of C. bracei and described it as a new subspecies, noting that it had a longer bill and a different plumage. Subfossil remains of three species of hummingbirds were discovered in a Pleistocene cave deposit on New Providence, Bahamas (Olson & Hilgartner 1982), and one of these species was identified as C. bracei (Graves & Olson 1987). This not only proved the validity of Brace’s Hummingbird, it also showed that the species had survived as a Pleistocene relict population on the Bahamas until the late 19th century. The reason for its extinction is unclear, but anthropogenic changes to the island were no doubt to blame.

Alfaro’s Hummingbird  Amazilia (Saucerottia) alfaroana

Underwood

Amazilia alfaroana Underwood p.441 (Volcán de Miravalles, Costa Rica) Specimens  The type specimen is in Tring. Status  Probably extinct. No records since the beginning of the 20th century. Range  Costa Rica. Description  10cm (4in). Similar to Blue–vented Hummingbird A. hoffmanni, but larger and bill more curved. Generally bronzegreen above with blue tint, more intense on crown and forehead; underside glittering green, with white thighs and blue vent; undertail-coverts steel-blue, with narrow white margin; tail blue-black; lower bill pale for one-third its basal length, the rest (and upper bill) black.

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.

Alejandro Selkirk Island Firecrown Sephanoides fernandensis leyboldi

(Gould)

Eustephanus leyboldi  Gould 1870, p. 406 (Alejandro Selkirk Island, Juan Fernández Archipelago) Specimens  Specimens are in Dublin and Tring. Status  Probably extinct. Last recorded in 1908. Range  Alejandro Selkirk Island and Juan Fernández Island, eastern Pacific. 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 introduced 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 laid in a small ­cup-­shaped nest typically 3–4m above ground, and almost exclusively in Luma apiculata. 207

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Bogotá Sun-angel Heliangelus zusii

Graves

Bogotá Sun-angel Heliangelus zusii

Heliangelus zusii Graves 1993, Auk 110, p. 4. (no locality) Specimens  The type specimen is in Philadelphia. Status  Extinct. Known only from a single specimen of unknown origin purchased in 1909 in Bogotá. Range  Possibly Colombia. Description  12cm (4.75in). A dark ­bluish-­black hummingbird with a deeply forked tail. Male general coloration dark bluish black; underside duller; probable white spot behind eye; frontlet and gorget bright green; lower back and rump deep ­greenish-­blue; primaries and secondaries dark brown with purplish tint; tail glittering dark purple; bill black; legs and feet brown.

The only known specimen (thought to be a male) of this sun-angel was purchased as a trade skin in Bogotá, so its provenance is uncertain (Graves 1993). There have been doubts as to the validity of this species, but mitochondrial DNA analysis has shown that it is valid, and possibly occupied a restricted distribution between the upper tropical and temperate zones of the northern Andes (Kirchman et al. 2010). Severe deforestation is widespread in the lower limits of the northern Andes, so it is likely that the population disappeared due to habitat destruction. Nothing was recorded about its habits.

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 thorny scrub. They all have restricted ranges, and have declined due to the severe deforestation that has taken place on Madagascar.

Ampoza ­Ground-­roller  Brachypteracias langrandi

Goodman

Brachypteracias langrandi Goodman 2000, p. 318–322 (Ampoza, Madagascar) Specimens  The type specimen is in New York. Status  Extinct. Known only from a single humerus. Date of extinction unknown. Range  ­South-­west Madagascar.

The Ampoza ­Ground-­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 ­Ground-­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.

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Halcyonidae

Halcyonidae (Kingfishers) The kingfishers are generally small to ­medium-­sized fish- and ­insect-­eating 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.

Mangareva Kingfisher  Halcyon gambieri gambieri

Oustalet

Halcyon Gambieri Oustalet 1895, p. 182 (Mangareva, Gambier Archipelago, Tuamotus) Specimens  The type specimen is in Paris. Status  Known only from the type, collected in 1841. Range  Mangareva 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.

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 Mangareva, about 6.4km long, and the highest peak, Mount Duff, is 380m high. Mangareva’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 Mangareva 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 1977), 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).

Mangareva Kingfisher Halcyon gambieri gambieri

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). 209

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Javan ­Blue-­banded Kingfisher  Alcedo euryzona euryzona

Temminck

Alcedo euryzona Temminck 1830, plate 508 (Java) Specimens  Subfossil remains are in New York and Washington, D. C. Status  Probably extinct. Last specimen taken in 1937. Range  Java, Indonesia. Description  20cm (8in). Male uniform dark blackish upperparts with ­silvery-­azure streak from upper back to rump; ­neck-­flash long, white with rufous tip; throat white; underparts pale ­rufous-­buff with bold blue band across chest. Female rich rufous underparts; lacks ­breast-­band; bill bluish black; iris dark brown; legs and feet ­orange-­red.

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 appears to always have been rare. It was a lowland riverine forest inhabitant, preferring ­slow-­moving streams and larger rivers, and suffered a severe population decline with the destruction of its habitat. The scale of deforestation in Java is alarming, and there is little hope for its continued survival.

Sangihe Dwarf Kingfisher  Ceyx fallax sangirensis

Meyer & Wiglesworth

Ceyx fallax sangirensis Meyer & Wiglesworth 1898, p. 278 (Sangihe Island, Indonesia) Specimens  A specimen is in Dresden. Status  Possibly extinct. Not seen since 1997. Range  Sangihe Island, Indonesia. Description  12cm (5.75in). Generally a uniform ­rufous-­orange; top of head blue; ­blue-­lilac on cheeks and hindneck; mantle ­rufous-­brown; ­wing-­coverts dark rufous brown; primaries and secondaries dark brown; lower back, rump and ­uppertail-­coverts bright ­violet-­blue; tail black; bill ­orange-­red; iris dark brown; legs and feet orange-red. Larger than nominate C. f. fallax, with larger blue markings on crown and more violaceous rump.

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.

Sakaraha Pygmy Kingfisher  Ceyx madagascariensis dilutus

(Benson)

Ispidina madagascariensis dilutus Benson, 1974, p.186 (Sakaraha, ­south-­west Madagascar) Specimens  A specimen is in Tring. Status  Probably extinct. Known from one specimen, taken in 1974. Range  Sakaraha, ­south-­west Madagascar. Description  13cm (5in). Generally bright rufous above; spot on side of neck; throat, breast and belly white; ­wing-­coverts dark ­rufous-­brown; primaries black with rufous edges; lower back, rump, ­uppertail-­coverts and tail deep rufous; bill dark blackish at base, red merging to light orange at tip; iris dark ­reddish-­brown; legs and feet orange-red. Compared to the generally darker nominate C. m. madagascariensis, lacks lilac wash on head, lower back and ­uppertail-­coverts.

The nominate race of Madagascar Pygmy Kingfisher C. m. madagascariensis is widely distributed in Madagascar, but it was replaced in the arid ­south-­west by the Sakaraha subspecies, which was only 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. 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. 210

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Upupidae

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 exclusively on the ground and has a strong odour, which acts as a defence against predators.

St Helena Giant Hoopoe  Upupa antaios

Olson

Upupa epops Ashmole 1963b, p. 405 Upupa antaios S.L. Olson 1975b, p. 32 (Prosperous Bay Valley, St. Helena) Specimens  Subfossil remains are in Tring and Washington, D. C. Status  Extinct since some time after 1502. Known only from subfossil remains. Range  Formerly occurred on St Helena.

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 noticeably more decurved than in Common Hoopoe. 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 ­pre-­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 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 Giant Hoopoe. It was probably exterminated by introduced rats and cats, severe deforestation, and perhaps loss of invertebrate prey items.

St Helena Giant Hoopoe Upupa antaios with a Giant Earwig Labidura herculeana

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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.

Ticao Tarictic Hornbill  Penelopides panini ticaensis

Hachisuka

Penelopides panini ticaensis Hachisuka 1930, p.169 (Ticao, Philippines) Specimens  A specimen is in Manila, Philippines. 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 ­over-­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 predominantly fruit, but ­small-­vertebrate and invertebrate prey was also consumed, particularly during the breeding season.

Lifou Hornbill  Aceros sp. Specimens  Subfossil remains are in Florida. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Lifou, Loyalty Islands, New Caledonia.

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.

Capitonidae (New World barbets) The New World barbets occur in Central and South America and are divided into two genera. They are generally brightly coloured, stocky birds with robust bills. Because they occur in humid forests, some species are endangered due to habitat destruction. 212

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Northern ­White-­mantled Barbet  Capito hypoleucus hypoleucus

Salvin

Capito hypoleucus Salvin 1897c, p.16 (Valdivia, State of Antioquia, Colombia) Specimens  Specimens are in Tring and Washington D. C. Status  Possibly extinct. Not recorded since the late 1940s. Range  Northern tip of the central Andes, Colombia. Description  19cm (7.5in). Forehead and crown scarlet-red; hindcrown and upper back black and white; sides of head and rest of upperparts ­bluish-­black; throat and upper breast white with diffuse dusky ­breast-­band; underparts ­yellowish-­white, more yellowish on flanks; bill pale ­yellow-­horn.

The ­White-­mantled Barbet is restricted to the ­north-­central Andes and western slope of the east Andes in Colombia. It is divided into three subspecies, with the nominate C. h. hypoleucus in the central Andes of ­north-­west Colombia (from Serranía de San Lucas in Bolivar to north Antioquia), C. h. carrikeri in the Botero area of Antioquia (see below), and C. h. extinctus in the Magdalena Valley (Graves 1986). The nominate northern form was rather restricted in range and has not been recorded since the late 1940s. The habitat in which it was once found has been almost entirely destroyed by commercial logging, agriculture (including the narcotics industry), clearance of forest for livestock, and mining (Stiles et al. 1999). If it still survives, its ­long-­term future is bleak. Habits Virtually nothing is known about the Northern ­White-­mantled Barbet. The only surviving subspecies of the ­White-­mantled Barbet, C. hypoleucus extinctus, is adaptable, occurring in lower montane humid forest at 400–2,000m, including anthropogenically disturbed areas, but it appears to prefer primary forest above 1,000m (Short & Horne 2002). The diet consists primarily of seeds, fruit and insects, and the bird appears to be nomadic in response to local fruit abundance (Stiles et al. 1999).

Botero ­White-­mantled Barbet  Capito hypoleucus carrikeri

Graves

Capito hypoleucus carrikeri Graves 1986, pp. 61–62 (Rio Porce, Antioquia) Specimens  The type specimen is in Washington D. C. Status  Possibly extinct. Not recorded since the 1950s. Range  Botero area of Rio Porce, Antioquia, Colombia. Description  19cm (7.5in). Differed from nominate C. h. hypoleucus in having more yellow in the white of the back and underparts.

The Botero ­White-­mantled Barbet has not been recorded since the 1950s, and may now be extinct. Its disappearance can be attributed to the severe deforestation that has taken place in Colombia. There may perhaps be very small populations still in existence, but there are no recent records. Of the three races of this species, only the southern population C. h. extinctus in the Magdalena Valley appears to survive; it has responded to secondary forest growth since the logging companies have departed. The Magdalena race was thought to be extinct in 1986, but it was rediscovered in 1990 (Short & Horne 2002).

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 ­Ivory-­billed Woodpecker C. bairdi; all are now almost certainly extinct.

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Guadalupe Flicker  Colaptes cafer rufipileus

Ridgway

Colaptes mexicanus rufipileus Ridgway 1876b, p. 191 (Guadalupe Island) Specimens  Specimens are in Berkeley, California; Chicago; Cambridge, Massachusetts; New York; Providence, Rhode Island; Tring; and Washington, D. C. Status  Extinct. Not seen since 1906. Range  Guadalupe Island, off Baja California, Mexico. Description  32cm (12.5in). Adult generally ­cinnamon-­brown with black bars on back and wings; forehead bright tawny; hindneck and rear neck grey; rump ­pinkish-­white merging to ­blackish-­brown tail; upper breast with black necklace; lower breast and belly barred black on white with ­buff-­yellow wash; bill black or slaty; iris deep brown; legs and feet grey. Males had black or red moustachial stripe.

The ­Red-­shafted Flicker C. c. 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. 83). 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 (1906) reported that the birds were very tame and fell easy prey to them. Thayer & Bangs (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.

Imperial Woodpecker  Campephilus imperialis

(Gould)

Picus imperialis Gould 1832, p. 140 (‘California’ = Jalisco, Mexico) Specimens  Known from about 120 museum specimens in many museums around the world including Berlin; Denver; Indiana; Leiden; London; Michigan; New York; Tring; Vienna; and Washington, D. C. Status  Almost certainly extinct. Last confirmed sighting in April or May 1956. Range  The Sierra Madre Occidentale of Mexico, in the states of Sonora, Chihuahua, Nayarit, Durango, Zacatecas, Jalisco and Michoacán. Description  56–60cm (22–24in). Similar to the Ivory-billed Woodpecker Campephilus principalis, but much larger, and lacking the white stripe down the side of the face and neck. Male almost uniform glossy greenish black apart from ­red-­sided crest, ­white-­tipped inner primaries; white secondaries; and white scapular stripe that does not extend on neck. Female similar but crest all black and (unlike C. principalis) recurved at the top; bill ­ivory-­white; iris ­golden-­yellow; legs and feet black.

At a meeting of the Zoological Society of London, held on August 14, 1832, specimens were exhibited of a previously 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: 214

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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 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.

Imperial Woodpecker Campephilus imperialis (female left, male right)

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 dedicated 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 delicacy 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 interviewed 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 agricultural poison to the trunks of foraged trees. No doubt this persecution, along 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 215

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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 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.

Ivory-billed Woodpecker  Campephilus principalis

(Linnaeus)

Picus principalis Linnaeus 1758, p. 113 (North America = South Carolina, based on Catesby 1731–43) Specimens   There are many skins in museums around the world including Cambridge and Springfield, Massachusetts; Chicago; Hamburg; New York; Paris; Princeton, New Jersey; Rouen; Springfield, Illinois; San Francisco; Syracuse, New York State; Tring; and Washington, D. C. Status  Probably extinct by the 1960s. Range  Formerly occurred over the ­south-­eastern lowlands of the United States, from North Carolina to Florida, and in the Mississippi Valley to southern Indiana and southern Illinois, extending along the coast of Texas to the Brazos River. In later years confined to a small area of forest in Louisiana. Description  A glossy black bird with purple reflections; white stripe commencing on each cheek continued down each side of neck meeting in middle of back; outer halves of the secondaries and ends of inner primaries white, forming a large white patch on rear of wing. Sexes alike except male had a prominent scarlet crest, female’s crest entirely black. Bill ­ivory-­white; iris bright yellow; legs and feet black.

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 populations 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. 216

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Picidae

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 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 undisputed observation of a living ­Ivory-­billed Woodpecker has been made since 1944. The species is now almost certainly extinct. Recent mitochondrial DNA analysis (Fleischer et al. 2006) has shown that the ­Ivory-­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 sedentary, 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 Ivorybillled 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.

Cuban ­Ivory-­billed Woodpecker  Campephilus bairdii

Cassin

Campephilus Bairdii Cassin 1863, p. 322 (Monte Verde, Cuba) Specimens  Specimens are in Berlin; Cambridge, Massachusetts; Chicago; Guantánamo and Havana, Cuba; New York; Philadelphia; and Washington, D. C. Status  Now probably extinct. Last reliably recorded in March 1987. Range  Cuba.

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Extinct Birds Description  Male similar to the male Ivory-billed Woodpecker C. principalis in having white nasal tufts and a white neckstripe, but differed in the neck-stripe being continued to the base of the bill, the black feathers of the crown not reaching to the tip of the red crest. The female was similar to the male, differing only in having the crest entirely glossy blue-black.

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 ­north-­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 population 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 ­Ivory-­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 mountains, 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.

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Acanthisittidae

A note on the Passeriformes (Passerines) The passerines form the largest order of birds. It contains around 5,000 species, more than all the other bird orders combined. The Passeriformes is divided into three suborders, the Tyranni (suboscines), Passeri (oscines) and the Acanthisitti (New Zealand wrens). Apart from the basal Acanthisitti, the suborders are further divided into a large number of families, of which the taxonomy and relationships of many remain unresolved. They are an extremely diverse group, with a vast array of sizes, colours and bill shapes. Cosmopolitan in distribution, the passerines are absent only from the extreme polar regions, and have populated even the most remote island archipelagos. The passerines appear to have suffered relatively few extinctions. Some families, such as the birds of paradise, have no confirmed losses, whereas the New Zealand wrens and Hawaiian finches have suffered almost complete species loss, and at least one family, the Hawaiian mohos or Mohoidae, is now entirely extinct. Because passerines are generally small, inconspicuous birds and thus are less likely to have been recorded in life or preserved in the fossil record, the actual number of extinctions that have taken place is likely to be much higher.

Acanthisittidae (New Zealand wrens) The New Zealand wrens are small, almost ­tail-­less birds that are considered distinct enough from all other passerines to be given suborder status. The family comprised seven species, of which five were flightless, unique amongst passerines except for the ­Long-­legged Bunting Emberiza alcoveri of the Canary Islands (see p. 322). Only two species now survive, the Rifleman Acanthisitta chloris, considered common on South Island but restricted on North Island, and the Rock Wren Xenicus gilviventris, which is endangered. The New Zealand wrens are generally terrestrial birds, foraging and nesting near or on the ground, which made them particularly susceptible to introduced predators. At least three were victims of Polynesian introductions, whereas the other extinct species disappeared after colonisation of the islands by Europeans and their commensal animals.

North Island Bush Wren  Xenicus longipes stokesii

Gray

Xenicus stokesii G. R. Gray 1862b, p. 219 (Rima-taka Hills, New Zealand) Specimens  Specimens are in Liverpool and Tring; subfossil remains are in Wellington, New Zealand. Status  Became extinct around 1955. It had been rare and restricted in range for many years. Range  North Island, New Zealand. Description  9cm (3.5in). Similar to the South Island Bush Wren X. l. longipes but distinguished by shining ­slate-­blue sides of neck; chest tinged, in certain lights, greenish; and patch of pure yellow feathers on the yellowish-green flanks.

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, 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 ­ground-­nesting birds.

Stead’s Bush Wren  Xenicus longipes variabilis

Stead

Xenicus longipes variabilis Stead 1936, p. 313. (islands ­south-­west of Stewart Island) Specimens  Specimens are in Canterbury and Wellington, New Zealand; Tring; and Washington, D. C. Status  Extinct; last recorded in 1972. Range  Formerly occurred on Stewart Island, Solomon Island, Big South Cape Island, and South Cape Island, New Zealand.

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Extinct Birds 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 undisturbed. 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 specimens 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 transferred 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 sometimes 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 disproportionately 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.

South Island Bush Wren  Xenicus longipes longipes

(Gmelin)

Long-­legged Warbler. Latham 1783, p. 465. Motacilla longipes Gmelin 1789, p. 979 (‘Nova Seelandiae’, restricted to Ducky Sound by Oliver 1955, p. 453) Specimens  Skins are in Liverpool and Tring; subfossil remains are in Wellington, New Zealand. Status  Probably extinct. Last certainly recorded in 1968. Range  Formerly widespread throughout South Island, New Zealand. Description  9cm (3.5in). Adult top of head, hind neck and face ­purplish-­brown; rest of upperparts and tail dark green tinged with yellow; ­lemon-­yellow patch on edge of wing bordered behind with a deep brown patch; outer webs of primaries ­olive-­green, inner ones brown; superciliary streak and chin white; breast and abdomen grey; sides of body and ­undertail-­coverts yellowish green.

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 (Collar et al. 1994). A few reports have subsequently been made from Fiordland and Nelson Lakes, but they remain unsubstantiated. Its disappearance was due to predation by feral cats, rats, weasels and stoats. 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 220

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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.

Stephens Island Wren  Xenicus lyalli

(Rothschild)

Traversia lyalli Rothschild 1894b, p. 10 (Stephens Island, New Zealand) Xenicus insularis Buller 1895, p. 236 and plate Specimens  Skins are in Canterbury, Dunedin, and Wellington, New Zealand; Cambridge, Massachusetts; Liverpool; New York; Philadelphia; Pittsburgh; and Tring; subfossil remains are in Wellington, New Zealand. Status  Extinct. Last recorded in 1894. Range  In historic times confined to Stephens Island, but known from Holocene subfossil bones, and found at several sites in North Island and South Island, New Zealand. Description  10cm (4in). Male smaller than female: mottled dark olive on upperside, with brown margins to feathers; wings at flexure ­olive-­green yellow; quills and tail ­olive-­green brown; abdomen and sides of body ­olive-­green; throat, foreneck and breast ­olive-­green yellow with dark margins to feathers. 

In 1894 the cat belonging to Mr. D. Lyall, the lighthouse keeper on Stephens Island, brought in 11 specimens of a little bird, which were given to H. H. Travers. Ten of these were forwarded to Lord 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. ­Over-­collecting by the lighthouse keepers, and a rush to obtain specimens by professional collectors, would also have decimated the population (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). 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 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. Rothschild (1907a) gave the only information, 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. The population of wrens on Stephens Island is now known to be relictual, as remains of the bird are fairly common in owl deposits in 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 the bird was flightless, and the peculiar 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 sometimes placed in its own genus Traversia (Millener 1989, Turbott 1990), but this is not universally accepted.

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South Island ­Stout-­legged Wren  Pachyplichas yaldwyni

Millener

Pachyplichas yaldwyni Millener 1988, p. 391 (Honeycomb Hill Cave, Oparara Valley, Buller District, South Island, New Zealand) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction uncertain. Range  South Island, New Zealand.

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 capabilities, 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.

North Island ­Stout-­legged Wren  Pachyplichas jagmi

Millener

Pachyplichas jagmi Millener 1988, p. 395 (Ruakuri Cave, Waitomo district, North Island, New Zealand) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from bones found in North Island, New Zealand. Date of extinction uncertain. Range  North Island, New Zealand.

The North Island ­Stout-­legged Wren was smaller than its South Island counterpart P. yaldwyni, but otherwise the birds were similar. Numerous subfossil remains of this species and P. yaldwyni have been found in Laughing Owl Sceloglaux albifacies cave deposits (Worthy & Holdaway 2002), indicating that the owl was probably 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. South Island Stout-legged Wren Pachyplichas yaldwyni beneath the feet of a moa

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Pittidae

Long-­billed Wren  Dendroscansor decurvirostris

Milliner & Worthy

Dendroscansor decurvirostris Milliner & Worthy 1991, p. 182 (Moonsilver Cave, Takaka State Forest Park, New Zealand) Specimens  Subfossil remains are in Wellington, New Zealand. Status  Extinct. Known only from bones found in South Island, New Zealand. Date of extinction uncertain. Range  South Island, New Zealand.

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 probably 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 acanthisittidid, 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 inhabitant of ­high-­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 ­Long-­billed Wren became extinct before the arrival of Europeans, and may have been an early casualty of the Polynesianintroduced Pacific Rat Rattus exulans.

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.

Bougainville ­Black-­faced Pitta  Pitta anerythra pallida

Rothschild

Pitta anerythra pallida Rothschild 1904, p.7. (Bougainville Island, Solomon Islands) Specimens  The type specimen is in Tring. Status  Possibly extinct. Last recorded in 1938. Range  Bougainville, Solomon Islands. Description  15cm (6in). Crown rich ­chestnut-­brown (sometimes absent); rest of head, hindneck and upper mantle black, tinged chestnut on nape; remaining upperparts bright green; lesser ­upperwing-­coverts turquoise, rest of coverts green; tail dull green with black base; underparts ­whitish-­ochre; bill black; iris dark brown; legs and feet buff to ­greyish-­slate. Larger than nominate P. a. anerythra, which has darker undersides, and on which chestnut nape is always present.

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 subsequent 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 birds, but the nominate P. a. anerythra from Santa Isabel is a wary bird that occurs in primary forest remnants, secondary forest, gardens and small plots. The call of the nominate is distinctive and has been described as a single or double rasping tooyiii.

Choiseul ­Black-­faced Pitta  Pitta anerythra nigrifrons

Mayr

Pitta anerythra nigrifrons Mayr 1935, p. 4 (Tauro, Choiseul Island, Solomon Islands) Specimens  The type specimen is in New York. Status  Possibly extinct; last recorded date uncertain. Range  Choiseul, Solomon Islands.

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Extinct Birds Description  15cm (6in). Differs from nominate P. a. anerythra in the greater extent of black in the crown; chestnut nape sometimes absent; and underparts deep ochre, not ­ochre-­buff. Green upperparts sometimes with dusky stripes.

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.

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.

Peruvian ­Scale-­throated Earthcreeper  Upucerthia dumetaria peruana Zimmer Upucerthia dumetaria peruana Zimmer 1954, p.189 (15 miles east of Juliaca, Dept. Puno, ­south-­eastern Peru) Specimens  Specimens are in Berkeley and Washington, D. C. Status  Possibly extinct. Not seen since the late 20th century. Range  Puno region, Andes, southern Peru. Description  21.5cm (8.5in). Generally dark dull brown to ­greyish-­brown above, with olivaceous tinge; breast greyish-white with dusky scalloping; ­undertail-­coverts dull ­grey-­brown; white superciliary stripe; indistinct rufous edges to flight feathers; tail dull ­greyish-­brown with outer ­tail-­feathers tipped white; bill blackish; iris dark brown; legs and feet dark brown. Has a longer bill and generally darker than nominate U. d. dumetaria, which also has distinctly outlined rufous in wing.

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 ­long-­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.

Northern ­Stripe-­crowned Spinetail  Cranioleuca pyrrhophia rufipennis Sclater

& Salvin

Synallaxis rufipennis Sclater & Salvin 1879, p.620 (Tilotilo, Bolivia) Specimens  The type specimen is in Tring. Status  Possibly extinct. Not seen since the 1950s. Range  Northern Bolivian Andes, in Tilotilo, La Paz, and ­north-­west Cochabamba, northern Bolivia. Description  14.5cm (5.75in). Generally ­brownish-­grey above; whitish below, tinged buff on flanks; crown streaked buff and blackish; broad white superciliary and black postocular line; upper bill black to grey, lower pinkish with dark tip; iris dark red to pale orange or dark brown; legs and feet greenish to ­yellowish-­brown. More streaking on crown than nominate C. p. pyrrhophia, which is less brown above.

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. 224

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Grallaridae

Grallaridae (Antpittas and ­ant-­thrushes) The antpittas were formerly included in the Formicariidae, but they are quite distinct and have now been placed in their own family. There are at least 32 species distributed in Central and South America. The antpittas are generally terrestrial forest birds with long legs, and some are specialist ­ant-­followers. A number of species are in danger of extinction due to habitat loss.

Northern Giant Antpitta  Grallaria gigantea lehmanni

Wetmore

Grallaria gigantea lehmanni Wetmore 1945, p.19 (San Marcos, Moscopán, Cauca, Colombia) Specimens  Specimens are in Popayan, Colombia; Los Angeles; and Washington, D. C. Status  Possibly extinct. Last seen in the 1940s. Range  Central and ­south-­west Colombia, at the head of the Magdalena Valley. Description  26.5cm (10.5in). Generally dusky ­olive-­brown above; ­ochraceous-­buff below, edges to throat and breast feathers black giving a barred appearance heaviest on flanks; forehead chestnut; hindcrown and nape ­slate-­grey; upper bill blackish, lower ­grey-­brown; iris dark brown; legs and feet grey. Back more ­olive-­brown than nominate G. g. gigantea, which also has darker ­ochraceous-­buff underparts and central underparts less heavily barred.

Sometimes considered synonymous with the nominate G. g. gigantea, the Northern Giant Antpitta G. g. 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 & Long 1995); the bird has not conclusively been seen since. Birds of this race are said to have been recorded at La Planada Nature Reserve in Nariño in 1988 and 1989, but the identity of these birds awaits verification (de Soye et al. 1997). The disappearance of this subspecies can be attributed to severe deforestation and changes in land use on both slopes of the central Andes, which has been almost completely cleared for agriculture and the production of narcotics. The future of the northern population of this antpitta, if it still survives, is very bleak indeed. Habits The Northern Giant Antpitta was confined to humid montane forest in the upper subtropical to temperate zones at an altitude of around 3,000m. It foraged close to the ground in leaf litter and muddy soil, hopping with springing bounds through the undergrowth and on low limbs (Ridgely & Tudor 2009). Diet comprised invertebrates, especially the giant Rhynodrylus earthworms, but slugs and beetle larvae were also consumed (Krabbe et al. 1994b).

Ecuadorian Giant Antpitta  Grallaria gigantea hylodroma

Wetmore

Grallaria gigantea hylodroma Wetmore 1945, p.18 (Gualea, Province of Pichincha, Ecuador) Specimens  Specimens are in Brussels; New York; Philadelphia; and Washington, D. C. Status  Possibly extinct. Last recorded in 1989. Range  Western slope (Pichincha and Cotopaxi) of Andes in Ecuador. Description  26.5cm (10.5in). Differed from nominate in being much brighter; face ­rufous-­chestnut; ­brownish-­olive above; head and underparts ­rufous-­chestnut with narrow barring; central belly and vent ochraceous buff.

The Ecuadorian Giant Antpitta may represent a distinct species, but is presently considered a subspecies of the Giant Antpitta Grallaria gigantea, pending more taxonomic work. It is known only from 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 severely reduced the population; the last individual was recorded in 1989. The Ecuadorian Giant Antpitta is fully protected in a number of forest reserves in Ecuador (Robbins & Stiles 1999), but there is no confirmation that it still survives. Active logging and deforestation are severe around the reserve areas, so its ­long-­term survival remains in doubt. The habits of this antpitta are the same as those of Northern Giant Antpitta G. g. lehmanni, but it was dependent on lower altitude ­cloud-­forest, from 1,200–2,000m. 225

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Táchira Antpitta  Grallaria chthonia

Wetmore & Phelps

Grallaria chthonia Wetmore & Phelps 1956, p. 6. (Río Chiquito, ­south-­west Táchira, Venezuela) Specimens  The type specimen is in Washington D. C. Status  Possibly extinct; not seen since 1956. Distribution  Hacienda La Providencia at the Rio Chiquito, ­south-­west Táchira, Venezuela. Description  17cm (6.75in). Adult upperparts brownish; crown and nape grey; black barring on mantle; throat and ­ear-­coverts brown; white malar stripe; lower underparts whitish, with grey barring on flanks and breast; bill, legs and feet bluish black.

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.

Antioquia ­Brown-­banded Antpitta  Grallaria milleri gilesi

Salaman et al.

Grallaria milleri gilesi Salaman et al. 2009, p. 7 (Santa Elena, 8km east of Medellín, Antioquia, Colombia) Specimens  The type specimen is in Tring. Status  Possibly extinct. Known only from the type, collected in September 1878. Range  Antioquia, Colombia. Description  18cm (6.25in). Lores white with dark brown feather tips; ­ear-­coverts warm ­cinnamon-­brown; ­ear-­coverts to neck sides and upper breast warm ­cinnamon-­brown forming a ­breast-­band; upperparts uniform ­rufous-­brown; throat white with warm ­cinnamon-­brown feather tips; underparts pale grey, slightly paler whitish and creamy on central belly; sides of throat and malar region pale ­cinnamon-­buff, extending around throat to form a distinctive upper ­breast-­band; bill dark horn, tipped yellowish horn; iris dark; legs and feet dark horn. Larger than nominate G. m. milleri, with longer wings and tail but shorter tarsus.

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. 450). The type locality of the Antioquia bird has undergone extreme ecological change – almost complete deforestation 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.

Tyrannidae (Tyrant flycatchers) An exclusively New World family, the tyrant flycatchers have been divided into 97 genera with more than 400 species, making it by far the largest passerine family. They are diverse in morphology, but are generally rather drab birds that feed mainly on insects, although some of the larger species will eat small vertebrates and fruit. Some species are in danger of extinction due to habitat destruction, especially those confined to islands or island groups.

Grenada Euler’s Flycatcher  Lathrotriccus euleri flaviventris

(Lawrence)

Blacicus flaviventris Lawrence 1887, p. 617 (St. Andrews, Grenada) Empidonax johnstonei Barbour 1911, p. 58 (Grand Étang, Grenada) Specimens  Subfossil remains are in New York and Washington, D. C. Status  Probably extinct. Not recorded since the 1950s. Range  Grenada, West Indies.

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Tyrannidae Description  12.5cm (5in). The Grenada Euler’s Flycatcher L. e. flaviventris was very similar to L. e. lawrencei of Trinidad and northern Venezuela, but with a narrower bill; upperparts very dark greenish olive; cap and cheeks dusky; throat more pure white.

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. 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.

San Cristóbal Vermilion Flycatcher  Pyrocephalus rubinus dubius

Gould

Pyrocephalus dubius Gould 1839, p. 46 (San Cristóbal, Galápagos Islands) Specimens  Specimens are in Tring and Washington D. C. Status  Possibly extinct. Last recorded in the mid-1980s. Range  San Cristóbal, Galápagos Islands. Description  11cm (4.25in). Differs from mainland subspecies in shorter wings and tail, red less vivid and different song. Male has crown glossy dark vermilion red; throat and breast intensive red shading to pale red on underparts; chin reddish shading to white; lores, ­ear-­coverts and upperparts are generally uniform dark brown; bill, legs and feet black. Female dark grey above; bright yellow below, throat and chin paler; tail ­blackish-­grey.

The Vermilion Flycatcher has a wide distribution across the Americas, with P. r. nanus occurring in the Galápagos Islands except for San Cristóbal, where the endemic subspecies P. r. dubius once occurred. The San Cristóbal Vermilion Flycatcher was abundant in 1929 when Fisher surveyed the island (Fisher & Wetmore 1931); it occurred along the arid western coast inland to the small settlement of El Progreso in the highlands, but by the 1980s it had become extremely rare (Steadman 1988). An extensive survey in 1998 failed to find any birds (Vargas & Bensted–Smith 2000), and it might now be extinct. The loss of suitable habitat due to introduced invasive plants appears to have been a major factor in its decline, as this impacted on its food supply, but ­mosquito-­borne avian diseases may have also been partly responsible for the bird’s demise (Wikelski 2004).

Bogotá Bearded Tachuri  Polystictus pectoralis bogotensis

Chapman

Polystictus pectoralis bogotensis Chapman 1915, p. 646 (Suba, western Colombia) Specimens  Specimens are in Bogotá; Cambridge, Massachusetts; New York; and Washington, D. C. Status  Possibly extinct. Not recorded with certainty since the 1950s Range  Suba marshes, Bogotá, western Colombia. Description  10cm (4in). Male crest ­earth-­brown with white bases; pale brownish yellow face and short superciliary stripe; mantle and back dull brown, uppertail-coverts tinged rufous; wings and tail ­blackish-­brown with two ­rufous-­buff wingbars; throat pale yellow; breast and flanks tawny; belly and ­undertail-­coverts yellow; bill black. Female lacks black on head and chin and throat ­buff-­white.

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 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 ­sight-­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. 227

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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 now appear to lie closest to the bowerbirds (Ptilonorhynchidae) (Olson et al. 1983; Christidis et al. 1996). The North and South Island piopios were traditionally regarded as races of a single species, but they are now known to be specifically distinct (Holdaway 1999; Worthy & Holdaway 2006).

South Island Piopio  Turnagra capensis capensis

(Sparrman)

Tanagra capensis Sparrman 1787, pl. 4 (Dusky Sound, South Island, New Zealand) Specimens  Skins are in Basle; Bremen; Brussels; Cambridge, England; Cambridge, Massachusetts; Chicago; Dresden; Exeter; Gothenburg; Liverpool; Stockholm; Tring; and Washington, D. C.; eggs are in Tring. Status  Extinct since 1963 (possibly since the early 19th century). Range  South Island, New Zealand. Description  28cm (11in). An ­olive-­brown bird above, ­yellowish-­white below: upperparts ­olive-­brown; forehead striped darker; ­wing-­coverts tipped rufous; ­uppertail-­coverts and tail rufous; throat and sides of neck olive, striped rufous; rest of underparts ­olive-­brown streaked pale yellow; bill and feet brown; iris pale yellow. Juvenile rufous on head and throat. Eggs white or pale pink, spotted and blotched with dark and light brown, forming a zone at the larger end.

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 originated from the Cape (Worthy & Holdaway 2002). Decline and extinction The South Island Piopio was formerly abundant in the forests, and in early gold-mining days it was a common bird on the west coast, but by 1870 it was disappearing fast (Buller 1887–88). According to Potts (1872), they were rare on the upper Rakaia River, whereas four or five years before they had been plentiful. They could still be found at that time in the bushes that skirted the Havelock River,

South Island Piopio Turnagra capensis capensis

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the upper Waimakariri River and the Bealey River, and in the gorge of the Otira. Near the River Arahura on the west coast, Potts found two or three specimens, where, three years previously, it had been ‘most abundant’. He concluded that they had been driven away from the Arahura by clearances for paddocks required for the ­west-­coast cattle trade. However, the piopio was still common on the west coast between Ross and Okarita. By 1888 the bird was considered to be the rarest of all the birds of New Zealand, and by the beginning of the 20th century it was virtually extinct. The last confirmed specimen was shot at Oharu in 1902, although alleged sightings continued to be made. Allison et al. (1949) provided unconfirmed reports of sightings in forest inland from Patea in 1923, from between Gisborne and Wairoa on May 7, 1947, from Southland District 17 in December 1947, and in the Nelson district in January 1948. The last supposed sighting was in 1963 (Knox & Walters 1994). 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.

Stephens Island Piopio  Turnagra capensis minor

Fleming

Turnagra capensis minor Fleming 1915, p. 121 (Stephens Island) Specimens  Specimens are in Bremen; Dresden; Liverpool; Pittsburgh; Toronto; and Tring. Status  Extinct. The last specimen was taken in January 1897. Range  Stephens Island, New Zealand. Description  28cm (11in). Similar to the nominate T. capensis except slightly smaller, with shorter wings, tail and tarsi, but bill was of equal size; back ­brownish-­olive, not raw umber; concealed bases of feathers of back lighter.

The Stephens Island population of piopio has generally been relegated to the synonymy of T. 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 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. 229

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The Stephens Island Piopio numbered in the hundreds in 1894 (Medway 2004a), but the cats introduced to the island during the establishment of its lighthouse had multiplied to hundreds by that time. This appears to have spelt the end for the resident piopios, as the last specimen was taken on January 7, 1897; they had certainly gone by 1898.

North Island Piopio (Toatoa)  Turnagra tanagra

(Schlegel)

Otagon tanagra Schlegel 1865, p. 190 (no locality = North Island, New Zealand) Turnagra hectori Buller 1869, p. 39 (North Island) Specimens  Specimens are in Cambridge, England; Chicago; Leiden; Liverpool; Philadelphia; Tring; and Wellington, New Zealand. Status  Probably extinct since 1955. Unconfirmed report in March 1970. Range  North Island, New Zealand. Description  28cm (11in). Differed from the South Island Piopio T. capensis in being pale and unstreaked below. Head and back ­olive-­brown; wings and tail rufous, with olive tinge on two central retrices; throat white; breast and abdomen ­ash-­grey, darker on breast; abdomen and ­undertail-­coverts tinged with yellow; sides of body ­olive-­brown, washed with yellow; bill and feet dark brown; iris yellow. Juvenile rufous on head, with band of same colour on breast.

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 1882). The North Island Piopio was considered common at the time of European arrival, but it soon began to decline, and by 1892 had nearly disappeared. The last specimens were obtained 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 November 11, 1952, near Lake ­Waikare-­iti on April 9, 1953, and the last observation, also at Lake ­Waikare-­iti, on April 9, 1955. Mead (1950) observed birds from March 14–17, 1950 on the Wanganui River, and Bell & Singleton (1974) reported a parent bird feeding young in March 1970, but this remains unconfirmed. The North Island Piopio disappeared for the same reasons as the South Island species, a combination of habitat loss and predation by introduced cats, dogs, rats and mustelids. It is generally now considered extinct. 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, ­pio-­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. Mead (1950) gave an excellent 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. 230

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Chatham Island Bellbird  Anthornis melanocephala

Gray

Anthornis melanocephala G.R. Gray 1843, p. 188 (Chatham Islands) Specimens  Skins are in Auckland and Canterbury, New Zealand; Cambridge, England; Liverpool; and Tring; eggs in Auckland and Tring; unique nest in Canterbury, New Zealand. Status  Extinct. Last observed in 1906. Range  Chatham Island, Pitt Island, Mangere Island and Little Mangere Island, Chatham Islands. Description  23cm (9in). Generally ­olive-­green, lighter on abdomen and sides; forehead and crown ­steel-­blue, becoming purplish on throat and sides of head; wings and tail ­blackish-­brown, margined with ­steel-­blue. Differed from the extant New Zealand Bellbird Anthornis melanura in that the male was larger; there was more ­steel-­blue on head; less yellow in the plumage generally; and no yellow patch on sides.

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 ­over-­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 1906b). 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 extirpated 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.

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 ­dull-­coloured, terrestrial birds, endemic to Australia. As they prefer coastal environments they are extremely vulnerable to anthropogenic disturbance.

Western Rufous Bristlebird  Dasyornis broadbenti litoralis

(Milligan)

Sphenura litoralis Milligan 1902, p. 69 (Ellensbrook, Western Australia) Specimens  A specimen is in Melbourne, Australia. Status  Extinct. Last recorded in 1908. Range  Coastal ­south-­west Western Australia, from Cape Naturaliste south to Cape Mentelle and Margaret River. Description  25–27cm (10–10.5in). Smaller and brighter than other subspecies, and apparently differed vocally. Head rich rufous, extending onto ­ear-­coverts; lores and rest of face ­off-­white; ­hind-­neck, back, uppertail and scapulars ­reddish-­brown, grading to ­olive-­brown with a ­reddish-­brown tinge on lower back and rump; upperwings brownish with rufous tinge; chin, throat and breast ­greyish-­white with marked scalloping, rest of underparts grey; upper bill ­greyish-­black; lower bill pale grey or pink; iris red; legs and feet dark brown or ­greyish-­brown.

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 231

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Victoria (Seymour et al. 2003). A western population, D. b. litoralis, was endemic to the ­south-­western coast of Western Australia, between Cape Mentelle and Cape Naturaliste (Johnstone & Storr 2004). It was considered 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 shrublands 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.

Acanthizidae (Australian warblers) The Australian Warblers are generally confined to Australia and New Guinea, but they also occur in Indonesia, New Zealand, and some Pacific islands. The gerygones have colonised New Zealand, the Chatham Islands, Norfolk and Lord Howe Islands. In general, the family is not at risk, but some island populations are either extinct or threatened, primarily due to predation from introduced mammals, especially rats.

Lord Howe Gerygone  Gerygone insularis

Ramsay

Gerygone insularis Ramsay 1878, (Lord Howe Island) Specimens  Specimens are in Liverpool; Sydney; and Tring. Status  Became extinct soon after 1918. Range  Formerly occurred on Lord Howe Island. Description  12cm (4.75in). A small, predominantly brown and greyish bird: top of head brown; ­ear-­coverts pale greyish; pale grey ­eye-­ring; throat and chin grey with yellowish tinge; many parts varied as to the degree of yellow, this being apparent in its bright yellow belly; neck and upperparts brown grading to ­yellow-­brown on rump; breast pale grey with yellowish tinge; belly bright yellow; tail ­olive-­brown at base, blackish toward end, with white spots near tip; iris pink; bill ­greyish-­black; legs and feet ­blue-­grey.

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 disappeared after Black Rats Rattus rattus came ashore from the wreck of the Mokambo, on Ned’s Beach in 1918. Owls were imported to kill the rats, but may have fed on the endemic birds as well. McAllan et al. (2004) suggest that disease introduced 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. 232

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Callaeidae

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 insectivorous, 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 ­dome-­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.

Huia  Heteralocha acutirostris

(Gould)

Neomorpha acutirostris Gould 1837a, pl. 11 (a female). Neomorpha crassirostris Gould 1837a (= male) Specimens  Many skins and some skeletons occur in museums around the world. They principally are in Amsterdam; Ann Arbor, Michigan; Auckland, Canterbury, Wellington and Christchurch, New Zealand; Basle; Berlin; Bremen; Brussels; Cambridge, Massachusetts; Chicago; Dresden; Edinburgh; Exeter, England; Geneva; Leiden; London; Melbourne; Munich; Newcastle; New York; Oslo; Philadelphia; Pittsburgh; Princeton, New Jersey; Stockholm; Tring; Vancouver; and Washington, D. C. A unique egg and subfossil remains are in Wellington. Status  Extinct. Last confirmed sighting was in December 1907. Range  North Island, New Zealand. Description  Male 45cm (18in); female 48cm (19in). General coloration black with green gloss; terminal tail-band white; facial wattles orange; bill ivory; iris brown; legs and feet black. Juvenile duller, suffused brown; terminal ­tail-­bar washed rufous. Maoris reported that birds from the Ruahine range had a broader white band on the tail than those from the Tararue range (implying subspecific differentiation), but this has not been substantiated. Leucistic Huias were known. Egg ­stone-­grey with spots and dots of dark purplish-grey.

Huia Heteralocha acutirostris (female left, male right)

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. 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 ornithologist John Gould, he believed that the sexes were different species, calling the female H. acutirostris and the male H. crassirostris (Gould 1888). 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 233

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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 ­over-­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 ­year-­round. Fuller (2000) also disagreed with Greenway’s suggestion, and noted that he had not taken into account the numbers taken just for their tail feathers, and specimens lost during collection; 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 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 collections. By the early 1900s, the Huia was effectively extinct. Myers (1923), with the help of R. H. D. 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. Reasons for extinction Myers (1923) was firmly of the opinion that collecting, first by Maoris and later by Europeans, was largely responsible for the bird’s demise, but that ticks and tick-borne diseases may also have been partly to blame. Mason (1921) reported finding ticks of two species of Oriental and African distribution on a Huia, both previously unrecorded in New Zealand. Both ticks have occurred on the Common Myna Acridotheres tristis in its native homeland; this species was introduced to New Zealand in 1875, where it quickly spread. It may have brought with it a disease fatal to the Huia. Ferrets, stoats and weasels were introduced to New Zealand in the 1880s to control rabbits, and they undoubtedly took their toll of the principally terrestrial Huias. Rats, of course, were already ­well-­established. Official protection was extended to the Huia in 1892 (Morris & Smith 1995), but the legislation had little effect, in spite of the ­well-­intentioned efforts of the Governor of New Zealand, the Earl of Onslow. Plans were made to create ‘reserves’, into which Huias could be introduced, but it appears that incompetence and inertia prevented anything happening (Phillipps 1963). Even Buller was not exempt from exploiting the Huia’s plight. According to ­Jayne-­Wilson (2004), Buller appropriated a live pair destined for a new bird sanctuary on Kapiti Island in 1893, and intentionally bent the law in order to take them back to England as a gift for Rothschild at Tring. Buller sold many New Zealand bird specimens to Rothschild, but the bitterness that ensued over the naming of the Stephens Island Wren Xenicus lyalli (see p. 221), and sale of a Laughing Owl Sceloglaux albifacies (see p. 199), was to irrevocably undermine their friendship. Unconfirmed sightings of the Huia persisted throughout the 20th century, even as late as 1961, but the Huia had almost certainly disappeared half a century before. Habits Buller (1887–88) gave the best accounts of the bird and recorded that: 234

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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 fed chiefly on insects, particularly the grub of the Huhu Beetle Prionoplus reticularis, and also 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 open apart the substrate with considerable force (Burton 1974). The 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 insert her long bill to extract a 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 1996). Apparently the birds were often 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.

South Island Kokako  Callaeas cinerea cinerea

(Gmelin)

Cinereous Wattlebird Latham 1781, p. 364. Glaucopis cinerea  Gmelin 1788, p. 363 (Nova Seelandia = New Zealand) Specimens  Specimens are in Christchurch and Dunedin, New Zealand; La Rochelle, France; Liverpool; Frankfurt; and Tring. Status  Extinct. Last confirmed report in 1967. Range  South Island, New Zealand. Description  38cm (15in). A uniform ­bluish-­grey bird with a black mask; bill, legs and feet black; iris unrecorded. Large fleshy wattles orange with blue base (all blue in North Island Kokako C. c. wilsoni), smaller and pink in juvenile.

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; 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 conservation effort underway to save it from extinction. The Kokako is a ­poor-­flying terrestrial bird that has suffered particularly from deforestation and introduced predators, especially cats, Stoat Mustela erminea and Black Rat Rattus rattus, and from competition and nest predation by the ­Brush-­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). 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 ­bell-­like duet. Buller (1873) gave a very good account of the bird when it was still common, but he also prophesied its impending doom: 235

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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, 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.

North Island Stitchbird (Hihi)  Notiomystis cincta cincta

(Du Bus)

Meliphaga cincta Du Bus de Gisignies 1839, p. 295 ­(Nouvelle-­Zélande = North Island, New Zealand) Specimens  Specimens of N. c. cincta are in Auckland, New Zealand; Cambridge, England; Liverpool; New York; and Tring. Status  Possibly extinct; last recorded in the 1880s. Range  North Island, New Zealand. Description  18cm (7in). Male head, back and upper breast ­velvet-­black, on breast underlined with ­golden-­yellow; erectile white ­ear-­tufts; rump and ­uppertail-­coverts dark ­brownish-­olive; wings black with white ­wing-­bars and ­golden-­yellow shoulder patches; rest of underparts pale brown; bill black; iris ­blackish-­brown; legs dark ­reddish-­brown to grey black; soles ­cream-­coloured. Female smaller, ­grey-­brown with white ­wing-­bar; ­ear-­tufts indistinct and ­grey-­brown.

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 translocation 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 considered 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 invertebrates (Angehr 1984; Castro et al. 1994b). Buller (1873) 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.

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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.

Mount Lofty Spotted ­Quail-­thrush Cinclosoma punctatum anachoreta

Schodde & Mason

Cinclosoma punctatum anachoreta Schodde & Mason 1999, p. 418 (Mount Lofty, South Australia) Specimens  The type specimen is in Adelaide. Status  Possibly extinct. Last recorded in 1984. Range  Mount Lofty Ranges, South Australia. Description  24–30cm (9.5–12in). Adult ­olive-­brown above with distinctive black streaks; large white supercilium extending to nape; face and throat black bordering a white patch on side of neck; ­wing-­coverts black tipped white; breast grey, belly white divided by irregular black bar; flanks ­yellow-­brown and distinctive black streaks and spots along sides of body; bill black; iris pale ­greyish-­white; legs and feet dull flesh. Male has more complex and more intensely coloured pattern than female.

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 part of the 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 (Close 1972), one in 1977 (Higgins & Peter 2002) one in 1981, when a single breeding observation was made (Garnett & Crowley 2000), and a final record in 1984 (Blakers et al. 1984). 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 likelihood 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; Close 1972). Eggs were laid from August to September (Ford 1983), with the birds nesting in Eucalyptus trees. 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.

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. 237

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Marinduque Blackish ­Cuckoo-­shrike  Coracina coerulescens deschauenseei  DuPont Coracina coerulescens deschauenseei duPont 1972, p. 9 (Marinduque, Philippines) Specimens  The type specimen is in Delaware. Status  Possibly extinct. Last seen in 1971. Range  Marinduque, Philippine Islands. Description  25–26cm (10–10.25in). Entirely black glossed purplish; rump slaty grey; female uniform dark ­ash-­grey, paler on rump; bill, legs and feet black; iris dark brown. Nominate C. c. coerulescens slightly more purple.

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 endangered. Nothing is known about its habits.

Cebu Blackish ­Cuckoo-­shrike  Coracina coerulescens altera

(Ramsay)

Edoliosoma alterum Ramsay 1881, p. 34 (Cebu) Specimens  The type specimen is in Tring. Status  Probably extinct. Not recorded since 1906, but an unconfirmed report was made in 2000. Range  Cebu, Philippine Islands. Description  25–26cm (10–10.25in). Entirely black, glossed green; rump slaty grey. Nominate C. c. coerulescens differs in being glossed purplish, and female much darker.

The Cebu Blackish ­Cuckoo-­shrike has not been seen since McGregor (1909) collected it in 1906. Its extinction 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.

Cebu ­Bar-­bellied ­Cuckoo-­shrike  Coracina striata cebuensis

(Ogilvie-Grant)

Artamides cebuensis Ogilvie-Grant 1896, p. 535 (Cebu) Specimens  Specimens are in Tring. Status  Probably extinct. Not recorded since 1906, but an unconfirmed report was made in 2000. Range  Cebu, Philippines. Description  30cm (12in). Male similar to the Mindoro subspecies, C. s. mindorensis, from which it differed in being larger; feathers covering the nostrils grey instead of black; tail black, slightly washed with grey towards base of middle feathers. Female was most like the female nominate C. s. striata but lower back, rump and ­uppertail-­coverts uniform grey, like the back.

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 (1909) 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 mountains, but also occurred in secondary growth (Rabor 1959).

Norfolk Island Triller  Lalage leucopyga leucopyga

(Gould)

Symmorphus leucopygus Gould 1838, p. 3 (‘New South Wales’ = error for Norfolk Island) Specimens  Specimens are in Leiden; Melbourne, Australia; Tring; and Vienna. Status  Extinct since 1942. Range  Formerly occurred on Norfolk Island.

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Oriolidae Description  17–18cm (6.5–7in). The Norfolk Island Triller was similar to other subspecies of the Long-tailed Triller, but was slightly larger with richer buff wash on underparts and rump; and broader white tip to outer rectrices. Adult male: greenish 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 ­wing-­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 ­wing-­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). 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 ­ground-­insects, or flying upwards, takes others on the wing. Owing to its habit of collecting an early breakfast amongst ­dew-­drenched grass, its otherwise ­snow-­white breast becomes discoloured with the reddish dust which settled on the ­grass-­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.

Cebu Oriole  Oriolus steerii assimilis

Tweeddale

Oriolus assimilis Tweeddale 1878, p. 760, pl. 76 (Cebu) Specimens  Specimens are in Tring. Status  Extinct since 1906. Range  Cebu, Philippine Islands. Description  20cm (8in). Adult dark ­greenish-­yellow on upperside and ­undertail-­coverts; face, throat and breast dark grey, tinged with ­greenish-­yellow on breast; abdomen, flanks and ventral region grey or white with broad dark streaks; wings dark grey; tail dark grey, the outer feathers with a yellow subterminal spot; bill dark sienna; iris crimson; legs and feet dark ­lead-­grey; claws black.

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 1878). 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 ‘exceedingly common in the small amount of forest left in Cebu’. The Cebu Oriole has not been seen since McGregor (1909) 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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Rhipiduridae ­(Fan-­tailed flycatchers) The ­Fan-­tailed Flycatchers are a ­wide-­ranging group, with around 40 species currently recognized, all placed in the single genus, Rhipidura. Some are extremely widespread and migratory, others are sedentary and restricted to single islands. The ­fan-­tailed flycatchers are ­long-­tailed, insectivorous birds that prefer rainforests, but they do occur in a variety of other habitats, including heavily modified areas. They are generally unthreatened, but an island species and subspecies became extinct due to introduced predators.

Lord Howe Fantail ­(Fawn-­breasted Fantail)  Rhipidura cervina

Ramsay

Rhipidura cervina Ramsay 1879, p. 340 (Lord Howe Island) Rhipidura macgillivrayi Sharpe 1881, p. 789, pl. 67 (Lord Howe Island); Sharpe’s name was given in error, unaware that Ramsay had already named the bird. Specimens  Specimens are in New York; Sydney; and Tring; eggs and nests are in Tring. Status  Extinct since between 1924 and 1928. Range  Lord Howe Island. Description  15cm (6in). Entire undersurface pale ­creamy-­yellow, with a pale throat, lacking the white throat and black band of the Grey Fantail Rhipidura flabelifera, also having a stouter bill and shorter tail. Head, neck and upperbody grey, with a pale ­creamy-­yellow supercilium; underparts pale ­creamy-­yellow, throat slightly paler; tail dark ­brownish-­grey with white tip; wings ­grey-­brown with two prominent pale ­wing-­bars; bill black; iris ­blackish-­brown; legs and feet ­greyish-­black.

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 (Hindwood 1940), and by 1928 it appears to have completely disappeared (Sharland 1929). Habits 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 ­nesting-­place had been chosen in a small prickly shrub, overgrown with clematis, the nest being almost hidden amongst the ­vine-­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.

Guam Rufous Fantail  Rhipidura rufifrons uraniae

Oustalet

Rhipidura uraniae Oustalet 1881, p. 75 (Guam, Marianas Islands) Specimens  Specimens are in New York; Paris; and Tring; a nest is in Honolulu. Status  Extinct. Last recorded in 1984. Range  Guam, Marianas Islands, ­south-­west Pacific. Description  14–16cm (5.5–6.25in). Adult upperparts ­ochraceous-­brown, becoming bright rufous on forehead, rump and ­uppertail-­coverts; tail shining ­dusky-­brown, almost black with pale tips; primaries and secondaries brown; throat and ­neck black, with white tips on breast; lower breast and belly white, with rufous tint; chin ­greyish-­white; cheeks, lores and ­ear-­coverts dusky; sides, flanks, thighs and ­undertail-­coverts rufous; bill dusky; iris dark hazel; legs and feet dark brown.

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 240

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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 ­north-­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 understorey 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. 244), 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 ­mucous-­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 ­brown-­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 morphologically that many earlier writers (e.g. Sharpe 1879) failed to distinguish between them.

Maupiti Monarch  Pomarea pomarea

(Garnot)

Muscicapa Pomarea Garnot 1828, pl. 17, figs A–C (2l June); Muscicapa Maupitiensis Garnot 1829, p. 592 (21 November) (Maupiti Island) Specimens  There is no surviving specimen. Status  Extinct. Known only from the type (now lost); not seen since 1823. Range  Maupiti Island, Society Islands, South Pacific. Description  Male uniform glossy black; bill, legs and feet black. A colour morph occurs where head, neck, mantle and upper breast glossy black, rest of plumage white, with a few brown feathers on ­wing-­coverts. Female ­chestnut-­brown above, white below; lower back and rump black; ­wing-­coverts and secondaries black with brown edges; primaries brown with blackish tips; tail brown.

Maupiti is a small, high island that was ­well-­wooded in the 19th century, but the ravages of human occupation 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.

Hiva Oa Monarch  Pomarea mendozae

Hartlaub

Pomarea mendozae mendozae Hartlaub 1854, p. 170 (St Christina Island [= Tahuata], Marquesas) Specimens  Specimens are in Liverpool; New York; and Tring. Status  Probably extinct. Last recorded in March 1975. Range  Hiva Oa and Tahuata, Marquesas Islands, South Pacific. Description  17cm (6.5in). Adult male uniform glossy black; bill pale ­bluish-­grey; legs and feet black. Female body white, tinged pinkish buff below; head and wings black except for white edges of tertials; tail white with black terminal spots. Juvenile ­reddish-­brown above, ­pinkish-­buff below with white ­eye-­ring.

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The monarch flycatchers of the Marquesas were originally considered a single species, but mitochondrial DNA analysis has shown them to represent four distinct species (Cibois et al. 2004). The Hiva Oa birds were common in 1921 and 1922, when some were collected by the Whitney South Seas Expedition, but they declined thereafter. A single individual was recorded in March 1975, but despite extensive surveys in the 1990s and in 2000 (Thibault & Meyer 2001) it has not been seen since. The Tahuata population was also considered common in 1922 (Murphy & Mathews 1928), but birds have not been recorded on the island since. The Hiva Oa Monarch was particularly sensitive to rat predation, with both Black Rat Rattus rattus and Pacific Rat R. exulans common on both islands. Overgrazing and burning of the lowland dry forest severely reduced the available habitat, and the introduced predatory Great Horned Owl Bubo virginianus and competition with the introduced Common Myna Acridotheres tristis must also have impacted on the bird’s numbers (Thibault & Meyer 2001). It is unlikely that this species still survives. 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 population surviving outside what would have been its normal range. The birds foraged for insects in the dense understorey.

Ua Pou Monarch  Pomarea mira

Murphy & Mathews

Pomarea mendozae mira Murphy & Mathews 1928, p. 4 (Ua Pou, Marquesas) Specimens  Specimens are in New York. Status  Probably extinct. Last recorded in 1922. Range  Ua Pou, Marquesas Islands, South Pacific. Description  17cm (6.5in). Adult male uniform glossy black; bill pale ­bluish-­grey; legs and feet black. Female differed from female P. mendozae in being black with white tail and wings black-and-white, with much white.

This species was restricted to forested valleys at the western end of the island in 1922 (Murphy & Mathews 1928), when it was discovered by the Whitney South Seas Expedition, but a search for it in 1975 found no birds (Holyoak & Thibault 1977b), and more recent surveys for it have been similarly unsuccessful (Thibault & Meyer 2001). It may now be extinct. The introduction of the ­tree-­climbing Black Rat Rattus rattus, plus the destruction of lowland habitat, appears to have caused its extinction. Nothing is known about its habits.

Nuku Hiva Monarch  Pomarea nukuhivae

Murphy & Mathews

Pomarea mendozae nukuhivae Murphy & Mathews 1928, p. 5 (Nuku Hiva, Marquesas) Specimens  Specimens are in New York and Tring. Status  Extinct. Last recorded in the 1930s. Range  Nuku Hiva, Marquesas Islands, South Pacific. Description  17cm (6.5in). Adult male uniform glossy black; bill pale ­bluish-­grey; legs and feet black. Female differed from female P. mendozae in having black hood; back, rump and tail white; wings ­black-­and-­white with much black.

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.

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Eiao Monarch  Pomarea fluxa

Murphy & Mathews

Pomarea iphis fluxa Murphy & Mathews 1928, p. 7 (Eiao, Marquesas) Specimens  Specimens are in New York. Status  Probably extinct. Last recorded in 1977. Range  Eiao, Marquesas Islands, South Pacific. Description  17cm (6.5in). Adult male mainly glossy black, mottled with white on back, lower breast and belly; ­undertail-­coverts black; bill dark ­bluish-­grey; legs and feet black. Female light brown above with white mottling; lores ­rufous-­brown; underparts white; throat heavily streaked black. Juvenile like female but more extensively streaked.

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. 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.

Negros Celestial Monarch  Hypothymis coelestis rabori

Rand

Hypothymis coelestis rabori Rand 1970, p. 363 (Negros, Philippines) Specimens  Specimens are in Chicago; Delaware; Hamburg; Luzon; New York; and Yale. Status  Possibly extinct. Not recorded since the late 1990s. Range  Negros and possibly Sibuyan, Philippine Islands. Description  18cm (7in). Adult almost uniform dark ­sky-­blue, washed lilac on cheeks and throat; belly duller ­grey-­blue; ­eye-­ring yellowish; erectile pale ­turquoise-­blue crown feathers. Female similar but slightly duller.

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.

Manu’a Shrikebill  Clytorhynchus vitiensis powelli

(Salvin)

Pinarolestes powelli Salvin 1879, p. 128 (Tutuila, Samoa) Specimens  The type specimen is in Tring. Status  Possibly extinct. Not recorded since the 1990s. Range  Manu’a, Ofu and Ollosega Islands, American Samoa, South Pacific. Description  18cm (7in). A ­medium-­sized bird with a heavy bill. Head, back, wings, and tail ­brownish-­black; three lateral feathers of tail tipped white; throat, breast and belly brownish slate; sides brownish.

The shrikebills of the genus Clytorhynchus occur on islands in Melanesia and western Polynesia. The Fiji Shrikebill C. vitiensis occurs in American Samoa, Fiji and Tonga, and it has been divided into 12 subspecies. C. v. powelli once occurred on three islands in American Samoa, Manu’a, Ofu and Olosega, but it gradually declined as the native forests were cut. Once the forests were fragmented, the shrikebill was more susceptible to cyclones, and there have been no reports of it since American Samoa was devastated by Cyclone Ofa in 1990 and Cyclone Val in 1991. 243

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Habits It was an aggressive, bold bird, and Salvin (1879) was told by Powell that it flew around people entering its territory, keeping up a continuous harsh cry. The shrikebill was insectivorous, using its large heavy bill to forage in matted dead leaves and decaying wood.

Guam Flycatcher (Guam Broadbill)  Myiagra freycineti

Oustalet

Myiagra freycineti Oustalet 1881, p. 73 (Guam, Mariana Islands) Specimens  Specimens are in Honolulu; Leiden; and Washington, D. C; nest and eggs are in Honolulu. Status  Extinct. Last known individual died in May 1984. Range  Guam, Mariana Islands, ­south-­west Pacific. Description  13cm (5in). Adult head and neck bluish with a metallic lustre; back and ­upperwing-­coverts ­greenish-­blue; rump ­greenish-­blue washed grey; chin and throat white; breast light cinnamon; tail ­bluish-­slate; bill, legs and feet black; iris dark brown.

The Guam Flycatcher is sometimes considered to be a subspecies of the ­wide-­ranging Oceanic Flycatcher Myiagra oceanica. It once occurred all over Guam bar the southern plains, but by 1971 it was restricted to the northern plateau, primarily below the cliff line in Artocarpus forests, where it was common (Jenkins 1983; Drahos 2006). A serious decline occurred over the next decade (Collar et al. 1994), and less than 100 birds were recorded in early 1983 in the Pajon Basin. By October 1983, only a single male was observed. This individual or another was captured for potential captive propagation but this effort was a failure as no other Guam Flycatchers could be found (Drahos 2006). The single male died of unknown causes on May 15, 1984. Disease may well have been a factor when the last population was virtually wiped out in the space of six months, but the critical factor in the species’ decline was almost certainly predation by the introduced Brown Tree Snake Boiga irregularis, which arrived on the island in the 1940s (Savidge 1987). The discovery of subfossil remains of an undescribed Myiagra species on Ua Huka in the Marquesas (Steadman 1989b) suggests that monarchs were much more widely distributed in the past, and many islands had more than one genus prior to the arrival of Polynesians. Guam Flycatcher Myiagra freycineti

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. 240), 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 ­rufous-­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. 244

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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.

Chatham Island Raven  Corvus moriorum

Forbes

Corvus moriorum Forbes 1892b, p. 252 (Chatham Islands) Paleocorax moriorum Forbes 1892c, p. 21 Specimens  Subfossil remains are in London; Canterbury and Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  Chatham Islands, New Zealand.

The discovery of the New Zealand and Hawaiian crows from subfossil remains has shown that the family was distributed widely across the Pacific, and that island populations were extremely vulnerable to anthropogenic changes. The Chatham Island Raven was larger than the mainland New Zealand ravens (see below), and its remains are commonly found in coastal sites on the Chatham Islands (Gill 2003). 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 fruit and invertebrates (Worthy & Holdaway 2002). Forbes (1892c) placed the Chatham Island Raven in its own genus, Paleocorax, based on cranial differences, but Worthy & Holdaway (2002) consider the taxonomic placement unwarranted. The bill of the Chatham Island Raven was long, broad and slightly curved. The bird reached a size of 72cm in total length, making it one of the largest passerines known. The arrival of the Morimori followed by the Maori, both of whom cleared the forests and introduced potential predators, led to the crow’s extinction.

North Island Raven  Corvus antipodum antipodum

(Forbes)

Paleocorax antipodum Forbes 1893c, p. 544 (North Island) Specimens  Subfossil remains are in London; and Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  North Island, New Zealand.

Despite being smaller than the Chatham Island Raven C. moriorum, the North Island race of New Zealand Raven was still a large bird, with a long but rather more broad and straight bill than its Chatham Islands congener (Gill 2003). They were morphologically typical corvids, but the tarsometatarsus was comparatively long, suggesting that these birds may have occupied a more terrestrial niche than other ravens. Like the Chatham Island and South Island birds, their remains are numerous in coastal fossil deposits, and they would have been extremely vulnerable to ­human-­induced changes of lowland habitat.

South Island Raven  Corvus antipodum pycrofti

Gill

Corvus antipodum pycrofti Gill 2003, p. 54 (Mussel Point, Marfell Beach, Marlborough) Specimens  The type specimen is in Wellington, New Zealand. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  South Island, New Zealand.

Gill (2003) made a morphological study of the extinct New Zealand ravens, and concluded that their closest relative is the Australian Raven C. coronoides, and that the New Zealand crows evolved directly from it or from a species ancestral to it. All of the New Zealand ravens were perfectly adept fliers, and a single bone now attributed to the South Island subspecies has been found on the remote Auckland Islands (Worthy & Holdaway 2002). However, it is not known whether this bone represented a resident population or was that 245

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of a vagrant to the islands. The South Island Raven probably disappeared for the same reasons as the other New Zealand ravens.

Puerto Rican Crow  Corvus pumilus

Wetmore

Corvus pumilus Wetmore 1920, p. 81 (Cueva San Miguel, Puerto Rico) Specimens  Subfossil remains are in New York. Status  Extinct, known only from subfossil remains. Date of extinction not known. Range  Puerto Rico and the Virgin Islands.

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.

Deep-­billed Crow  Corvus impluviatus

James & Olson

Corvus impluviatus James & Olson 1991, p. 12 (Barbers Point, Oahu) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct, known only from subfossil remains. Date of extinction not known. Range  Oahu, Hawaiian Islands.

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 illustrates 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 extant Hawaiian Crow C. hawaiiensis, or something similar to it, also occurred on Maui. The ­Deep-­billed Crow was a large species with a pronounced high and deeply arched bill. It was the largest of the Hawaiian crows, although there was some overlap in size between this species, C. viriosus (see below) and C. hawaiiensis. The bill is reminiscent of that of the ­White-­billed Crow C. woodfordi from the Solomons, and of the Bougainville Crow C. meeki, but broader and deeper. These western Pacific corvids are forest inhabitants that feed on fruit and insects (Madge & Burn 1993), and it is likely that the ­Deep-­billed Crow had a similar ecology. This species would have been susceptible to forest destruction after the arrival of the Polynesians. Loss of habitat and Pacific Rat Rattus exulans predation are almost certainly the causes of its extinction.

Deep-billed Crow Corvus impluviatus

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Robust Crow  Corvus viriosus

James & Olson

Corvus viriosus James & Olson 1991, p. 19 (Barbers Point, Oahu) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct, known only from subfossil remains. Date of extinction not known. Range  Oahu and Molokai, Hawaiian Islands.

The fossil record shows that a second large species of crow formerly occurred on Oahu, where it was sympatric with the ­Deep-­billed Crow C. impluviatus, and also on Molokai. 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 ­post-­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.

Petroicidae (Australasian robins) The Australasian robins occur in Australia, New Guinea, New Zealand and a number of Pacific islands. The family has been divided into 15 genera with approximately 45 species. They prefer woodlands but are generalists, and occupy a broad range of habitat types. Some species are threatened; for example the New Zealand Black Robin Petroica traversi was down to just five individuals in 1980, but numbers have now climbed to more than 250 through intense conservation management. Only one subspecies is known to have become extinct.

Tiwi Islands Hooded Robin  Melanodryas cucullata melvillensis

Zietz

Melanodryas cucullata melvillensis Zietz 1914, p. 15 (Melville Island) Specimens  Specimens are in Adelaide. Status  Possibly extinct. Last recorded January 1992. Range  Tiwi Islands, off Northern Territory, Australia. Description  16cm (6.25in). Male head, back and tail black; wings black with white ­wing-­bars; chest, belly, and ­undertail-­coverts white, belly area washed grey; bill, legs, feet and iris black. Female ­grey-­brown above; pale grey throat, paler below washed brown; wings dark brown with white wing-bars. Smaller with a longer bill than nominate M. c. cucullata, which is washed grey (male) or brown (female).

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.

Mohoidae (Mohos) One of the most extraordinary discoveries of recent years was the resolution of the true relationships of the Hawaiian ’o’os (pronounced ­oh-­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 247

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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. 251), 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.

Kaua’i ’O’o  Moho braccatus

(Cassin)

Mohoa braccata Cassin 1855, p. 440 (Sandwich Islands = Kaua’i) Specimens  Skins are in Ann Arbor; Berlin; Bremen; Cambridge, Massachusetts; Cambridge, England; Dresden; Edinburgh; Honolulu; Liverpool; Oslo; Paris; Philadelphia; Pittsburgh; New York; Stockholm; Tring; and Washington, D. C; subfossil remains are in Washington, D. C. Status  Extinct. Last certainly recorded in 1985. Range  Kaua’i, Hawaiian Islands. Description  20cm (8in). Smaller and duller than other ’o’os. Male head black with a few streaks of white; tail black with the central feathers longer than the others; wings black with a white patch at shoulder; rest of upperparts ­slaty-­brown, turning to russet on rump and flanks; throat and breast black barred with white; rest of underparts streaky grey; thighs yellow; iris light yellow; bill, legs and feet black. Female similar except throat feathers more extensively barred white. Immature lighter underparts; more brownish bill and legs; and lacking the yellow thighs and white ­shoulder-­patch.

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 (Munroe 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 destruction 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 ­sea-­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 cockroaches, 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 polymorpha, 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).

Oahu ’O’o  Moho apicalis

Gould

Moho apicalis Gould 1860b, p. 380 (‘Owhyie’ = Oahu) Specimens  Skins are in Berlin; Cambridge, Massachusetts; New York; Paris; Tring; and Vienna; subfossil remains are in Washington, D. C.

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Mohoidae Status  Extinct. The last specimens were collected in 1837, but another may have been taken in September, 1840. Range  Mountain forests of Oahu, Hawaiian Islands. Description  30–36cm (12–14in). A striking black, brown, white and yellow bird. Predominantly sooty black; tail brown with outer feathers tipped white; the two central ­tail-­feathers lacking the white tips, and narrower and tapered; sides and ­undertail-­coverts yellow; ­underwing-­coverts white; bill, legs and feet black. Females smaller, with shorter tails.

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 1996). 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 1996): 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 1994), 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 ­over-­hunting for its valuable yellow plumes.

Hawaii ’O’o  Moho nobilis

(Merrem)

Yellow-­tufted ­Bee-­eater. Latham 1782, p. 683 Gracula nobilis Merrem 1786, p. 7 (Hawaii) Specimens  Specimens are in Berlin; Bremen; Brussels; Cambridge, Massachusetts; Dresden; Honolulu; Hamburg; Leiden; Liverpool; Los Angeles; New York; Paris; Stockholm; Tring; and Washington, D. C. Status  The last specimen was taken on May 13, 1902, but there was an unconfirmed record in 1934. Range  Mountain forests of Hawaii, Hawaiian Islands. Description  32cm (12.5in). Head, body, wings, and central pair of ­tail-­feathers glossy black, the latter elongated and twisted at tips; remaining rectrices had varying amounts of white or yellow at base; black shading to brown on abdomen; axillary feathers elongate, bright yellow and forming tufts; ­undertail-­coverts yellow; bill, legs and feet black; iris ­reddish-­brown. Female smaller with shorter tail; immature lacked axillary tufts.

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 ­over-­hunting was the ultimate reason for its extinction. How long the bird survived is not clear, but it probably became extinct early in the 20th century. However, Munro (1944), a trusted and veteran Hawaiian ornithologist, was certain he heard one in 1934 on the slopes of Mauna Loa. There have been no other reports. 249

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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 mountains of the Hawaiian Islands, ­bare-­foot and on 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):

Hawaii ’O’o

Moho nobilis 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 ­nectar-­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 ­undertail-­coverts of the same colour can be distinctly seen . . . it presents a very fine appearance.

Molokai ’O’o (Bishop’s ’O’o)  Moho bishopi

(Rothschild)

Acrulocercus bishopi Rothschild 1893b, p. 41 (Molokai) Specimens  Specimens are in Bremen; Cambridge, Massachusetts; Honolulu; Molokai; New York; Stockholm; and Tring. Status  Extinct. Last seen in 1904. Range  Mountain forests of Molokai, Hawaiian Islands.

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Mohoidae Description  29cm (11.5in). Uniform black, with metallic gloss on crown, neck, breast and flanks; elongated ­ear-­coverts, axillary tufts and ­undertail-­coverts golden yellow; tail black with narrow fringe of white; central rectrices long, pointed and curved upwards at tips; bill, legs and feet black; iris brown.

The Molokai ’O’o was last observed by Munro in 1904 (1944), 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 ’O’o  Moho sp. Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known from subfossil remains and accounts. Probably disappeared around the early years of the 20th century. Range  Maui, Hawaiian Islands.

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. 312). 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 ­north-­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.

Kioea  Chaetophila angustipluma

(Peale)

Entomiza angustipluma Peale 1848, p. 147 (Hawaii) Specimens  Specimens are in Cambridge, England; Honolulu; New York; and Washington, D. C.; subfossil remains are in Washington, D. C. Status  Extinct. Last seen in 1859. Range  Hawaii, Hawaiian Islands.

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Extinct Birds Kioea Chaetophila angustipluma

Description  32cm (13in). A large bird with a black face mask, ­greenish-­brown wings and tail, and a heavily streaked pattern of brown and white on the head, upper back and underparts. Top of head and neck ­blackish-­brown striped ­greenish-­yellow; lores, sides of face and ­ear-­coverts dull black; chin and throat dull white; ­wing-­coverts and back ­greenish-­brown, ochraceous on rump; tail deep brown, outer margins yellow; breast and abdomen dull white, striped longitudinally dark brown; bill, legs and feet dark ­blackish-­brown.

This large, attractive member of the Mohoidae is known from just four skins, the last taken by the collector Mills in 1859 (Munro 1844). 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 coloration. 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. 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’.

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Macrosphenidae

Narrow-­billed Kioea  Chaetoptila sp. Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Range  Maui, Hawaiian Islands.

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 specimens 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.

Hirundinidae (Swallows and martins) The Hirundinidae has a worldwide distribution, except in the polar regions and more remote islands, and the family has been divided into 19 genera, with about 83 species. They are characterised by a short, flat ­insect-­catching bill, weak feet and long, pointed wings. Most are not threatened, but a number of forest and island forms are vulnerable due to habitat destruction.

Jamaican Golden Swallow  Tachycineta euchrysea euchrysea

(Gosse)

Hirundo euchrysea Gosse 1847, p. 68 (Jamaica) Specimens  Specimens are in Washington, D. C. Status  Possibly extinct. Not recorded with certainty since 1989. Range  Jamaica, West Indies. Description  12cm (4.75in). Male upperparts (including ­ear-­coverts, malar area and chin) iridescent ­bronze-­green, especially on mantle; primaries and tail darker, dusky ­bronze-­green; underparts white; bill, legs and feet black. Female occasionally indistinctly mottled ­grey-­brown below. Juvenile overall less glossy, more mottled below, with dull grey sides to head.

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 (Stejneger 1893), 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 ­sea-­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 ­cane-­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.

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 subspecies may have become extinct in recent times. 253

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Chapin’s Crombec  Sylvietta leucophrys chapini

Schouteden

Sylvietta chapini Schouteden 1947, p. 193 (Djugu, Lendu Plateau, DR Congo) Specimens  Specimens are in Tuvuren, Belgium. Status  Possibly extinct. Last recorded in 1942. Range  Lendu Plateau, DR Congo. Description  9cm (3.5in). Adult: top and sides of head and nape rich ­chestnut-­brown; mantle and back brownish; rest of upperparts ­olive-­green; ­flight-­feathers brown conspicuously fringed bright yellowish green; chin, throat and midline of abdomen white; rest of underparts ­brownish-­grey; ­underwing-­coverts, ­undertail-­coverts and thighs ­lemon-­yellow; undertail brown; bill greyish pink or ­flesh-­coloured, darker or brownish below; iris pale brown or chestnut; legs ­greyish-­pink to flesh.

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).

Cettiidae (Cettias and stubtails) The Cettiidae contains a number of problematic genera that were formerly placed in the Sylviidae. The family is presently divided into about nine genera with about 34 species. They are widely distributed and occur in Europe, Africa, and Asia eastwards to islands in the South Pacific. Many are forest birds, and some forest and island species are threatened due to deforestation.

Babar Stubtail  Urosphena subulata advena

(Hartert)

Orthnocichla subulata advena Hartert 1906, p. 298 (Babar Island, east of Timor) Specimens  Specimens are in New York. Status  Possibly extinct. Not recorded since the mid-20th century. Range  Babar Island, east of Timor Island. Description  10cm (4in). Entire upperparts plain ­olive-­brown washed rufescent, distinct long, broad white superciliary and dark eye stripe; greyish below, washed ­greyish-­brown on breast; upper bill blackish, lower bill pale; iris blackish; legs and feet ­pinkish-­white. Smaller and more rufescent on upperparts than nominate U. s. subulata, which has light brown upper bill.

The Timor Stubtail Urosphena subulata is a poorly known species, with the nominate occurring on the island of Timor. A race, U. s. advena, formerly existed on Babar Island, to the east of Timor. The population may have disappeared due to the intensive deforestation that has taken place on the island, especially since the late 1980s, when logging began on a large scale. Human encroachment and invasive plant species exacerbate the situation. The Babar Island Stubtail has not been recorded since the 1950s, but if it still exists it must be critically endangered. Habits Little is known about the habits of this skulking bird, but on Timor the extant race inhabits primary and secondary monsoon forest and dry evergreen forest, including scrubby hillsides and gullies, spending most of its time on the ground foraging for invertebrates (Bairlein 2006).

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Acrocephalidae

Phylloscopidae (Leaf warblers) The leaf warblers were originally placed in the Sylviidae. The family contains two genera with about 66 species. They are widely distributed and occur in forest or scrub across Eurasia, Africa and Indonesia. Most species are not threatened, but some island populations are at risk from habitat destruction. Only one subspecies is known to have become extinct.

Lanzarote Chiffchaff  Phylloscopus canariensis exsul

Hartert

Phylloscopus collybita exsul Hartert 1907, p. 505 (Lanzarote, Canary Islands) Specimens  Specimens are in Tring and New York. Status  Probably extinct. Last recorded in 1986 or probably earlier. Range  Lanzarote, and doubtfully Fuerteventura, eastern Canary Islands. Description  10cm (4in). Generally ­olive-­brown upperparts washed with chestnut; breast and flanks ­rufous-­buff; ­undertail-­coverts dull white; pale superciliary stripe; bill, legs and feet blackish. Lighter above and less rufous below than nominate P. c. canariensis.

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. The exact reasons for its disappearance are not known, but much of its habitat was cleared for agriculture, 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.

Acrocephalidae (Reed warblers) The acrocephalid warblers occur widely across Eurasia, with some species in Africa and on Indian Ocean islands, and they have colonised many islands and island archipelagos in the Pacific. The family is divided into four genera with about 50 species. They are generally dull olivaceous or brown birds above with yellowish or whitish underparts. Continental populations are not generally threatened, but the Pacific island species in particular have suffered from habitat destruction, and from predation by introduced mammals. It is likely that prior to colonisation of the Pacific islands by Polynesians, almost every island large enough to support forest had a species of acrocephalid warbler inhabiting it.

Aldabra Brush Warbler  Nesillas aldabranus

Benson & Penny

Nesillas aldabranus Benson & Penny 1968, p. 102 (Aldabra) Specimens  Skins and eggs are in Tring. Status  Extinct. Last recorded in October 1983. Range  Aldabra, ­south-­west Indian Ocean. Description  18–20cm (7–8in). Top of head and upperparts warm brown, more rufous on rump; lores and supercilium ­greyish-­white; mantle and scapulars dark brown, darker on primaries and secondaries; alula and outer feathers of ­lesser ­coverts, chin, throat and centre of abdomen white; ­underwing-­coverts creamy buff; rest of underparts white tinged with buff; flanks pale brown; upper bill dark horn, lower bill light horn; iris brown; legs and feet grey; soles yellowish.

The Aldabra Brush Warbler was restricted to a tiny part of Gionnet, at the west end of Middle Island, Aldabra ­(Pryfls-­Jones 1979). ­Pryfls-­Jones estimated that its greatest distribution was never more than a 50m wide strip extending a total of 9km along the coast – perhaps one of the smallest ranges of any known bird species. 255

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An adult female with its nest and three eggs were collected December 11, 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 ­Pryfls-­Jones’s survey of 1974–1976, when only five individuals were found – three males and two females (Pryfls-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. In February 1977, only two males were definitely known still to survive. In October 1983, just one male was found (Collar & Stuart 1985), and this was the last record. The species is now extinct. 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 responsible for the extinction of the Aldabra Brush Warbler. Goat introductions may also have played a part, as their browsing opens up otherwise dense scrub areas, which are probably less suitable for ­brush-­warbler habitation ­(Pryfls-­Jones 1979). Habits The warbler occurred in areas of rugged pavé and champignon limestone rock, covered by dense mixed scrub including stands of Pandanus, and ‘Pemphis scrub’ communities ­(Pryfls-­Jones 1979). It was insectivorous, and obtained its food by picking, ­leap-­snatching, and probing in leaf litter. Breeding probably took place in October or November, and the call was a harsh chirr or a ‘short, scolding chatter’ (Benson & Penny 1968).

Astrolabe Warbler  Acrocephalus (?luscinia) astrolabii

Holyoak & Thibault

Acrocephalus luscinia astrolabii Holyoak & Thibault 1978b, p. 125 (suggested locality = Yap) Specimens  Specimens are in Paris. Status  Probably extinct. Known only from two specimens collected on the voyage of the Astrolabe (1826–29). Range  Unknown locality but perhaps the Gambier Islands, South Pacific, where no such bird now occurs. Description  17cm (7in). Generally dull ­greyish-­olive above with pale yellowish eyebrow and underparts; dark line through eye; bill dark brown above, buff below; iris dark brown; legs and feet grey. Female slightly smaller than male. The Astrolabe birds are very similar to A. l. luscinia, but differ in their larger size and relatively shorter bill, and their extraordinarily stout feet and claws.

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 Mangareva, Gambier Islands, and the other specimen from 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 specimens. 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 luscinia, which is extant (if critically endangered) in the northern Marianas. 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. 256

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Acrocephalidae

Pagan Nightingale Warbler  Acrocephalus luscinius yamashinae

(Takatsukasa)

Conopoderas yamashinae Takatsukasa & Yamashina 1931, p. 485 (Pagan Island) Specimens  Specimens are in Paris; Tokyo; and Washington, D. C. Status  Probably extinct. Last recorded in the 1970s. Range  Pagan, Marianas Islands, ­south-­west Pacific. Description  17cm (7in). Similar to nominate A. l. luscinia except overall much duller and more brownish; less ­olive-­rufous on back, rump and tail; tail nearly square, less rounded; bill shorter and more curved.

Pagan Island once harboured a distinct population of Nightingale Warbler A. luscinius, 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). The race had been reduced to a very small population during the 1960s, and appears to have been restricted to the area around the island’s two lakes. Severe habitat modification was the primary threat to the warblers as wetlands were drained for agriculture and feral livestock destroyed the understorey, essential for the warbler’s 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 vegetation was almost gone by 1979, and volcanic eruptions in 1981 destroyed much of the remaining woody vegetation. There seems little chance that this race survives. The Nightingale Reed Warbler is an inhabitant of thickets, reed marshes, wetlands, and Tangantangan Leucaena leucocephala scrub (Mosher & Fancy 2002), and can most often be located by its song, which is complex, loud and melodious. The birds are strictly territorial, and remain in the same territory throughout the season. The two remaining races of this species are critically endangered.

Moorea Warbler  Acrocephalus longirostris

(Gmelin)

Long-billed Thrush. Latham 1783, p. 67 (Eimeo & York Island = Moorea, Society Islands) Turdus longirostris Gmelin 1789, p. 823 (Moorea) Specimens  Specimens are in Leiden; New York; and Vienna. Status  Probably extinct since 1987, but one recent unconfirmed report. Range  Moorea, Society Islands. Description  24cm (9.5in) Upperparts pale mottled ­olive-­brown, more yellowish on rump; eyebrow and undersides pale brimstone yellow; ­wing-­coverts and primaries edged yellow; tail brown edged yellow; iris dark brown; upper bill ­horn-­brown, lower bill ­flesh-­coloured; legs and feet ­greenish-­slate.

The Moorea Warbler was first collected by William Anderson during Cook’s third voyage to the South Seas in 1777 (Stresemann 1950). It was long considered to be a subspecies of Tahiti Warbler A. caffer, but the bird was recognised as distinct and given specific status by Cibois et al. (2008). It appeared to be confined to native montane forests, at elevations of between 800m and 900m (King 1981). Three individuals were collected by the Whitney South Seas Expedition in 1921, but even then it was scarce and very restricted in range (King 1981). By 1973, only two pairs could be found, and in 1987 Thibault failed to locate any at all (Anon. 1987; Thibault 1998). There has been one recent unconfirmed report (BirdLife International 2011). The reasons for this bird’s extinction are unclear, but it probably disappeared due to predation by introduced mammals and severe deforestation. However,

Moorea Warbler Acrocephalus longirostris

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since the 1970s, avian malaria has been reported on Moorea, and the Moorean birds are susceptible to the disease (King 1981). This new threat may have extirpated the last of the birds.

Huahine Warbler  Acrocephalus caffer garretti

Holyoak & Thibault

Tatare longirostris Gräffe 1873, p. 49–51 Acrocephalus otatare Tristram 1889, p. 152 Acrocephalus caffer garretti Holyoak & Thibault 1978b, p. 122 (Huahine) Specimens  Specimens are in Dresden; Hamburg; Leiden; New York; and Tring. Status  Extinct. Known from five specimens; not seen since c.1870. Range  Probably occurred on Huahine in the Society Islands, South Pacific. Description  20cm (8in). The nominate Tahiti Reed Warbler is a generally pale yellow bird, mottled with ­brownish-­olive above. The Huahine race was very similar but substantially larger, and the pale cream edges to the feathers of the upperparts of the body were slightly broader.

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 (1978) 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, 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).

Raiatea Warbler  Acrocephalus caffer musae

(Forster)

Conopoderas caffra G. Forster (1773), unpublished watercolour in The Natural History Museum, no. 55 (Tahiti) Oriolus musae J. R. Forster 1844, p. 163. Acrocephalus caffer musae Holyoak & Thibault 1984, p. 124 Specimens  A specimen is in Bremen. Status  Extinct. Disappeared some time after 1873. Known only from J. R. Forster’s description and a recently discovered skin. Range  Raiatea, Society Islands, South Pacific.

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 description 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: 258

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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 ­now-­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.

Laysan Millerbird  Acrocephalus familiaris familiaris

(Rothschild)

Tatare familiaris Rothschild 1892, p. 109 (Laysan) Specimens  Skins are in Bremen; Cambridge, Massachusetts; Chicago; Denver; Honolulu; New York; Pittsburgh; Tring; and Washington, D. C.; and eggs are in Tring. Status  Extinct since between 1913 and 1923. Range  Laysan, Hawaiian Islands. Description  13cm (5in). Generally ­greyish-­brown above, warmer on rump, paler on sides of head; wings and tail dark brown, feathers margined ­buff-­brown; chin, throat and undersides ­buffish-­white; bill brown; legs and feet ­flesh-­brown.

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 vegetation, 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 ­bluish-­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.

Sylviidae (Sylvia warblers) The Sylvia warblers are a complex family of birds of which the taxonomy is under frequent review. Morpohological and genetic evidence has moved a number of genera into different families, while the genus Bowdleria, the fernbirds, is tentatively placed here pending further study. Some authorities place this in the Megaluridae with the thicketbirds. Sylvia warblers are widespread and generally not threatened. Only one species is known to have become extinct. 259

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Chatham Islands Fernbird (Mata)  Bowdleria rufescens

(Buller)

Sphenoeacus rufescens Buller 1869, p. 38 (‘small rock isle, a satellite of Chatham Island’) Specimens  Specimens are in Auckland and Christchurch, New Zealand; Berlin; Cambridge, Massachusetts; Chicago; Liverpool; New York; Paris; Pittsburgh; and Tring. Status  Extinct. Not seen since 1900. Range  Formerly occurred on Pitt Island and Mangere Island, in the Chatham Islands, New Zealand. Description  18cm (7in). Upperparts, sides and tail dark ­rufous-­brown, brightest on crown and hind neck; sides of head and ­ear-­coverts marked with black; feathers of shoulders and sides centred with black; primaries dusky black, margined with ­rufous-­brown; supercilium, throat, breast and abdomen pale fawn to white; bill light brown, with culmen ridge black; iris ­orange-­brown; legs and feet dark brown. The feathers of the tail were extremely modified, being ­hair-­like and spiny.

The type of this species was collected by a conchologist, Charles Traill. He had found it on Mangere Island during a conchological expedition, but he was unable to say anything about the bird’s habits. Traill observed it flitting among grass and stunted vegetation, ‘and succeeded in knocking it over with a stone’ (Buller 1869). The natural vegetation of the Chatham Islands was devastated during the latter part of the 19th century, when the islands were settled. This was partly accomplished by burning and partly by the grazing of rabbits and goats. Before that, cats and rats had been accidentally introduced. Rothschild (1907a) believed that the Fernbird was exterminated by cats, rats and weasels Mustela nivalis. Henry Palmer had collected fourteen specimens for Rothschild, and Travers and Dannefaerd collected most of the others known. The last specimen was shot by one of Rothschild’s collectors on northern Mangere Island on a very steep scrubby precipice, said to have been inaccessible to cats; the species has not been recorded since 1900 (Stattersfield et al. 1998). Fleming (1939b) was of the opinion that this area was undisturbed and could still harbour birds, but Greenway (1967) considered this unlikely. It is certainly extinct today. Habits Almost nothing is known about the Chatham Island Fernbird’s habits, but 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 invertebrate 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.

Locustellidae (Grasshopper warblers) The grasshopper warblers, which would be more appropriately called grassbirds, are a complex group of passerines that were originally placed in the Sylviidae. Their relationships are poorly understood and some genera probably warrant familial status. At present, the family is divided into 12 genera with about 50 species. The grasshopper warblers have a wide distribution across Eurasia, Africa, Asia and many island groups. The island populations are vulnerable to anthropogenic changes, but only two subspecies are known to have become extinct.

Bismarck Thicketbird  Megalurulus grosvenori

(Gilliard)

Cichlornis grosvenori Gilliard 1960, p. 1 (Wild Dog Range, Whiteman Mountains, central New Britain) Specimens  The type specimen is in New York. Status  Possibly extinct, but data deficient; known only from two specimens collected in 1958, and never seen since. Range  New Britain, Bismarck Archipelago, ­south-­west Pacific. Description  17cm (6.75in). Mantle dark ­olive-­brown, becoming darker on lower back and rump; crown bordered laterally by prominent ­ochraceous-­tawny superciliary stripes; broad black ­face-­mask; undersides generally ochraceous-tawny, paler, more

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Cisticolidae pale ­ochraceous-­buff on chin, throat, and central abdomen, becoming darker, more tawny, on chest, sides of neck, and flanks; sides of lower abdomen darker, more washed with dark cinnamon brown; wings blackish, tail sharply graduated, the shafts black, stiff, and with long, ­spine-­like tips; bill brownish black; iris dark brown, with dull green rim; legs and feet dark ­smoke-­brown.

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 limestone 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.

Vanua Levu ­Long-­legged Thicketbird  Megalurulus rufa cluniei

(Kinsky)

Trichocichla rufa cluniei Kinsky 1975, p. 100 (Delanacau Mountains, Vanua Levu, Fiji) Specimens  The unique type is in Wellington, New Zealand. Status  Possibly extinct. Last seen in June 1974. Range  Vanua Levu, Fiji. Description  17cm (6.75in). Differs from the nominate T. r. rufa in having supercilium, chin and belly more conspicuous white. Adult upperparts ­rufous-­brown with distinct long white supercilium extending to back of neck; chin and throat white; ­breast-­sides and flanks ­rufous-­buff, fading into an ­off-­white belly; bill black; legs and feet bluish.

The rediscovery of the nominate ­Long-­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).

Cisticolidae (Cisticolas and allies) The Cisticolidae is divided into 30 genera with about 110 species. Africa is the centre of distribution, but species occur in tropical Asia and Australasia, with one species in Europe. Many are poorly known, and their generally drab plumage and elusive nature makes them difficult to locate. The birds are generally found in open country, but some occur in forest. Two subspecies, both ­forest-­dwellers, may have become extinct. 261

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Northern ­White-­winged Apalis  Apalis chariessa chariessa

Reichenow

Apalis chariessa Reichenow, 1879, p. 114 (Mitole, Tana River, Kenya) Specimens  A specimen is in Tring. Status  Possibly extinct. Last recorded in 1961. Range  Mitole, lower Tana River, eastern Kenya. Description  15 cm (6in). Male upperparts glossy black; bright, ­golden-­yellow underparts with orange lower breast and belly; throat white; ­breast-­band black; conspicuous white ­wing-­panels; tail long and black with white tips; bill black; legs and feet ­pinkish-­brown; iris ­yellowish-­brown. Female duller; head dark grey; underparts ­olive-­green; throat and ­breast-­band dark grey.

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.

Western Turner’s Eremomela  Eremomela turneri kalindei

Prigogine

Eremomela turneri kalindei Prigogine 1958, p. 147 (Kailo, Belgian Congo) Specimens  The type specimen is in Tervuren, Belgium. Status  Possibly extinct. Not recorded since the late 1970s. Range  DR Congo and the Nyondo forest, ­south-­west Uganda. Description  8–9cm (3.25–3.5in). Generally grey above, white below. Forehead rufous; black ­breast-­band; flanks grey; bill blackish; legs and feet flesh brown. Juvenile ­olive-­brown above, paler underparts; and only slight ­breast-­band.

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 ­mid-­altitudinal montane forests, and preferred the canopy of large trees, especially ­mid-­size and mature species such as Croton megalocarpus (Otienio et al. 2011).

Timaliidae (Babblers) The Timaliidae is an extremely diverse group of birds whose relationships are unclear. Up to 56 genera have been included in the family. The babblers have a wide distribution, and occur in a range of habitats. These birds are generally not considered threatened.

Black-­browed Babbler  Malacocincla perspicillata

(Bonaparte)

[Cacopitta] perspicillatum ‘Temminck Museum Lugd’ Bonaparte 1850, p. 257 (Java = error for Borneo) Specimens  The type specimen is in Leiden. Status  Possibly extinct. Not recorded since the type was collected in the 1840s. Range  Uncertain, but thought to be southern Borneo. Description  16cm (6.25in). Forehead and supercilium black; rest of head grey; upperparts, wings and tail ­olivaceous-­brown, becoming warmer on rump; chin, lores and throat white; rest of underparts ­plumbeous-­grey streaked with white; ­undertail-­coverts ­rufous-­brown; bill blackish; legs and feet pinkish.

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Black-browed Babbler Malacocincla perspicillata

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.

Irrawaddy Jerdon’s Babbler  Moupinia altirostris altirostris

(Jerdon)

Chrysomma altirostre Jerdon 1862, p. 22 (‘Islands on the Burrampootra River in Upper Burmah’; type from an island in the Irrawaddy of Thayetmyo, according to Sharpe 1883) Specimens  The type specimen is in Tring. Status  Not reported since 1941 and possibly extinct. Range  South-central Burma. Description  16cm (6.25in). Upperparts pale ­reddish-­brown, deepest on wings and tail; forehead and streak over eye hoary grey; undersides whitish, tinged on lower breast, abdomen, and flanks pale ­tawny-­brown; primaries and tail feathers slightly dusky on inner webs; ­underwing-­coverts pale ferruginous; bill fleshy horn; legs ­flesh-­coloured; iris dark brown with outer circle of white; eyelids pale ­sulphur-­yellow.

There are two extant races of Jerdon’s Babbler Moupinia altirostris, M. a. griseigularis in the Himalayas from Bhutan east to Assam and north-east Burma, and M. a. scindica in the Indus basin. There was a third race, the nominate, which formerly occurred on grasslands that covered the ­Irrawaddy-­Sittang plains of southern and central Burma. Smythies (1953) remarked: the [nomino-]typical race is known only from the Irrawaddy-Sittang plain; here it was widely distributed in the vast areas of high grass that covered the plain in Oates’s day, and we were able to prove in 1941 that it still exists in the small remnant of the great Myitkyo swamp . . . it has not been observed anywhere else in recent years. Habitat destruction was probably responsible for its disappearance. 263

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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 Ocean, Gulf of Guinea and the western Pacific. Island endemicity is high. Some island populations are seriously threatened by habitat destruction and introduced mammalian predators.

Chestnut-­flanked White-eye  Zosterops (mayottensis) semiflava

Newton

Zosterops semiflava E. Newton 1867, p. 354 (Marianne Island, Seychelles) Specimens  Specimens are in Cambridge, England, and Tring. Status  Probably extinct. Range  Known with certainty only from Marianne Island, Seychelles. Description  10cm (4in). A greenish-yellow bird with chestnut flanks and a prominent white eye-ring. Forehead and supercilium yellow; top of head and back yellow olive; wings and tail black; underparts pale yellow. Smaller than Z. m. mayottensis, overall slightly less yellow; frontal bar duller.

Edward Newton’s account (1867) 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. I was told that this bird was also found at La Digue, Praslin, and Silhouette. I am pretty certain I saw it once at Mahé, but I did not succeed in getting a shot at it. Other than Newton’s account, nothing else is known about it. 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 1870 and 1900. The decline of this bird was due to deforestation, competition with introduced birds, and predation by the Black Rat Rattus rattus. If it did occur on Mahé in the past, it would have been sympatric with the endangered Seychelles ­White-­eye Z. modestus, itself once thought to have been extinct. The nominate Z. m. mayottensis is still relatively common on Mayotte in the Comoros Islands (BirdLife International 2011).

Guam Bridled ­White-­eye  Zosterops conspicillatus conspicillatus

(Kittlitz)

Dicaeum conspicillatum Kittlitz 1833, p.15, pl 19, fig. 1 (Guam) Specimens  Specimens are in Honolulu and Washington, D. C. Status  Extinct. Last recorded in 1983. Range  Guam, Mariana Islands, ­south-­west Pacific. Description  12cm (4.75in). Adult upperparts bright ­olive-­green; ­ear-­coverts olivaceous; wings and tail dark brown with ­greenish-­yellow edges; chin and throat ­yellowish-­white; breast and abdomen dingy ­yellowish-­white; orbital ring white; upper bill yellow, lower bill lighter; iris light amber; legs and feet dark olive-grey. Female generally paler.

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 individuals, 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 264

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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).

Robust Silvereye (Lord Howe ­White-­eye)  Zosterops strenua

Gould

Zosterops strenuus Gould 1855, p. 166. (Lord Howe Island) Specimens  Specimens are in Cambridge, England; Leiden; New York; Sydney; and Tring. Status  Extinct since c.1918. Range  Endemic to Lord Howe Island. Description  7.6cm (3in). Larger than the other endemic ­white-­eye on Lord Howe Island, Z. tephropleura (see p. 456), and distinguished by its bright yellow (not greenish) throat. Separated from the Norfolk Island Z. albogularis by white (not yellow) belly. Adult head and upperparts bright ­olive-­green; grey band across shoulders; wings and tail ­slaty-­brown margined with greenish olive; chin and throat yellow; flanks pale ­vinaceous-­brown; ­underwing-­coverts and axillaries white; ­undertail-­coverts pale yellow; bill, legs and feet ­bluish-­black; iris light brown.

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. 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 materials, and placed among the masses of fibre clothing the undersides of kentia palms, or in shrubs overgrown with vines. 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, and remarked that ‘at Lord Howe Island, its powerful song makes music all day long in the ­palm-­glades and on the wooded hillsides’.

Robust Silvereye Zosterops strenua

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White-­breasted ­White-­eye (Norfolk Island ­White-­eye) Zosterops albogularis

Gould

Zosterops albogularis Gould 1837, p.75 (’Murrumbidgee River’ = error for Norfolk Island) Specimens  Specimens are in Tring and New York. Status  Possibly extinct. Last recorded in March 2000. Range  Norfolk Island. Description  13–14cm (5–5.5in). Adult head bright green; back ­olive-­green; underparts white; sides flushed rufous; ­undertail-­coverts lemon yellow; ­eye-­ring white; bill, legs and feet blackish.

The ­White-­breasted ­White-­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 in 2000 is the last confirmed report (Garnett & Crowley 2000). 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). 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, occasionally 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.

Mukojima ­White-­eye (Meguro)  Apalopteron familiare familiare

(Kittlitz)

Ixos familiaris Kittlitz 1831, p. 235, pl. 13. (Boninshima; restricted to Mukojima by Yamashina 1930) Specimens  Specimens are in Frankfurt and St. Petersburg. Status  Extinct since 1941. Range  Formerly confined to Mukojima and ­Nakōdo-­jima, and probably Chichijima, in the northern Ogasawara (Bonin) Islands, Japan. Description  13.5cm (5.25in). The plumage was long and fluffy. Forehead and eye surrounded by black; yellow spot near base of bill; throat, cheeks and underside yellow, tinged ­greenish-­grey on sides of abdomen; upperparts grey, becoming greenish on base of tail, and yellow on crown; quills and tail dark brown with yellowish green margins to outer webs; bill and feet blackish; iris light brown.

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). 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 1890), 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. 266

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Regulidae

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.

Guadalupe ­Ruby-­crowned Kinglet  Regulus calendula obscurus

Ridgway

Regulus calendula obscurus Ridgway 1876b, p. 184 (Guadalupe Island) Specimens  Specimens are in Washington D. C. Status  Probably extinct. Not recorded since the 1990s. Range  Guadalupe Island, off Baja California, Mexico. Description  9–11cm (3.5–4.25in). ­Grey-­green upperparts, ­olive-­buff underparts; two white ­wing-­bars, the larger bordered by a dark band; white broken ­eye-­ring. Male and female identical except male had a scarlet ­crown-­patch, which was usually hidden. Eggs white, dotted and splashed with ­reddish-­brown.

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 & Martin 2006). This tiny bird had managed to survive the ­human-­induced catastrophic events that eliminated 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 ­cup-­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 & Martin 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: 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’.

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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 distributed 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.

San Benedicto Rock Wren  Salpinctes obsoletus exsul

Ridgway

Salpinctes obsoletus exsul Ridgway 1903, p. 169 (Isla San Benedicto, Islas de Revillagigedo) Specimens  Specimens are in Washington D. C. Status  Extinct since 1952. Range  San Benedicto, in the Revillagigedo Islands, Mexico. Description  12cm (4.75in). Resembled nominate S. o. obsoletus, but the lateral rectrices were more numerously barred with ­cinnamon-­buff; wing averaged shorter; tail longer; and bill and feet larger. General coloration ­greyish-­brown with ­vinous-­cinnamon rump; most of upper surface marked with ­wedge-­shaped dark spots; supercilium pale; underparts dull white, merging into pale ­cinnamon-­buff on flanks.

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.

San Clemente Wren  Thryomanes bewickii leucophrys

Anthony

Thryomanes leucophrys Anthony 1895a, p. 52 (San Clemente Island, California) Specimens  Specimens are in New York; Tring; and Washington, D. C. Status  Extinct since 1968. Range  San Clemente, Channel Islands, California. Description  14cm (5.5in). Predominantly brown above with a grey wash; ­greyish-­white below, paler on throat and ­undertail-­coverts; white ­eye-­stripe passing through and behind eye; wings and tail barred light and dark ­blackish-­brown; ­undertail-­coverts barred ­brownish-­black; bill ­blackish-­brown, lighter at base of mandible; iris brown; legs and feet dark brown. ­Longer-­billed than nominate T. b. bewickii, which also has more heavily barred ­undertail-­coverts and lacks grey wash on upperparts.

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). 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.

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Troglodytidae

Guadalupe Wren  Thryomanes bewickii brevicauda

Ridgway

Thryomanes brevicauda Ridgway 1876b, p. 186. (Guadalupe) Specimens  Specimens are in Cambridge, Massachusetts; Chicago; Tring; and Washington, D. C. Status  Extinct since 1897. Range  Guadalupe Island, off Baja California, Mexico. Description  14cm (5.5in). Similar to most other Bewick’s Wren races, from which it differed only in minor details of the barring of the tail. Greenway (1967) stated that it was closest to T. b. correctus of southern California, differing in having narrower black bars on the tail feathers, and in being much smaller.

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 specimen, 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.

Martinique House Wren  Troglodytes aedon martinicensis

(Sclater)

Thryothorus martinicensis Sclater 1866, p. 321 (Martinique) Specimens  Specimens are in Cambridge, Massachusetts; Chicago; New York; and Tring. Status  Probably extinct, the last specimen being taken in 1886. Range  Martinique, West Indies. Description  11–13cm (4.25–5in). The West Indian island races were all fairly similar in appearance, greyish or tawny brown above, white or pale cinnamon below. The Martinique race was larger than T. a. guadeloupensis of Guadeloupe, but with smaller tail; duller in colour; flanks pale cinnamon; and ­undertail-­coverts barred black and tipped white.

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: 269

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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 thought extinct but was rediscovered in 1969 (Kroodsma & Brewer 2005); 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.

Mimidae (Mockingbirds and thrashers) The mockingbirds and thrashers are an exclusively New World family of birds, divided into 12 genera with about 30 species. They are ­medium-­sized terrestrial birds that forage in low undergrowth or on the ground. Some species are widely distributed, but others are restricted to islands, where they are threatened by habitat destruction and predation by introduced mammals.

Barbados ­Scaly-­breasted Thrasher  Margarops fusca atlantica

(Buden)

Allenia fusca atlantica Buden 1993a, p. 82 (Barbados, West Indies) Specimens  Specimens are in Tring and New York. Status  Possibly extinct. Not recorded since 1990. Range  Barbados, West Indies. Description  23cm (9in). Upperparts ­blackish-­brown; underparts white with ­blackish-­brown scaling; thin white wing-bars; rump rufous brown; tail dark with white terminal spots to rectrices; bill black; iris yellow; legs and feet ­pinkish-­grey.

The ­Scaly-­breasted Thrasher M. fusca 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 probably 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).

Cozumel Thrasher  Toxostoma guttatum

(Ridgway)

Harporhynchus guttatus Ridgway 1885c, p. 21 (Cozumel Island) Specimens  Specimens are in Washington D. C. Status  Possibly extinct. Not recorded since December 2007. Range  Cozumel Island, off ­north-­eastern Yucatán Peninsula, Mexico. Description  23cm (9in). A predominantly brown and white bird with a long, decurved bill. Adult upperparts rich ­chestnut-­brown with two ­blackish-­bordered white ­wing-­bars; face greyish with paler supercilium; underparts white heavily streaked with black; bill, legs and feet black; iris ­reddish-­brown.

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Sturnidae

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. Habits This thrasher was a skulking bird that foraged on or near to the ground, which made it vulnerable to introduced 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.

Réunion Crested Starling (Huppe, Hoopoe Starling) Fregilupus varius

(Boddaert)

? ‘Tiuouch’. Flacourt 1658, p. 166 La Huppe noire et blanche, du Cap de Bonne Espérance. De Montbeillard, in Buffon 1770–1783, p. 463 Huppe du Cap de Bon Espérance. Daubenton 1771–86, pl. 697 Madagascar Hoopoe. Latham 1782, p. 690 Upupa varia Boddaert 1783, p. 43 (no locality = Réunion) Upupa capensis Gmelin 1788, p. 466 La Huppe grise. Audebert & Vieillot 1802, p. 12, pl. 3 Le Mérops Huppé. Levaillant 1807, p. 43, pl. 18 Upupa madagascariensis Shaw 1811, p. 140 Upupa capensis Cuvier 1817, p. 407 Coracias tivouch Vieillot 1817, p. 3 Pastor upupa Wagler 1827, p. 90 Specimens  Specimens are in Brussels; Cambridge, England; Cambridge, Massachusetts; Florence; Genoa; Leiden; Lausanne; Mauritius; New York; Paris; Pisa; Réunion; Stockholm; Tring; and Troyes. Status  Extinct. Last recorded in the mid-19th century. Range  Réunion, Mascarene Islands. Description  30cm (12in). Male head and hindneck light ­ash-­grey, paler (almost white) on underside; back ­ash-­brown, lower back, rump and ­uppertail-­coverts washed rufous; wings and tail ­blackish-­brown; primary coverts white with brown tips; beak, legs and feet yellow; iris pale ­whitish-­blue. Female similar except smaller, with a straighter bill; juvenile browner, crest smaller.

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. 271

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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 (1873) examined its osteology in great detail, and confirmed its relationship to the starlings. A recent mitochondrial DNA analysis (Zuccon et al. 2007) 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 (see also Cheke & Hume 2008). 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: [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 ­game-­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, and this is probably correct. Perhaps this was contracted from introduced Common Mynas Acroditheres tristris, though these birds had been on Réunion on and off since 1759 (Cheke & Hume 2008). Deforestation accelerated from the 1830s, especially as coffee plantations began to give way to sugar cane, and the emancipation of the slave population from 1848 resulted in severe degradation of the comparatively untouched mountain cirques. Maybe all of these events combined to send the Huppe to extinction. Habits It was a forest bird, living in large flocks in lowland and montane forests. 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) received four live birds from his friend Sauzier in May 1835. Desjardins noted that they would ‘eat anything’, and that two escaped into the Mauritian forest and might populate it. Desjardins (in Oustalet 1897) wrote: We do not hesitate today to report that the Huppe, which lives in the forests of Bourbon and which is kept as a ­cage-­bird, may have become naturalised in Mauritius. Mr Autard has presented us one which had recently been 272

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killed in the District of Savane, and we ourselves had 4 which our colleague the elder Mr Sauzier, long term correspondent from Bourbon, had brought us from that neighbouring island, and which, after several months in captivity, took flight, and made off towards the woods, now so rare, in the District of Flacq. Thus, at least five birds escaped on Mauritius and the unfortunate fate of one is known. What happened to the rest cannot now be established.

Rodrigues Starling  Necropsar rodericanus

Günther & Newton

Necropsar rodericanus Günther & Newton 1879, p. 427 (Rodrigues, Mascarene Islands) Specimens  Subfossil remains are in Cambridge, England; London; Paris; Rodrigues; and Tring. Status  Extinct. Known from one account and subfossil remains. Extinct since the mid-18th century. Range  Rodrigues, Mascarene Islands. Description  25–30cm (10–12in). A large starling, similar in appearance to Fregilupus varius but lacking the distinctive crest. General colour white or ­greyish-­white; wings and tail ­blackish-­brown; bill, legs and feet yellow.

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. 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 wedgeshaped 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. Rodrigues Starling Necropsar rodericanus scavenging a juvenile Rodrigues Giant Tortoise Cylindraspis peltastes

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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 1726 ­(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 (from A. Newton’s translation, 1875a): A little bird is found which is not very common, for it is not found on the mainland. One sees it on the islet au Mât [= Île Gombrani], which is to the south of the main island, and I believe it keeps to that islet on account of the birds of prey which are on the mainland, as also to feed with more facility on the eggs of the fishing birds which feed there, for they feed on nothing else but eggs or turtles dead of hunger, which they well know how to tear out of their shells. These birds are a little larger than a blackbird [Réunion Bulbul Hypsipetes borbonicus], and have white plumage, part of the wings and tail black, the beak yellow as well as the feet, and make a wonderful warbling. I say a warbling, since they have many and altogether different notes. We brought up some with cooked meat, cut up very small, which they eat in preference to seeds. Tafforet’s account describes the starling’s ability to tear dead (juvenile?) turtles or tortoises out of their shells, and a strong gape, as seen in the cranial anatomy, would have been useful for this purpose. No doubt the birds utilised a variety of food items, but Tafforet stated that they could be easily reared using meat for food. He never saw them on the mainland, which infers that he was able to procure young birds from a breeding population on Île Gombrani and take them back with him. Leguat (1708) mentioned that the pigeons also only bred on the islets, due to rat persecution, so it is likely that the starling was doing the same. It was either absent or very rare on mainland Rodrigues in 1726, and when Pingré (1763) visited Rodrigues to observe the Transit of Venus in 1761, the starling was extinct. Rats could have arrived nearly a century before Leguat’s sojourn in 1692–93, as at least one Dutch fleet surveyed the island in 1601 (Moree 2001), which may have allowed these rodents to escape. The islets provided the only refuge, but when rats eventually colonised them, the bird’s fate was sealed. 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.

Norfolk Island Starling  Aplonis fusca fusca

Gould

Aplonis fusca Gould 1836, p. 73 (Murrumbidgee River, Australia = error for Norfolk Island) Specimens  Skins are in Leiden and Tring; eggs are in Tring; and subfossil remains are in Norfolk Island. Status  Extinct since 1925. Range  Norfolk Island. Description  20cm (8in). Male dark grey glossed on head with green spreading to the throat and breast, duller on the upper parts of body; underparts grey becoming white on ­undertail-­coverts; bill and feet black; iris ­reddish-­orange. Female generally lighter; greenish gloss less obvious; throat and breast shading to brown on sides and flanks; centre of lower breast washed ochraceous; and lower abdomen and ­undertail-­coverts ­yellowish-­white.

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 (1836) 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 274

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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 ­pine-­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 ­pine-­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 (Mathews 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 ­bluish-­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.

Lord Howe Starling  Aplonis fusca hulliana

Mathews

Aplonis fuscus hullianus Mathews 1912b, p. 451 (Lord Howe Island) Specimens  Skins are in Leiden; Liverpool; New York; Sydney; and Tring; eggs are in Tring. Status  Extinct. Disappeared some time after 1918. Range  Formerly occurred on Lord Howe Island. Description  18cm (7in). Differed from the nominate in being grey below, darker above, the head slightly glossy, with the bill stouter. It was also smaller. Head, neck, shoulders, chin and throat glossy green; back dark grey overlaid with a faint greenish tinge; rump and underparts grey; tail grey tipped with brown; wings brown; bill, legs and feet black; iris ­orange-­red.

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).

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Kosrae Starling  Aplonis corvina

(Kittlitz)

Lamprotornis corvina Kittlitz 1833, p. 12, pl. 15 (Kusaie Island = Kosrae, Caroline Islands) Specimens  Specimens are in Leiden and St. Petersburg. A skin formerly in Frankfurt has disappeared. Status  Extinct. Disappeared during the mid-19th century. Range  Kosrae, Caroline Islands, ­south-­west Pacific. Description  25cm (10in). A large glossy black starling, with a long, deep ­crow-­like bill and long tail. Generally glossy black with faint ­steel-­green and purple shimmer; eyebrow shimmering ­purple-­red; bill and feet black; iris bright red. Juvenile ­yellowish-­white with black and brown speckles, especially on upperparts; tail brown with ­yellowish-­white base; bill black, buff at base; iris light brown.

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: 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. The bird inhabited mountain forests on the island, but Kittlitz found it rare and solitary. 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 occasionally 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.

Mauke Starling (Mysterious Starling)  Aplonis mavornata

Buller

? Sturnus Mautiensis Bloxam, unpublished MS Aplonis mavornata Buller 1887–88 p. 25 (locality unknown) Aplonis inornata Sharpe 1890, p. 135 and footnote (? Raiatea = error, no locality) Specimens  The type specimen is in Tring. Status  Extinct. Known only from a unique specimen, without locality data. Range  Mauke, Cook Islands. Description  18.5cm (7.25in). Uniform dull dark brown, the edges of the feathers lighter, giving a scalloped effect; slight purplish gloss on head; bill legs and feet dull yellowish; iris yellow.

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. 278), and therefore the 276

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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: N.2. Sturnus. L[ength] 7 and a half inch. Color a light brownish black all over, the feathers edged round with a lighter shade of brown. Bill strong, 1 inch long. Lower m[andible] straight, upper m[andible] compressed, rather curved & slightly notched at the tip. Nostrils at the base oval. Tongue at the extremity bifid [forked]. Tail short, 12 equal [feathers], rounded at the tips. Legs strong, outer toe not connected with the middle. Iris yellow. A draft list of 121 specimens, mainly from South America, which Bloxam presented to the Admiralty on his return, included the three birds collected on Mauke: the starling was no. 32 on the list, taken from the label attached to the bird. Olson compared the type of mavornata with Bloxam’s description and found that they agreed well, so he considered that they were one and the same. The other two Bloxam Mauke Starling Aplonis mavornata specimens, the Mauke Kingfisher Halcyon tuta mauke and Mauke Fruit Dove Ptilinopus rarotongensis ‘byronensis’ (see p. 158), are either lost or have perished. Mauke was not ornithologically surveyed until 1973 (Holyoak 1974a), and by this time the Mauke Starling had long since disappeared. Bloxam (in Olson 1986c) in 1825 mentioned the numerous presence of what were probably Brown Rats Rattus norvegicus, which may have reached the island just two years before. They would more than likely have exterminated the starling population. Nothing is known about the bird, other than it was ‘shot hopping about in a tree’.

Huahine Starling  Aplonis diluvialis

Steadman

Aplonis diluvialis Steadman 1989c, p. 164 (Fa’ahia Archaeological Site, Huahine, Society Islands) Specimens  The type specimen is in Honolulu. Status  Extinct. Known only from subfossil remains. Range  Huahine, Society Islands, South Pacific.

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 disappeared 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. 277

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Steadman (1989c) suggests (with some authority) that the mysterious Bird of Raiatea ‘Turdus’ ulieitensis, known only from a painting by Georg Forster executed in 1774, is an Aplonis species related to the Huahine bird. We agree with this hypothesis.

Bay Thrush  Aplonis (‘Turdus’) ulietensis

(Gmelin)

Bay Thrush. Latham 1783, p. 35 Turdus ulietensis Gmelin 1789, p. 815 (Ulietea = Raiatea) Turdus badius J.R. Forster 1844, p. 239 (‘Oriadea insula’ = Raiatea); painting by G. Forster, plate 146 in the NHM Status  Known only from a painting and descriptions. Not seen since May 1774. Range  Raiatea, Society Islands, South Pacific. Description  22cm. Head dusky, general colour variegated brown, the small feathers fuscous margined brown. Upperparts fuscous, feathers edged with rufescent brown; underside ochraceous; wings fuscous, outer edges of primaries and ­wing-­coverts edged brown; ­underwing-­coverts fuscous edged with brown, rest of underside of wing ­dusky-­grey; tail fuscous, slightly edged with brown; bill ­ashy-­horn; iris dusky ­reddish-­yellow; legs and feet black; claws ­flesh-­coloured. The tongue was arrow-like with the tip bifid (forked) and ciliated.

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 specimen 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 reddishpearl 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. Forster’s artistic style After examination of the original painting by JPH, it is evident that the illustration was a ‘field sketch’ that was intended to be worked up at a later date. Georg Forster started with a pencil outline of the bird, followed by an attempt at making a natural perch, but probably time (and the large number of specimens that were being procured) was against any such background luxury. The basic colours were then added, with a rather crude wash. No doubt the availability of paint pigments on an 18th century voyage was limited, so determining any diagnostic colour characters from a field illustration alone can be problematic. Forster also made an error in his original pencil sketch, as he drew the legs and feet too large compared with the rest of the body. The legs and feet were subsequently redrawn, but the original pencil error can still be seen. This implies that the bird was not laid on the paper and an outline drawn around the specimen – a common technique to get the proportion right – but rather an attempt was made to portray the specimen in a more ­life-­like pose. This was no mean feat, bearing in mind the arduous conditions of an 18th century voyage. Taxonomic confusion It was Tristram (1879), Seebohm (1881) and Sharpe (1890, 1906) 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 disregarded as it bears little or no resemblance to the original. Sharpe (1890, 1906) twice suggested that it was the same as the erstwhile Mysterious Starling (now 278

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known as Mauke Starling Aplonis mavornata, see p. 276), but Wiglesworth (1892) noted that the measurements 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 (1986) has shown that the provenance 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 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 ­scale-­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 ­Eboona-­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. ­Mosquito-­borne diseases have also contributed to the almost complete extinction of the endemic Myadestes Hawaiian thrushes.

Kittlitz’s Thrush  Zoothera terrestris

Kittlitz

Zoothera [Aegithocichla] terrestris Kittlitz 1830 (1831), p. 245 (Boninsima = Chichijima, Ogasawara Islands) Specimens  Skins are in Frankfurt; Leiden; St. Petersburg; and Vienna. Status  Extinct. Known from four specimens collected in 1828. Range  Chichijima (formerly Peel Island), Ogasawara (Bonin) Islands, Japan. Description  16.5cm (6.5in). A ­medium-­sized olive-brown thrush; upperparts shading into chestnut on rump, uppertail-coverts and tail; inside web of each feather much darker, approaching black on back; pale eye-stripe; wings olive-brown, coverts mottled darker; chin, throat and centre of belly pale, nearly white, shading into brown on remainder of underparts, with large but faint dark spots on breast; undertail-coverts brown with white tips; bill dark brown.

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 ­ground-­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. 283) became extinct within a few years of rats invading Lord Howe Island, so it is conceivable that Kittlitz’s Thrush disappeared some time during the 1830s, for the same reason.

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Kibale ­Black-­eared Thrush  Zoothera cameronensis kibalensis

Prigogine

Zoothera kibalensis Prigogine 1978, p. 489 (Kibale Forest, ­south-­western Uganda) Specimens  Specimens are in Los Angeles and Tervuren, Belgium. Status  Possibly extinct. Known only from the ­co-­types, collected in December 1966. Not recorded since. Range  ­South-­western Uganda. Description  16.5–18cm (6.5–7in). Larger than nominate Z. c. cameronensis; face pattern more defined on a pale ­buff-­brown; bare skin behind eye dark grey; crown and upperparts more rufous; nape slightly paler; throat, breast and flanks bright orange; centre of belly to ­undertail-­coverts white; base of bill more compressed; and lower bill grey.

Three subspecies of the ­Black-­eared Thrush Z. cameronensis are generally recognised, Z. c. cameronensis from Cameroon and Gabon, Z. c. graueri from DR Congo, and Z. c. kibalensis from ­south-­western Uganda (Clement & Hathaway 2000). The Kibale race Z. c. kibalensis is sometimes regarded as a full species, or sometimes as a weakly differentiated subspecies. 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). The species in general is declining due to deforestation, and much of the Kibale forest area has already been lost to logging and agriculture. If it survives, it must be critically endangered. Habits Little is known about the Kibale ­Black-­eared Thrush. It seems to have favoured montane herbaceous ground vegetation, in pristine mature forests of ironwood and mahogany, but may also have occurred at lower elevations. It probably foraged for invertebrates on the ground. The ­co-­types were taken in thick secondary forest (King 1981).

Choiseul ­Russet-­tailed Thrush  Zoothera heinei choiseuli

(Hartert)

Turdus dauma choiseuli Hartert 1924, p. 273 (Choiseul Island, Solomon Islands) Specimens  The type specimen is in New York. Status  Possibly extinct since the 1940s. Known from only one specimen, collected in January 1904. Range  Choiseul, Solomon Islands. 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. 154). The thrush was never reported again. 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 during the 1940s. Nothing else is known about it.

Isle of Pines Solitaire  Myadestes elisabeth retrusus

Bangs & Zappey

Myadestes elisabeth retrusus Bangs & Zappey 1905, p. 208 (Pasadita, Isle of Pines, Cuba) Specimens  The type specimen is in Cambridge, Massachusetts. Status  Last seen in 1934, and probably extinct. Range  Confined to the Isle of Pines, near Cuba. Description  19cm (7.5in). Generally upperparts ­greyish-­olive; whitish ­eye-­ring; ­pre-­ocular supercilium pale buff; lores and outer tail feathers whitish; underparts pure white; upper bill blackish, lower bill ­horn-­coloured with dusky tip; legs and feet ­yellowish-­brown. Less ­brownish-­olive above than nominate M. e. elisabeth, which is dull white below and grey across breast, and has larger, wider and blunter bill.

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 280

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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 specimen was ever collected. The reasons for its disappearance are unknown.

Oahu Thrush (Amaui)  Myadestes woahensis

Bloxam

‘Turdus woahensis’ Bloxam, unpublished MS ‘variety of Turdus sandwichensis’ Bloxam 1826, p. 250; nomen nudum. Gives no description Phaeornis oahensis Wilson & Evans 1899 (1890–99), p. xiii (Oahu) Myadestes lanaiensis woahensis Wilson & Evans 1890–1899 Specimens  The probable type is in Tring; another, discovered by Storrs Olson, is in Philadelphia (pers. comm.); subfossil remains are in Washington, D. C. Status  Extinct. Last seen in 1825. Known only from the type, possibly one other specimen, and subfossil remains. Range  Oahu, Hawaiian Islands. Description  Upperparts olive-brown, edges of feathers much lighter giving a slightly scalloped effect; tail and wings brown; undersides ­greyish-­white; bill blackish; iris dark brown; legs and feet ­horn-­brown.

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 Oahu Thrush of specimens of Hawaiian birds, and among Myadestes woahensis 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. Munro then quotes directly from Bloxam’s diary of May 4, 1825: We soon began to ascend the pass the sun rising at the time amid the chirping of small birds and the melodious notes of a brown thrush, the only songster on the islands. Munro (1944) inferred from this that the bird was common at that time, and had a beautiful song. Nothing more seems to be known about it. Munro believed that no specimens existed, but Olson (1994, 1996) discovered one in Tring, without data, which he believed to be Bloxam’s type. No one since Bloxam has seen the bird, and the date of its extinction is unknown.

Lanai Thrush (Lanai Oloma’o)  Myadestes lanaiensis lanaiensis

(Wilson)

Phaeornis lanaiensis Wilson 1891, p. 460 (Lanai) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Dresden; Edinburgh; Honolulu; Leiden; New York; Paris; Philadelphia; Tring; Dresden; and Washington, D. C.

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Extinct Birds Status  Extinct. Last seen in 1931. Range  Lanai, Hawaiian Islands. Description  18cm (7in). Dusky ­olive-­brown above, lighter on the head, and almost white below. It was smaller than the Omao Myadestes obscurus, the only surviving Hawaiian Thrush, and the outer tail feathers had slight white markings at the tip.

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.

Molokai Thrush (Molokai Oloma’o)  Myadestes lanaiensis rutha

(Bryan)

Phaeornis rutha Bryan 1908, p. 86 (Molokai) Specimens  Skins are in Cambridge, England; Cambridge, Massachusetts; Honolulu; New York; Ottawa; Tring; and Washington, D. C.; subfossil remains are in Washington, D. C. Status  Probably extinct. Not officially recorded since 1980. Range  Molokai, Hawaiian Islands. Description  18 cm (7in). Predominantly brown above, and pale grey below, darkest on the throat; ­undertail-­coverts pale buff.

The Molokai Thrush formerly occurred on the central Hawaiian Islands of Lanai, Molokai, Maui and probably 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 young. The montane ­mosquito-­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 1999). 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.

Kama’o (Large Kaua’i Thrush)  Myadestes myadestinus

(Stejneger)

Phaeornis myadestina Stejneger 1887b, p. 90 (Kaua’i, Hawaiian Islands) Specimens  Specimens are in Auckland, New Zealand; Cambridge, England; Cambridge, Massachusetts; Dresden; Leiden; Los Angeles; New York; Paris; Philadelphia; Seattle; Tring; and Washington, D. C. Status  Probably extinct. Not recorded since 1989. Range  Kaua’i, Hawaiian Islands.

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Turdidae Description  20cm (8in). Generally ­reddish-­brown above and pale grey below, with slightly darker mottling on breast and flanks; bill horn black; iris ­brownish-­black; legs and feet dark ­horn-­brown. Juvenile dark ­chocolate-­brown above heavily spotted with buff; and greyish below heavily scalloped with dark brown.

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 unconfirmed 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 ­epiphyte-­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 ­high-­pitched note. It fed primarily on fruits and berries, especially the fruit of the ’ie’ie Freycinetia arborea, and also on insects.

Doctorbird (Lord Howe Thrush)  Turdus poliocephalus vinitinctus

(Gould)

Merula vinitincta Gould 1855, p. 165 (Lord Howe Island) Specimens  Skins are in Berlin; Leiden; New York; Sydney; Tring; and Washington, D. C; eggs are in Tring. Status  Extinct soon after 1918. Range  Endemic to Lord Howe Island. 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 ­outer-­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 ­over-­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). Habits Hull (1909) was on Lord Howe Island in October 1907, and recorded what little is known about the bird’s habits: 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 derived 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 but 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. 283

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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 scattered over the whole surface.

Norfolk Island Thrush  Turdus poliocephalus poliocephalus

Latham

Turdus poliocephalus Latham 1801a (Norfolk Island) Specimens  Skins are in Liverpool; New York; and Tring; eggs are in Tring. Status  Extinct. Last confirmed sighting in 1975. Range  Norfolk Island. Description  21cm (8.25in). Predominantly dark brown, with pale ­grey-­brown head and upper breast; bill yellow; orbital ring yellow; legs and feet yellow.

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. philomelos, and interbreeding with the European Blackbird T. merula (which yielded sterile young), finally caused 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.

Lifou Thrush  Turdus poliocephalus pritzbueri

Layard

Turdus pritzbueri Layard 1878, p. 374 (Lifu) Specimens  Specimens are in Berkeley; Liverpool; New York; and Tring. Status  Probably extinct. Last seen prior to 1939. Range  Lifou, Loyalty Islands, western Pacific. Description  20cm (7.75in). Male head, neck and breast creamy grey, clearly defined from remainder of dark brown, almost black, plumage; bill pale yellow. Female differed in having head, neck and breast pale ­earth-­brown, shading into dark brown of the rest of body, less dark than male. Immature top of head brown, spotted with rufous; underparts deep warm rufous, spotted and irregularly marked with small broken bars of dark sepia.

There are three principal Loyalty Islands, Maré, Lifou and Ouvéa, with a number of smaller islands in between. Lifou is thickly forested coral island, flat on top with sheer cliffs. The first European visitor to the Loyalty Islands is thought to have been Captain Butler, either in HMS Walpole in 1800 or in HMS Britannia in 1803. From 1827, they were regularly frequented by trading vessels. The Lifou Thrush, a race of the widespread Island Thrush T. poliocephalus, was last collected in 1878, 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 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 Lifou Thrush. 284

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It was also said to occur on the island of Tanna, but this is most unlikely. This was based on a skin in Tring from the Eyton Collection, allegedly collected by John MacGillivray at Port Resolution on Tanna in 1860. Another skin from Layard (1878) is said to have been collected on Tanna. It is unrealistic to suppose that the same race could occur on the widely separated islands of Lifou and Tanna, especially as a different race, T. p. mareensis, formerly occurred on the geographically intermediate island of Maré (Layard & Tristram 1879; see below). The ‘Tanna’ specimens were probably traded skins. Habits Information on the bird is scanty. Layard (in Tristram 1879) reported that it was eaten frequently by local people: On taking it to a resident of the Loyalty Islands, who is staying with my ­next-­door neighbour (his ­brother-­in-­law), his wife exclaimed, ‘Oh! we eat lots of those, they are splendid gibier!’ and her servant, a Lifu man, standing by, added, ‘Him scrape on ground, all same fowl, we call him Wassasa’.

Maré Thrush  Turdus poliocephalus mareensis

Layard & Tristram

Turdus mareensis Layard & Tristram 1879, p. 472 (Maré) Specimens  Specimens are in Liverpool and Tring. Status  Probably extinct. Not recorded since 1939. Range  Maré, Loyalty Islands, western Pacific. Description  20cm (7.75in). Male almost uniform dark brown, nearly black above; blackish on back; indistinct pale brown eyestripe; underparts dark brown, shading into ­near-­white on centre of belly: undertail-coverts brown with pale chestnut-brown tips and shafts; bill yellow with orange tip; iris dark brown; orbital ring yellow; legs and feet bright yellow. Female browner, less blackish above; underparts paler.

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 (1900) noted that the alarm note was ‘like a Blackbird’s siss siss’.

Grand Cayman Thrush  Turdus ravidus

(Cory)

Mimocichla ravidus Cory 1886b, p. 499 (Grand Cayman, West Indies) Specimens  Specimens are in Berlin; Cambridge, Massachusetts; Chicago; Philadelphia; Pittsburgh; Tring; and Washington, D. C. Status  Probably extinct, last seen in 1938. Range  Grand Cayman Island, West Indies. Description  28cm (11in). Almost uniform ­ash-­grey plumage, except for white lower abdomen and ­undertail-­coverts ; variable, but three to four outer ­tail-­feathers tipped with white on internal side; eye-ring, bill and feet ­coral-­red. Immature uniform buff upperparts, and underparts devoid of spots; abdomen and ­undertail-­coverts white; legs and feet ­buff-­brown.

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 (Patricia Bradley pers. comm.). It was the only full species endemic to the Cayman Islands, and had been declining for some time before the final sighting. Bangs (1916), in discussing a collection of birds made in the Cayman Islands by W. W. Brown (who shot fifteen Grand Cayman Thrushes) between April and July, said: The Thrush is now extremely rare and local in Grand Cayman. Brown covered the whole island and found it only in two remote patches of woodland. Each of these tracts of rather heavier forest than is usual in the island now-a-days was inhabited by a few pairs of thrushes, which Brown believes to be the entire population of the island. In each of these woods Brown was careful to leave birds enough to perpetuate the species, if it is not gradually becoming extinct from some natural cause, as seems to be the case. Habits Johnston (1969) interviewed the oldest resident on Grand Cayman, who provided information on the bird from his youth. Johnston wrote: 285

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It was my good fortune to interview on 28 December 1965, Bunyon Whittaker, probably the oldest (72) living resident on Grand Cayman. Still living at Old Man Bay, Mr. Whittaker clearly recalled his experiences in 1916 when W. W. Brown sailed around to Northside from Georgetown and remained in the home of Willie Tatum for several weeks, during which time Brown paid the small boys one dollar apiece for bird specimens. Mr. Whittaker helped his brother collect the thrushes and other species with slingshots. 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.

Cauca ­Black-­hooded Thrush  Turdus olivater caucae

(Chapman)

Planesticus caucae Chapman 1914, p. 182 (La Sierra, Central Andes, Cauca, Colombia) Specimens  Specimens are in New York and Washington D. C. Status  Possibly extinct. Not recorded since the late 1980s. Range  Cauca, central Andes, Colombia. Description  23–24cm (9–9.5in). Similar to nominate T. o. olivater except greyer, less suffused with brown, male with black of nape more sharply defined from grey of back; breast greyish, upper part lightly streaked black; sides of throat black; centre of throat and chin streaked black; female crown distinctly darker than back; underparts less brown; throat black, shaft-streaks weakly defined; bill bright yellow, maxilla tipped with blackish; legs and feet brownish. Juvenile male has black of nape less pronounced; throat and breast streaks duller.

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 subspecies – which has sometimes been given specific status, but there are too few morphlogical differences to support this – has suffered from severe deforestation for agriculture and the narcotics industry, and has not been seen for some decades. It may be extinct. Habits Little was recorded about its habits. It preferred subtropical moist montane forest at an elevation of between 800–1,600m, where it foraged on the forest floor or in the low understorey. The type was taken in deep forest (Chapman 1914).

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 madagascariensis, 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 286

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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.

Réunion Fody  Foudia delloni

Cheke & Hume

“Sparrow”. Du Bois 1897, p. 70–72. Foudia delloni Cheke & Hume 2008, p. 42–3, pl. 9 Status  Extinct. Last recorded in 1672. Known only from travellers’ accounts. Range  Réunion Island, Mascarenes. Description  13cm (5in). Differed from Mauritius Fody F. rubra in having ­wing-­coverts red, not ­grey-­brown; and from Madagascar Fody F. madagascariensis in restricted areas of red in male. Breeding male head, neck, throat, and upperparts of wings bright red; back brown, paler on belly; tail brown. Female and eclipse male head, neck and wings brown instead of red; paler on throat, otherwise similar.

The Réunion Fody was first mentioned by Gabriel Dellon (1668), 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 them fall into pots and platters, and burning their wings on fires, which are lit out of doors, the sun providing enough heat for [even] the coolest houses. 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 afterwards 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.

Estrildidae (Waxbills, munias and allies) The estrildid finches are generally small passerines with short, stout bills and short legs; these birds are very diverse in coloration. The family has been divided into 28 genera with about 140 species. They occur in the tropical regions of Africa and Australasia. The estrildid finches have been widely collected for the bird trade and have suffered from habitat destruction, which has resulted in some species becoming endangered. Two subspecies and a one species may be extinct. 287

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Eastern Star Finch  Neochmia ruficauda ruficauda

(Gould)

Amadina ruficauda Gould 1837a, pl. 10, fig. 2 (‘New Holland’ = interior of New South Wales, Australia) Specimens  Specimens are in Tring and Victoria, Australia. Status  Probably extinct; not positively seen since the 1990s. Range  Central Queensland to New South Wales, Australia. Description  11–12cm (4.25–4.75in). Adult facial mask red; upperparts ­olive-­green; tail dark reddish black with ­pinkish-­white spots or bands; breast olive-grey; belly and vent pale cream; white spots on face, throat, breast, and flanks; bill ­bright red; legs and feet buff, ­yellow-­brown or ­pinkish-­brown; ­eye-­ring dull red; iris bright orange. Sexual dimorphism is apparent in other subspecies, but no details are available for the eastern birds.

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 (fringilline 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.

Makawehi Finch  Telespiza persecutrix

James & Olson

Telespiza persecutrix James & Olson 1991, p. 30 (Barber’s Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Oahu and Kaua’i, Hawaiian Islands.

The Makawehi Finch is ­so-­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, 288

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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 catastrophic. It is unlikely that the main island populations survived to the historic period. 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.

Maui Nui Finch  Telespiza ypsilon

James & Olson

Telespiza ypsilon James & Olson 1991, p. 35 (Ilio Point, Molokai, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Molokai and Maui, Hawaiian Islands.

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 ­smaller-­billed bird was found at a much higher altitude of 808m; this may represent another distinct Telespiza species (James & Olson 1991).

Lanai ’Alauahio  Paroreomyza montana montana

(Wilson)

Himatione montana Wilson 1889 (1890), p. 446 (Lanai) Specimens  Specimens are in Berlin; Cambridge, England; Cambridge, Massachusetts; Dresden; Honolulu; Liverpool; New York; Ottawa; Paris; Pittsburgh; Tring; and Washington, D. C. Status  Extinct. Range  Mountain forests of Lanai, where none has been seen since 1937. Description  11cm (4.25in). A small bird, ­yellowish-­green above and lemon yellow below. Male forehead, sides of face and throat deep ­lemon-­yellow shading to lighter yellow on breast and abdomen, lower part white; ­undertail-­coverts deep ­lemon-­yellow; upperparts dull ­greenish-­yellow; rump yellow; primaries and tail ­ash-­brown, edged dull yellow; bill, legs and feet light pink. Female similar to male but duller yellow below; immature duller still.

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. 289

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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.

Kakawahie (Molokai ’Alauahio)  Paroreomyza flammea

(Wilson)

Loxops flammea Wilson 1889 (1890), p. 445 (Kalae, Molokai) Specimens  Specimens are in Berlin; Cambridge, England; Cambridge, Massachusetts; Dresden; Frankfurt; Honolulu; Ithaca; Leiden; Liverpool; New York; Ottawa; Paris; Tring; and Washington. Status  Last seen in 1963, and probably now extinct. Range  Molokai, Hawaiian Islands. Description  14cm (5.5in). Male scarlet, in several shades; upper bill ­bluish-­grey, lower bill dark; legs and feet ­brownish-­black. Female brown with scarlet markings. Immature males intermediate and variable between male and female plumages.

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 chipping wood, hence the bird’s name Kakawahie, which means ‘to break up firewood’. 290

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Oahu ’Alauahio  Paroreomyza maculata

(Cabanis)

Himatione maculata Cabanis 1850, p. 100 (footnote) (Oahu) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Dresden; Leiden; Los Angeles; New York; Philadelphia; Tring; and Yale. Status  Probably extinct. Last recorded December 12, 1985. Range  Oahu, Hawaiian Islands. Description  11cm (4.25in). Male generally ­olive-­green above, yellow below, with dark lores merging into olive ­eye-­stripe; forehead and supercilium yellow. Female ­greenish-­grey above, pale ­yellowish-­white below; forehead and lores pale yellow; eye-stripe dark; and with two prominent, pale ­wing-­bars.

The Oahu ’Alauahio was considered common in the last decade of the 19th century, despite having disappeared 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. 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.

Lanai Hookbill  Dysmorodrepanis munroi

Perkins

Dysmorodrepanis munroi Perkins 1919, p. 251 (Kaiholena Valley, Lanai) Specimens  The type specimen is in Honolulu. Status  Extinct. Known only from the unique type, collected on Lanai in February 1913. Range  Lanai, Hawaiian Islands. Description  15cm (6in). The type specimen of the Lanai Hookbill was thought partially albinistic (Greenway 1939), but this does not appear to be the case, though some specimens of ’O’u from Lanai show this tendency. None, however, has the yellow tinge on the lores and supraocular region. Upperparts light greenish olive; underside whitish yellow, merging to white on ­undertail-­coverts; supercilium pale yellow; white ­wing-­patches formed by the broad tips and inner vanes of secondaries. Upper and lower bill curve towards each other, so that tip of lower is the only part that touches the upper, leaving an opening between them like a pair of pincers.

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, 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 291

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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 ­south-­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 ­thick-­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 countless wild goats, and it was also full of wild pigs and cats that had run wild. The latter, as I have elsewhere recorded, were destroying native birds wholesale. Only on the sheer sides of the mountain and on a very small part of the narrow backbone was the forest in a natural condition. Neither Lord Rothschild’s collectors nor myself ever found a specimen of this bird. Hybrid or mutant? Greenway (1939) considered D. munroi to be an aberrant specimen of the ’O’u Psittirostra psittacea (see p. 307) and probably a young male, or possibly a hybrid. The most distinguishing character of D. munroi is the bill, in which the lower mandible is curved and touches the upper mandible only at the tip. Greenway thought this to be a malformation, due either to an accident or a mutation, and as the population of ’O’u on Lanai was small (and probably by the time he wrote extinct) deformed specimens might well occur. Greenway also laid great stress on the fact that Munro was unable to find another specimen, despite diligent searches. The type specimen was ­re-­examined, and the skull removed from the skin to study the cranial osteology by James et al. (1989). No evidence was found of any deformity, and moreover the cranium and mandible showed modifications that were concordant with the unusual bill form. The Lanai Hookbill was, as Munro had stated, a valid species after all. 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.

Greater ’Amakihi (Green Solitaire)  Hemignathus sagittirostris

(Rothschild)

Viridonia sagittirostris Rothschild 1892, p. 112 (lower slopes of Mauna Kea, Hawaii) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Dresden; Honolulu; Los Angeles; New York; Tring; and Washington, D. C. Status  Extinct. Last recorded in 1901. Range  Hawaii, Hawaiian Islands. Description  15cm (6in). Similar to Common ’Amakihi H. virens but larger and more olive-green. Upperparts bright ­olive-­green, slightly yellower on forehead, sides of head, ­uppertail-­coverts, throat, breast and abdomen; wings ­blackish-­brown, primaries edged with olive; bill black; iris reddish hazel; legs and feet black. Sexes alike.

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 292

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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: 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.

Kaua’i ’Akialoa (Greater ’Akialoa)  Hemignathus (ellisianus) stejnegeri(Wilson) Hemignathus procerus Cabanis 1890, p. 331 Hemignathus stejnegeri Wilson 1889, p. 400 (this name has precedence over Cabanis’s procerus) Akialoa stejnegeri Olson & James 1995, p. 385 Specimens  Specimens are in Auckland; Berlin; Cambridge, England; Dresden; Frankfurt; Honolulu; Leiden; Liverpool; New York; Ottawa; Paris; Philadelphia; Pittsburgh; Seattle; Tring; and Washington, D. C. Status  Probably extinct. Believed to have survived in very small numbers until 1965. Range  Mountain forests of Kaua’i. Description  20cm (8in). Male front and top of head dark ashy olive, shading into olive on back, brighter on rump; face olive with an indistinct superciliary stripe; light yellow below, washed with olive on flanks; ­undertail-­coverts ­olive-­yellow; wings and tail brown edged on outer webs with olive; secondaries and ­wing-­coverts brown broadly edged with olive; iris dark hazel; bill black; legs and feet bluish black. Female yellowish buff above, feathers of head with dark centres; superciliary stripe more distinct; and underside ­olive-­buff.

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 historically 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 (see p. 303). 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) rediscovered 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 ­woodpecker-­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 ­brush-­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 293

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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.

Maui Nui ’Akialoa  Hemignathus (ellisianus) lanaiensis

Rothschild

Hemignathus lanaiensis Rothschild 1893a, p. 24 (Lanai) Specimens  Specimens are in New York and Tring. Status  Extinct. The last definite sighting was in 1894. Range  Mountain forests of Lanai, Hawaiian Islands. Description  16.5cm (6.5in). Adult male olivaceous green on upperside; underside dirty yellow; undertail-coverts creamy. Female greyish olive, more yellowish on abdomen. Bill black. Breeding plumage of the male was apparently yellow.

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.

Lesser ’Akialoa (Hawaii ’Akialoa)  Hemignathus (ellisianus) obscura

(Gmelin)

Hook-billed Green Creeper. Latham 1782, p. 703, pl. 33, fig. 1 ? Akaiearooa. King 1784, p. 119. Certhia obscura Gmelin 1788, p. 470 (Sandwich Islands = Hawaiian Islands) Specimens  Specimens are in Berkeley; Cambridge, England; Cambridge, Massachusetts; Dresden; Honolulu; Leiden; Liverpool; Los Angeles; New York; Ottawa; Paris; Philadelphia; Pittsburgh; San Francisco; Sydney; Tring; Vienna; Washington, D. C.; and Yale. Status  Extinct. Last recorded September 1903. Range  Mountain forests of Hawaii, Hawaiian Islands.

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Fringillidae Description  15cm (6in). Male dull greenish olive on head and body, brighter on rump, paler on abdomen and ­undertail-­coverts; lores dusky with yellow mark above eye; wings and tail ash-brown edged with dull olive; iris dark hazel; bill black; legs and feet brown. Female similar but duller.

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 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 unconfirmed 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 ­mosquito-­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 (1892) 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 material 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.

Oahu ’Akialoa  Hemignathus (ellisianus) ellisianus

(Gray)

Drepanis (Hemignathus) ellisianus Gray 1859a, p. 9 (Oahu) Hemignathus lichtensteinii Wilson 1889, p. 401 (Oahu) Specimens  Specimens are in Berlin and Philadelphia. Status  Extinct. Last reported in 1894. Range  Mountain forests of Oahu. Description  16.5cm (6.5in). Dull ­olive-­green, paler on throat; wings and tail ­brownish-­olive, edged with light olive; bill, legs and feet brown.

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 295

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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 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.

Hoopoe-­billed ‘Akialoa  Hemignathus upupirostris

James & Olson

Hemignathus upupirostris James & Olson 1991, p. 60 (Makawehi dunes, Kaua’i, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  Kaua’i and Oahu, Hawaiian Islands.

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.

Kaua’i Nukupu’u  Hemignathus lucidus hanapepe

Wilson

Hemignathus hanapepe Wilson 1889, p. 401 (Kaua’i) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Honolulu; Liverpool; New York; Philadelphia; Ottawa; and Tring. Status  Probably extinct. Possibly seen in 1995. Range  Mountain forests of Kaua’i, Hawaiian Islands. Description  14cm (5.5in). The male was bright yellow on head and breast, shading to white on abdomen; upperparts yellowish green; lores, ­eye-­ring and bill black; legs and feet ­blackish-­brown. Female paler and duller, ­greenish-­grey above; primrose on breast and dull white on abdomen; chin, upper throat and supraloral patch yellow.

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). Its extinction can be attributed to forest clearance, opening of the understorey by cattle and pigs, and ­mosquito-­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 caterpillars, 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’. 296

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Oahu Nukupu’u  Hemignathus lucidus lucidus

Lichtenstein

Hemignathus lucidus Lichtenstein 1839, p. 451, pl. 5, figs 2–3 (Oahu) Heterorhynchus olivaceus Lafresnaye 1839, pl. 10, 2 unpaginated pp. text Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Frankfurt; Leiden; New York; Tring; and Vienna. Status  Extinct. Last recorded around 1893. Range  Mountain forests of Oahu, Hawaiian Islands. Description  14cm (5.5in). It was generally ­olive-­green above and yellow below; head yellowish; lores blackish; wings and tail ­olive-­brown, edges of feathers light olive; bill brown; legs and feet black. Female and immature similar except less yellowish and duller.

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 (1890–99; 1894) provides an interesting summary: While Deppe was in the Sandwich [Hawaiian] Islands he was joined by the still better known American naturalist 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 specifically 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 undoubtedly peculiar to the island of Oahu, where Deppe informed Lichtenstein that it frequented the ­plantain-­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.

Maui Nukupu’u  Hemignathus lucidus affinis

Rothschild

Hemignathus affinis Rothschild 1893d, p. 112 (Maui) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Dresden; Liverpool; New York; Ottawa; Philadelphia; and Tring. Status  Probably extinct. Last collected in 1896; last observed in 1995. Range  Forested slopes of Haleakala, Maui, Hawaiian Islands. Description  14cm (5.5in). Head, throat and upper breast intense golden yellow, fading into yellow on lower breast; vent and ­undertail-­coverts yellowish-green; upperparts ­yellowish-­green; lores, ­eye-­ring and bill black; legs and feet ­blackish-­brown.

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 subsequent 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.

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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.

Giant ­Scimitar-­billed Nukupu’u  Hemignathus vorpalis

James & Olson

Hemignathus vorpalis James & Olson 2003, p. 972 (Petrel Cave, Hualalai Quadrangle, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Hawaii, Hawaiian Islands.

The ­dual-­purpose bills of the Nukupu’u H. lucidus and Akiapola’au H. munroi are truly amazing adaptations for foraging, but the discovery of the Giant ­Scimitar-­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 ­flat-­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 ­ground-­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.

Giant Scimitar-billed Nukupu’u Hemignathus vorpalis

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Hawaii Nukupu’u  Hemignathus lucidus ssp. Specimens  The type specimen is in Washington D. C. Status  Extinct. Known from a unique type taken in 1840 or 1841. Range  Hawaii, Hawaiian Islands. Description  14cm (5.5in). The plumage appears to be that of an immature moulting into yellower adult plumage. Dorsum, crown and wings dull olive with greyish cast; superciliary stripe faint yellow; cheeks, midline of throat and sides of upper breast yellow; rest of underparts creamy whitish, more olivaceous on flanks; bill and feet probably blackish.

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 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 1992).

Kiwi ­Shovel-­billed Finch Vangulifer mirandus

James & Olson

Vangulifer mirandus James & Olson 1991, p. 63 (Puu Naio Cave, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  Maui, Hawaiian Islands.

The discovery of a large number of subfossil species of Hawaiian finches has shown that the true extent of radiation within the group is far greater than anyone could ever have expected. The Kiwi ­Shovel-­billed Finch was characterised by its long, broad and rather deep bill, culminating in a very blunt rounded tip, the function of which is unknown. James & Olson (1991) comment that the bill seems too long and weak to crack seeds, too deep and broad to be used for probing in bark, and too blunt to be ­nectar-­feeding apparatus. The ventral surface and lateral edges of the upper bill were richly supplied with blood vessels, which is a very unusual characteristic otherwise limited to kiwis.

Kiwi Shovel-billed Finch Vangulifer mirandus

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These presumably supplied sensory neurons, indicating that the bird had a sensitive bill that it used to detect food items.

Pololei ­Shovel-­billed Finch  Vangulifer neophasis

James & Olson

Vangulifer neophasis James & Olson 1991, p. 65 (Puu Naio Cave, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  Maui, Hawaiian Islands.

The differences between this species and the Kiwi ­Shovel-­billed Finch (see above) are substantial, and the birds could be construed as belonging to separate genera (James & Olson 1991). The Pololei ­Shovel-­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.

Straight-­billed Gaper  Aidemedia chascax

James & Olson

Aidemedia chascax James & Olson 1991, p. 67 (Barber’s Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  Oahu, Hawaiian Islands.

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 ­Straight-­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.

Sickle-­billed Gaper  Aidemedia zanclops

James & Olson

Aidemedia zanclops James & Olson 1991, p. 69 (Barber’s Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  Oahu, Hawaiian Islands.

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. 295). James & Olson (1991), with some degree of caution, suggest that the bill differences seen between the ­Sickle-­billed Gaper A. zanclops and ­Straight-­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. 233), which exhibited extreme sexual dimorphism, 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. 300

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Maui Nui Gaper  Aidemedia lutetiae

James & Olson

Aidemedia lutetiae James & Olson 1991, p. 71 (Moomomi dunes, Molokai, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction not known. Range  Maui and Molokai, Hawaiian Islands.

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). 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 population of one island colonising the other by crossing a channel. This was also the case with the Maui Nui Finch Telespiza ypsilon (see p. 289) and Maui Nui ’Akialoa Hemignathus (ellisianus) lanaiensis (see p. 294). 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.

Oahu ’Akepa  Loxops wolstenholmei

Rothschild

Fringilla rufa Bloxam 1826, p. 250 (Oahu) Loxops wolstenholmei Rothschild 1893c, p. 56 (Oahu) Specimens  Specimens are in Berlin; Bremen; Cambridge, England; Cambridge, Massachusetts; Kassel, Germany; Liverpool; New York; Philadelphia; St. Petersburg; Tring; and Vienna. Status  Extinct. The last specimen was taken in May 1893. Range  Mountain forests of Oahu, Hawaiian Islands. Description  10cm (4in). Adult male rufous orange above with a brownish wash; wings ­blackish-­brown; greenish orange on outer webs; reddish orange below, paler on the abdomen and ­undertail-­coverts. Female and immature ­greenish-­grey.

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.

Maui ’Akepa  Loxops ochraceus

Rothschild

Loxops ochracea Rothschild 1892, p. 16 (no description or type locality) Loxops ochracea Rothschild 1893d, p. 112 Specimens  Specimens are in Cambridge, Massachusetts; Dresden; Ithaca; Leiden; Liverpool; New York; Philadelphia; San Francisco; and Tring.

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Extinct Birds Status  Probably extinct. Last seen in 1988. Range  Maui, Hawaiian Islands. Description  10cm (4in). Similar to Oahu ’Akepa L. wolstonholmei except two distinct colour forms of adult male; first was ochraceous or ­brownish-­orange, fading to yellowish chestnut on lower breast and vent; ­undertail-­coverts dull ­orange-­yellow, tail plumbeous, edged with ochraceous yellow on outer edges; second male predominantly ­mustard-­yellow (see Pratt 2005). Female similar to L. coccinea but upperside more ­blackish-­green; underparts more olivaceous and duller.

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 Hawaii ’Akepa today (Camp et al. 2009). The last record of the Maui bird was in 1988 (Engilis 1990). 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.

Greater Koa Finch  Rhodacanthis palmeri

Rothschild

Rhodacanthis palmeri Rothschild 1892, p. 111 (Kona, Hawaii) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Honolulu; Liverpool; New York; Ottawa; Philadelphia; Tring; and Washington, D. C. Status  Extinct. Last seen in March 1896. Range  Mauna Loa, Hawaii, Hawaiian Islands. Description  22–23cm (8.5–9in). The largest passerine in the Hawaiian Islands; male forehead bright orange-red; throat washed with orange; back dark olive tinged orange on rump; breast and belly pale yellowish; wings and tail ­blackish-­brown, feathers margined yellow on outer edges; bill ­bluish-­grey; iris dark brown; legs and feet blackish. Female forehead yellow; upperparts ­olive-­brown; ­yellow-­green on rump and ­uppertail-­coverts; throat, breast light ­olive-­green merging to whitish green on belly. Immature like female except dark spotting on undersides.

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 (1999) suggests that the persistence of Rhodacanthis and Kona Grosbeak Chloridops kona (see p. 304) 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. 302

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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: 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 ­lava-­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 ­full-­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.

Lesser Koa Finch  Rhodacanthis flaviceps

Rothschild

Rhodacanthis flaviceps Rothschild 1892, p. 111 (Kona, Hawaii) Specimens  Specimens are in New York and Tring. Status  Extinct. Last seen on October 16, 1891. Range  Mauna Loa, Hawaii, Hawaiian Islands. Description  15cm (6in). Differed from the Greater Koa Finch R. palmeri in having yellow (not orange) on the head and neck; also much smaller. Female similar but smaller.

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 carefully 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 (see p. 304), 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 ­yellow-­headed bird, nor did I hear any song or call note that could be referred to a species unknown to me. 303

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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.

Scissor-­billed Koa Finch  Rhodacanthis forfex

James & Olson

Rhodacanthis forfex James & Olson 2005, p. 533 (Koloa Quadrangle: Makauwahi Cave, Kaua’i) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Kaua’i and Maui, Hawaiian Islands.

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 probably disappeared as a result of deforestation and ­over-­hunting following the arrival of the Polynesians, but prior to the arrival of Europeans in the 1770s.

Oahu Koa Finch  Rhodacanthis litotes

James & Olson

Rhodacanthis litotes James & Olson 2005, p. 536 (Ewa Quadrangle, Barber’s Point, Oahu) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Oahu and possibly Maui, Hawaiian Islands.

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.

Kona Grosbeak  Chloridops kona

Wilson

Chloridops kona Wilson 1888, p. 218 (Kona, Hawaii) Specimens  Specimens are in Cambridge, England; Cambridge, Massachusetts; Dresden; Honolulu; Leiden; Liverpool; New York; Ottawa; Paris; Tring; and Washington, D. C. Status  Extinct. Last seen in September 1892. Range  Mountain forests of Hawaii, Hawaiian Islands. Description  18cm (7in). Large with a huge hawfinch-like bill. Uniform bright olive-green; throat more golden green; lores blackish; abdomen whitish; bill dull flesh; iris dark hazel; legs and feet dark ­blackish-­brown. Sexes alike.

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 304

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below 1,400m (Olson 1999a). It was discovered by Wilson (1888) on June 21, 1887, when he obtained two specimens. Palmer (in Munro 1944) collected it again in September 1891. A specimen taken in September 1892 by Perkins (not in 1894 as often cited, e.g. Munro 1944) was the last known (Banko 1986; Olson 1999a). Even in 1892, Perkins considered it rare and highly localised; its historical existence lasted for only five years and six months (Olson 1999). 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 anthropogenic changes. Habits Those ornithologists who saw the Kona Grosbeak found it in Naio or Bastard Sandalwood trees Myoporum sandwicense, which grow on old lava flows, and occasionally in Sandalwood trees Santalum sp., where the birds fed on the dry seeds and occasionally took caterpillars. Munro (1944) noted that the birds seldom strayed from the more recent lava flows, where there was no undergrowth and the trees were of ­medium-­size. 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.

Wahi Grosbeak  Chloridops wahi

James & Olson

Chloridops wahi James & Olson 1991, p. 36 (Barber’s Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Oahu and Maui, Hawaiian Islands.

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 genera Chloridops once occurred throughout the Hawaiian Islands. The Wahi Grosbeak occurred on Oahu and Maui. 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 smaller and shorter (James & Olson 1991). 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. Two more probable grosbeak species have been recovered from fossil localities on Kaua’i and Maui respectively (Olson & James 1982). The Maui species, 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. A series of jaw remains found on Kaua’i are referable to Chloridops, but differ in certain characteristics from C. wahi, and may also represent a distinct taxon.

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King Kong Grosbeak  Chloridops regiskongi

James & Olson

Chloridops regiskongi James & Olson 1991, p. 40 (Barber’s Point, Oahu, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Oahu, Hawaiian Islands.

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.

Mauka Grosbeak  Orthiospiza howarthi

James & Olson

Orthiospiza howarthi James & Olson 1991, p. 47 (Lua Manu, Kipahulu Valley, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Maui, Hawaiian Islands.

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.

Pila’s Palila  Loxioides kikuchi

James & Olson

Loxioides kikuchi James & Olson 2006, p. 338 (Koloa Quadrangle, Makauwahi Cave, Kaua’i) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. May have survived until the late 18th or early 19th centuries. Range  Kaua’i, Hawaiian Islands.

The Palila Loxioides bailleui is confined to ­high-­elevation open Mamane Sophora chrysophylla forest on the slopes of Mauna Kea on Hawaii (Banko et al. 2002b), but subfossil remains have been found at ­sea-­level on Oahu (Burney et al. 2001; James & Olson 2006). The discovery of a lowland population of L. bailleui, or something very similar, on both Kaua’i and Oahu is extremely interesting, as it shows that the ­present-­day montane occurrence of the Palila on Hawaii is an artefact of human habitat alteration (Olson & James 1982). Furthermore, the species was sympatric on Kaua’i with a second species, Pila’s Palila L. kikuchi. The larger size of Pila’s Palila (which was named after the Kikuchi family of Kaua’i, in particular William, or Pila, who assisted with excavations on Kaua’i), which also differed from L. bailleui in lacking a hooked bill, meant the birds fed on different food items and thus could coexist in similar habitats. The drastic changes caused by farming in the dry, lowland forests on Kaua’i, which intensified after settlement by Europeans, was probably the reason for this bird’s extinction. James & Olson (2006) suggest that Pila’s Palila may have survived until comparatively recently, but that it was overlooked during the early ornithological surveys of the island.

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’O’u  Psittirostra psittacea

(Gmelin)

Parrot–billed Grosbeak. Latham 1783 (‘in insula Sandwich’ = Hawaii) Loxia psittacea Gmelin 1789, p. 844 based on Latham. Psittirostra psittacea deppei Rothschild 1905a, p. 45 (O’ahu) Psittirostra psittacea oppidiana Bangs 1911, p. 30 (Molokai) Specimens  Specimens are in Ann Arbor, Michigan; Auckland; Berkeley; Berlin; Cambridge, England; Cambridge, Massachusetts; Dresden; Frankfurt; Honolulu; Leiden; Liverpool; Los Angeles; New York; Ottawa; Paris; Philadelphia; Pittsburgh; San Francisco; Tring; Vienna; and Washington, D. C.; and Yale. Status  Probably extinct. Last reliably recorded in February 1989. Range  Kaua’i, Oahu, Lanai, Molokai, Maui and Hawaii, Hawaiian Islands. Description  17cm (6.75in). This was a robust, ­paunch-­bellied bird with a slightly hooked bill. Male uniform ­olive-­green, darker on the back; head sharply defined deep yellow; ­undertail-­coverts white; bill, legs and feet pink; iris hazel. Female similar except lacked yellow head; and greyer on throat and upper breast.

The ’O’u formerly occurred throughout the Hawaiian Islands, and it was considered one of the commonest of the Hawaiian finches (Perkins 1903). However, Perkins only saw a single pair in March 1893 on Oahu, shortly after which it disappeared. Rothschild (1905a) considered the Oahu population 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. 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 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). 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 ­mass-­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, beginning with three or four distinct and penetrating whistled notes. Perkins noted that they would sometimes sing in concert on the wing.

Cone-­billed Finch  Xestospiza conica

James & Olson

Xestospiza conica James & Olson 1991, p. 53 (Makewehi dunes, Kaua’i, Hawaiian Islands) Specimens  Subfossil remains are in Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Kaua’i, Hawaiian Islands.

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). 307

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The bill of the ­Cone-­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. 312), 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. 195), which was probably the finch’s main predator.

Ridge-­billed Finch  Xestospiza fastigialis

James & Olson

Xestospiza fastigialis James & Olson 1991, p. 55 (Crystal Cave, Manawainui Gulch, Maui, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Maui, Molokai and Oahu, Hawaiian Islands.

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 reminiscent 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. 307). Despite the wide distribution of the ­Ridge-­billed Finch, it disappeared before European settlers arrived, probably as a result of habitat destruction and predation by introduced mammals.

Mamo  Drepanis pacifica

(Gmelin)

Great Hook-billed Creeper. Latham 1782, p. 703 (Friendly Islands = error for Hawaii) ? Hoohoo (partim). King 1784, p. 119 Certhia pacifica Gmelin 1788, p. 470. (‘in insulis amicis’ = error for Hawaii) 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 ­undertail-­coverts; primary and ­underwing-­coverts white; four central ­tail-­feathers black, rest brown with patch of dull white near tip; bill black; legs and feet ­blackish-­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. 249). Perkins considered it extremely rare in the 1890s, and Wilson (1890–99; 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 information 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 308

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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. The tale of the captive Mamo In 1892, Rothschild’s collector, Henry Palmer, who was laid up after being kicked by a horse, sent his assistant Wolstenholme and Ahulu, a native ­bird-­catcher, to the forested slopes of Mauna Loa, above Hilo (Rothschild 1893–1900). There, in the Olaa forest, they managed to catch a live specimen on April 16, the first seen since Théodore Ballieu obtained the two unique juvenile specimens at Pulehua on the leeward slopes of Mauna Loa around 1876 (Olson & Hume 2009). The Mamo was fed sugar and water and happily roosted on a stick in its captor’s tent. The party returned triumphantly to Palmer with the Mamo still alive, as Wolstenholme was unable to bring himself to kill it. Palmer, who was less sentimental, killed and skinned it immediately. Wolstenholme gave Palmer a photograph (see Pratt 2002), the only one known, of the live Mamo perched on his finger. Knowing its value and just to make sure that the bird did not fly off, a noose was tied to its leg. Henshaw (1902) saw a family group in July 1898 near Kaümana above Hilo, where he shot one but failed to retrieve the specimen. He described the flight as ‘not rapid, but smooth and well sustained’, reminding him of the flight of a Common Cuckoo Cuculus canorus. None has been recorded since. 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 ­black-­and-­yellow Mamo was said to forage with the similarly striking black, red and grey ’Ula-’ai-Hawane Ciridops anna (Pratt 2002; see p. 313), 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.

Black Mamo (Hoa)  Drepanis funerea

Newton

Drepanis funerea A. Newton 1893, p. 690 (Molokai) Specimens  Specimens are in Bremen; Cambridge, England; Cambridge, Massachusetts; Honolulu; Molokai, Hawaiian Islands; New York; and Tring. Status  Extinct. The last recorded specimens were taken in 1907. Range  Mountain forests of Molokai, and possibly Maui, Hawaiian Islands.

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 devastated 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 distinguish each separate movement. It had not so silvery an appearance 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 309

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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 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 Black Mamo Drepanis funerea 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. The last specimen The Black Mamo is now extinct, as ornithologists have searched the remaining forest up to the top of Mount Olokui, without success. No doubt habitat destruction, introduced rats and ­mosquito-­borne diseases were responsible for its extinction. Perkins (1895, 1903) found it to be a bird of forest undergrowth, and noted that imported deer and cattle had destroyed most of this habitat. Bryan (1908) records the fate that awaited the last specimens taken in May, 1907: Actively hopping and flitting from limb to limb, scarcely stopping a second, eyeing me sharply all the while, it persisted in keeping so close to me that I could not with safety fire even my auxiliary barrel. Yet the restlessness of the bird made it apparent that I must fire soon or lose my chance, for if the bird got fifty feet from me in the tangle of trees and vines on the side of the ridge, it would, in all probability be gone from sight and forever. I pulled the trigger and, to my consternation my gun missed fire. It was after all little wonder that gun and ammunition failed me, for both had been soaking wet for three days. The next few seconds were moments of painful indecision. To break and reload my gun would surely frighten the bird; to use the heavy charge in the left barrel was almost criminal, but my only recourse. Without further delay I availed myself of the first opportunity when the bird was screened by limbs and fired. The feathers flew: my first Black Mamo had been shot. Although historically known only from Molokai, subfossil remains collected on Maui indicate that the Black Mamo, or a bird osteologically indistinguishable from it, also once occurred there (James & Olson 1991). 310

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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.

Laysan ’Apapane (Laysan Honeycreeper) Himatione sanguinea freethii

Rothschild

Himatione fraithii Rothschild 1892, p. 109 (Laysan Island) Specimens  Specimens are in Ann Arbor, Michigan; Bonn; Cambridge, England; Cambridge, Massachusetts; Chicago; Denver; Honolulu; Los Angeles; New York; Ottawa; Paris; Philadelphia; Pittsburgh; San Francisco; Tring; Washington, D. C.; and Yale. Status  Extinct. Not seen since 1923. Range  Laysan, Hawaiian Islands. Description  13cm (5.1in). Head, throat, breast and upper abdomen bright scarlet vermillion, with faint tinge of golden orange; rest of upperparts orange scarlet; lower abdomen ­dusky-­grey fading to ­brownish-­white; ­undertail-­coverts greyish; wings, tail, bill, legs and feet dark brown; iris black with brown outline. Immature brown, paler below, with green edges to ­wing-­coverts. Nominate H. s. sanguinea of the main Hawaiian Islands differed in being generally ­blood-­red, wing and tail black, ­undertail-­coverts whiter and bill longer.

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 decimated so many of its congeners. A race, H. s. freethii, once occurred on the remote island of Laysan, which differed only slightly from the nominate form. 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 Captain 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. 311

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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.

Po’ouli  Melamprosops phaeosoma

Casey & Jacobi

Melamprosops phaeosoma Casey & Jacobi 1974, p. 219 (Maui, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and New York. Status  Probably extinct. Last recorded in November 2004. Range  Maui, Hawaiian Islands. Description  14cm (5.5in). Adult male has broad black mask extending behind eye; crown grey merging to dark brown on back; scapulars and ­wing-­coverts dark ­rufous-­brown; primaries and secondaries dark brown; rump and ­uppertail-­coverts ­rufous-­brown; chin black; throat and breast white shading to ­greyish-­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 disappeared in 2003 and 2004, 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). 312

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Reasons for extinction The rapid disappearance of the Po’ouli was due to a number of factors, including ­mosquito-­borne avian diseases, habitat loss, and predation by and competition with introduced exotic animals. The decline in native tree snails due to predation by rats may have also impacted on the bird population (Groombridge et al. 2004), as the snails comprised the bulk of the Po’ouli’s diet. Habits Although the Po’ouli was restricted to ­high-­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 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.

’Ula-’ai-Hawane (Hawaii Palmcreeper)  Ciridops anna

(Dole)

Fringilla anna Dole 1878, p. 49 (Hawaii) Specimens  Specimens are in Cambridge, Massachusetts; Honolulu; New York; and Tring. Status  Extinct. Last specimen collected in February 1892. Range  Mountain forests of Hawaii and perhaps also on Molokai, Hawaiian Islands. Description  13cm (5in). Male forehead and lores black, crown and upper back pale grey; ­wing-­coverts and rump red; throat and sides of head black streaked with grey; breast black; belly red; wings black with outer webs of secondaries pink; tail black; bill yellowish; iris dark hazel; legs and feet ­pinkish-­brown. Female generally brownish with a ­blue-­grey breast; wings and tail black, and back ­greenish-­brown.

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 February 20, 1892 near the headwaters of the Awini River on Mount Kohala. 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. Munro (1944) thought that he saw one in 1937 but could not be sure. Olson (1992b) has shown that three of the five known specimens are from Hawaii; the other two (which are at Harvard) are of unknown origin, but circumstantial evidence suggests that they may have been from Molokai. The evidence for this is sound, and further supported by the discovery of subfossil remains of a Ciridops on Molokai, which may or may not be conspecific with C. anna (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). The ­green-­plumaged specimen is almost certainly an adult female (and not an immature 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). Habits The ’Ula-’ai-Hawane was to be found in groves of the Loulu Palm Pritchardia sp., on the fruit and blossoms of which it was reputed to feed; it was wild and shy, but pugnacious (Emerson in Perkins 1903). The Loulu Palm has a patchy distribution, it being generally found in clumps in dense forests, so the ’Ula-’ai-Hawane may have had an irregular distribution as well. Analysis of the skeleton of C. anna has shown that bill is shorter and the pelvic region is much stouter than the other ­red-­and-­black Hawaiian finches, especially in the femur and tibiotarsus (James & Olson 1991). This may be an adaptation for foraging in the matted mass of Loulu Palm inflorescences, in which the legs were used to move tough material to obtain food. 313

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Kaua’i Palmcreeper  Ciridops tenax

James & Olson

Ciridops tenax James & Olson 1991, p. 74 (Makawehi dunes, Kaua’i, Hawaiian Islands) Specimens  Subfossil remains are in Honolulu and Washington, D. C. Status  Extinct. Known only from subfossil remains. Date of extinction unknown. Range  Kaua’i, Hawaiian Islands.

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 ­over-­hunting for its feathers (which were probably red and black). The subfossil remains were discovered in a Kaua’i ­Stilt-­legged Owl Grallistrix auceps pellet (see p. 195), 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.

Bonin Islands Finch  Chaunoproctus ferreirostris

(Vigors)

Coccothraustes ferreirostris Vigors 1828, p. 354 (Bonin-Sima = Chichijima) Specimens  Specimens are in Berlin; Frankfurt; Leiden; New York; St. Petersburg; Tokyo; and Tring. Status  Extinct. Disappeared between 1828 and 1854. Range  Chichijima (formerly Peel Island), Ogasawara (Bonin) Islands. Description  18cm (7in). These were largish finches with massive bills. Male forehead, supercilium, cheeks, throat and upper breast reddish orange or scarlet; lower breast paler, becoming gradually white on belly; rest of plumage ­olive-­brown, striped on back with darker brown. Female differed in being uniform brown or ­olive-­brown except for yellowish forehead and dark brown tips to flank feathers, giving a mottled appearance. Bill blackish horn; legs dull ­yellowish-­brown.

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). Habits Kittlitz (1832) recorded the only known details about the bird: 314

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Bonin Islands Finch Chaunoproctus ferreirostris

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.

McGregor’s House Finch  Carpodacus mexicanus mcgregori

Anthony

Carpodacus mcgregori Anthony 1897, p. 165 (San Benito Island, Baja California) Specimens  Specimens are in Pittsburgh and Tring. Status  Extinct since 1938. Range  San Benito and Isla Cedros, off Baja California, Mexico. Description  13cm (5in). Head, neck and shoulders variably coloured orange; breast, belly, back and tail brown, sometimes streaked on breast and belly; deep grey wash on wing feathers; bill ­horn-­coloured; iris black; legs and feet dark brown. Female brown above and brown streaked dark brown below. Most similar to C. m. amplus but slightly smaller; differed from the nominate C. m. mexicanus in the red areas being orange. However, degree and extent of this coloration extremely variable.

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: 315

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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. 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 ‘miscellaneous’ constructions of bark, twigs and fibre.

Trías Greenfinch  Carduelis triasi

Alcover & Florit

Carduelis triasi Alcover & Florit 1987, pp. 78–79 (Cuevas de los Murciélagos, Los Sauces, La Palma) Specimens  Subfossil remains are in Tenerife. Status  Extinct. Date of extinction not known. Range  La Palma, Canary Islands.

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. 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.

Slender-­billed Greenfinch  Carduelis aurelioi

Rando et al.

Carduelis aurelioi Rando et al. 2010, p. 2 (Cueva del Viento, Tenerife) Specimens  Subfossil remains are in Tenerife and Mallorca. Status  Extinct. Date of extinction not known. Range  Tenerife, Canary Islands.

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.

Parulidae (New World warblers) The New World warblers occur in both North and South America, with the greatest diversity in Central America. The family has been divided into 28 genera with more than 100 species. They are generally small, insectivorous birds, and a number of species are migratory. In general, the New World warblers are not threatened, but habitat destruction has reduced some populations. Two species and one subspecies are probably extinct.

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Bachman’s Warbler  Vermivora bachmanii

(Audubon)

Fringilla bachmanii Audubon, 1833, pl. 165 (near Charleston, South Carolina) Specimens  Specimens are in Boston; Cambridge, Massachusetts; Michigan; New York; Philadelphia; Pittsburgh; Tring; and Washington, D. C. Status  Probably extinct. Last recorded in 1988. Range  Bred in Missouri, western Kentucky, Alabama, and coastal South Carolina, United States; wintered in Cuba and occasionally Florida. Description  12cm. A small warbler with a slender, decurved bill. Male forecrown black; ­hind-­crown and nape grey; forehead, orbital ring, lores, supercilium and throat yellow; underparts yellow with black patch on upper breast; undertail white; upperparts ­olive-­green; wings grey with olive fringes to feathers; lesser ­wing-­coverts yellow; tail grey with white spots on inner webs except for central rectrices; feet and legs brownish; soles of feet yellowish. Female duller, lacking black on upper breast and with whitish orbital ring. Juvenile generally ­brownish-­buff on underside, throat whiter; two buff ­wing-­bars.

Bachman’s Warbler was discovered in 1832 by the Rev. John Bachman, and described by Audubon the following year (Audubon 1833). It was an elusive bird even then, and its distribution was not known until the 1880s. Its main breeding range was in the ­south-­eastern United States: in coastal South Carolina, and in the Gulf States along the Mississippi, especially central Alabama, ­south-­eastern Missouri and western Kentucky. It may once have bred in Virginia, and as far south as Georgia (Dunn 1997). Bachman’s Warbler migrated to Cuba and occasionally Florida in the winter (Hamel 1988), but was most often recorded in Florida and the Florida Keys during the winter migration. It was one of the commonest migrants along the Suwannee River (Ehrlich et al. 1992). Spring migration began in late February, and the birds appeared in the South Carolina breeding grounds in ­mid-­March and in ­south-­eastern Missouri in ­mid- ­to late April. The winter migration to Cuba began in July, peaking by the end of August (Dunn 1997). It was frequently seen on the breeding grounds from the 1880s until about 1910 (King 1981), after which the number of sightings gradually decreased. From the 1930s there were increasingly long periods with no sightings at all. The last specimens collected were taken in Mississippi on February 28, 1940 (a female) and in March 21, 1941 (a male) (Dunn 1997). There are records for Fairfax County, Virginia, in the 1950s. A bird was photographed in South Carolina in May 1958 by Jerry A. Payne, and one was observed in the I’on Swamp near Charleston, South Carolina in April 1962 (Shuler 1977). A female was recorded in Brevard County, Florida, on March 30 1977, and individuals were recorded in Cuba in 1981 (Ripley & Moreno 1980) and 1984 (Hamel 1988). The last official record was in Louisiana in August 1988 (Anderson 1996). There is supposed film footage taken on January 14, 2002 at Guardalavaca, Cuba, but this awaits confirmation. Breeding habitat destruction appears to be the main factor in this bird’s demise. In the late 18th and early 19th centuries, selective logging actually benefitted Bachman’s Warbler (Dunn 1997) by increasing its favoured dense scrub and canebrake habitat. But with the introduction of clear felling during the 1920s, drainage of marshlands, and habitat loss in its wintering grounds (Ehrlich et al. Bachman’s Warbler 1992; Collar et al. 1992), the population Vermivora bachmanii 317

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crashed. Concordant with its increasing rarity was an increase in demand from collectors, and this may have accelerated its decline in the later years. Although there is still hope that Bachman’s Warbler still survives, the lack of recent sightings suggests that it is now probably extinct.

Semper’s Warbler  Leucopeza semperi

Sclater

Leucopeza semperi Sclater 1876, p. 14, pl. 2 (St Lucia, Lesser Antilles) Specimens  Specimens are in Liverpool; Tring; and Washington, D. C. Status  Probably extinct. Not seen since 1961. Range  St Lucia, West Indies. Description  14.5cm (5.75in). Adult a distinctive warbler, fairly large and stocky, with a long, ­deep-­based pointed bill. Head and upperparts dark grey; underparts whitish with grey flanks; bill yellow at base, blackish toward tip; legs and feet pale yellow. ­First-­year birds had ­olive-­brown wash to upperparts; underparts pale ­brownish-­buff, with white throat.

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), unconfirmed 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 ­Fer-­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.

New Providence Yellowthroat  Geothlypis rostrata rostrata

Bryant

Geothlypis rostratus Bryant 1867, p. 67 (Nassau, New Providence, Bahamas) Specimens  Specimens are in Boston and Washington D. C. Status  Possibly extinct. Last recorded in the 1990s. Range  New Providence, Bahamas. Description  13.6cm (5.25in). Upperparts bright yellowish olive; a black mask from forehead to behind ­ear-­coverts, bordered posteriorly with pearl grey, shading to olive of back; underparts bright ­chrome-­yellow; flanks shading to olive on back; bill black; iris dark brown; legs and feet ­flesh-­coloured. Female duller and lacking black mask.

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 mammalian predators has impacted on many bird populations. 318

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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. 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.

Icteridae (New World orioles) The New World orioles have a wide distribution. Most occur in the tropics of North and South America, with the greatest breeding diversity in Colombia and southern Mexico. The family is divided into 27 genera with up to 100 species. They are ­medium-­sized passerines, often displaying bright colours, and some species, like the meadowlarks, are ‘gapers’ (see p. 300). New World orioles are generally not threatened, but some island populations are threatened by anthropogenic disturbance. One continental species and one island subspecies have become extinct.

Grand Cayman Oriole  Icterus leucopteryx bairdi

Cory

Icterus bairdi Cory 1886b, p. 500 (Grand Cayman, West Indies) Specimens  Specimens are in Cambridge, Massachusetts; Tring; and Washington, D, C. Status  Extinct since 1938. Range  Grand Cayman Island, West Indies. Description  18.5cm (7.25in). Front of face and throat black; underparts bright yellow; back dull yellow, washed with faint olive on upper back; wings and tail black; lesser ­wing-­coverts bright yellow; greater ­wing-­coverts white, forming a ­wing-­patch; some of inner edges of primaries and secondaries white; bill and feet black.

The Jamaican Oriole I. leucopteryx occurs on Jamaica and San Andrés Island (off Colombia); it is not considered threatened. However, the Grand Cayman subspecies I. l. bairdi appears to have been affected by severe deforestation, and has not been recorded since 1938 (Johnston 1969). Little was recorded about the Grand Cayman Oriole. On Jamaica, the nominate race occupies subtropical lowland and montane forests, but it can survive in heavily degraded secondary forest.

Slender-billed Grackle  Quiscalus palustris

(Swainson)

Scaphidurus palustris Swainson 1827a, p. 437 (marshes and borders of lakes around City of Mexico) Specimens  Specimens are in Liverpool; Tring and Washington, D. C. Status  Extinct. Last collected in 1910. Range  Marshes of the headwaters of the Río Lerma Santiago, and in Xochimilco in the Valley of Mexico. Description  38cm (15in). Male similar to Boat-tailed Grackle Q. mexicanus but much smaller, with a more slender bill, and glossed violet rather than green. Female cinnamon brown with paler throat and belly. Juvenile similar to female but paler and more buff, less cinnamon.

The ­Slender-­billed Grackle was first described by William Swainson (1827) in a paper devoted to a collection of specimens that William Bullock brought back from his trip to Mexico in 1822. Swainson’s original description is brief: Glossy blue black; thighs brown; bill slender, commissure [line between upper and lower bill] straight; legs slender, claws long, slightly curved. Inhabits the marshes and borders of the lakes round Mexico, in flocks. Total length, 15 inches: bill, 1 and ­seven-­tenths; wings 6½; tail, 7½; tarsi, 1¼. M. Vieillot’s name for this group, Quiscalus, being already used in botany, I propose to call it Scaphidurus, as expressive of the singular ­boat-­shaped tail common to most, if not all, of the species. Very little is known about it, and its true habitat was not known for certain until Dickerman (1965) investigated the circumstances surrounding the last known series of specimens, collected by W. W. Brown in 1910. 319

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He also examined a previously unreported series taken by E. A. Goldman in 1904, which contains the only known juvenile of the species, and as a result Dickerman was able to describe this plumage for the first time. Dickerman also corrected Swainson’s type locality from the marshes and lakes around Mexico City to the headwaters of the Río Lerma Santiago. He claimed that the bird had never occurred in the Valley of Mexico. Peterson (1998), however, overturned this, and demonstrated that Swainson’s original locality was correct. The ­Slender-­billed Grackle almost certainly occurred in both localities. It disappeared due to habitat destruction; persecution due to the damage it did to crops would have hastened its demise. Habits Haemig (2009) 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.

Emberizidae (Buntings and New World sparrows) The relationships of the Emberizidae are in need of review, and some genera, especially the New World sparrows, may not belong in this family. They have a wide distribution, occurring in North and South America and Eurasia, with about 70 genera currently recognized. The family is generally not considered threatened, and remarkably few species have become extinct. One of these, the ­Long-­legged Bunting Emberiza alcoveri, is one of the very few passerines that had evolved flightlessness.

Dusky Seaside Sparrow  Ammodramus maritimus nigrescens

Ridgway

Ammodromus [sic] maritimus nigrescens Ridgway 1873, p. 198 (Indian River, Florida) Specimens  Specimens are in Chicago; Tring; and Washington, D. C. Status  Extinct since June 17, 1987. Range  Merritt Island and adjacent mainland, within a ­ten-­mile radius of Titusville, Brevard County, Florida. Description  13–14cm (5–5.5in). A very dark subspecies of Seaside Sparrow A. maritimus. Adult generally blackish above, undersides white heavily streaked with black; throat, belly and vent white; yellow streak above eye; bill, legs and feet grey; iris black.

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 traditionally nested began to disappear in the lower parts of the marsh. Delany (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.

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Decline and extinction The Dusky Seaside Sparrow was considered threatened with extinction in 1966 and officially classed as endangered under the 1973 Endangered Species Act. There were approximately 900 birds in 1968 (Sharpe 1970), but the population steadily declined. In 1977 there were only 28 pairs, and in 1979 just 13 remained (Sykes 1980). By 1981 only five males and no females were left, and these last surviving birds were taken into captivity. In 1986 two birds remained. The last Dusky Seaside Sparrow died on June 16, 1987 in Walt Disney World’s Discovery Island Zoological Park in Orlando, Florida (Anon 1987). The disastrous attempt to hybridise the last male Dusky Seaside Sparrows with females of the subspecies A. m. peninsulae (Zink & Kale 1995) is another example of human apathy, bad politics and arrogant taxonomic philosophy, all of which conspired to doom this bird to extinction (see Walters 1992; Rising 2005). 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.

Guadalupe Towhee  Pipilo erythropthalmus consobrinus

Ridgway

Pipilo maculatus consobrinus Ridgway 1876b, p. 189 (Guadalupe) Specimens  Specimens are in Cambridge, Massachusetts; New York; and Washington, D. C. Status  Extinct. Last seen in June 1897. Range  Guadalupe Island, off the coast of Baja California, Mexico. Description  80cm (7in). Resembled the Oregon Spotted Towhee P. e. oregonus of western North America, but back browner; black areas of male duller; larger hind claw; and wing and tail shorter. Head black; back and ­wing-­coverts ­blackish-­brown; ­wing-­coverts with outer margins invariably marked white, giving a speckled appearance; throat and upper breast ­blackish-­brown; sides and flanks dark ­buffish-­brown; belly white; tail brown, outer feathers tipped with white.

The Eastern Towhee P. erythropthalmus is widely distributed across the 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 1876). 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 responsible 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 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).

Santa Barbara Song Sparrow  Melospiza melodia graminea

Townsend

Melospiza fasciata graminea Townsend 1890, p. 139 (Santa Barbara Island, California) Specimens  Specimens are in Washington D. C. Status  Extinct since about 1960. Range  Endemic to Santa Barbara, off the California coast.

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Extinct Birds Description  13cm (5in). ­Ash-­grey on upperparts, especially on hind neck; small dark streaks on back; white underneath with dark streaking; dark brown spot in middle of breast; brown cap and tail; face grey with dark streak through eye; bill, iris and legs and feet brown. Differed from the nominate in being overall lighter, giving an ­ash-­grey appearance.

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.

Todos Santos ­Rufous-­sided Sparrow  Aimophila ruficeps sanctorum  van Rossem Aimophila ruficeps sanctorum van Rossem 1947, p. 55 (Todos Santos Islands, off Ensenada, Baja California) Specimens  The type specimen is in Berkeley. Status  Extinct. Last recorded in the 1970s. Range  Todos Santos Islands, off Baja California, Mexico. Description  13cm (5.25in). Adult; generally brown above with dark streaks; grey undersides; crown rufous; face and supercilium grey with brown or rufous streak extending from each eye; thick black malar streak; throat white, with dark stripe; bill yellow; legs and feet ­pinkish-­grey. Much darker than nominate A. r. ruficeps.

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.

Antioquia ­Brush-­finch  Atlapetes blancae

Donegan

Atlapetes blancae Donegan 2007, p. 257 (Llano de Ovejas, Antioquia, Colombia) Specimens  Specimens are in Bogotá and Medellin. Status  Possibly extinct. Survived until at least 1971. Range  Antioquia, Colombia. Description  18cm (7in). Adult forehead, crown and nape bright rufous with black face mask; indistinct dark moustachial stripe; back and ­uppertail-­coverts ­slate-­grey; darker ­soot-­grey flight feathers with a small white ­wing-­panel at base of central primaries; underside pale whitish; bill black; legs and feet probably pale ­pinkish-­brown. Juvenile lacks wing-panel; overall browner.

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 ­Brush-­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. 322

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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 elsewhere, 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. 457), so there is some hope for the Antioquia birds.

Long-­legged Bunting  Emberiza alcoveri

Rando et al.

Emberiza alcoveri Rando et al. 1999, p. 2 (Cueva del Viento, Tenerife) Specimens  Subfossil remains are in Tenerife. Status  Extinct. Date of extinction unknown. Range  Tenerife, Canary Islands.

The discovery of a flightless bunting on Tenerife, Canary Islands, has shown that flightlessness has occurred in a number of passerine genera, although it is a rare phenomenon. The ­Long-­legged Bunting had limb proportions similar to the extinct and similarly flightless Stephens Island Wren Xenicus lyalli (see p. 221) and ­Long-­billed Wren Dendroscansor decurvirostris (see p. 223) of New Zealand. Like those birds it was probably a terrestrial inhabitant of the forest floor. The musculature of the bill suggests that it was capable of feeding on harder seeds than other members of the genus (Rando et al. 1999). It may have been an inhabitant of the ­herb-­rich understorey associated with the laurel forests. Two waves of human colonisation on the islands, the first about 2,000 bp and the second in the 15th century, caused severe habitat destruction (Rando et al. 1997). This, along with introduced mammals, including cats and rats, was almost certainly the cause of the bunting’s extinction.

Long-legged Bunting Emberiza alcoveri

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 ­medium-­sized birds, many being brightly coloured, but the family includes the dull ­blackish-­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 tanagers are threatened by habitat destruction and introduced mammalian predators. Two species and four subspecies have become extinct, including one subspecies of Galápagos finch.

Darwin’s Large Ground Finch  Geospiza magnirostris magnirostris

Gould

Geospiza magnirostris Gould 1837b, p. 5 (Galápagos Islands) Specimens  Specimens are in Cambridge, England and Tring. Status  Extinct. Last recorded in 1957. Range  Known only from Floreana (Charles Island) and San Cristóbal (Chatham Island).

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Extinct Birds Description  15cm (6in). A large Geospiza with an extremely large bill. Male uniform black except ­undertail-­coverts white or white–tipped. Female generally dark brown, olive edges to feathers giving a mottled appearance; throat and breast with whitish buff edges to feathers; lower breast and belly whitish buff speckled with dark brown; bill black in male, variably black or brownish horn with darker tip in female; iris dark brown; legs and feet black.

The taxonomy of the Large Ground Finch Geospiza magnirostris is extremely confused. The Large Ground Finch has been divided into two subspecies, nominate G. m. magnirostris, endemic to Floreana (Charles Island) and San Cristóbal (Chatham Island), and G. m. strenua, which occurrs on several of the other islands. The Floreana race had the largest bill of any of the Galápagos finches, being slightly larger than G. m. strenua. However, there has been considerable confusion regarding the validity of the nominate form. It was first collected by Charles Darwin and others supposedly 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 (Hellmayr 1938). Furthermore, Swarth (1931) argued that as the ­largest-­billed individuals of G. m. strenua came from the north of the archipelago, including James Island, it is likely that Darwin obtained his specimens of G. m. magnirostris from there. This was given further support when subsequent ornithological surveys to Floreana and San Cristóbal failed to locate the birds (Lack 1947). Either the nominate Large Ground Finch had become extinct since the time of Darwin’s visit, or it never occurred on Floreana or San Cristóbal in the first place. 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, including G. m. magnirostris, and collected specimens of this form on both Floreana and San Cristóbal. Therefore, it is beyond doubt that Darwin’s Large Ground Finch once occurred on these two islands, and that it was a valid but ­now-­extinct taxon. Bowman (1961) collected a single bird on Floreana in September 1957, and also saw other individuals. But Sulloway (1982) showed that Bowman’s specimen is morphologically more similar to G. m. strenua than to G. m. magnirostris, and that Bowman had in fact encountered a small population of G. m. strenua that had recently colonised the island. This population subsequently died out. Habits Almost nothing is known about the bird’s habits. Darwin (in Gould 1841) records that ‘he seldom saw them in the upper and damp regions’, inferring that they usually inhabited the lowlands. The extinction of Darwin’s Large Ground Finch must surely be a direct result of human activity on the Galápagos Islands. It was an extremely tame bird, and Darwin saw a small boy killing them with a stick near a well on Floreana (Greenway 1967). The introduction of goats and pigs would have destroyed its habitat and introduced rats would have been serious predators.

St Kitts Bullfinch  Loxigilla (portoricensis) grandis

Lawrence

Loxigilla grandis Lawrence 1881, p. 204 (St Christopher = St Kitts) Specimens  Specimens are in Cambridge, England; Chicago; New York; Philadelphia; and Washington, D. C. Status  Extinct. Last collected in July 1929. Range  St Kitts, West Indies. Description  15cm (6in). Generally a uniform dull glossy black, with terracotta crown, throat, and ­undertail-­coverts; anal coverts with black speckles; bill, legs and feet black. Juvenile forehead, area above eyes, and throat with few red feathers; underparts brownish beige, tinged with cinnamon; vent feathers dark grey; back and rump brownish, tinged ­olive-­reddish. Larger and glossier than nominate L. p. portoricensis, which has a less extensive upper ­breast-­patch.

The Puerto Rican Bullfinch Loxigilla portoricensis occurs on Puerto Rico, where it is considered not threatened. 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, when Ober collected the type series (Lawrence 1881). 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 324

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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.

Samaná Eastern Chat Tanager  Calyptophilus frugivorus frugivorus

(Cory)

Phoenicophilus frugivorus Cory 1883, p. 45 (Samaná Peninsula, Dominican Republic) Specimens  Specimens are in Washington D. C. Status  Possibly extinct. Last recorded in 1982. Range  Samaná Peninsula, ­north-­eastern Dominican Republic. Description  17–20cm (6.75–8in). Adult top of head brown, shading to ­ash-­brown on neck; rest of upperparts, wings and tail ­brownish-­olive; throat and breast white, becoming ­ash-­brown on sides; flanks ­brownish-­olive; carpal area and loral spot yellow; distinct yellow broken ­eye-­ring; upper bill dark brown; lower bill ­yellowish-­brown, darker at base. Female smaller.

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. 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 325

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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 tanager’s large, strong feet were adapted for searching through the leaf litter (Latta et al. 2006).

Gonâve Eastern Chat Tanager Calyptophilus frugivorus abbotti

Richmond & Swales

Calyptophilus frugivorus abbotti Richmond & Swales 1924, p. 106 (Gonâve Island, Haiti) Specimens  Specimens are in Puerto Rico and Washington D. C. Status  Probably extinct. Last recorded in 1977. Range  Gonâve Island, Haiti. Description  17–20cm (6.75–8in). This subspecies was similar to nominate in coloration, except it was overall paler, and slightly smaller in size.

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 extensive 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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Hypothetical Birds The accounts

Megapodiidiae (Megapodes) Anderson’s Megapode  Megapodius andersoni

Gray

Tetrao australis Anderson MS from Captain Cook’s second voyage; in Banks’s MS, no. 81 ? Megapodius ? andersoni Gray 1862a, p. 291 (New Caledonia) Megapodius sp. Seebohm c. 1895 MS Specimens  One egg from New Caledonia, formerly in the Gould Collection. This egg is no longer in Tring. Status  No living megapode survives on New Caledonia. May represent an extinct species. Range  Formerly occurred on New Caledonia.

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. 96). Balouet & Olson (1989) mention, but do not overly support, Anderson’s account based on the available evidence, but recent work (Steadman 2005) has shown that another (as yet undescribed) megapode once occurred alongside the extinct ­Pile-­builder Megapode Megapodius molistructor (see p. 35). The possibility exists that Anderson saw either one of these.

Kermadec Megapode  Megapodius sp. Megapodius sp Cheeseman 1890, p. 219 (Sunday Island, Kermadec Islands) Status  Known only from the accounts of T. F. Cheeseman (1890) and J. J. Lister (1911). Range  Believed to have been wiped out in a volcanic eruption in 1876, but the status of the form is unclear.

Cheeseman (1890) 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 (1890) 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 327

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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 zoogeographic 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 populated by megapodes (Steadman 2005).

Phasianidae (Grouse and quails) Two-banded Argus  Argusianus bipunctatus

(Wood)

Argus bipunctatus Wood 1871, p. 281 (no locality) Specimens  The unique type is held at Tring. Status  Known only from a portion of a single primary feather, of unknown origin. Presumed to represent a now extinct species, but possibly an artefact or mutation. Range  Unknown. Description  The feather differs considerably from that of Great Argus A. argus; shaft very thin (suggesting that it comes from near the tip of a very long feather); general colour deep blue with short black bars on both sides of reddish chestnut; minutely spotted band on either web, followed by a buff band spotted dark brown; both webs resemble the inner web of Great Argus.

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. Mysterious origins and ecology A number of authorities have managed to confuse rather than clarify the origins and ecology of this enigmatic bird. Based on the length and width of the feather, and compared with the Great Argus, Davison 328

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(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 provenance, 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.

Anatidae (Ducks, geese and swans) Newfoundland Duck (Brown Duck)  Anas fuscescens

Gmelin

Brown Duck. Pennant 1785, p. 565 Anas fuscescens Gmelin 1789, p. 534 Status  Known only from the descriptions. Never identified, but possibly represents an extinct species. Range  Newfoundland.

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 ­rust-­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 labradorius (see p. 53), the Newfoundland Duck may have existed but disappeared even earlier.

Cathartidae (Vultures) Painted Vulture  Sarcorhamphus sacer

(Bartram)

The Warwoven or Indian Vulture. Albin 1734, p. 4, pl. 4 (Albin’s name is usually quoted as one of the joint sources of Linnaeus’s Vultur (i.e. Sarcorhamphus) papa, but the white tail suggests it was more likely to have been this form) Vultur sacra Bartram 1791, p. 148 (Florida) Status  Probably extinct since the mid-18th century. Known only from Bartram’s account, and Albin’s plate. Range  Formerly inhabited the ­south-­eastern United States.

The case history of this bird is remarkable, and highlights the extraordinary reluctance of ornithologists to admit the possibility that birds could have become extinct in historical times in such seemingly ­well-­known areas as the ­south-­eastern United States (e.g. Allen 1871).

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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. What was this vulture? Bartram’s description of the Painted Vulture agrees with that of the King Vulture Sarcorhamphus papa, except for the tail, which in the King Vulture is black; in the Painted Vulture it was white with a black tip. Furthermore, the King Vulture is not known to occur north of Mexico; Bonaparte (1828) stated that it occurred ‘occasionally in Florida during the summer’, though this statement may have been based on Bartram’s account. Harper (1936), who examined the case in detail, and McAtee (1942), both considered that Bartram had genuinely observed a population of the King Vulture (or something closely related to it), which was restricted to Florida but had died out at some point in the mid-18th century. There are some biogeographical grounds to support this; a handful of Neotropical species maintain populations in Florida (Crested Caracara Polyborus cheriway, for example). With regard to the vulture, in the absence of specimens, nothing more can be said about it.

Rallidae (Rails and coots) Solomon Islands ­Bare-­eyed Rail  Gymnocrex plumbeiventris intactus

(Sclater)

Rallus intactus Sclater 1869a, Proceedings of the Zoological Society of London, p. 123, pl. 10 (Solomon Islands) Specimens  The unique type is in Tring. Status  Extinct, or possibly survives in very low numbers. Known only from the type. Range  Solomon Islands. Description  30 cm (12 in) Head and neck vinous chestnut, blushed with maroon, paler on throat; upperparts ochraceous ­olive-­brown; ­wing-­coverts, lower back, rump, ­uppertail-­coverts and tail black; primaries rufous, shaded ­olive-­brown at tips; secondaries ­olive-­brown; underside ­silver-­grey, darker on lower flanks and ­undertail-­coverts; bill ­olive-­yellow, darker at tip; iris red; legs and feet ­coral-­red.

The unique type specimen of this bird was obtained along with a number of others during a voyage to the Solomon Islands in 1872 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 ­Bare-­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. 330

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Dark Rail  Porzana (tabuensis) tenebrosa

(Gray)

Rallus tenebrosus. Latham unpublished MS Dark Rail. Latham 1824, p. 378 (Norfolk Island) Rallus tenebrosus G. R. Gray 1862b, p. 239 footnote (Norfolk Island) Porzana tabuensis tabuensis Ripley 1977, p. 230 Status  Known only from Latham’s description. Range  Norfolk Island. Description  18–20 cm (8–9 in). Upperparts dusky black with deep bluish tinge; back and wings deep brown; ­underwing-­coverts black with darker spots; bill dusky; iris red; legs and feet pale red.

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.

Miller’s Crake (Miller’s Rail)  Porzana nigra

(Miller)

Rallus nigra Miller 1784, pl. 50, fig. B (Tahiti) Status  May represent an extinct species. Known only from illustrations and descriptions. Range  Tahiti. 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 1989). It appears in two illustrations, 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 & Iredale1921; Lysaght 1956b). There are also later descriptions by Latham and Gmelin. Lysaght (1956b) believed Miller’s plate to depict the bird now known as Porzana 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). Peters (1934), after examination of Miller’s illustration, tentatively placed P. nigra in the synonymy of the Henderson Island Crake P. atra, but commented: Miller’s plate represents a wholly black rail somewhat larger than tabuensis; it cannot be identified with any of the known forms of tabuensis and possibly represents the bird later named Porzana atra, or at least a bird closely allied to it. Peters overlooked the fact that the provenance of Miller’s Rail was Tahiti, however. Considering the speciation of rails on Pacific oceanic islands, it is more than likely that Miller’s Rail represents a ­now-­extinct population, perhaps related to P. tabuensis, or a species akin to P. atra. Mangaia Recent palaeontological work on Mangaia, the southernmost of the Cook Islands, has shown that populations of Porzana rails occurred sympatrically there, one related to Porzana tabuensis (or indeed a race of it), and the other related to P. atra (Mangaia Crake P. rua; see p. 103) (Steadman 1985). Steadman adds significantly: The Mangaian fossils suggest that perhaps two species of Porzana once inhabited many of the islands in eastern Polynesia, the smaller being P. tabuensis or its derivative, and the larger, more flightless one being similar to P. atra. 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 331

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might refer to P. 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) and Walters (1989) fully support the validity of Porzana nigra, and coupled with the discovery of so many new but extinct rallids throughout the Pacific, subfossil remains of a Porzana rail on Tahiti will almost certainly be found at some point in the future.

Scolopacidae (Snipes and sandpipers) Ellis’s Sandpiper  Prosobonia ellisi

Sharpe

Prosobonia ellisi Sharpe 1906b, p. 86. Status  Extinct. Known only from two illustrations. Range  Moorea, Society Islands, South Pacific. Description  Differed from the White-winged Sandpiper P. leucoptera in having a rufous patch or streak over the eye, not whitish; greater coverts pale rufous; legs yellowish not green; and bill slightly downcurved, not straight.

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 ­White-­winged Sandpiper P. leucoptera of Tahiti (see p. 129) 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 population 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.

Kiritimati Sandpiper  Prosobonia cancellata

(Gmelin)

Barred Phalarope. Latham 1785, p. 274 (Christmas Island, Pacific Ocean; based on an unpublished figure by W. W. Ellis, no. 64 in NHM) Tringa cancellata Gmelin 1789, p. 675 (‘in insula Nativitatis Christi’ = Christmas Island). Based on Latham Status  Known only from the type, now lost. Generally considered conspecific with Tuamotu Sandpiper P. parvirostris (in which case this species takes the name cancellata). Range  Kiritimati (Christmas Island), Kiribati, South Pacific.

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 recent 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). 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). 332

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Columbidae (Pigeons and doves) Vava’u Pigeon  Columba vitiensis ssp.

Latham

Pacific Pigeon. Latham 1823, p. 45 (Howe’s Island) Caloenas sp. Gray 1859a, p. 45. Status  Known only from Latham’s description. Never satisfactorly identified but possibly representing an extinct form. Range  Vava’u Islands, Tonga, South Pacific.

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.

Strigopidae (Kakas, keas and kakapos) Lord Howe Island Kaka  Nestor sp. According to Hindwood (1940), two observers, Bowes (1788) and Watts (in Phillip 1789), refer to parrots as well as parakeets on Lord Howe Island. Neither of these men visited Norfolk Island, where a parrot Nestor productus and parakeet Cyanorhamphus cookii ­co-­existed, but both men landed on Lord Howe Island at the same time. No one else has ever mentioned parrots on Lord Howe, but if one existed it was quite likely to be a Nestor.

Psittacidae (Parrots and macaws) Moseley’s Parrot  Psittrichas sp. Dasyptilus pequetti Moseley 1879, p. 479. (Admiralty Islands) Status  Known only from Moseley’s description. Range  Admiralty Islands, Bismarck Archipelago, ­south-­west Pacific.

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 population 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 Lorikeet  Charmosyna samoensis

Hartlaub

Papagaien. Kotzebue 1830, p. 154 (Olajava = Samoa). Charmosyna samoensis Hartlaub 1867, p. 21 Status  Known only from Kotzebue’s description. Never identified, but possibly a ­now-­extinct species. Range  Samoa, South Pacific.

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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, yellowtipped 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 specifically distinct.

Tanna Parrot  Prosopeia sp. Parroquets. G. Forster 1777, p. 334 (Tanna). Status  Known only from Georg Forster’s account. Range  Tanna, Vanuatu, South Pacific.

Georg Forster mentions this bird only in passing: The next morning we came ashore again, and immediately walked into the woods on the plain. We saw a great number of large and beautiful parroquets, of black, red and yellow plumage; but they kept on the tops of the highest fig trees, where they were wholly out of reach of small shot, guarded by the thick foliage. No one else has ever seen these parrots, nor has any reference been made of them except to quote the substance of Forster’s account. There seems little reason to doubt that parrots formerly occurred on Tanna, the southernmost of the large islands in the New Hebrides. The likely relationship of this species would have been with the Prosopeia shining parrots, a genus comprising three species occurring in Fiji and Tonga (Juniper & Parr 1998). These birds are predominantly black, yellow, red and green in colour. How the Tanna birds, if indeed they were an indigenous population, differed is now impossible to say.

Westerman’s Eclectus Parrot  Eclectus roratus westermani

Bonaparte

Eclectus westermani Bonaparte 1850, p. 4 (no locality) Specimens  Specimens are in Amsterdam; Leiden; and Tring. Status  Known from 16 captive birds, this enigmatic subspecies is often considered an aviary hybrid. Last recorded in 1899. Range  Possibly Indonesia, but provenance in doubt. Description  35 cm (14 in). The Eclectus Parrot E. roratus is generally green in the male, sides of body red; rostrum coral–red with yellow tip; mandible black; iris orange; legs and feet grey. Female generally red, darker and more brownish on back and wings; lower breast, abdomen and band across upper back dull purple; bill black; iris ­yellowish-­white; feet and legs grey. Male E. r. westermani: similar to E. r. roratus except smaller; overall duller, more bluish; less red on sides of body. Female: smaller.

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.

Rose-fronted Parrot  Psittacula sp. Rose-fronted Parrot. Latham 1822, p. 186 (no locality) Specimens One in Liverpool. Status  Known only from Latham’s description of the type, still extant in Liverpool Museum. It is clearly a Psittacula, but cannot be identified with any known species. Range  Unknown. Description  22 cm (8.5 in). A green, ­long-­tailed parakeet. Upperparts green, more yellowish on underparts; forehead and around bill dull ­rose-­buff; cheeks and chin dusky pale green; rump and ­underwing-­coverts ­blue-­green; primaries dusky edged with green; outer tail feathers green, the two central feathers blue with pale tips; bill dusky ­pale-­flesh; legs and feet brown.

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Salvadori (1891a) thought that Latham’s ­Rose-­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.

Obscure Parrot  Psittacus obscurus

Linnaeus

Psittacus obscurus Linnaeus 1758, p. 97. Based on Hasselquist 1757, p. 236 (Africa) Status  Known only from Linnaeus’s description, based on that of Hasselquist. Range  Africa.

The identity of this parrot remains a mystery. Linnaeus (1766) synonymised this name with the extinct Mascarene Parrot Mascarinus mascarinus, but the descriptions totally disagree. It was described from a specimen 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.

Jamaican ­Yellow-­headed Macaw  Ara gossei

Rothschild

Yellow-headed Macaw. Gosse 1847, p. 260. Ara tricolor Clark 1905d, p. 348 Ara gossei Rothschild 1905b, p. 14 (Mountains of Hanover Parish, about ten miles east of Lucea, Jamaica) Status  Known from Gosse’s description of a specimen, now lost, and a contemporary painting. Range  Jamaica. May have survived until the late 18th century. Description  Forehead, crown and back of neck bright yellow; sides of face around eyes, rest of neck and back scarlet; ­wing-­coverts and breast deep red; primaries and ­primary-­coverts light blue; tail red and yellow intermixed; basal half of upper bill black, apical half ­ash-­coloured, lower bill black with ­ash-­coloured tip; legs and feet black.

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 335

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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 impossible to ascertain.

Guadeloupe Parakeet  Aratinga labati

(Rothschild)

Perriques. Labat 1742, p. 218 Conurus labati Rothschild 1905b, p. 13 (Guadeloupe) Status  Extinct, probably since the late 18th century. Known only from Labat’s description and an account and illustration in Du Tertre (1667). Range  Guadeloupe, West Indies.

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. 338) and Guadeloupe Macaw (see p. 183). 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.

Martinique Parakeet  Aratinga pertinax ssp. Brown-throated Parrakeet. Edwards 1751, p. 177, pl. 177 Psittacus aeruginosus Linnaeus 1758, p. 98 (‘America’ = West Indies) Psittaca martinica Brisson 1760, p. 356 (based on Edwards) 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 ­Brown-­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 ­Brown-­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.

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Linnaean problems The application of the name aeruginosus has posed a number of nomenclatural problems. Peters (1937) 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 ­ orth-­western Venezuela, but Edwards’s plate does not agree with birds from these areas either. Clark n (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 ­Brown-­throated Parakeet subspecies that once occurred on Martinique. It was not recorded again, and the reasons for its disappearance are unknown. No parakeet survives on Martinique today.

Ober’s Parakeet  Aratinga sp. Parrot. Ober 1878, p. 64 (no description) Status  Probably represents an extinct population. Range  Formerly occurred on Dominica.

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 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.

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Barbados Parakeet  Aratinga sp. Parakite. Hughes 1750, p. 73 Status  Known only from the account of Hughes (1750). Range  Probably once inhabited Barbados.

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. 336), 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.

Guadeloupe Amazon  Amazona violacea

(Gmelin)

Perroquet de la Guadeloupe. Du Tertre 1654, p. 298 Perroquets. de Rochefort 1658, p. 157 Perroquets. Labat 1742, p. 214 Psittacus violaceus Gmelin 1788, p. 337 (‘Insulae aquarum Lupiarum’ = Guadeloupe). Ruff-necked Parrot. Latham 1822 p. 217 Status  Extinct since the 18th century; known only from the descriptions of Du Tertre, de Rochefort and Labat, and an illustration in Du Tertre. Range  Guadeloupe, West Indies. Description  Most similar to the Imperial Amazon A. imperialis, both being characterised by a large ruff around the neck, but while A. imperialis is primarily green, A. violacea seems to have been ­slaty-­blue (Labat described this as slate; Du Tertre as violet). The ­slaty-­blue was intermixed with a few green and black feathers, the back completely green, and the wings green, yellow and red.

The surviving West Indian amazons are found on Cuba and the Bahamas, Jamaica, Hispaniola and Puerto Rico, and three islands in the Lesser Antilles, Dominica, St Vincent and St Lucia (Juniper & Parr 1998). They are all considered endangered, primarily due to trapping for the pet trade, ­over-­hunting for food and habitat destruction (BirdLife International 2011). Amazons were also mentioned from Guadeloupe, the largest island in the Lesser Antilles, and Martinique, which is situated between Dominica and St Lucia (see below). The possible existence of these parrots is based on accounts only and not supported by physical evidence of any kind. However, both were mentioned by trusted observers, and zoogeographically at least it is very likely that they once existed. 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 flavor of its flesh is excellent, but changeable, according to the kind of food. If it eats cashew nuts, the flesh has an agreeable flavor of garlic; if ‘bois des inde’ it has a flavor 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 flavor. When it feeds on guavas it is at its best, and then the French commit great havoc among them. 338

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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 it 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 the pressures on the birds would have been greater. The endemic amazon had disappeared by the end of the 18th century. 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.

Martinique Amazon  Amazona martinicana

Clark

Perroquets. Labat 1742. p. 214 Psittacus leucocephalus var. Vieillot 1817, p. 332 (Martinique) Amazona martinicana Clark 1905c, p. 343 (Martinique) Status  Extinct probably since the early 18th century. Known only from Labat’s brief description of 1742. Range  Martinique, West Indies.

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 (1905) 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). Rothschild’s (1907) plate does not resemble Labat’s description, and since A. arausiaca has a blue head, it is possible that Labat saw the bird in the trees, where, in the shadows, this feature was not obvious. It seems therefore that A. martinicana differed from A. arausiaca only in having some red feathers on the head. There seems no reason to doubt the existence of this parrot; subfossil remains will confirm its former existence on the island.

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 ­ lue-­headed Hummingbird Cyanophaia bicolor, a species that still occurs on the island. He described the B adult male as follows: 339

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Upperparts rich iridescent metallic green, becoming deep peacock blue or ­verditer-­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 outermost 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 beautiful structure of felted ­tree-­fern down, ornamented with spider webs, beetle elytra, and bits of ­blue-­green 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 specimen’ was correct, the bird was most likely an aberrant ­Blue-­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) Latham’s Black-capped Kingfisher  Halcyon sp. Black-capped Kingfisher, var. B. Latham 1781, p. 626 (‘South Seas’) Status  Known only from Latham’s description, based on a specimen in the Leverian Museum, now lost. Never satisfactorily identified, but perhaps representing an extinct species from the South Pacific. Range  Possibly one or more South Pacific islands.

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 represent a ­now-­extinct kingfisher from somewhere in the South Pacific.

Miyako Kingfisher (Ryukyu Kingfisher) Halcyon (Todiramphus) miyakoensis

Kuroda

Halcyon miyakoensis Kuroda 1919, p. 231 (Miyakoshima, Ryu Kyu Islands) Specimens  The type specimen is in Tokyo.

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Hypothetical Birds Status  Now extinct. Known only by the type, a male, collected in 1887. Range  ­Miyako-­jima, Ryukyu Islands. Description  24cm (9.5 in). Lores black; small white patch over eye; narrow ­greenish-­blue stripe under eye and widening to side of nape; rest of head and neck cinnamon; back and scapulars dark greenish blue; lower back and ­uppertail-­coverts cobalt blue; primaries brownish black; ­wing-­coverts and tail ultramarine tinged green; underparts cinnamon, slightly darker on breast; iris not recorded; feet dark red; claws brown. Differs from Micronesian Kingfisher H. cinnamominus in lack of a black ­nape-­band and red (not black) legs and feet.

The Ryukyu Islands stretch from 225km ­south-­west of Okinawa to 290km from the northern tip of Taiwan. The Miyako Kingfisher was collected on February 5, 1887 on ­Miyako-­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 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) Klages’s Jacamar  Brachygalba lugubris phaeonota

Todd

Brachygalba phaeonota Todd 1943, p. 10 (Tonantins, Rio Solimões, Brazil) Specimens  The type specimen is in Pittsburgh. Status  Known only from the type; taxonomic position is in doubt. Range  Tonantins, Rio Solimões, Brazil. Description  18 cm (7 in). Upperparts sooty black, glossed ­greenish-­black; wings and tail blackish with slight blue sheen; shafts of chin and throat whitish with rufescent tips; breast, flanks and ­undertail-­coverts ­rufescent-­brown; lower breast and belly ­dirty-­white; bill black; legs and feet black; iris dark brown.

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.

Paradisaeidae (Birds of paradise) Duivenbode’s Riflebird  Craspedophora (Paryphephorus) duivenbodei

Meyer

Craspedophora duivenbodei Meyer 1890, p. 419 (Mountains of New Guinea) Specimens  Specimens in Tring and New York; the Dresden type was lost during World War II. Status  Possibly a valid rare or extinct species. Known from three specimens, the last taken November 1, 1937. Range  ­Deva-­Deva and Foula, near Yule Island, south-eastern New Guinea. Description  24 cm (9.5 in). Two slightly different plumage variations. New York specimen velvet black with purplish gloss; crown metallic green; sides of face and throat black with purple and bronze reflections; ­hind-­neck black with a fanlike frill of black feathers glossed with violet; triangular breast shield metallic green showing more blue towards the margins; tail black with the two central feathers metallic green: underparts black washed with bronze and violet on flanks; bill, legs and feet black; iris probably dark brown. Tring specimen differs in having crown metallic blue and purple, not green; sides of face black with a slight golden olive tinge; breast shield bluish green widening to a purplish band; underparts completely black; and wings and tail black, the latter glossed with blue. Cape on upper back in both specimens indistinct.

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The type of this bird was obtained by Messrs van Duivenbode, Dutch merchants on Ternate in the Moluccas, and sent to Dresden Museum, where it was described by Meyer (1890). It came from ­north-­western New Guinea, but neither the exact locality nor the collector appears to be known. The specimen was lost in World War II. Two other specimens were collected, but the loss of the type is important, for it differed significantly from the others in having a cape of black feathers on the upper back. The last two specimens came from the mountains inland from Yule Island, off the coast of south-eastern New Guinea. The first of these was taken in 1903 by Antwerp Edgar Pratt at Foula at about 1,300m. He was on a collecting expedition, and the skin was sent to Walter Rothschild at Tring, eventually passing to the American Museum of Natural History with the rest of Rothschild’s collection. The collector Fred Shaw Meyer obtained another specimen (now at Tring) at ­Deva-­Deva, no more than a mile from Pratt’s camp, on November 1, 1937. The fact that these two specimens were found so close together is of great significance, as Fuller (1995) was quick to point out. Stresemann (1930) had considered Duivenbode’s Riflebird to be a hybrid of the Magnificent Riflebird Ptiloris magnificus and Superb Bird of Paradise Lophorina superba, two ­wide-­ranging species in New Guinea (Frith & Frith 2010 reached a similar conclusion). This was before the discovery of the second specimen. If hybridisation had occurred between the Magnificent Riflebird and Superb Bird of Paradise, their hybrids might be expected to occur widely, but this seems not to be the case, although Frith & Frith (2010) dispute this. Earlier, Fuller (1979) had pointed out that the Tring specimen does not show incontrovertible characters of the putative parents, and that Gilliard (1969) maintained that the Superb Bird of Paradise could hardly hybridise with the Magnificent Riflebird as their altitudinal ranges do not meet. It is possible of course that in ­south-­eastern New Guinea they come into more regular contact. Conversely, Duivenbode’s Riflebird may be a rare or extinct species confined to the mountains near Yule Island.

Elliot’s Bird of Paradise  Epimachus ellioti

Ward

Epimachus ellioti Ward 1873, p. 742 (Dutch New Guinea = western New Guinea) Specimens  Specimens are in Dresden and Tring. Status  Possibly a valid rare or extinct species. Known from two specimens, the last taken before 1893. Range  North-­western New Guinea. Description  55 cm (21.5 in). Resembles more the sicklebills in having tufts of feathers forming a pair of ‘false wings’, which are presumably raised in display. General coloration dark, almost black with purple and violet iridescence, markedly so on crown in contrast to the black forehead; on cheeks the gloss is green; throat velvet ­blackish-­brown merging into wash of ­olive-­green on breast; broad band of dull ­grape-­red crosses this area; false wings tipped with blue.

Elliot’s Bird of Paradise is an enigmatic bird. Both specimens originated somewhere in ­north-­western New Guinea, but the precise location is not known. It was suggested that it might have come from Waigeu, on the strength of a statement made by Labillardière (1800): I killed a great many scarce birds, among others the species of promerops, which Buffon calls the promerops of New Guinea. The ‘promerops’ referred to here was a sicklebill, no species of which is known to occur on Waigeu. Perhaps, therefore, Elliot’s Bird of Paradise is an extinct form from that island, but the link is tenuous. Stresemann (1930; followed by Frith & Frith 2010) suggested that Epimachus ellioti was a hybrid of a Black Sicklebill Epimachus fastuosus and an Arfak Astrapia Astrapia nigra, which seems unlikely, as E. ellioti differs in a number of characteristics (Fuller 1995). Fuller (1979) discussed E. ellioti at some length. He remarked that the pairing of E. fastuosus and A. nigra had been used to account for another alleged hybrid, and stated: A distinctly different bird known as Epimachus astrapioides Rothschild had previously been put forward as the offspring of these putative parents, and to account for the differences in plumage Stresemann proposed that ellioti be considered the reciprocal or reversed cross to astrapioides. Rothschild, in Stresemann (1930), indicated that he did not find this solution convincing and appended his own set of suggestions to Stresemann’s paper . . . It seems extraordinary that such conclusions have not been challenged. Two taxonomists, both with considerable experience of the Paradisaeidae, were unable to agree on the ancestry of one form. Fuller and other authorities on the subject saw no reason to doubt its validity as a species. There is a distinct possibility that Elliot’s Bird of Paradise is either rare or extinct, occurring in a restricted montane range in New Guinea. Until more specimens come to light or perhaps mitochondrial DNA analysis reveals its taxonomic status, the bird must remain an enigmatic and hypothetical species. 342

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False-lobed Astrapia  Pseudastrapia lobata

Rothschild

Pseudastrapia lobata Rothschild 1907b, p. 25. (Dutch New Guinea = western New Guinea) Specimens  A specimen is in New York. Status  Possibly a valid rare or extinct species. Known only from the type. Range  ­North-­west New Guinea. Description  54 cm (21.25 in). Middle rectrices sharply pointed and elongate as in Epimachus; bill almost straight, as in Astrapia, Loborhamphus, and other genera, and not long and ­sickle-­shaped; feathering of forehead continues along bill and conceals nostrils; and, as in Loborhamphus, there is a curious fleshy lobe above and below angle of mouth. General coloration dull black; forehead with ­bottle-­green gloss; elongate middle rectrices with ­steel-­blue gloss, especially on the outer webs; bill, legs and feet black.

This mysterious form is known only from the type specimen, which bears no locality data other than ­‘north-­western New Guinea’. It is sometimes considered to be a hybrid ­Long-­tailed Paradigalla Paradigalla carunculata × Black Sicklebill Epimachus fastuosus, but it may be the immature form of Elliot’s Bird of Paradise Epimachus ellioti (Fuller 1995).

Bensbach’s Bird of Paradise  Janthothorax bensbachi

Büttikofer

Janthothorax Bensbachi Büttikofer 1895a, p. 163 (Arfak Mountains, New Guinea) Specimens  A specimen is in Leiden. Status  Possibly a valid rare or extinct species. Known only from the type, collected before 1895. Range  Arfak Mountains, New Guinea. Description  33 cm (13 in). Head metallic green; hind neck, back, rump and upperwing- and ­tail-­coverts ­velvet-­black with a purplish gloss, shining green under a certain light, many of the feathers broadly tipped ochraceous brown; quills ­earth-­brown, faintly glossed ­steel-­green; tail ­earth-­brown; central pair of ­tail-­feathers nearly double length of outer pairs, ­velvet-­black and richly glossed metallic green; chin glossy bluish green, upper throat metallic green; entire chest and breast purplish blue, ­underwing-­coverts, abdomen, vent and ­undertail-­coverts ­earth-­brown; from flanks a rich tuft of straight feathers, ­earth-­brown basally, sooty brown on terminal half; iris red; bill and feet black.

The type specimen of Janthothorax bensbachi was presented to Leiden Museum by a Mr. Bensbach, a Dutch resident on Ternate in the Moluccas. Stresemann (1930) considered it to be a hybrid of Lesser Bird of Paradise Paradisaea minor and Magnificent Riflebird Ptiloris magnificus, and briefly remarked: One can . . . remain in no doubt that it is the hybrid of Paradisaea minor and Ptiloris magnificus . . . Completely blackened undersides and the quality of shimmer in various parts of the plumage exclude Seleucidis ­[Twelve-­wired Bird of Paradise] and point towards Ptiloris [Magnificent Riflebird]. Although Frith & Frith (2010) agreed with Streseman, Fuller (1979) considered the bird to be unlikely to be a hybrid between these genera, and later (Fuller 1995) elaborated on this: Possibly Bensbach’s Bird of Paradise is exactly what Stresemann believed it to be, but his cryptic explanation is unsatisfactory and its virtually unquestioned acceptance is perplexing. Quite why the blackish underparts should point to a riflebird is unclear; no less than ­twenty-­one bird of paradise species show this particular feature. Stresemann’s vague remark on the ‘quality of shimmer’ is no more helpful, one of the trademarks of birds of paradise being their bewildering array of shimmers, glosses, sheens and iridescent glows. The status of this bird still remains unclear. It may be a hybrid, or a rare or extinct species, but the fact that many feathers are tipped ­ochre-­brown suggest the single specimen has not quite assumed the plumage of the adult bird.

Rothschild’s Lobe-billed Bird of Paradise  Loboramphus nobilis

Rothschild

Loboramphus nobilis Rothschild 1901, p. 34 (Dutch New Guinea) Specimens  Specimens are in New York. Status  Possibly a valid rare or extinct species. Known from two specimens, the last taken before 1907. Range  Western New Guinea. Description  30 cm (12 in). Crown purple; neck, back, and rump ­velvet-­black, with a bronze gloss; wings and tail black, with a purple sheen on the exposed webs; sides of neck and head ­copper-­bronze; chin and throat dark ­bronze-­green; pectoral shield

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Extinct Birds glittering purple, with metallic blue reflexions, some of the feathers of the lateral tufts have a metallic blue border; below pectoral shield an ­ill-­defined ­bronze-­green band; rest of underparts black washed purple; bill and legs black; two yellow fleshy folds form wattles around basal third of bill.

This curious bird was described by Rothschild (1901) at a meeting of the British Ornithologists’ Club on December 18, 1901. It was sent to him by Renesse van Duivenbode from an unspecified locality in Dutch New Guinea, and Rothschild set up the genus Loboramphus to accommodate his new species. At a subsequent meeting of the club in November 1907, Rothschild (1907b) exhibited a second specimen. Stresemann (1930) supposed that this bird was a hybrid between a ­Long-­tailed Paradigalla Paradigalla carunculata and a ­Twelve-­wired Bird of Paradise Seleucides melanoleuca, but it has nothing in common with the latter, and only the wattles in common with the former. Stresemann’s reasoning for referring to many enigmatic birds of paradise as hybrids is justifiable, but this bird remains one of the few that cannot be easily explained away. It seems doubtful that it is a hybrid, and it is possible that it is a rare or extinct species. Fuller (1995) agreed that Stresemann’s suggested parents are most unlikely.

Sharpe’s Lobe-billed Bird of Paradise  Loborhamphus ptilorhis

Sharpe

Loborhamphus ptilorhis Sharpe 1908, p. 67. (Dutch New Guinea) Specimens  There is a specimen in Tring. Status  Possibly a valid rare or extinct species. Known only from the type, collected prior to 1908. Range  Western New Guinea. Description  33 cm (13 in). General coloration ­velvet-­black, glossed with coppery purple; ­breast-­plate iridescent pink, turquoise and violet covering the breast; crown metallic violet; a tuft of black feathers above bill with patch of iridescent green behind; chin dark green; flanks and abdomen barred chocolate and cream; small fleshy wattle at gape.

This alleged species is known from a single specimen at Tring, purchased from one D. K. Dunstall. It may be a hybrid between two ­well-­known species: the Long-tailed Paradigalla Paradigalla carunculatus and the Arfak Parotia Parotia sefilata have been suggested (Stresemann 1930; Frith & Frith 2010). The suggested parentage is problematic, because the two suggested parents are not particularly close and are thus unlikely to hybridise, so it is just as likely to be an excessively rare or perhaps extinct species. This bird remains a mystery.

Ruys’s Bird of Paradise  Neoparadisea ruysi

Van Oort

Neoparadisea ruysi Van Oort 1906, p. 129 (near Warsembo, west coast of Geelvink Bay, New Guinea) Specimens  A specimen is in Leiden. Status  Possibly a valid rare or extinct species. Known only from the type, a male, collected in 1905. Range  Geelvink Bay, western New Guinea. Description  29 cm (11.5 in). Head and upper neck brownish yellow with greenish black margins to feathers becoming more ­orange-­brown on lower neck; forehead greenish black; chin and throat black with ­purple-­green gloss; lower neck ­blackish-­brown with ­purplish-­blue gloss; breast deep ­brownish-­purple; wings brown with purplish blue gloss; abdomen and ­undertail-­coverts ­blackish-­brown, the latter with lighter centres; tuft of loose feathers on flanks sooty brown; rump and ­uppertail-­coverts ­olive-­brown; tail brown with purplish gloss; legs dull ­bluish-­black.

A rather obscure character named H. H. Ruys spent four years on the ­north-­western coast of New Guinea in the early years of the 20th century. During this time he obtained the skin of a bird that was collected in August or September 1905 by Papuan hunters. It was sent to Leiden where it was described by van Oort (1906). Stresemann (1930; followed by Frith & Frith 2010) believed this specimen to be a hybrid between the Lesser Bird of Paradise Paradisea minor and the Magnificent Bird of Paradise Diphyllodes magnificus. However, this may be unrealistic. Fuller (1979) considered that it was not likely to be a hybrid of these species, remarking: The probability of the Lesser Bird of Paradise mating with the much smaller Magnificent does not . . . seem very great, particularly as the smaller species has evolved a breeding system that in certain respects approaches that of the bowerbirds. It is possible, therefore, that this specimen may represent a very rare or extinct species.

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Troglodytidae (Wrens) Daitō Wren  Troglodytes troglodytes orii

Yamashina

Troglodytes troglodytes orii Yamashina 1938b, p. 227 (Minami ­Daito-­shima, Borodino Islands) Specimens  The type specimen is in Tokyo. Status  Extinct. Not seen since 1938 and known only from the type. Range  Confined to Minamidaitō, in the Daitō (or Borodino) Islands, Japan. Description  10 cm (4 in). The type of the Daitō Wren is very similar to the Izu Wren T. t. mosukei from the southern Izu Islands.

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.

Timaliidae (Babblers) Astley’s Leiothrix  Leiothrix (lutea) astleyi

Delacour

Liothrix [sic] astleyi Delacour 1921, p. 115 (‘China’) Specimens  Specimens are in Tring. Status  Possibly extinct. Known only from aviary specimens of unknown origin. Range  China. Description  15cm (6 in). Male similar to male of L. lutea but with forehead, crown and eyebrows strongly tinged with orangescarlet; ear-coverts (which are plain grey in L. lutea) bright orange-scarlet; breast scarlet. Female paler; ear-coverts yellowish-orange.

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 specimens were obtained. If this is a genuine taxon, it may have disappeared through ­over-­trapping.

Fringillidae (Finches) Sillem’s Mountain Finch  Leucosticte sillemi

Roselaar

Leucosticte sillemi Roselaar 1992, p. 226 (southern Xinjiang, China) Specimens  Specimens are in Amsterdam. Status  Known from an adult male and juvenile collected on September 7, 1929. Range  Southern Xinjiang, China. Description  18 cm (7.25 in). A rather drab rosefinch. Generally ­greyish-­brown above, more rufous on head, paler below; bill ­blackish-­brown; legs and feet blackish.

Sillem’s Mountain Finch is sometimes listed as an extinct species, but it may be an aberrant Brandt’s Mountain Finch L. brandti, a hybrid, or a vagrant from an ­as-­yet undiscovered locality. It was described from two specimens collected on September 7, 1929 on a plateau at 5,125m at Kushku Maidan, in southern 345

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Xinjiang, China, and has never been recorded since. As one of the specimens was a juvenile with the wings not fully grown, it has been suggested that the birds were breeding near to where they were collected or possibly on peaks of the nearby Kunlan Shan range (BirdLife International 2011). No other information is available.

Cardinalidae (Cardinals) Townsend’s Bunting  Spiza townsendi

(Audubon)

Emberiza townsendii (later amended to townsendi) Audubon 1834, p. 183 (New Garden, Chester County, Pennsylvania) Specimens  A specimen is in Washington, D. C. Status  Known only from the type and description of Audubon. Considered by some to be an extinct species, but almost certainly a colour variant of the Dicksissel Spiza americana, or (less likely) a hybrid. Range  Chester County, Pennsylvania. Description  16.5 cm (6.5 in). The upperparts, head, neck, sides and forepart of breast ­slate-­blue; back and upper parts of wings tinged with ­yellowish-­brown, streaked black on the interscapular area; ­eye-­stripe, chin, throat, central line of underparts and edge of wing white; ­black-­spotted line from lower corner of mandible down the side of throat, connecting with a crescent of streaks on upper edge of ­slate-­blue breast.

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 ­colour-­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 specimen (see Fuller 2000), contrive to give a more distinctive plumage than the specimen actually merits.

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Appendix 1 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.

Bornean Baillon’s Crake  Porzana pusilla mira

Riley

Porzana pusilla mira J. H. Riley 1938, p.95 (Tanggorong, Mahakkam River, east Borneo) Specimens  The type is in Washington, D. C. Status  Data deficient. Possibly extinct. Known only from the type specimen, an adult female, collected in 1912. Range  Borneo. Description  17 cm (6.75 in). Baillon’s Crake is a rich brown above with black streaks bearing numerous white markings, and pale grey below. The iris is bright red and the legs and feet greenish. The Bornean bird was the smallest race and most similar to nominate P. p. pusilla, but it differed in having the brown of the upperparts more rufous; supercilium and cheeks lighter grey; throat and breast white; chest very pale grey; bill olive-yellow, base and culmen ­medal-­bronze not olivaceous black.

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.

Sinú ­Brown-­throated Conure  Aratinga pertinax griseipecta

Meyer de Schauensee

Aratinga pertinax griseipecta Meyer de Schauensee 1950, p.6 (Tierra Alta, Bolivar, Colombia) Specimens  Specimens are in Philadelphia and Washington, D. C. Status  Data deficient. Possibly extinct. Known only from two specimens collected in 1949. Range  Sinú valley, Córdoba, ­north-­east Colombia. Description  25 cm (10 in). Generally green; crown ­greenish-­blue; cheeks, throat and upper breast ­olive-­grey merging into green of lower breast, rest of underparts paler ­greenish-­yellow; forehead, lores and sides of head ­orange-­yellow; throat and upper breast pale ­olive-­brown; outer webs of primary feathers green, dull blue at tips; tail green with ­blue-­green tip; bill ­greyish-­black; bare skin around eye ­buff-­white; iris yellow; legs and feet grey.

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. 347

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Extinct Birds

Red-­throated Lorikeet  Charmosyna amabilis

(Ramsey)

Trichoglossus amabilis Ramsey 1875 (Ovalau, Fiji) Specimens  Specimens are in Delaware; Fiji; Liverpool; Melbourne; Philadelphia; Sydney; Tring; Vienna; and Washington, D. C. Status  Data deficient. Possibly extinct. Last confirmed sighting in August 1993. Range  Viti Levu, Vanua Levu, Taveuni and Ovalau, Fiji. Description  18 cm (7 in). Uniform green except for red cheeks, throat and thighs; red throat bordered by yellow on upper breast; undertail and tips of tail ­mustard-­yellow; bill legs and feet orange; iris yellow. Immature birds duller; yellow bordering on throat reduced; and thighs dull ­greyish-­mauve tinged with red.

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. 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 ­old-­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, ­high-­pitched squeaks (Forshaw 1989).

Bahia ­Rufous-­vented Ground Cuckoo  Neomorphus geoffroyi maximiliani

Pinto

Neomorphus geoffroyi maximiliani Pinto 1962, pp. 299–301 (Rio Gongogi, Bahia, Brazil) Specimens  The type specimen is in São Paolo. Range  Atlantic forests of southern Bahia, ­north-­eastern Brazil. Status  Data deficient. Possibly extinct. Not seen since the 1950s. Description  50cm (16 in). Adult dark bronze green above; crest black; forehead buff; tail purplish; breast with semicircular bands; narrow black band on chest; underparts buff with rufous vent; bare facial skin blue; bill yellowish green; iris brown; feet and legs dark grey.

Little was recorded about the Bahia ­Rufous-­vented Ground Cuckoo. The species occurs in lowland evergreen 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 ­Rufous-­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. 348

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Appendix 1: Data-deficient taxa

Greater Crested Coua  Coua (cristata) maxima

Milon

Coua cristata maxima Milon 1950, p.65 (Tolagnaro, ­south-­east Madagascar) Specimens  The type specimen is in Paris. Status  Data deficient. Probably extinct. Known from one specimen taken in 1950. Range  South-­east Madagascar, near Tolagnaro. Description  42cm (16.5 in). Upperparts greenish grey; back greyish blue; crest grey; innermost secondaries distinctly blue; uppertail intense ­violet-­blue; throat ­bluish-­grey; lower breast tawny brown merging with darker tawny brown on upper breast; belly and ­undertail-­coverts white.

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 cristata 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.

Peleng Masked Owl  Tyto rosenbergii pelengensis

Neumann

Tyto rosenbergii pelengensis Neumann 1939, p. 92 (Peleng Island, Indonesia) Specimens  The type specimen is in Cambridge, Massachusetts. Status  Data deficient. Possibly extinct. Known from one specimen, taken in 1938. Range  Peleng Island, Indonesia. Description  43–46cm (17–18 in). Differed from nominate Tyto rosenbergi rosenbergi in being smaller; feathers of lower throat and upper breast with ­sub-­terminal incomplete black edges, giving a scaled appearance; otherwise similar.

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).

Buru Masked Owl  Tyto sororcula cayelii

(Hartert)

Strix cayelii Hartert 1900, p. 228 (Buru, Indonesia) Specimens  The type specimen is in New York. Status  Data deficient. Possibly extinct. Last recorded in 1921. Range  Buru, Indonesia. Description  21–31 cm (8.25–12.25 in). Similar to nominate Lesser Masked Owl Tyto sororcula, but slightly larger; more ­tawny-­buff; upperparts more ­blackish-­brown, white mottlings replaced by brownish ones, except on middle and greater ­wing-­coverts; tail ­yellowish-­brown, narrowly barred black.

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.

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Extinct Birds

Natuna Bay Owl  Phodilus badius arixuthus

Oberholser

Phodilus badius arixuthus Oberholser 1932, p. 40 (Bunguran Island, north Natuna Islands) Specimens  The type specimen is in Washington, D. C. Status  Data deficient. Possibly extinct. Known only from the type. Range  Bunguran Island, Natuna Islands, Borneo. Description  23–33 cm (9–12 in). Forehead white washed russet; crown, occiput and nape tawny with a few pale feathers on occiput; rest of upperparts tawny with a few brown and large white spots; tail russet, inner webs barred ­blackish-­brown; primaries light russet with buff tips and barred inner webs; orbital ring tawny with facial ruff silvery white, washed with tawny; chin and throat white, slightly tinged tawny; underside ­cinnamon-­buff sparingly marked with small brown spots.

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. arixuthus of Bunguran Island in the Natuna archipelago (Bruce 1999). The reasons for the possible disappearance of the Natuna subspecies are unclear, but owls are notoriously cryptic, and Bunguran is not commonly visited by ornithologists, so a population may still survive.

Siau Scops Owl  Otus manadensis siaoensis

(Schlegel)

Scops siaoensis Schlegel 1873b, p. 13 (Siau, Indonesia) Specimens  The type specimen is in Leiden. Status  Data deficient. Possibly extinct. Not recorded since 1866. Range  Siau Island, Indonesia. Description  17cm (6.5 in). A small, eared owl. Generally dark brown above, pale below, with dark and light streaks; wings barred brown and white; face whitish. Much smaller than the nominate, with much smaller ­ear-­tufts; wings and tail shorter with narrower barring on wings.

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).

Virgin Islands Screech Owl  Megascops nudipes newtoni

(Lawrence)

Gymnoglaux newtoni Lawrence 1860, p. 259 (St. Croix) Specimens  A specimen is in New York. Status  Data deficient. Probably extinct, but it may survive on Guana Island, British Virgin Islands. Range  Vieques, Culebra, St Thomas, St Kitts and St. Croix, Virgin Islands, West Indies. Description  23–25 cm (9–10 in). Generally ­rufous-­brown above; underside ­light-­brownish to white, with thin brown bars; eyebrows white; lores and chin whitish; bill ­greenish-­yellow; iris ­reddish-­brown; feet yellow. More ­rufous-­brown above than nominate; also narrower wavy dark brown bars on neck and upper back; underparts of nominate more uniform.

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. Habits This owl preferred forests with large hollowed trees, and would call ­year-­round, mostly at dawn, and from dense foliage. Its local name in the Virgin Islands is ­‘cuckoo-­bird’, derived from the loud ­coo-­coo call. The diet comprised mainly large insects, particularly orthopterans, but also small birds, geckos and small rodents. 350

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Cayenne Nightjar  Antrostomus maculosus

(Todd)

Nyctipolus maculosus W. E. Todd 1920, p. 74 (Tamanoir, Cayenne) Specimens  The type specimen is in Pittsburgh. Status  Data deficient. Possibly extinct. Known only from the type, a male, collected in 1917. Range ­North-­west French Guiana. Description  23cm (9 in). A dark nightjar, the primaries black with a white spot on the inner web of the four outermost; outermost ­tail-­feather virtually uniform, next two tipped white; upperparts ­greyish-­brown, broadly streaked ­blackish-­brown on crown and nape; narrow, indistinct tawny collar; ­wing-­coverts ­greyish-­brown, heavily spotted with buff; scapulars ­blackish-­brown broadly edged with buff; large white patch on either side of throat; underparts buff, heavily barred with brown.

Little is known about the origin of the sole specimen of this species. Todd (1920) says merely that exploration 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.

Itombwe Nightjar (Prigogine’s Nightjar)  Caprimulgus prigoginei

Louette

Caprimulgus prigoginei Louette 1990, p. 349 (Maleuge, DR Congo) Specimens  The type is in Tervuren, Belgium. Status  Data deficient. Possibly extinct. Known from a single specimen collected in 1955. Range  Malenge, Itombwe Mountains, DR Congo, and possibly Congo Brazzaville, ­north-­east Gabon, ­south-­east Equatorial Guinea and ­south-­east Cameroon. Description  19 cm (7.5 in). Small brown nightjar. Generally brown variegated with black and buff; ­wing-­coverts spotted buff; narrow white tips to two outermost tail feathers; bill ­blackish-­brown; legs and feet brown.

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 indicates 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 agriculture and for livestock grazing makes the ­long-­term survival of this elusive species doubtful.

Nechisar Nightjar  Caprimulgus solala

Safford et al.

Caprimulgus solala Safford et al. 1995, p. 301 (Nechisar Plains, south Ethiopia) Specimens  The type specimen is in Tring. Status  Data deficient. Probably extant. Known from a single specimen (a wing from a road-kill). Range  Nechisar Plains, south Ethiopia. Description  Length unknown. Probably a large, variegated brown bird. ­Wing-­coverts brown, spotted buff; primaries and secondaries barred with black, buff and grey; a broad white band on four outermost primaries toward tip; and white tips to at least two outermost tail feathers.

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.

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Vaurie’s Nightjar  Caprimulgus centralasicus

Vaurie

Caprimulgus centralasicus Vaurie 1960a, p. 1, figs 1 & 2 (western Xinjiang, West China) Specimens  The type specimen is in Tring. Status  Data deficient. Probably extant. Known from one specimen taken in 1929. Range  Xinjiang, western China. Description  19 cm (7.5 in). This is a generally ­sandy-­buff nightjar: lacks collar around ­hind-­neck and white markings in wings; throat patches ­buffish-­white; two outermost tail feathers with ­buffish-­white tips.

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.

New Caledonia ­Owlet-­nightjar  Aegotheles savesi

Layard & Layard

Ægotheles savesi Layard & Layard 1881, p. 132, pl. 5 (Tongué, near Noumea, New Caledonia) Specimens  Specimens are in Liverpool and Rome. Status  Data deficient. Probably extinct, but there have been recent possible sight records. Range ­South-­west New Caledonia. Description  28cm. This was a large and very distinctive owlet-nightjar with short rounded wings, long, slightly rounded tail and relatively long, stout legs; general coloration a uniform brownish black, each feather having several irregular transverse bars of ­greyish-­white; bill black; iris ­sienna-­yellow; legs and feet dark brownish yellow.

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 colonists (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. 203).

Malaita Variable Kingfisher  Ceyx lepidus mal

Mayr

Ceyx lepidus malaitae Mayr 1935, p. 2 (Malaita, East Solomons) Specimens  The type specimen is in New York. Status  Data deficient. Probably extant. Range ­Malaita, Solomon Islands, western Pacific. Description  28cm. 14 cm (5.5 in). Head and wings blackish spotted with dark blue; white bar on side of neck; dazzling methylblue back; rump silvery blue; chin and throat buffy white; rest of underparts yellow ochre; upper bill black, lower bill orange-red; iris dark brown; legs and feet orange or orange-yellow.Nominate C. l. lepidus darker with ochraceous orange underparts and deep purplish-blue back.

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 difficult 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). 352

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Luzon Whitehead’s Swiftlet  Aerodramus whiteheadi whiteheadi

(Ogilvie-­Grant)

Collocalia whiteheadi ­Ogilvie-­Grant 1895, p. 459 (Mount Data, Luzon, north Philippines) Specimens  The type specimen is in Tring. Status  Data deficient. Possibly extinct. Not recorded since 1895. Range  Mount Data, Luzon, Philippines. Description  14cm (in). Generally ­greyish-­brown in coloration; upperparts sooty; lacked white on abdomen and greyish band across rump compared with other ­South-­east Asian Aerodramus swiftlets; tail distinctly forked. Luzon race differs from A. w. origenis in being lighter above and below.

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 ­cliff-­faces (BirdLife International 2011). A nest of the Luzon race was taken in July 1895; otherwise nothing is known of these birds.

Sierra Leone Black Swift Apus barbatus glanvillei 

Benson

Apus barbatus glanvillei Benson 1967, p. 125. (Great Scarcies River, Rokupr, Sierra Leone) Specimens  The type specimen is in Tring. Status  Data deficient. Possibly extinct. Range  Sierra Leone. Description  16–18cm (6.5–7 in). A robust bird, almost uniform ­blackish-­brown, except for a small white or pale grey patch on chin. Slightly darker than the nominate Black Swift A. b. barbatus.

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, 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.

Turquoise-­throated Puffleg  Eriocnemis godini

Bourcier

Eriocnemis godini Bourcier 1851, p. 96 (Río Guaillabamba, south of Perucho, Pichincha, north Ecuador) Specimens  Specimens are in Tring. Status  Data deficient. Probably extinct. Known from six ­19th-­century specimens, and an unconfirmed sighting in 1976. Range  The type was collected in ravines of the Río Guaillabamba, south of Perucho, Pichincha, north Ecuador. Description  10–11cm (4–4.25 in). This was a predominantly green hummingbird with ­violet-­blue ­undertail-­coverts. Male upperparts and most of underparts shining golden green; rump, ­uppertail-­coverts bluish green; throat pale blue; ­undertail-­coverts ­violet-­blue; tail bluish black; bill black. Female plumage generally duller, more golden on belly; lacking ­violet-­blue throat patch.

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 353

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unconfirmed 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).

White-­chested Tinkerbird  Pogoniulus makawai

Benson & Irwin

Pogoniulus makawai Benson & Irwin 1965, p. 6 (Mayau, ­north-­west Zambia) Specimens  The type specimen is in Tring. Status  Data deficient. Possibly extinct. Known only from the type. Range  Mayau, ­north-­west Zambia. Description  11cm (4.25 in). Upperparts from crown to back ­bluish-­black; chin black, flecked centrally white; ­wing-­coverts and secondaries black with narrow yellow edges; throat and upper chest creamy white, fading to pale yellow on lower chest; central belly black; bill whitish at base, otherwise black; legs and feet pale brown.

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 ­north-­west Zambia and neighbouring Angola. The bird has a large, comparatively robust bill, which may be an adaptation for feeding on the berries of mistletoes (Collar & Fishpool 2006).

Kinglet Calyptura (Kinglet Cotinga)  Calyptura cristata

(Vieillot)

Pardalotus cristatus Vieillot 1818, p. 528 (Rio de Janeiro, Brazil) Specimens  Specimens are in Berlin; Brussels; Cambridge, England; Cambridge, Massachusetts; Copenhagen; Florence; Kiel, Germany; Liverpool; New York; Paris; Philadelphia; Tring; Vienna; and Washington, D. C. Status  Data deficient. Possibly extinct, but may survive in small numbers. Not recorded since 1996. Range  Restricted to Rio de Janeiro, Brazil, but may have occurred in southern Espirito Santo. Description  8 cm (3 in). Male generally bright ­olive-­green above with bright yellow rump; forehead yellow; crown bright ­orange-­red enclosed by broad black lateral crown stripes; wings brown with white tips on median and greater ­wing-­coverts; throat, sides of face and undersides olive, becoming yellower from upper breast to ­undertail-­coverts; bill, legs and feet dark. Female duller, lacking yellow forehead, red of crown paler and more reduced.

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 354

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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 actually 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).

Archer’s Lark Heteromirafra archeri Clark Heteromirafra archeri Clark 1920, 40, p. 64 (Jifa, western frontier of British Somaliland) Specimens  The type is in Tring. Status  Data deficient. Last recorded in 1955, but is likely to survive. Range  Wajaale plains, Somalia. Description  14 cm (5.5 in). Upperparts brown with dark edges to feathers producing a scaled effect; underparts ­buff-­brown, heavily streaked dark brown on breast; tail brown with white outer tail feathers; bill pale greyish; legs and feet pale ­yellowish-­buff; iris blackish.

Archer’s Lark was first collected on September 23, 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 Waajale clay plains from Jifa Medir to Ban Wujaleh near Hargeisa, north-west Somalia, with the last record within the type locality in 1922. However, a small population was reported in 1955, 100km to the north-west at Buramo (Collar & Stuart 1985), but it has not been seen since. Surveys in the type locality between the 1970s and 2010 have failed to locate the population, but an unconfirmed report was made in 2003, and a further report was made in 2004 from neighbouring eastern Ethiopia (BirdLife International 2011). The bird’s decline was due to human habitat disturbance, and more recently the spread of invasive weeds, especially from the noxious American weed Parthenium hysterophorus. Archer’s Lark is a secretive bird that is difficult to locate, so there is a good possibility that it still survives. Habits This lark occurred in open grasslands and open rocky areas with scattered bush and grass cover, where it skulked in dense vegetation, never flying across open ground, even when pressed (Ash & Miskell 1998). Breeding presumably began in April or May, as nests were recorded in June. The nest was ­funnel-­shaped, extending deep into a grass clump, and where three eggs were laid (Archer & Godman 1961). Nothing else is known about the bird’s habits.

Red Sea Swallow Hirundo perdita

Fry & Smith

Hirundo perdita Fry & Smith 1985, Ibis 127, p. 2 (Sanganeb lighthouse, north-east of Port Sudan, Sudan) Specimens  The type is in Tring. Status  Data deficient. Probably extant. Known only from the type found dead in May 1984 at Sanganeb lighthouse, ­north-­east of Port Sudan, Sudan. Range  Type locality Port Sudan, Sudan. Description  14cm (5.5 in). Forehead, lores, crown, nape, sides of neck, mantle and wings dark ­blackish-­brown with dark ­steel-­blue gloss; ­uppertail-­coverts dark glossy brown; throat and upper breast ­bluish-­black; lower breast and belly ­silky-­white; vent ­pale-­grey, strongly washed pale orange; bill, legs and feet black.

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 355

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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.

White-­eyed River Martin  Eurochelidon sirintarae

(Kitti)

Pseudochelidon sirintarae Kitti 1968, 1, p. 1–10 (Bung Boraphet, Amphoe Muang, Nakhon Sawan Province, central Thailand) Specimens  The type is in Bangkhen, Thailand. Status  Data deficient, possibly extinct. Last recorded February 3, 1978. Range  Winters in central Thailand, summer distribution unknown. Description  15 cm (5.75 in). Generally uniform blackish with ­blue-­green gloss; wings black; rump ­silvery-­white; tail black with green gloss, each feather tipped pale brown; two central tail feathers form long, narrow racquets; bill bright ­green-­yellow, black at tip of upper bill; iris and edges of ­eye-­lids white; legs and feet ­flesh-­coloured. Juvenile browner, with paler throat, and lacks tail streamers.

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 juveniles 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 crepuscular, 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).

Sangihe ­White-­eye  Zosterops nehrkorni

Blasius

Zosterops nehrkorni Blasius 1888a, p. 85 (near Manganitu, Sangihe, Indonesia) Specimens  The type is in Braunschweig. Status  Data deficient, possibly extinct. Not recorded since 1999. Range  Restricted to Sangihe Island, northern Indonesia. Description  12 cm (4.75 in). Forehead black; broad white ­eye-­rings; upperparts ­olive-­green, with contrasting ­yellow-­green rump; wings ­olive-­green, primaries ­blackish-­green edged with olive; tail ­greenish-­black; chin, throat and ­undertail-­coverts bright yellow; rest of underparts white with grey flanks; bill, upper bill dark brown, lower bill light brown; iris ­red-­brown; legs and feet light brown.

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. 356

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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 ­White-­eye Z. atrifrons of Sulawesi (Riley 1997).

Colombian Whiskered Wren  Thryothorus mystacalis macrurus

(Allen)

Thryothorus macrurus Allen 1889, p. 137 (Vicinity of Bogota, Colombia) Specimens  Specimens are in Washington, D. C. Status  Data deficient. Possibly extinct. Last recorded in the late 20th century. Range  East slope of central Andes and western slope of east Andes, Colombia. Description  16 cm (6.5 in). Nominate has ­greyish-­white supercilium from bill to above ­ear-­coverts; dull black ­ear-­coverts speckled white; lore dull black; crown greyish; back, shoulders and rump bright chestnut; primaries and secondaries ­blackish-­grey; chin and throat dull white; black malar strip edged white; chest grey becoming more olivaceous on belly; upper bill black, lower grey or ­greyish-­brown; iris brown; legs and feet grey. Race T. m. macrurus differs in having tail dingy brown with longitudinal barring on the rectrices.

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).

Rueck’s Blue Flycatcher  Cyornis ruckii

(Oustalet)

Siphia Ruckii Oustalet 1881, p. 78 (Malacca, Malaysia = Sumatra) Specimens  Specimens are in Leiden and Paris. Status  Data deficient. Last collected February 1918. Locality probably in error as two of the four known specimens were almost certainly derived as trade imports from Sumatra. Range  Possibly Sumatra, Indonesia. Description  17cm (6.75 in). Male uniform dark blue, paler on forehead, rump and lower breast; belly white; rump iridescent blue; bill, legs and feet black; iris brown. Female ­rufous-­brown; breast, rump and tail rusty; throat and belly ­buff-­white. Juvenile almost uniform brown, spotted with buff; breast rufous; whitish centres on underparts.

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).

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Black-­lored Waxbill  Estrilda nigriloris

Chapin

Estrilda nigriloris Chapin 1928, p. 1 (Lualaba River, DR Congo) Specimens  Specimens are in Tervuren, Belgium; Brussels; and New York. Status  Data deficient. Possibly extinct. Last collected in 1950. Range  Lualaba River and Lake Upemba, DR Congo. Description  11–13cm (4.25–5 in). Adult male generally ­grey-­brown with hindcrown, upperparts, sides of chest, and flanks barred with dark brown; abdomen and ­undertail-­coverts black; loral region and small area behind eye black, extending for 4mm, with streak of pale pinkish or whitish above. Along the upper loral region runs a streak of pale pinkish or whitish; bill dull scarlet; iris very dark brown; feet dusky brown. Adult female similar but duller; black of face almost entirely restricted to lores, and a faint black streak behind eye.

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).

Short-­toed Nuthatch Vanga (Blüntschli’s Vanga)  Hypositta perdita

Peters

Hypositta perdita Peters 1996, p. 8 (Fort Dauphin, Madagascar) Status  Data deficient. Possibly extinct. Range ­South-­eastern Madagascar.

The ­Short-­toed Nuthatch Vanga is known from two recently fledged specimens, collected in 1931 in primary forest and grassland near Eminiminy, a small village situated north of Fort Dauphin (now Taolanaro) in ­south-­east Madagascar (Peters 1996; Hawkins & Goodman 1999). It differs from the similar Nuthatch Vanga H. corallirostris, which occurs in the same area, in having longer tarsi and shorter toes. Extensive field studies in the type locality have failed to relocate the birds. Peters (1996) suggested that its foot morphology pointed towards the bird being a forest- and ­scrub-­inhabiting species, but not specialised in climbing on tree trunks and branches. This implies that the ­Short-­toed Nuthatch Vanga and the Nuthatch Vanga were adapted to different ecological niches and could therefore share the same habitat (Peters 1996; Hawkins & Goodman 1999). No more recent information is available, but if the ­Short-­toed Nuthatch Vanga still survives, its numbers must be perilously low (see BirdLife International 2011).

Hooded Seedeater  Sporophila melanops

(Pelzeln)

Spermophila melanops Pelzeln 1870, p. 224, 331 (based on Natterer’s MS) (Registro do Araguaia, Brazil) Specimens  The type specimen is in Vienna. Status  Data deficient. Possibly extinct. Known only from the type, collected in 1823. Range  South-­central Brazil. Description  11cm (4.25 in). Adult male upperparts olive; underparts dingy buff; ­undertail-­coverts whitish; head and throat black, forming a hood; bill ­bluish-­black; legs and feet black.

The Hooded Seedeater is an enigmatic bird. 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 (BirdLife International 2011), it is sometimes considered to be an aberrant or hybrid ­Yellow-­bellied Seedeater S. nigricollis. This bird may have been a victim of deforestation, but despite surveys that failed to locate the species, the surrounding forests are poorly surveyed. Nothing else is known about it.

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Appendix 2 Doubtful and invalid taxa

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 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 genuinely extinct species, but their correct status can now never be determined.

Kalinowski’s Tinamou  Nothoprocta kalinowskii

Berlepsch & Stolzmann

Nothoprocta kalinowskii Berlepsch & Stolzmann 1901, p. 192 (Licamachay, Cuzco, Peru) Status  Invalid taxon. Range ­North-­central and ­south-­east Peru.

Now considered a synonym of Ornate Tinamou Nothoprocta ornata branickii (Krabbe & Schulenberg 2005).

Pernambuco Solitary Tinamou  Tinamus solitarius pernambucensis

Berla

Tinamus solitarius pernambucensis Berla 1946, p.2 (município de Igaraçu, Pernambuco) Status  Invalid taxon. Range  East Brazil.

Considered a synonym of the nominate race of Solitary Tinamou Tinamus s. solitarius (do Amaral & Silveira 2004).

Levaillant’s Dwarf Ostrich  Struthio bidactylus

(Gray)

Petit Gralle bidactyle, Autruchon [small ­two-­toed plover, ostrich]. Temminck, fide G. R. Gray 1841 Charadrius bidactylus Gray 1847, p. 544 Status  Doubtful taxon. Putative range ­North-­east Africa.

Known only from hearsay comments by Sclater (1862) and Heuglin (1859) and an unpublished picture by Levaillant. Based on the available evidence, there is nothing to distinguish this bird from the North African Ostrich S. c. camelus or Syrian Ostrich S. camelus syriacus (see p. 18) except for its small size, so it may have been a juvenile of either of these forms. Another supposed subspecies, S. c. spatzi, was described by Stresemann (1926) from a series of eggs and chicks obtained at the Rio de Oro in the former Spanish Sahara (see Walters 1982). This subspecies should be regarded as a nomen dubium, as bird eggs and chicks lack diagnostic characters.Now considered a synonym of Ornate Tinamou Nothoprocta ornata branickii (Krabb & Schulenberg 2005).

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Extinct Birds

Dwarf Rhea  Rhea nana

Lydekker

Rhea nana Lydekker 1894a, p. 654 (No locality) Specimens  Known from an egg in the Museo de La Plata, from which a wax cast was produced. This is at Tring. Status  Invalid taxon. Putative range  Patagonia.

Described from a wax cast taken from an egg in the Museo de La Plata (Lydekker 1894b), which lacks diagnostic characters. This wax cast is at Tring.04).

Andaman Megapode  Megapodius andamanensis

Walters

Megapodius andamanensis Walters 1980, p. 33 (based on Rothschild’s manuscript) (no locality = Andaman Islands) Status  Doubtful taxon. Putative range  Andaman Islands.

May represent a ­now-­extinct species, but at present best regarded as a nomen dubium, mainly because the holotypical specimens (eggs only) lack determinable characters.

Burnaby’s Megapode  Megapodius burnabyi

Gray

Megapodius sp. Gray 1859a, p. 46 Megapodius burnabyi Gray 1861, p. 290 (Hapace Islands, near Tongataboo) Status  Doubtful taxon. Putative range  Samoa, South Pacific.

May represent a ­now-­extinct species, but at present best regarded as a nomen dubium, mainly because the holotypical specimens (eggs only) lack determinable characters.

Lord Howe Island Megapode  ? Megapodius sp. ? Peacock. Thomas Gilbert 1789, Voyage from New South Wales to Canton in the year 1788, p. 12. (Lord Howe Island). Megapodius sp. Seebohm c.1895 (Lord Howe Island). The eggs are no longer in Tring. Status  Invalid taxon. Putative range  Lord Howe Island.

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).

New Guinea Quail  Coturnix novaeguineae

(Gmelin)

La Caille de la Nouvelle Guinée. Sonnerat 1776, p. 170, pl. 105 New Guinea Quail. Latham 1783, p. 789 Oriolus cothurnix Scopoli 1786, p. 87 (based on Sonnerat) Tetrao novae-guineae Gmelin 1789, p. 764 (based on Sonnerat) Perdix novae-guineae Latham 1790, p. 655 Status  Invalid taxon. Putative range  New Guinea.

Known only from Sonnerat’s description, from which all others derive. Unidentifiable and now rejected (Mayr 1963).

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Appendix 2: Doubtful and invalid taxa

Florida Pheasant  ?Phasianus sp. ‘pheasant’ Stork 1767, p. 51. Status  Invalid taxon. Known only from the accounts of Stork (1767) and Le Conte (1854). Unidentifiable. Putative range  Florida.

Superb Pheasant  Phasianus superbus  Phasianus superbus Linnaeus 1767–71, p. 526 (? China) Status  Invalid taxon. Putative range  China.

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.

Italian Partridge  Perdix perdix italica

Hartert

Perdix perdix italica Hartert 1917, p.283 (Badia de Passignano, Italy) Status  Doubtful taxon. Putative range  Chianti, Italy.

Considered synonymous with nominate Grey Partridge Perdix perdix perdix (Violani et al. 1988; McGowan 1994).

Reichenow’s Quail  Lophortyx leucoprosopon

Reichenow

Lophortyx leucoprosopon Reichenow 1895, p. 11 (origin unknown; description based on a living pair in a private aviary, believed to have been bought from a sailor arriving at Hamburg, Germany. Status  Doubtful taxon. Range  Unknown.

Known only from the types. Believed to be a hybrid between Gambel’s Quail Lophortyx gambelii and Douglas’s Quail L. douglasii (Peters 1934).

Key West Bobwhite  Colinus virginianus insularis

Howe

Colinus virginianus insulans Howe 1904, p. 168 (Key West, Florida) Status  Doubtful taxon. Range  Population formerly occurred on Key West, and perhaps other islands in the Florida Keys.

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.

Snow’s Canada Goose  Branta canadensis ssp. Branta canadensis (? asiatica) Aldrich 1946b, p. 95 (Bering Island) Status  Doubtful taxon. Known only from eggs and immature birds. Synonymous with B. c. asiatica or B. c. hutchinsii. Putative range  Northern Kuril Islands.

Cream-coloured Goose  Anser lacteus

Latham

Cream Coloured Goose. Latham 1824, p. 296 (New South Wales, Australia) Status  Doubtful taxon. Known only from Latham’s inadequate description. Putative range  New South Wales.

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Extinct Birds

Great Goose  Anser grandis

(Gmelin)

Great Goose. Pennant 1785, p. 570 (the bird referred to as the Great Goose by Pallas 1771–76 does not refer to this) Great Goose. Latham 1785, p. 446 Anas grandis Gmelin 1789, p. 504 Status  Doubtful taxon. Known only from the inadequate account of P. S. Pallas. Putative range  Eastern Siberia.

David’s Swan  Cygnus davidi

Swinhoe

Cygnus (Coscoroba) davidi Swinhoe 1870, p. 430 (Tientsin, China) Status  Doubtful taxon. Known from a single specimen, now lost. Probably a leucistic Bewick’s Swan C. bewickii (Kear, in Scott 1972). Putative range  Tientsin, China.

Polish Swan  Cygnus immutabilis

Yarrell

Cygnus immutabilis Yarrell 1838, p. 19 (Medway, Kent) Status  Invalid taxon. Formerly believed to be a full species, but now considered a morph or mutation of the Mute Swan Cygnus olor (Kear, in Scott 1972). Putative range  Eurasia.

Gattair Duck  Anas gattair

Gmelin

Anas gattair Forskål 1775, p. 3 (Alexandria) Anas gattair Gmelin 1789, p. 542 (Alexandria, Egypt) Status  Invalid taxon. Known only from Forskål’s description. Synonymised under the Ferruginous Duck Aythya nyroca by Salvadori (1895). Putative range  Egypt.

Gmelin’s Wandering Duck  Anas peregrina

Gmelin

Anas lurida S. G. Gmelin 1770, p. 70 Anas peregrina S. G. Gmelin 1774, p. 183, tab. 16 Status  Invalid taxon. Known only from S. G. Gmelin’s description. Synonymised under the Ferruginous Duck Aythya nyroca by Salvadori (1895). Putative range  Southern Russia.

Lappmark Duck  Anas latirostra

Brünnich

Anas latirostra Brünnich 1764, p. 21 (Christiansöe) Status  Invalid taxon. Putative range  Lappmark and Denmark. Known from various early accounts. Synonymised under the the Ferruginous Duck Aythya nyroca by Salvadori (1895)

Gaulaund Duck  Mergus borealis

(Gmelin)

Gaulaund Duck. Pennant 1785, p. 572 Anas borealis Gmelin 1789, p. 512 Status  Doubtful taxon. Putative range  Iceland.

Known only from the inadequate descriptions of Pennant and Latham. Considered to be conspecific with the Common Eider Somateria mollissima by Salvadori (1895). 362

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Appendix 2: Doubtful and invalid taxa

Larsen’s Penguin  Eudyptes sp. Penguin sp. Donald 1894, p. 176 Eudyptes sp. Ogilvie-Grant 1898, p. 625 Status  Doubtful taxon. Putative range  South Orkneys, South Atlantic.

Known only from Donald’s account, but unidentifiable.

Commerson’s Penguin  Aptenodytes torquata

Forster

Le Manchot à Collier de la Nouvelle Guinée. Sonnerat 1776, p. 180, pl. 114 Aptenodytes torquata J. R. Forster 1781, p. 146 Collared Penguin. Latham 1785, p. 571 Apterodita platirhingos Scopoli 1786, p. 91 Chrysocoma torquata Stephens 1826, in Shaw’s General Zoology, 3, p. 60 Status  Doubtful taxon. Putative range  Unknown.

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.

Molina’s Penguin  Aptenodytes chilensis

(Molina)

Diomedea chilensis Molina 1782, pp. 238, 334 Aptenodytes chiloensis & A. chilensis Gmelin 1788, p. 559 (based on Molina) Pinguinus chilensis Bonnaterre 1790, p. 30 Aptenodytes molinae Latham 1790, p. 881 Three-toed Penguin. Latham 1824, p. 393 Status  Invalid taxon. Putative range  Chile.

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.

Diablotin  Pterodroma diabolica

(Lafresnaye)

Diabloton [sic] Attwood 1791, p. 30 (quoted in Carte 1866, p. 95) Procellaria diabolica L’Herminier 1844, p. 168 Status  Doubtful taxon. Putative range  Guadeloupe, perhaps also Dominica.

Considered to be either a dark morph of the ­Black-­capped Petrel Pterodroma hasitata, a population of the Jamaican Petrel Pterodroma caribbaea (see p. 59), or a mistaken identification of Audubon’s Shearwater Puffinus lherminieri (Collar et al. 1992; BirdLife International 2011).

Hindwood’s Petrel  Pterodroma hindwoodi

(Whitley)

Cookilaria hindwoodi Whitley 1938, p. 197 (Norfolk Island) Status  Invalid taxon. Putative range  Norfolk Island.

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). 363

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Extinct Birds

Agile Petrel  Pterodroma agilis

(Mathews)

Procellaria agilis Mathews 1912a, p. 152 (based on Solander’s manuscript) (36°49’S, 111°30’W. Near Easter Island) Status  Doubtful taxon. Putative range  Possibly Easter Island, East Pacific.

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. 60).

Sordid Petrel  Pterodroma sordida

(Mathews)

Procellaria sordida Mathews 1912a, p. 162 (based on Solander’s manuscript) (25°21’S, 129°W, March 21 1769; and 39° 49’S 111°30’W, 3 March 1769) Status  Doubtful taxon. Putative range  Pacific Ocean.

Known only from Solander’s inadequate description.

Mourning Petrel  Pterodroma atrata

(Mathews)

Procellaria atrata Mathews 1912a, p. 163 (based on Solander’s manuscript) (25°21’S, 129°W) Status  Doubtful taxon. Putative range  Pacific Ocean.

Known only from Solander’s inadequate description.

Sailing Petrel  Pterodroma velificans

(Mathews)

Procellaria velificans Mathews 1912a, p. 161 (44° 35’S, 109°2’W, February 23 1769; 36°49’S, 111° 30’W, March 3 1769) Status  Doubtful taxon. Range  Pacific Ocean.

Known only from Solander’s inadequate description.

Kuril Petrel  Procellaria curilica

Pallas

? Great Black Peteril. Edwards 1743–51, p. 89, pl. 89 (Cape of Good Hope; according to Dabbene 1923, Edwards’s type came from South Georgia) Kuril Petrel. Latham 1785, p. 399 (Kuril Islands and Kamschatka) Kuril Black Petrel. Pennant 1785, p. 536 (based on Pallas’s manuscript) Procellaria curilica Pallas 1811, p. 314 Status  Doubtful taxon. Putative range  Kuril Islands and Kamchatka.

Known only from a description by P. S. Pallas. Unidentifiable.

Two-­coloured Petrel  ?Oceanodroma bicolor

(Tschudi)

Puffinus bicolor Tschudi 1856, p. 187 Status  Doubtful taxon. Putative range  Easter Island and Isla Sala y Gomez.

Known only from Tschudi’s description. Unidentifiable.

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Appendix 2: Doubtful and invalid taxa

Samoan Petrel  Nesofregetta fuliginosa moestissima

(Salvin)

Fregetta moestissima Salvin 1879, p. 130 (Samoa) Status  Doubtful taxon. Range  Polynesian ­Storm-­petrel occurs widely in the ­south-­central Pacific.

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.

Gadow’s Grebe  Podiceps gadowi

Hachisuka

Podicepes [sic] sp. Newton & Gadow 1893, p. 289 (Mauritius) Podiceps gadowi Hachisuka 1953, p. 123 Status  Invalid taxon. Putative range  Mauritius.

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.

Galeata Stork  Euxenura galeata

(Molina)

Ardea galeata Molina 1782, pp. 235, 344 (Chile) Status  Invalid taxon. Putative range  Chile.

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.

Chilean Jabiru  Euxenura pilla

(Molina)

Tantalus pillus Molina 1782, pp. 243, 344. (Chile) Status  Doubtful taxon. Putative range  Chile.

Known only from Molina’s inadequate description. Unidentifiable.

Blue-­­headed Heron  Ardea cyanocephala

(Molina)

Ardea cyanocephala Molina 1782 pp. 235, 344. (Chile) Status  Doubtful taxon. Putative range  Chile.

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’.

Miller’s Heron  Ardea naevia

Miller

Ardea naevia Miller 1782, pl. 36 Status  Known only from Miller’s plate and Shaw’s description of it. Putative range  South America.

Unidentifiable. 365

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Extinct Birds

Red-headed Heron  Ardea erythrocephala

Molina

Ardea erythrocephala Molina 1782, pp. 235, 344. (Chile) Status  Doubtful taxon. Putative range  Chile.

Known only from Molina’s description. Never identified, and ignored by Sharpe (1898) and Hellmayr & Conover (1948).

Rusty-­crowned Heron  Ardea rubiginosa

Gmelin

Rusty-crowned Heron. Pennant 1785, p. 452. (North America) Ardea rubiginosa Gmelin 1789, p. 632 Status  Doubtful taxon. Putative range  North America.

Known from the descriptions of Pennant and Latham. Considered unidentifiable by McAtee (1963).

Red-billed Heron  Ardea aequinoctialis

Linnaeus

The Little White Heron. Catesby 1731, p. 77 Ardea aequinoctialis Linnaeus 1766, p. 240 Little White Heron. Latham 1785, p. 93 Red–billed Heron. Pennant 1785, p. 66 Le crabier bec rouge. Buffon 1770–86, p. 401 Status  Doubtful taxon. Putative range  Carolina, United States.

Known only from Catesby’s description, from which the other accounts derive. Never identified, but may represent a ­now-­extinct form. Sharpe (1898) identified Linnaeus’s Ardea aequinoctialis as the white morph of the Reddish Egret Egretta rufescens.

Catesby’s Great Crested Heron  Ardea cristata

Catesby

Ardea cristata maxima americana. Catesby 1754, pl. 10 (Virginia) Great Heron. Latham 1785, p. 85 Status  Doubtful taxon. Putative range  Virginia, United States.

Known only from Catesby’s description. Ridgway (1878) treated it as fictitious, or the same as the Great Blue Heron Ardea herodias.

New Zealand Pelican  Pelecanus (conspicillatus) novaezealandiae

(Scarlett)

Pelecanus conspicillatus novaezealandiae Scarlett 1966, p. 209 Status  Invalid taxon. Putative range  North and South Islands, New Zealand.

Now considered synonymous with the Australian Pelican Pelecanus conspicillatus (Worthy 1998).

Sharpe’s Pelican  Pelecanus sharpei

Bocage

Pelecanus sharpei Bocage 1870, pp. 173, 409 (Angola) Status  Uncertain. Putative range  Angola, Lower DR Congo and Togo.

Known from a number of specimens from tropical Africa. Usually dismissed as an aberration ­(Ogilve-­Grant 1898; Bannerman 1930a). 366

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Appendix 2: Doubtful and invalid taxa

Violet Cormorant  Phalacrocorax violaceus

(Pennant)

Violet Corvorant (Pelecanus Violaceus) Pennant 1785, vol. 2, p. 584 (Kamtschatka and the islands) Violet Shag. Latham 1785, p. 600 Status  Doubtful taxon. Putative range  Kamchatka and nearby islands.

Known only from Pennant’s brief description. Unidentifiable (Hellmayr & Conover 1948).

Forbes’s Shag  Phalacrocorax novaezelandiae

Forbes

Phalacrocorax novaezelandiae var. major Forbes 1892d, p. 189 (New Zealand) Status  Invalid taxon. Putative range  New Zealand.

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).

White-tailed Cormorant  Phalacrocorax leucurus

Audubon

Phalacrocorax leucurus Audubon 1849 (based on a MS by Townsend), p. 336 (Cape Disappointment, Mouth of Columbia River, Washington State, USA) Status  Doubtful taxon. Putative range  Washington State, United States.

Known only from Audubon’s description (1849), based on specimens seen by Townsend. Considered to be Pelagic Cormorant P. pelagicus ­(Ogilvie-­Grant 1898).

White-rumped Cormorant  Phalacrocorax leuconotus

Audubon

Phalacrocorax leuconotus Audubon 1849 (based on a MS by Townsend) p. 336 (Cape Disappointment, Mouth of Columbia River, Washington State, USA) Status  Doubtful taxon. Putative range  Washington State, United States.

Known only from Audubon’s description (1849), based on specimens seen by Townsend. Considered to be Pelagic Cormorant P. pelagicus ­(Ogilvie-­Grant 1898).

Steller’s White Cormorant  Phalacrocorax albidus

Pallas

Phalacrocorax albidus Pallas 1811 [ex Steller MS], Zoographia Rosso-Asiatica, vol. 2, p. 305. (Bering Island). Status  Doubtful taxon. Putative range  Bering Island.

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.

White-winged Vulture  Gymnogyps sp. White-winged Vulture. Latham 1821, p. 8 (no locality) Status  Doubtful taxon. Putative range  Unknown.

Known only from Latham’s description. Unidentifiable.

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Extinct Birds

Sonnerat’s Secretary Bird  Saggitarius philippensis

(Ogilby)

Bird from the East Indies. G. Edwards 1771, p. 55, pl. 2 (Philippine Islands). Gypogeranus Philippensis Ogilby 1835, p. 105. Status  Invalid taxon. Putative range  Possibly the Philippine Islands.

Known only from the account of Sonnerat. Almost certainly a ­dreamed-­up bird.

Ash-coloured Vulture  Neophron leucocephalus

(Latham)

Ash-coloured Vulture. Latham 1781, p. 13, var. A Vultur leucocephalus Latham 1790, p. 2 Status  Doubtful taxon. Putative range  North Africa and Europe.

Known only from early descriptions. Synonymised under Egyptian Vulture Neophron percnopterus (Strickland 1855).

Hare Vulture  Aegypius cristatus

(Gmelin)

Vultur leporarius. Gesner 1555 Hare Vulture. Willughby 1676, p. 67 Vultur leporarius Ray 1713, p. 10 Vultur cristatus Brisson 1760, p. 460 Status  Doubtful taxon. Putative range  Germany.

Known only from Gesner’s description, from which all others cited. Unidentifiable.

Chincou Vulture  Aegypius chincou

(Daudin)

Le Chincou. Levaillant 1799, pl. 12 Vultur chincou Daudin 1800, p. 12 Status  Doubtful taxon. Putative range  China.

Known only from Levaillant’s description. Synonymised under the Eurasian Black Vulture Aegypius monachus (Sharpe 1874).

Heude’s Eagle  Haliaeetus niger

Heude

Haliaeetus niger Heude 1887, p. 95 (‘Mer de Tartarie’ = seas near Korea) Haliaeetus branickii Taczanowski 1888, p. 451, fig. 1 (Korea) Status  Invalid taxon. Range  The morph was known from the coasts of the Korean Peninsula, and perhaps also in Ussuriland in Russia. It may well ­re-­appear.

This has generally been considered as either a subspecies (‘niger’) of Steller’s Sea Eagle Haliaeetus pelagicus (for example, ­Ferguson-­Lees & Christie 2005), or a ­range-­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.

Maritime Eagle  Haliaeetus maritimus

(Wurmberg & Lichtenstein)

Falco maritimus Wurmberg 1787, p. 6 (Java) Status  Known only from the brief description by Wurmberg (1787), from which various others derive. Putative range  Coasts of Java.

Gray (1869) treated it as a synonym of the ­White-­bellied Sea Eagle H. leucogaster. 368

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Appendix 2: Doubtful and invalid taxa

Macarran Eagle  Aquila dicronyx

Rafinesque

Aquila dicronyx Rafinesque 1832, p. 63 (Buenos Aires) Status  Doubtful taxon. Putative range  South America.

Known only from the description of Rafinesque. Considered to be an aberrant Bald Eagle Haliaeetus leucocephalus (Rhoads 1911), which make no sense on zoogeographical grounds. Unidentifiable.

European White Eagle  Aquila alba

(Gmelin)

Aquila alba sive Cygnea. Gesner 1555, p. 199 Aigle toute blanche. Belon 1555, p. 89 Aquila alba seu cycnea. Aldrovandus 1599, p. 231 Aquila alba. Rzaczynski 1721, p. 299 Aquila alba, seu Aquila Cygnea Aldrovandi. Rzaczynski 1745, p. 362 Aquila alba Cygnea. Klein 1750, p. 42 Aquila alba. Brisson 1760, p. 424 White Eagle. Latham 1781, p. 36 Falco albus Gmelin 1788, p. 257 Falco cygneus Latham 1790, p. 14 Status  Uncertain. Range  This morph formerly occurred in the Alps and in the mountains near the banks of the Rhine in Germany.

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.

Black-cheeked Eagle  ?Aquila sp. ? Black-bellied Falcon. Forster 1771, p. 8 (no description) Black-cheeked Eagle. Latham 1781, p. 35 Status  Doubtful taxon. Putative range  North America.

Known only from the descriptions of Pennant and Latham. Strickland (1855) synonymised the ­Black-­cheeked Eagle with the North American race of Osprey Pandion haliaetus carolinensis.

Tiger Eagle  Aquila tigrina

(Beseke)

Falco tigrinus Beseke 1792, p. 10 Tiger Eagle. Latham 1821, p. 57 Status  Doubtful taxon. Putative range  Courland (now part of Latvia).

Known only from Beseke’s account. Unidentifiable.

Courland Eagle  Aquila germanicus

(Latham)

Der Rothlichweisse Falke. Beseke 1792, p. 11 Falco germanicus Latham 1801b, p. 3 Courland Falcon. Latham 1802, p. 19 Status  Doubtful taxon. Putative range  Courland (now part of Latvia).

Known only from Beseke’s account. Unidentifiable.

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Louisiana White Eagle (Conciliating Eagle)  Aquila candidus

(Gmelin)

White Eagle. Du Pratz 1758, p. 75 (possibly p. 109) White Eagle. Pennant 1785, p. 197 Louisiana White Eagle. Latham 1781, p. 36 Falco candidus Gmelin 1788, p. 258 Falco Conciliator Shaw 1809, p. 77 Status  Known only from the account of Du Pratz. Strickland (1855) placed the Louisiana White Eagle in the synonymy of the Bald Eagle Haliaeetus leucocephalus, with a note that Du Pratz was inaccurate. Putative range  North America.

Genoese Eagle  Aquila sp. Genoese Eagle. Latham 1822, p. 53 (Genoa and parts adjacent) Status  Doubtful taxon. Putative range  Genoa and adjacent areas.

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.

Fierce Eagle  Accipiter ferox

Gmelin

Accipiter ferox Gmelin 1771, p. 442, pl. 10 (Astrakhan) Fierce Eagle. Latham 1781, p. 33 Falco astracanus Shaw 1809, p. 85 Status  Known only from Gmelin’s description. Putative range  Astrakhan, Russia.

Considered unidentifiable by Mayr (1944).

Cooper’s Buzzard  Buteo cooperi

Cassin

Buteo cooperi Cassin 1856, p. 253 (Near Mountain View, Santa Clara County, California) Status  Known only from the type. Putative range  California.

Generally believed to be an abnormal specimen of ­Red-­tailed Hawk Buteo jamaicensis harlani (Ridgway 1884, 1885a).

Mauritius Sparrowhawk  Accipiter alphonsi

(Newton & Gadow)

Astur sp. Milne-Edwards 1874a, p. 25–6, pl. 15, fig. 2 Astur alphonsi Newton & Gadow 1893, p. 285–6, pl. 33, figs 9–10 (Mauritius) Circus maillardi ­Mourer-­Chauviré et al. 2004, p.168 Specimens  Subfossil remains of the Mauritius population are in Cambridge, England and Paris. 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 indistinguishable 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 370

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forest, a similar evolutionary trait exhibited by the Hawaiian Harrier Circus dossenus (see p. 79). The harrier was possibly mentioned on Mauritius in 1602 (Cheke & Hume 2008), but never again, and probably disappeared as a result of persecution and deforestation.

Cape Verde Kite  Milvus milvus fasciicauda

Hartert

Milvus milvus fasciicauda Hartert 1914, p. 89 (Cape Verde Islands) Status  Almost certainly invalid. Range  Cape Verde Islands.

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).

D’Orbigny’s Caracara  Daptrius gymnocephalus

(D’Orbigny)

Ibycter gymnocephalus D’Orbigny 1835, p. 2 (Cochabamba, Bolivia) Status  Doubtful taxon. Putative range  Cochabamba, Bolivia.

Known only from the original description with no surviving specimens. Considered referable to the Black Caracara Daptrius ater (Hellmayr 1921).

Latham’s Brown Falcon  Falco fuscus

Gmelin

Vultur pÿgargus. Frisch 1734–63, tab. 76 Falco fuscus. Brisson 1760 p. 331 Brown Falcon. Latham 1781, p. 68 Falco fuscus Gmelin 1788, p. 271 (not Falco fuscus Gmelin 1788, p. 280, which = Accipiter striatus velox) Status  Doubtful taxon. Putative range  Europe.

Known only from the plate in Frisch, on which the descriptions of Brisson and Latham are based. Unidentifiable.

Salt’s Falcon  Falco sp. Falcon. Salt 1814, p. xlii-xliii. Status  Doubtful taxon. Range  Abyssinia.

Known only from the description by Henry Salt following his voyage to Abyssinia. Unidentifiable.

Rusty Falcon  Falco rubiginosus

Latham

Falco rubiginosus. Piller 1783, p. 29 Falco rubiginosus Latham 1790, p. 27 Rusty Falcon. Latham 1802, p. 36 Falco aeruginosus, Var A. Latham 1821, p. 88 Status  Doubtful taxon. Range  Slovenia.

Known only from Piller’s description. Strickland (1855) placed it in the synonymy of the Hen Harrier Circus cyaneus. 371

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Starry Falcon  Falco stellaris

Gmelin

Falco cyanopus. Charleton 1668, p. 73 Blue-footed Falcon. Willughby 1678, p. 82 Falco cyanopus. Klein 1750, p. 52 Falco stellaris. Brisson 1760, p. 359 Starry Falcon. Latham 1781, p. 79 Falco stellaris Gmelin 1788, p. 274 Status  Doubtful taxon. Putative range  Europe.

Known only from a description in Albertus Magnus. Strickland (1855) placed it in the synonymy of Saker Falcon Falco cherrug.

White-headed Falcon  Falco leucocephalus

Gmelin

Vultur subluteus, capite albo, Kauh-fuss geyer; gelbrauner Geÿer. Frisch 1734–1763, tab. 75 Falco leucocephalus. Brisson 1760, p. 325 White-headed Falcon. Latham 1781, p. 66 Falco leucocephalus Gmelin 1788, p. 270 Status  Invalid taxon. Putative range  Europe.

Known only from Frisch’s plate. Bechstein (1793) identified it as the ­Rough-­legged Buzzard Buteo lagopus.

White Falcon  Falco albus

Gmelin

Falco albus. Gesner 1555 Falco albus Gmelin 1788, p. 270 (not Falco albus, p. 257 which = Aquila alba Gmelin) Status  Invalid taxon. Range  Europe.

Known from several early accounts, but it is unclear whether they all refer to the same bird. Unidentifiable.

Speckled Partridge Hawk  Falco sacer

Forster

Falco sacer. Forster 1772, p. 383, 423 American Sacre or Speckled Partridge Hawk. Latham 1781, p. 78 Status  Invalid taxon. Putative range  North America.

Known only from Forster’s description. Synonymised under the Gyrfalcon Falco rusticolus (Hellmayr & Conover 1948).

Rougri Falcon  Falco desertorum

Daudin

Le Rougri. Levaillant 1799, p. 49, pl. 17 Falco desertorum Daudin 1800, p. 164 Buteo desertorum Smith 1830, p. 382 Status  Doubtful taxon. Putative range  South Africa.

Known only from Levaillant’s description. Considered unidentifiable (Hartert & Neumann 1914).

Norton Sound Bustard  Otis sp. Norton Sound Bustard. Pennant 1785, p. 321 Status  Doubtful taxon. Putative range  North America.

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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.

Peter Mundy’s Rail  Kuinia mundyi

Hachisuka

Mauritius Henne. Peter Mundy 1608–67 (in Temple 1914), p. 352 Kuinia mundyi Hachisuka 1937a, p. 156 (Mauritius) Status  Invalid taxon. Putative range  Mauritius.

Based on the account of Peter Mundy. Probably referable to a juvenile Mauritius Red Rail Aphanapteryx bonasia (Olson 1977b) (see p. 108).

Sharpe’s Rail  Gallirallus sharpei

(Büttikofer)

Stictolimnas sharpei Büttikofer 1893, p. 274 Gallirallus sharpei Olson 1986a, p.263 Status  Invalid taxon. Range  Unknown but possibly the Greater Sundas, Indonesia.

Known only from the type. Now considered a colour morph of ­Buff-­banded Rail G. philippensis, based on unpublished genetic work (R. Dekker in litt. 2008), and removed from the 2010 IUCN Red List (Bird & Butchart 2008).

Red-throated Wood Rail  Aramides gutteralis

Sharpe

Aramides gutteralis Sharpe 1894, p. 57, pl. 5 Status  Doubtful taxon. Putative range  Believed to have come from Lima, Perú.

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 ­Grey-­necked Wood Rail A. cajanea (Meyer de Schauensee 1966), or a possible subspecies of it (Taylor & van Perlo 1998).

Leguat’s Giant  Leguatia gigantea

Schlegel

Le Geans. Leguat 1708, p. 171. Gallinula (Leguatia) gigantea Schlegel 1857, p. 116, 142; translated in 1866, p. 146–168 Status  Invalid taxon. Putative range  Mauritius and Rodrigues.

Based only on the account of Leguat (1708), who almost certainly confused it with a flamingo Phoenicopterus sp. (Cheke & Hume 2008).

Notable Rail  Gallirallus forbesi

Forbes

Ocydromus insignis Forbes 1892 p. 188 (Middle Island, New Zealand) Status  Dubious taxon. Range  New Zealand.

Known only from a single element with an insufficient description (Olson 1977b).

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Short-winged Rail  Rallus ecaudotus

King

Rallus ecaudotus King 1784, p. 119 Pennula ecaudata [sic] Hartlaub 1892, p. 396 Status  Invalid taxon. Putative range  Hawaii.

Known only from an inadequate description in King (1784), and now a synonym of Hawaiian Crake Porzana sandwichensis (see p. 103).

Blue-bellied Rail  Rallus ecaudatus

Miller

Rallus pacificus (part) J. Forster, unpublished drawing, no. 127, in Banksian Library Collection in NHM. Rallus ecaudata Miller 1783, pl. 47 (Otaheitee) Rallus ecaudatus Shaw & Miller 1796, pl. 47 Status  Invalid taxon. Putative range  Supposedly Tahiti, but probably Hawaii.

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 Porzana sandwichensis (see p. 103)

Gilbert Rail  Tricholimnas conditicius

Peters & Griscom

Tricholimnas conditicius Peters & Griscom 1928, p. 102 (Apiang Island, Gilbert Group = error, probably for Ebon Atoll in the Marshall Islands) Status  Invalid taxon. Range  Gilbert Islands (Kiribati) in error. Type probably from Lord Howe.

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 Tricholimnas sylvestris (Olson 1992a).

Herbert’s Hen  Didus herberti

Schlegel

‘A Hen’. Herbert 1634, p. 212 Didus herberti Schlegel 1854, p. 256 (Mauritius) Status  Invalid taxon. Putative range  Mauritius.

Based on the extremely poor probable rendition of Aphanapteryx bonasia in Herbert (1634; see Olson 1977b).

Little Woodhen  Gallirallus minor

(Hamilton)

Ocydromus minor Hamilton 1893, p. 103 (Castle Rocks, New Zealand) Gallirallus minor Oliver 1930, p.594 Status  Doubtful taxon. Range  Known from deposits from Takaka, Lake Grassmere, Pyramid Valley, Forest Hill, Hamilton Swamp, Line Hills and Castle Rocks in New Zealand.

Known only from subfossil material, which has never been properly characterised or illustrated (see Olson 1977b).

Bruner’s Rail  Cacroenis inornatus

Bruner

Cacroenis inornatus Bruner 1972, p. 47 (Tuamotu Islands) Status  Doubtful taxon. Putative range  Tuamotu Archipelago, South Pacific.

Known only from an inadequate account in Bruner (1972). 374

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Colenso’s Coot  Fulica novazealandiae

Colenso

Fulica Nova-Zealandiae Colenso 1845, p. 283 (Ngaruawahie, New Zealand) Status  Invalid taxon. Putative range  New Zealand.

Known only from Colenso’s account. Now thought to be a misidentification of the New Zealand Grebe Poliocephalus rufopectus.

Mysterious Rail  Porzana circoleps

Lesson

Râle écaudé. Cuvier unpublished MS in Paris Museum Porzana circoleps Lesson 1831, p. 538 Rallus ecaudatus Cuvier (according to Bonaparte) Ortygometra cercoleps ‘Temminck’. Gray 1871, p. 64; nomen nudum Rallus caudatus Cuvier (according to Gray) Status  Invalid taxon. Putative range  Philippine Islands.

Based on the inadequate account of Lesson (1831).

Luzon Sarus Crane  Grus antigone luzonica

Hachisuka

Grus antigone luzonica Hachisuka 1941b, p. 83 (Luzon) Status  Invalid taxon. Range  Luzon, Philippine Islands.

Considered probably extinct (Dickinson et al. 1991). However, now synonymised under Grus antigone sharpii (Dickinson 2003).

Audubon’s Great White Crane  Grus sp. Great White Crane. Nuttall 1834, p. 39 (Florida) Status  Doubtful taxon. Putative range  Florida.

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.

Marco Polo’s Crane  Grus polii

Yule

Grus polii Yule 1871, p. 262. Status  Invalid taxon. Putative range  Unknown.

Known only from Marco Polo’s description, as translated by Yule. Synonymised under Siberian Crane Grus leucogeranus (Blaauw 1897).

Ross’s Shore Plover  Thinornis rossii

Gray

Thinornis rossii G. R. Gray 1845, p. 12, pl. 11 (Auckland Island = ? Campbell Island) Specimens  The unique specimen is in Tring. Status  Invalid taxon. Putative range  Auckland Islands, New Zealand, though this may be in error.

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). 375

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Black-crowned Plover  Charadrius atricapillus

Gmelin

Black-crowned Plover. Latham 1785 p. 210 Charadrius atricapillus Gmelin 1789, p. 686 Status  Known only from the descriptions of Latham and Pennant. Putative range  New York State, United States.

Unidentifiable.

Uniform Sandpiper  Tringa uniformis

Gmelin

Keildu-suin, Islandis. Müller 1776, n. 205. Uniform Sandpiper. Latham 1785, p. 173 Tringa uniformis Gmelin 1789, p. 678 Status  Doubtful taxon. Putative range  Iceland.

Known only from Müller’s account, perhaps based on an earlier work by Ólafsen (1774–75). Never identified, and almost certainly unidentifiable.

Waved Sandpiper  Tringa undata

Brünnich

Tringa undata Brünnich 1764, p. 55, no.183 Waved Sandpiper. Pennant 1785, p. 481 Tringa sordide flava subtus alba. Beseke 1786, p. 463 Status  Doubtful taxon. Putative range  Possibly Denmark and Norway.

Known only from Brünnich’s description, from which others derive.

Sakhalin Sandpiper  Tringa meleagris

Krusenstern

Tringa meleagris Krusenstern 1814, table 86 of Atlas Tringa Sakhalmi Vieillot 1819, p. 471 Status  Doubtful taxon. Putative range  Unknown.

Known only from Hartert’s comments on Krustenstern’s plate. Probably fictitious, based on a mixture of species.

Cooper’s Sandpiper  Calidris cooperi

(Baird)

Tringa cooperi Baird 1860, p. 716 Specimens  The type is in Washington D.C. Status  Known only from the unique type, believed to be a male in summer plumage, collected on Long Island, New York, on May 24, 1833, by William Cooper. Range  North America.

Now considered to probably represent a hybrid between Curlew Sandpiper Calidris ferruginea and ­Sharp-­tailed Sandpiper C. acuminata (Cox 1989, 1990).

Cox’s Sandpiper  Calidris paramelanotos

Parker

Calidris paramelanotos Parker 1982, p. 63 (Saint Vincent’s Gulf, South Australia) Status  Invalid taxon. Range  South and south-east Australia, Massachusetts, United States and Japan.

DNA analysis has shown that Cox’s Sandpiper is a hybrid Curlew Sandpiper C. ferruginea x Pectoral Sandpiper C. melanotos (Christidis et al. 1996). 376

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Kamchatkan Tern  Sterna camtschatica

Pallas

Larus Martyschka. Steller unpublished MS Kamtschatkan Tern. Pennant 1785, p. 525 Sterna camtschatica Pallas 1811, p. 335 Status  Invalid taxon. Putative range  Kamchatka, Alaska, Japan and Aleutian Islands.

Originally known only from Pallas’s description, taken from Steller’s manuscript. Synonymous with the extant Aleutian Tern Onychoprion aleuticus.

Cloven-footed Tern  Sterna naevia

Linnaeus

Cloven-footed Gull. Albin 1738, p. 75, pl. 82 Rallus cinereus facie Lari. Klein 1750, p. 103 ? La Guissette. Buffon 1770–83, p. 339 ? Rallus lariformis Linnaeus 1758, p. 153 Sterna naevia Brisson 1760. p. 216, fig. 2 Sterna naevia Linnaeus 1766, p. 228 Status  Invalid taxon. Putative range  Picardy Coast, France.

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.

Black Garefowl  Pinguinus sp. Gairfowl. Martin 1698, p. 27 Status  Invalid taxon. Putative range  St Kilda.

Based on an inadequate description, and probably confused in part with a cormorant.

Collared Garefowl  Pinguinus sp. Anser magellanicus seu Pinguini. Wormius 1655, p. 300–1 Status  Invalid taxon. Putative range  Faroe Islands.

Description based on an aberrant individual, or possibly confused with Great Northern Diver Gavia immer.

Réunion White Dodo  Raphus solitarius

(de Sélys-Longchamps)

Apterornis solitarius de Sélys-Longchamps 1848, p. 293 Didus apterornis Schlegel 1854, p. 232 Ornithaptera borbonica Bonaparte 1854a, p. 2 Victoriornis imperialis Hachisuka 1937b, p. 71 Status  Invalid taxon. Existence based entirely on contemporary pictures and travellers accounts. Putative range  Réunion, Mascarene Islands.

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. 67). Large numbers of bird subfossil remains have now been 377

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recovered 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.

Mexican Blue Dove  Columba caerulea

Gmelin

Tlacahoilotl. Hernandez 1651, p. 46 (Mexico) Columba caerulea mexicana. Brisson 1760, p. 139 Pigeon bleu du Mexique. Buffon (1770–1783), p. 525 Blue Pigeon. Latham 1783, p. 634 Columba caerulea Gmelin 1788, p. 776 Status  Invalid taxon. Putative range  Mexico.

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).

Indian Blue Dove  Columba dorsocaerulea

Temminck & Knip

Columba caerulea Temminck & Knip 1811, pl. 37 Status  Invalid taxon. Putative range  Bengal, India.

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 (Sharpe 1895).

Sonnerat’s Pigeon  Columba cinerea

Scopoli

Tourterelle centrée de l’Isle de Luçon. Sonnerat 1776, p. 52, pl. 22 (Luzon, Philippine Islands) Luzonian Turtle. Latham 1783, p. 646, var. c Columba cinerea Scopoli 1786, p. 94, no. 93 Columba turtur, delta, Turtur luzoniensis. Gmelin 1788, p. 786 Tourterelle de l’Isle de Luçon. Bonnaterre 1790, Tp. 248, pl. 81 Columba turtur delta. Latham 1790, p. 606 Columba phoenicorhyncha Wagler 1829, p. 745 Status  Doubtful taxon. Putative range  Luzon, Philippine Islands.

Known only from Sonnerat’s description, the basis of Scopoli’s name Columba cinerea. Almost certainly referable to Madagascar Turtle Dove Streptopelia picturata (Walden 1875), and considered doubtful (Wagler 1827).

San Domingo Dove  Columba dominicensis

Latham

Tourterelle de St. Dominigue. Daubenton 1771–86, no. 487 Columba dominicensis Latham 1790, p. 615, no. 79 Status  Invalid taxon. Putative range  Hispaniola, West Indies.

Known only from Daubenton’s plate, on which Latham’s (1790) description was based. Almost certainly fictitious (e.g. Wagler 1827; Salvadori 1893).

Mexican Dove  Columba mexicana

Gmelin

Chehoilotl. Hernandez 1651, p. 42 Columba mexicana Brisson 1760, p. 99 Columba mexicana Gmelin 1788 p. 777

378

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Known only from the inadequate description of Hernandez (1651).

Rose-coloured Dove  Columba rosea

Miller

Columba rosea Miller & Shaw 1796, p.105, (India) Status  Doubtful taxon. Putative range  India.

Known only from Miller’s plate and inadequate description. Never identified, and considered a doubtful species (Wagler 1827; Bonaparte 1854a).

Hesler’s Dove  Columba sumatrensis

Hesler

Columba sumatrensis Hesler 1854, p. 287 (no locality) Status  Invalid taxon. Putative range  Unknown.

Known only from Hesler’s note, with inadequate description.

Gmelin’s Dove  Columba fusca

Pallas

Columba minor fusca Gmelin ‘the elder’ (i.e. J.G. Gmelin) MS. Columba fusca Pallas 1811, p. 567 Status  Doubtful taxon. Putative range  Krasnoyarsk on the upper Yenisei, Siberia.

Known only from Pallas’s inadequate ­second-­hand description, based on Gmelin’s manuscript.

Seba’s Dove  Columba brasiliensis

(Linnaeus)

Cuculus brasiliensis. Seba 1734, p. 102, pl. 66, fig. 2 Columba adfinis Möhring 1752, p. 103 Cuculus brasiliensis cristatus ruber Brisson 1760, no. 25 Cuculus brasiliensis Linnaeus 1766, p. 171. Red-cheeked Cuckow. Latham 1781, p. 545 Status  Invalid taxon. Putative range  Unknown.

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).

Siamese Pigeon  Columba sp. Pigeon from Xiengmai. Schomburgk 1864, p. 250 Siamese Pigeon. Blyth 1867, p. 149 Status  Known only from the description of Schomburgk (1864). Putative range  Chiang Mai, Thailand.

Unidentifiable.

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Hottentot Pigeon  Goura hottentotta

(Temminck & Knip)

Columbi-caille. Levaillant 1808, p. 83, pl. 283 Columba hottentotta Temminck & Knip 1808–1811, p. 26, pl. 15 Goura hottentota Stephens 1819, p. 131 Status  Doubtful taxon. Putative range  Great Namaqualand, South Africa.

Apparently known only from Levaillant’s description. Considered fictitious (Sundervall in Layard 1867).

Burnes’s Pigeon  Psammoenas burnesii

Blyth

Psammoenas Burnesii Blyth 1845, p. 860 footnote (Cabul) Status  Invalid taxon. Putative range  Afghanistan.

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.

Levaillant’s Dove  Verrulia carunculata

(Temminck & Knip)

Colombi–Gallinae. Levaillant 1808, p. 98, pl. 278 Columba carunculata Temminck & Knip 1808–11, p. 19, pl. 11 Goura carunculata Stephens 1819, p. 123 Verrulia carunculata Fleming 1822, p. 232. Status  Invalid taxon. Putative range  Africa.

Known only from Levaillant’s plate and description, from which all other accounts derive. Considered a fictitious bird (Salvadori 1893).

Callao’s Dove  ‘Columbicolin laure’

Bonaparte

Columbicolin laure Bonaparte 1854a, p. 51 Status  Invalid taxon. Putative range  Unknown.

Known only from Bonaparte’s inadequate description. Never satisfactorily identified.

Spotted-­necked Turtle Dove  Streptopelia sp. Spotted Necked Turtle. Latham 1785, p. 645 Columba turtur var. beta. Gmelin 1788, p. 786 Status  Invalid taxon. Putative range  From England to South Africa, China and India.

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.

Senegal Cuckoo Dove  Macropygia macerona

(Müller)

Tourterelle à large queue du Sénégal. Daubenton 1771–86, no. 329 Tourocco. Buffon 1771 p. 553 (Senegal) Great-tailed Turtle. Latham 1783, p. 667 Columba macerona Müller 1776b, p. 134, no. 43 Columba macroura Gmelin 1788, p. 790 Macropygia macroura Salvadori 1893, p. 335–6 footnote (Senegal; Sri Lanka according to some = error)

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Appendix 2: Doubtful and invalid taxa Status  Doubtful taxon. Putative range  Senegal, West Africa or perhaps Sri Lanka.

Known only from Daubenton’s plate and Buffon’s description. Never satisfactorily identified. No other Macropygia occurs in Africa, but provenance uncertain.

China Dove  Geopelia sinica

(Linnaeus)

China Dove. Albin 1740, p. 43, pl. 46 Columba sinica Linnaeus 1758, p. 164 Status  Doubtful taxon. Putative range  China.

Known only from Albin’s plate and description. Provenance uncertain and the plate differs from the description.

King’s White Pigeon  Ducula sp. Large White Pigeon. King 1784, p. 120 Status  Invalid taxon. Putative range  Hawaiian Islands.

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.

Indian Pigeon  ?Treron asiatica

Latham

Indian Pigeon. Latham 1787, p. 202 Columba asiatica Latham 1790, p. 597, no 14 (India) ? Treron asiatica Gray 1844a, p. 467 Status  Doubtful taxon. Putative range  India.

Known only from Latham’s description. Considered indistinct (Stephens 1819) and an unidentifiable Treron (Salvadori 1895).

Lefèbvre’s Pigeon  ?Treron phasianella

(Lefèbvre)

Columba phasianella Lefèbvre 1845–51, p. 142 (Abyssinia). Status  Doubtful taxon. Putative range  Abyssinia.

Known only from Lefèbvre’s inadequate account. Never identified.

Brown Pigeon  Hemiphaga brunnea

(Latham)

Columba brunnea Latham 1790, p. 603, no. 38 (New Zealand) Brown Pigeon. Latham 1802, p. 267 Status  Doubtful taxon. Putative range  New Zealand.

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). 381

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Extinct Birds

Chamisso’s Dove  Columba australis

Chamisso

Columba australis Chamisso 1821, p.113 (Radack or Marshall Islands) ?Ducula oceanica ratakensis Takatsukasa & Yamashina 1932, p. 221 (Aruno Atoll) Status  Doubtful taxon. Putative range  Marshall Islands, South Pacific.

Known only from a passing reference by Chamisso, without adequate description.

Earl’s Pigeon  Columba sp. Pigeon of New Zealand with ‘The Head and neck White, the former Crested’. Earl 1844 (in Gray & Sharp), p. 10 Status  Doubtful taxon. Putative range  New Zealand.

Known only from Percy Earl’s inadequate description. The provenance may be in error (Gray 1862b).

Gloger’s Dove  Rhagorhina auricularis

(Temminck & Knip)

Columba auricularis Temminck & Knip 1808–11, p. 54, pl. 21 Columba temminckii Wagler 1827, sp. 40 Rhagorhina auricularis Gloger 1842, p. 360 Status  Doubtful taxon. Putative range  Supposedly from a Pacific island.

Known only from Temminck’s description, based on a captive bird in Amsterdam. Provenance uncertain, and now unidentifiable.

Portugal Dove  Streptopelia lusitanicus

(Latham)

Portugal Dove. Albin 1738, p. 44, pl. 48 Turtur lusitanus Brisson 1760, p. 98 Columba turtur Gmelin 1789, p. 786 Turtur lusitanicus Latham 1823, p. 53 Turtur lusitanus Salvadori, 1893, p. 646 Status  Doubtful taxon. Putative range  Portugal.

Known only from Albin’s description and plate. Considered conspecific with European Turtle Dove Streptopelia turtur (Salvadori 1893).

Bougainville’s Pigeon  Ducula sp. Large Deep Blue Pigeon. Bougainville 1772, p. 247 (Otaheite) Status  Doubtful taxon. Putative range  Tahiti.

Known only from a passing inadequate reference by Bougainville (1772).

Hook-billed Pigeon  ?Treron curvirostra

(Gmelin)

Hook-billed Pigeon. Latham 1783, p. 632, no. 23, pl. 59 (Tanna) Columba curvirostra Gmelin 1789, p. 777 (Tanna) Columba aromatica var a. Temminck 1813, p. 441 Status  Doubtful taxon. Putative range  Supposedly Tanna, New Hebrides, though perhaps in error, and also the Malay Peninsula.

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. 382

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Appendix 2: Doubtful and invalid taxa

Purple-breasted Pigeon  Gallicolumba eimeensis

(Gmelin)

Purple breasted Pigeon. Latham 1783, p. 629 (Eimeo = Tahiti) Columba eimeensis Gmelin 1789, p. 784. Based on Latham Status  Doubtful taxon. Putative range  Moorea, Society Islands, South Pacific.

Known only from Latham’s description, and doubtfully distinct. Salvadori (1893) listed this bird as an unidentified form, and Peters (1937) said that it is ‘doubtless’ the same as the Polynesian Ground Dove G. erythroptera.

Jamaican Blue Ground Dove  ?Claravis plumbea

(Gosse)

Blue Partridge. Gosse 1847, p. 324 Zenaida? plumbea Gosse 1849, pl. 85. Status  Doubtful taxon. Putative range  Jamaica, but there is some doubt about the provenance.

Known only from Gosse’s brief and inadequate account and a contemporary illustration.

Fermin’s Dove  Columba surinamensis

Gmelin

Tourterelle de Surinam. Fermin 1769, p. 165 (Surinam). Surinam Turtle. Latham 1783, p. 647 Columba surinamensis Gmelin 1789, p. 787 Columba Fermin Temminck 1813, p. 375. Status  Doubtful taxon. Putative range  Surinam.

Known only from Fermin’s description, from which the others derive. No such bird has been reported since.

Tschudi’s Dove  Columba melancholica

Tschudi

Columba melancholica Tschudi 1844, p. 306 (Peru) Status  Invalid taxon. Putative range  Peru.

Known only from Tschudi’s description of the type, formerly in the Museum of Neuchâtel, but now apparently lost. Referred to the synonymy of the ­Maroon-­chested Ground Dove Claravis mondetoura (Taczanowski 1884–86; Ridgway 1916).missing.

Immaculate Tambourine Dove  Turtur virgo

(Hartlaub)

Tympanistria virgo Hartlaub 1886, p. 2 (Djanda, Eastern Equatorial Africa) Status  Doubtful taxon. Putative range  Djanda, Central African Republic.

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.

Long-­billed Kaka  Nestor norfolcensis

Pelzeln

Long-billed Parrakeet. Latham 1822, p. 171 Nestor norfolcensis Pelzeln 1860a, p. 322 & pl. (Norfolk Island) Status  Invalid taxon. Putative range  Norfolk Island.

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Extinct Birds

Known only from Pelzeln’s lost type with an abnormally developed bill, Latham’s apparently normal specimen (also lost), and another specimen in Liverpool, which lacks the bill. Considered to be an aberration and thus synonymised under Nestor productus (see p. 166).

Small Kaka  Nestor sp. Nestor sp. Scarlett 1955a, p. 263 (Pyramid Valley, New Zealand) Status  Doubtful taxon. Putative range  South Island, New Zealand.

Known only from a very brief statement by Scarlett (1955): ‘many bones of a small form [of Kaka] have been recovered’. Whether or not this represents an extinct species seems not to have been decided. No further information is available.

Masséna’s Kaka  ?Nestor esslingi

Souancé

Nestor esslingi Souancé 1856, p. 223 (no locality) Status  Invalid taxon. Range  New Zealand.

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).

Mamberiok Lory  Chalcopsitta spectabilis

van Oort

Chalcopsitta spectabilis van Oort 1908, p. 127 (Mambrioe = Mamberiok Peninsula, ­north-­western New Guinea) Status  Doubtful taxon. Range  Mamberiok Peninsula, ­north-­western New Guinea.

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) 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.

Stavorini’s Lory  Chalcopsitta stavorini

(Lesson)

Psittacus stavorini Lesson 1826, p. 355, 628 (Waigiou) Status  Doubtful taxon. Putative range  Waigeo, western New Guinea.

Known only from Lesson’s description, based on the unique type, now lost. Considered doubtful (Salvadori 1891a) or unidentifiable (Peters 1937).

Red-­and-green Lory  Eos bornea

(Linnaeus)

Long-­tailed Scarlet Lory. Edwards 1751, p. 173, pl. 173 Psittacus borneus Linnaeus 1758, p. 97 (‘Borneo’) Status  Doubtful taxon. Putative range  Perhaps Borneo.

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). 384

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Appendix 2: Doubtful and invalid taxa

Unicoloured Lory  Eos unicolor

(Shaw)

Le Perroquet Lori unicolor. Levaillant 1805a, p. 131, pl. 125 (Moluccas) Psittacus unicolor Shaw 1811, p. 538. Based on Levaillant Unicoloured Lory. Latham 1822, p. 227 Eos unicolor G.R. Gray 1849, appendix, p. 20 Status  Known only from Levaillant’s description and plate, based on two specimens, now lost, but formerly in the Temminck Collection in Amsterdam. Putative range  Moluccas.

Never identified.

Blue-tailed Lory  Eos sp.

(Levaillant)

Le Perroquet Lori à queue blue. Levaillant 1805a, pl. 97, p. 43 (Borneo = error, ?Moluccas) Status  Doubtful taxon. Putative range  Perhaps Borneo.

Known only from Levaillant’s plate and description, based on a specimen, now lost, formerly in the collection of Raye de Breukelvaert in Amsterdam. Provenance in doubt.

Levaillant’s Lory  Eos coccinea

(Brisson)

Psittaca indica coccinea Brisson 1760, p. 376–8, pl. 25, fig. 2 Le Lori Perruche violet et rouge. Daubenton 1771–86, no 143 Status  Doubtful taxon. Putative range  Unknown.

Known only from the descriptions of Brisson, Daubenton and Levaillant, apparently all based on a specimen (now lost) in the collection of the Abbé Aubry. Salvadori (1891a) put this bird in the synonymy of ­Red-­and-­blue Lory Eos histrio.

Goodfellow’s Lory  Eos goodfellowi­Ogilvie-­­Grant Eos goodfellowi ­Ogilvie-­Grant 1907, p. 102 (Obi) Status  Invalid taxon. Putative range  Moluccas.

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.

Variegated Lory  ?Eos variegata

(Gmelin)

Variegated Lory. Latham 1781, p. 220 Psittacus variegatus Gmelin 1788, p. 319 Status  Doubtful taxon. Putative range  Moluccas.

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).

Layard’s Mysterious Lory  Trichoglossus sp. Trichoglossus (Lorius) sp. inc. Layard 1878, p. 274 (Santo, New Hebrides) Status  Doubtful taxon. Putative range  New Hebrides.

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Extinct Birds

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.

White-collared Lory  Psittacus (Trichoglossus) multicolor

(Gmelin)

White-collared Parrot. Pennant 1773, p. 59–60, pl. 2 (East Indies?) Psittacus multicolor Gmelin 1788, p. 328 Psittacus semicollaris Latham 1790, p. 103 Status  Doubtful taxon. Putative range  East Indies.

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.

Gray’s Lory  Trichoglossus coccineifrons

Gray

Trichoglossus coccineifrons G. R. Gray 1858, pp. 183, 194 Status  Invalid taxon. Putative range  Aru Islands, Indonesia.

Known only from the type collected by Alfred Russel Wallace, an adult male that Salvadori (1891) considered either a hybrid Rainbow Lorikeet T. [haematodus] nigrogularis x ­Yellow-­streaked Lory Chalcopsitta sintillata, or an immature of something else.

Verreaux’s Lory  Trichoglossus verreauxius

Bonaparte

Trichoglossus verreauxius Bonaparte 1854b, p. 157 (Australasia) Status  Invalid taxon. Putative range  Unknown.

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.

Aru Islands Rainbow Lorikeet  Trichoglossus haematodus brooki­Ogilvie-­­Grant Trichoglossus brooki ­Ogilvie-­Grant 1907, p. 102 Status  Invalid taxon. Putative range  Spirit Island, off Trangan, Aru Islands.

Now considered to be immature of the race T. h. nigrogularis (Juniper & Parr 1998).

Green-thighed ­Yellow-­collared Lory  Lorius sp. Le Perroquet Lori à Collier Jaune – Variety. Levaillant 1805, pl. 95 (no locality) Status  Invalid taxon. Putative range  Unknown.

Known only from Levaillant’s plate and inadequate description. Synonymised by Salvadori (1891a) with Purple-naped Lory Lorius domicella.

Walpole’s Lory  Lorius sp. Scarlet Lory. Edwards 1751, p. 172, pl. 172 Psittacus garrulus (part) Linnaeus 1758, p. 100 (Asia = East Indies)

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Appendix 2: Doubtful and invalid taxa Status  Doubtful taxon. Putative range  East Indies.

Known only from Edwards’s 1751 description. Never satisfactorily identified.

Red-­and-­gold Lory  Lorius rex

(Bechstein)

Le Perroquet lori Radhia. Levaillant 1805, p. 39, pl. 94 (Moluccas) Psittacus rex Bechstein 1811, p. 92 Psittacus raja Shaw 1811, p. 537, pl. 82 Psittacus radhea Vieillot 1817, p. 337 Rajah Lory. Latham 1822. p. 226–7 Status  Doubtful taxon. Putative range  Moluccas.

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).

Little Red-winged Lory  Lorius incarnatus

(Gmelin)

Little Red-winged Parrakeet. Edwards 1758, p. 53, pl. 236 Psittaca indica. Brisson 1760, p. 341 Perruche à gorge rouge. Buffon 1779 p. 157 Psittacus incarnatus Gmelin 1788, p. 327 Red-winged Parrakeet. Latham 1822, p. 171 Lorius incarnatus Stephens 1826, in Shaw, p. 132 (no description) Palaeornis incarnatus Wagler 1832, p.517 Status  Doubtful taxon. Putative range  Moluccas.

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).

Jay-winged Lory  Lorius sp. Jay Winged Lory. Latham 1822, p. 130 (Van Diemen’s Land = Tasmania) Status  Invalid taxon. Putative range  Tasmania.

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.

Crested Mexican Parrot  Lorius mexicanus

(Gmelin)

Avis de coco, Psittaci species, Mexicana vario colore. Seba 1734, p. 94, fig. 2 Psittacula Mexicana cristata Brisson 1760, p. 405 Crested Mexican Parrot. Latham 1781, p. 322 Psittacus mexicanus Gmelin 1788, p. 352, no. 141 Status  Doubtful taxon. Putative range  Mexico.

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.

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Extinct Birds

Albin’s Lory  Lorius brasiliensis

(Linnaeus)

The Laurey from the Brazils. Albin 1738, p. 13, pl. 13. Psittacus brasiliensis Linnaeus 1758, p. 100, in synonymy of Psittacus Domicella. Omitted by Linnaeus 1766, and thereafter ignored Status  Doubtful taxon. Putative range  Brazil, but this may be in error.

Known only from Albin’s description, based on a live specimen, now lost, which he saw in London. Unidentifiable.

Meyen’s Lory  Lorius cyanolenus

(Meyen)

Psittacus cyanolenus Meyen 1834, p. 95 (China) Status  Doubtful taxon. Putative range  China.

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.

Jamrach’s Lory  Lorius tibialis

Sclater

Lorius tibialis Sclater 1871b, pp. 499, 544, pl. 40 (no locality; type a specimen in the London Zoo bought alive in Calcutta by the dealer, Jamrach). Status  Invalid taxon. Range  Provenance unknown.

Known only from the unique type in Tring. Believed it to be an aberrant specimen of Lorius domicella (Forshaw 1971).

Stresemann’s Lory  Lorius amabilis

Stresemann

Lorius amabilis Stresemann 1931, p. 182 (Nakanai, New Britain) Status  Invalid taxon. Range  New Britain.

Known only from the unique type, a female, considered by Forshaw (1971) to be an aberrant specimen of the Purple-bellied Lory Lorius hypoinochrous.

Wilson’s Lorikeet  Vini cyanescens

(Wilson)

?Psittacus cyaneus Sparrman 1787, pl. 27 (Tahiti) Coriphilus cyaneus Wilson 1907, p. 378–9, pl. 8 (Bora-Bora) Coriphilus cyanescens Wilson 1907, p. 653 (fide Salvadori 1907) Status  Known from Wilson’s specimen and description. Putative range  Tahiti.

Generally considered to be the immature plumage of the Blue Lory Vini peruviana (Finsch 1868; Townsend and Wetmore 1919).

Bontius’s Lory  Charmosyna bontii

(Latham)

De Psittaco parvo. Bontius, 1658, p. 63 with fig Psittacus parvus Bontii. Willughby 1676, p. 81 Crested Red Parakeet. Latham 1781, p. 233, no. 25. Based on Bontius Psittacus javanicus Gmelin 1788, p. 319 Psittacus bontii Latham 1790, p. 92

388

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Appendix 2: Doubtful and invalid taxa Status  Doubtful taxon. Range  Java.

Known only from the description of Bontius. Salvadori (1891a) stated that both Wagler and G.R. Gray identify it with the Papuan Lorikeet Charmosyna papuensis.

Latham’s Lory  Charmosyna sp. Papuan Lory. var. D. Latham 1822, p. 126 (no locality) Papuan Lory, var. (= Charmosyna sp) Salvadori 1891a, p. 607 Status  Doubtful taxon. Putative range  Unknown.

Known only from Latham’s description. Never identified.

Andai Parrot  Geoffroyus dorsalis

Salvadori

Geoffroyus dorsalis Salvadori 1875, p. 758 (near Andai, New Guinea) Status  Doubtful taxon. Putative range  New Guinea.

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.

Golden Parrot  ?Geoffroyus aureus

(Shaw)

Le Perroquet d’Or. Levaillant 1805a, pl. 138 (?Brazil) Psittacus aureus Shaw 1811, p. 543 Status  Doubtful taxon. Putative range  Africa or Brazil.

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.

Scopoli’s Parrot  ?Geoffroyus simplex

(Kuhl)

Petite Perruche de l’Isle de Luçon. second espece. Sonnerat 1776, p. 76, pl. 38 upper fig. (Luzon) Psittacus leucophthalmus Scopoli 1786, p. 87 Psittacus simplex Kuhl 1820a, p. 66 Status  Doubtful taxon. Putative range  Luzon.

Known only from Sonnerat’s inadequate description. Unidentifiable.

Rufous-tailed Parrot  Tanygnathus heterurus

Salvadori

Tanygnathus heterurus Salvadori 1912, p. 328 (Celebes) Status  Doubtful taxon. Putative range  Celebes.

Known only from the unique type of unknown locality. Forshaw (1973) and Collar (2006) examined it and believed it to be an aberrant specimen of Muller’s Parrot T. sumatranus.

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Extinct Birds

Amboina Parrot  Tanygnathus sp. ?Psittacus gramineus Gmelin 1788, p. 338 (Amboina = error for Buru) Le Perroquet a calotte bleue. Levaillant 1805a, pl. 121 (Amboina) Status  Doubtful taxon. Putative range  Amboina.

Known only from Levaillant’s plate. Never satisfactorily identified.

Crimson-crowned Parakeet  ?Alisterus sp. Crimson-crowned Parrakeet. Shaw 1811, p. 419. Crimson-crowned Parrakeet. Latham 1822, p. 184 Status  Doubtful taxon. Putative range  Australia.

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.

Mrs Bligh’s Rosella  Alisterus sp. Crimson and Yellow Parrot. Latham 1822, p. 143 Status  Doubtful species. Putative range  Australia.

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.

Bastard King Parrot  Aprosmictus insignissimus

Gould

Aprosmictus insignissimus Gould 1874, p. 500 Status  Doubtful taxon. Range  Queensland.

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) identified it as a hybrid King Parrot x ­Red-­winged Parrot Aprosmictus erythropterus, while Ramsay (1878) thought it was simply an aberrant King Parrot.

Lady Tavistock’s Parrot  Barnardius crommelinae

Mathews

Barnardius crommelinae Mathews 1925, p. 21 Status  Doubtful taxon based on an aviary specimen. Putative range  Unknown.

Known only from the type, a female, now in Tring. Peters (1937) considered that it represented a specimen of Barnard’s Parrot Barnardius barnardi in which the yellow pigment is largely lacking.

Mysterious Rosella  Platycercus erythropeplus

Salvadori

Platycercus erythropeplus Salvadori 1891c, pl. 12, p. 130 Status  Invalid taxon. Putative range  Australia.

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). 390

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Appendix 2: Doubtful and invalid taxa

Fiery Rosella  Platycercus ignitus

Leadbeater

Platycercus ignitus Leadbeater 1837, p. 8 Status  Doubtful taxon. Range  Australia.

Known from two specimens in Tring. Peters (1937) concluded that it is an aberration of Eastern Rosella P. eximius.

Masters’s Rosella  Platycercus masterianus

Ramsay

Platycercus masterianus Ramsay 1877, p. 27 (‘Interior of New South Wales’) Status  Doubtful taxon. Putative range  New South Wales, Australia.

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).

Chlorotic Parrot  ?Platycercus sp. Chlorotic Parrot. Latham 1822, p. 144 Putative range  New South Wales.

Known only from Latham’s description. Finsch (1868) considered the Chlorotic Parrot to be a variety of some species of the genus Platycercus. Unidentifiable.

Sulphur Parakeet  Platycercus sp. La Perruche souffre. Levaillant 1801, pl. 43, p. 86 Sulphur Parrakeet. Shaw 1811, p. 428 Status  Invalid taxon. Putative range  Unknown.

Known from Levaillant’s plate and description. Usually considered to be a lutino (yellow variety) of the ­Rose-­ringed Parakeet Psittacula krameri.

Crimson-vented Parrot  Platycercus leverianus

(Gmelin)

Crimson vented Parrot. Latham 1781, p. 229 (East Indies or China) Psittacus leverianus Gmelin 1788, p. 322 Psittacus erythropygius Latham 1790, p. 94 Status  Doubtful taxon. Putative range  East Indies or China.

Known only from a specimen, formerly in the Leverian Museum but now lost, on which Latham’s description was based. Unidentifiable.

Paradise Rosella  Platycercus paradiseus

Russ

Platycercus paradiseus Russ 1871, p. 236 Status  Invalid taxon. Putative range  Australia.

Known only from an inadequate reference by Russ (1871), probably in error for the Paradise Parrot Psephotus pulcherrimus. 391

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Extinct Birds

Bechstein’s Horned Parrot  ? Platycercus cornutus Psittacus cornutus Bechstein 1811–12, p. 78 Status  Doubtful taxon. Putative range  Unknown.

Apparently known only from Bechstein’s description. Considered to be a Platycercus (Salvadori 1891a).

Kuhl’s Parrot  Cyanoramphus erythronotus

(Kuhl)

Psittacus erythronotus Kuhl 1820a, p. 45 (New Holland) Status  Doubtful taxon. Putative range  Unknown.

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).

Mysterious Kakariki  Cyanoramphus sp. Psittacus novae Zeelandiae var. 1. Kuhl 1820a, p. 44 Status  Doubtful taxon. Putative range  New Zealand.

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. 172). Kuhl described it as resembling ­Red-­fronted Parakeet C. novaezelandiae in all respects, except that it has a brown forehead.

Intermediate Kakariki  Cyanoramphus intermedius

Reichenow

Cyanoramphus intermedia Reichenow 1881, p. 44 Status  Doubtful taxon. Putative range  New Zealand.

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).

Rowley’s Kakariki  Cyanoramphus rowleyi

(Buller)

Platycercus rowleyi Buller 1874, p. 219 (Otago) Status  Doubtful taxon. Range  New Zealand.

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 ­Red-­fronted Parakeet on the Auckland Islands, C. novaezelandiae aucklandicus.

Red-topped Kakariki  Cyanoramphus verticalis

(Latham)

Psittacus verticalis Latham 1802, pp. 22, 369, no. 9 (Port Jackson, New Holland = Australia) Status  Doubtful taxon. Putative range  Australia.

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. 392

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Appendix 2: Doubtful and invalid taxa

Red-hooded Kakariki  Cyanoramphus capitatus

(Shaw)

Lori Perruche de la Mer du Sud. Virey 1805, p. 162 (‘One of the South Sea Islands’) Psittacus capitatus Shaw 1811, p. 466 Red-hooded Parrot. Latham 1822, p. 136 Status  Doubtful species. Putative range  South Pacific.

Known only from Virey’s account (in Sonnini 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.

Large-billed Kakiriki  ‘Psittacus’ magnirostris

Forbes & Robinson

Cyanoramphus magnirostris Forbes & Robinson 1897, p. 21 (Tahiti) Status  Doubtful taxon. Putative range  Provenance uncertain. Tahiti almost certainly in error.

Known only from the type in the Liverpool Museum. Synonymised with the ­Black-­fronted Parrot C. zealandicus by Peters (1937). Considered conspecific with C. cooki of Norfolk Island (Sims 1953).

Lineated Parrakeet  ‘Psittacus’ lineatus

Linnaeus

Psittacus lineatus Linnaeus 1768, Appendix p. 223 (no locality) Lineated Parrakeet. Latham 1781, p. 252 Status  Doubtful taxon. Putative range  Unknown.

Known only from Linnaeus’s description with provenance unknown.

Peregrine Parrot  ‘Psittacus’ peregrinus

Latham

Psittacus peregrinus Latham 1790, p. 105 (South Sea Islands) Peregrine Parrot. Latham 1802, p. 62 Putative range  Unknown.

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.

Black-necked Parakeet  ‘Psittacus’ nigricollis

(Latham)

Psittacus nigricollis Latham 1802, supplement 2, p. xxii Black-necked Parrakeet. Latham 1802, p. 90 Status  Doubtful taxon. Putative range  Brazil.

Known only from Latham’s description. Salvadori (1891a) stated that ‘Dr. Finsch thinks this is an artificial specimen’.

Pale Parakeet  ‘Psittacus’ pallidus

Shaw & Nodder

Psittacus pallidus Shaw & Nodder 1796, pl. 258 and text (Australia) Pale Parrakeet. Latham 1822, p. 145 Status  Doubtful taxon. Putative range  Australia.

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. 393

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Extinct Birds

Japanese Parakeet  ‘Psittacus’ japonicus

Linnaeus

Psittacus erythrochlorus macrourus Aldrovandus 1599, p. 678, p. 681 Psittaca japonensis Brisson 1760, p. 362 Psittacus japonicus Linnaeus 1766, p.141 (Japan) Perruche verte et rouge. Buffon 1779, p. 159 Japanese Parrot. Latham 1781, p. 209 Status  Invalid taxon. Putative range  Japan.

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).

Marcgrave’s Parrot  ?Psittacus paraguanus

(Gmelin)

Paragua. Marcgrave 1648. p. 207 Lorius brasiliensis. Brisson 1760, p. 229 Paraguayan Lory. Latham 1781, p. 277 Psittacus paraguanus Gmelin 1788, p. 336 Status  Doubtful taxon. Putative range  Africa or Brazil.

Known only from Marcgrave’s description, which has been the subject of discussion but never satisfactorily identified. Provenance uncertain.

Red-winged Parrot  Psittacus vaillanti

Rüppell

Psittacus levaillantii Rüppell 1835, p. 63 Pionus vaillanti Rüppell 1845, p. 94 Status  Doubtful taxon. Putative range  Ethiopia.

Known only from Rüppell’s inadequate description.

Red-and-white Parrot  Psittacus erythroleucus

Linnaeus

Psittacus erythroleucos. Aldrovandus 1599, pp. 675, 677 (no locality) Red and White Parrot of Aldrovandus. Willughby 1678, p. 144 Cacatua alis et cauda rubris. Brisson 1760, p. 214. Based on Aldrovandus. Psittacus erythroleucus Linnaeus 1766, p. 144 Red and White Parrot. Latham 1785, p. 260 Psittacus erithacus var [beta]. Salvadori 1891a, p. 378 Status  Doubtful taxon. Putative range  Unknown.

Known only from descriptions derived from Aldrovandus. Generally considered to be a variant of the African Grey Parrot Psittacus erithacus.

Crested Red-and-green Parakeet  ?Psittacus erythrochlorus

(Gmelin)

Psittacus erythrochlorus cristatus. Aldrovandus 1599, p. 682 (with figure) Psittacula cristata. Willughby 1676, p. 78 Crested red and green Parrakeet. Latham 1781, p. 321 Psittacus erythrochlorus Gmelin 1788, p. 352 Status  Doubtful taxon. Putative range  Unknown.

Probably known only from Aldrovandus’s description, from which all others derive. Salvadori (1891a) dismissed it as ‘a fabulous bird’. 394

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Appendix 2: Doubtful and invalid taxa

Nooffy Parakeet  ‘Psittacus’ sp. Noofy Parrakeet. Latham 1822 (Nooffy, interior of Africa) Status  Invalid taxon. Putative range  Africa.

Known only from Latham’s inadequate account, based on Lyon (1821).

Cayenne Parakeet  ?Psittacus sosove

(Gmelin)

Petite Perruche de Cayenne. Daubenton 1771–86, no. 456, fig. 2 Psittacus cajenneus Müller 1776b, supplement, p. 81 Le Sosové. Buffon 1779, p. 280 Cayenne Parrakeet. Latham 1781, p. 320 Psittacus notatus Boddaert 1783, p. 27 (based on Daubenton) Psittacus sosove Gmelin 1788, p. 352 Conurus tuipara var. notatus Gray 1859b, p. 46 Conurus tuipara var. cayenneus Gray 1870, p. 150 Status  Doubtful taxon. Putative range  French Guiana.

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.

Short-­tailed Parakeet  ‘Psittacus’ sp. Short-tailed Parrakeet. Latham 1822, p. 274 (no locality) Status  Doubtful taxon. Putative range  Unknown.

Known only from Latham’s description. Unidentifiable.

Taraba Parrot  ‘Psittacus’ taraba

Gmelin

Tarabe brasiliensibus. Marcgrave 1648, p. 207 Psittacus Brasiliensis erythrocephalus. Brisson 1760, p. 240 La Maracana à tête rouge. Azara 1809 Tarabé ou Amazone à tête rouge. Buffon 1779, p. 211 Red-headed Parrot. Latham 1781, p. 295 Psittacus taraba Gmelin 1788, p. 344 Status  Known only from the accounts of Marcgrave and Azara. Putative range  Brazil.

Unidentifiable.

Blue-­faced Parakeet  ‘Psittacus’ sp. Blue-faced Parrakeet. Latham 1822, p. 271 (no locality) Status  Doubtful taxon. Putative range  Unknown.

Known only from Latham’s description. Unidentifiable.

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Extinct Birds

Crimson-­banded Parakeet  ‘Psittacus’ sp. Crimson-banded Parrot. Latham 1822, p. 254 (no locality) Status  Doubtful taxon. Putative range  Unknown.

Known only from Latham’s description. Unidentifiable.

Dubious Parrot  ‘Psittacus’ dubius

Latham

Dubious Parrot. Latham 1787, p. 62 Psittacus dubius Latham 1790, p. 97 Status  Doubtful taxon. Putative range  Unknown.

Known only from Latham’s description of a specimen formerly in the British Museum. Unidentifiable.

Ruby-capped Parrot  Poicephalus rubricapillus

(Forbes & Robinson)

Poeocephalus rubricapillus Forbes & Robinson 1898, p. 15 (West Africa) Status  Invalid taxon. Range  West Africa.

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).

Heuglin’s Parrot  ‘Poicephalus’ sp. Palaeornis? sp. Heuglin (on the authority of Finsch 1868, p. 933) Status  Doubtful taxon. Putative range  East Africa.

Known only from Heuglin’s inadequate description. Finsch (1868) and Salvadori (1891a) both suggested that it was conspecific with the Senegal Parrot Poicephalus senegalus.

Green-­crested Parrot  ‘Poicephalus’ sp. Mittelgrosser Papagei ‘grün mit grosser Haube’. Heuglin (fide Finsch 1868) (Kitsch-Neger Land, Africa) Status  Invalid taxon. Putative range  Sudan, North Africa.

Based on an inadequate description.

David Livingstone’s Parrot  ‘Poicephalus’ sp. Parrot. Livingstone 1857, p. 239 (River Leeambye, South Africa) Papagei. Finsch 1868, p. 932 Status  Doubtful taxon. Putative range  South Africa.

Known only from David Livingstone’s inadequate account.

Hecquard’s Parrot  ‘Poicephalus’ sp. Grün Papagei mit rothem Kopfe. Hecquard 1854, p. 242; id. Finsch 1868, p. 932 Status  Invalid taxon. Putative range  West Africa.

Based on an inadequate description. 396

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Appendix 2: Doubtful and invalid taxa

Alexander’s Parrot  Poicephalus kintampoensis

(Alexander)

Poeocephalus kintampoensis Alexander 1901, p. 10 (Kintampo, Gold Coast = Ghana) Status  Doubtful taxon. Range  Kintampo, Ghana.

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.

Leona Lovebird  Agapornis picta

Hartlaub

Leona Parrakeet. Latham 1822, p. 263 Poliopsitta picta Hartlaub MS in Bonaparte 1856b, no. 227 (nomen nudum) Agapornis picta Hartlaub 1857, p. 169 Status  Known only from Latham’s description. Putative range  Sierra Leone.

Both Hartlaub (1879) and Salvadori (1891a) considered this bird to be probably the same as the Blackcollared Lovebird Agapornis swinderniana.

Red-naped Parrot  ?Loriculus cervicalis

(Latham)

Red-naped Parrakeet. Latham 1787, p. 66 Psittacus cervicalis Latham 1790, p. 130 Le Perroquet Langlois. Levaillant 1805a, p. 105, pl. 136 Psittacus nuchalis Shaw & Nodder 1789–1813, pl. 913 (based on Levaillant) Psittacus erythrogaster Bechstein 1811–12, p. 102, pl. 19 (based on Levaillant) Psittacus Langloisi Vieillot 1817, p. 312 Eos cervicalis Wagler 1832, p. 561 Status  Doubtful taxon. Putative range  Unknown.

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’.

Salvadori’s Hanging Parrot  Loriculus salvadorii

Hachisuka

Loriculus apicalis Gray 1859, p. 56, specimen ‘a’ (Mindanao) ? Coryllis hartlaubi (part) Finsch 1868, p. 712, p. 721 Loriculus sp. Salvadori 1891a, p. 530 footnote Loriculus salvadorii Hachisuka 1930, p. 163 (Mindanao) Status  Invalid Taxon. Range  Mindanao, Philippines.

Known from two specimens. It is now considered to be an aberration of Loriculus philippensis apicalis (Peters 1937; Forshaw 1989; Juniper & Parr 1998).

Edwards’s Long-tailed Green Parakeet  Psittacula rufirostris

(Linnaeus)

Long-tailed Green Parrakeet. Edwards 1751, p. 175, pl. 175 (West Indies) ? Psittace torquata. Brisson 1760, p. 323 Psittacus rufirostris Linnaeus 1766, p. 143 (female) Status  Doubtful taxon. Putative range  West Indies?

Known only from Edwards’s description. Salvadori (1891a) placed it in the synonymy of the ­Ring-­necked Parakeet Psittacula krameri. 397

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Extinct Birds

Jonquil Parakeet  Psittacula narcissus

(Latham)

Psittacus narcissus Latham 1801b, supplement, p. 21 Jonquil Parrakeet. Latham 1802, supplement, p. 83, pl. 123 Status  Invalid taxon. Putative range  Bengal, India.

Known only from Latham’s description. Salvadori (1891a) called this bird a lutino (yellow) variety of the ­Plum-­headed Parakeet Psittacula cyanocephala.

Blue-headed Parakeet  Psittacula indica

(Latham)

Blue-headed Parrakeet. Edwards 1760, p. 175, pl. 292, lower figure Psittacus indicus Latham 1790, p. 86 La Perruche à épaulettes jaunes. Levaillant 1801, pl. 61 Psittacus xanthosomus Bechstein 1812, p. 74 (based on Levaillant) Psittacus ternatensis Vieillot 1817, p. 346 (based on Levaillant) Status  Invalid taxon. Putative range  East Indies.

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.

Intermediate Parakeet  Psittacula intermedia

Rothschild

Psittacula intermedia Rothschild 1895b, p. 492 Status  Invalid taxon. Range  Northern India.

This parakeet was known from a few specimens and its status was surrounded in mystery. It is now considered to be a captive-bred hybrid between the ­Slaty-­headed Parakeet Psittacula himalayana and the ­Plum-­headed Parakeet P. cyanocephala (Rasmussen & Collar 1999a, b).

Mauritian Parrot  Necropsittacus francicus

Rothschild

Necropsittacus francicus Rothschild 1905c, p. 197 Nomen nudum Hume 2007, p. 35 Status  Invalid taxon. Putative range  Mauritius.

Known only from Rothschild’s account based on Dubois’s of 1674, in which Rothschild completely confused the provenance.

Humboldt’s Macaw  Cyanopsitta sp. Ara sp. Humboldt 1860, p. 6 Status  Doubtful taxon. Putative range Rio Casquiare, Orinoco basin, South America.

Known only from Humboldt’s inadequate description.

Black Macaw  Anodorhynchus ater

(Gmelin)

Ararauna ou Machao. De Laet 1630, p. 490 (Guiana) Black Macaw. Latham 1781, p. 206 Psittacus ater Gmelin 1788, p. 314 (based on Latham). Status  Invalid taxon. Putative range  Guiana.

Based on inadequate descriptions. 398

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Appendix 2: Doubtful and invalid taxa

Violet Macaw  Anodorhynchus purpurascens

Rothschild

? Le Gros Perroquet de la Guadloupe. Don de Navaret 1838, p. 425 Anodorhynchus purpurascens Rothschild 1905b, p. 13 (Guadeloupe Island, Lesser Antilles) Status  Invalid taxon. Putative range  Guadeloupe, West Indies.

The Violet Macaw is known only from Rothschild’s 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 – its status must be considered dubious.

Orange-­bellied Macaw  Ara martinica

(Rothschild)

Les Aras. Bouton 1640, p. 71–72 Anodorhynchus martinicus Rothschild 1905b, p. 14. (Martinique) Ara martinicus Rothschild 1907a, pl. 14 Ara erythrura Rothschild 1907a, pl. 15 (‘One of the West Indian Islands’) Status  Doubtful taxon. Putative range  Martinique, West Indies.

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.

Satin Macaw  Ara erythrura

Rothschild

Des Arras. De Rochfort 1658, p. 154 Anodorhynchus coeruleus Rothschild 1905b, p. 15 Ara erythrura Rothschild 1907a, Extinct Birds, p. 54, pl. 15 (‘One of the West Indian Islands’) Status  Invalid taxon. Putative range  One of the West Indian islands.

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).

Red-­headed Macaw  Ara erythrocephala

Rothschild

Ara militaris Gosse 1847, p. 261 (Jamaica) Ara erythrocephala Rothschild 1905b, p. 14 (Mountains of Trelawny and St. Anne’s, Jamaica) Status  Invalid taxon. Putative range  Jamaica.

Based on one hearsay observation only (see Greenway 1967). Only one species of Jamaican macaw, the Jamaican ­Yellow-­headed Macaw A. gossei (see p. 335) 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 ornithologists without a shred of physical evidence, epitomises the ­over-­zealous passion often associated with extinct birds.

Jamaican ­Blue-­and-­yellow Macaw  Ara sp. Psittacus maximus cyanocroceus. Sloane 1725, p. 296 The blue Mackaw of Edwards. Browne 1756 Ara jamaicensis cyanocrocea. Browne 1789, p. 472 Ara sp? Gosse 1847, p. 261

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Extinct Birds Status  Doubtful taxon. Putative range  Jamaica.

Almost certainly relates to an escaped, mainland ­Blue-­and-­yellow Macaw A. ararauna.

Albin’s Macaw  Ara sp. The Maccaw from Jamaica. Albin 1738, p. 16, pl. 17 Status  Doubtful taxon. Putative range  Jamaica.

Known only from Albin’s (1738) description. Almost certainly referable to the ­Red-­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.

Hispaniolan Macaw  Ara tricolor haitius

Hoppe

L’Ara Rouge. Buffon 1779 p. 183 Ara tricolor (part) Clark 1905c, d, p. 337, 348 (Haiti) Macaw. Rothschild 1905b, p. 14 Ara tricolor haitius Hoppe 1983 (Hispaniola) Status  Invalid taxon. Putative range  Hispaniola.

Known only from Buffon’s and other early accounts. Records for Hispaniola are in error (see Walters 1995; Olson 2005).

Seaforth’s Macaw Ara sp. Red and Yellow Macaw. Latham 1822, p. 107, pl. 21 Ara macao (part) Clark 1905a, p. 271 Status  Invalid taxon. Putative range  Trinidad.

Based on a good description by Latham (1822) but almost certainly referable to a colour variety of Scarlet Macaw Ara macao (Finsch 1867).

Marcgrave’s Macaw  Ara sp. Maracana II. Marcgrave 1648, p. 207 Brasilian Green Macaw. Edwards 1758, p. 41, pl. 229 Status  Doubtful taxon. Putative range  Brazil.

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.

Marcgrave’s Parakeet  Aratinga sp. Tui. prima species. Marcgrave 1648, p. 207 Status  Invalid taxon. Putative range Brazil.

Known only from a vague, inadequate account of Marcgrave.

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Appendix 2: Doubtful and invalid taxa

Azara’s Parakeet  Aratinga flaviventris

(Wagler)

Maracana cabeza y encuentro roxos. Azara 1805, p. 418 Maracana à tete et pli de l’aile rouges. Azara 1809, p. 59 Sittace flaviventris Wagler 1832 p. 658 (based on Azara) Status  Doubtful taxon. Putative range  Paraguay.

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.

Guaicaramo ­White-­eyed Parakeet  Aratinga leucophthalma nicefori

Schauensee

Meyer de

Aratinga leucophthalma nicefori Meyer de Schauensee 1946, p. 2 Status  Invalid taxon. Putative range  Meta, Colombia.

This supposed subspecies is known from one individual. Juniper & Parr (1998) regard it as an aberrant A. leucophthalma callogenys.

Whitley’s Parakeet  Cyanoliseus whitleyi

(Kinnear)

Aratinga whitleyi Kinnear 1926, p. 82 (based on a living specimen ‘probably from South America’) Status  Doubtful taxon. Putative range  South America.

Known only from the type, of unknown origin, now in Tring. Peters (1937) considered it a hybrid, as did Forshaw (1989), who thought it was between the Burrowing Parakeet Cyanoliseus patagonus and ­Scarlet-­fronted Parakeet Aratinga wagleri.

‘The conure that never was’  Pyrrhura beryllina

Bonaparte

Pyrrhura beryllina ? Masséna & Souancé, on the authority of Bonaparte 1856, p. 352 Status  Invalid taxon.

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.

Yellow-sided Parakeet  Pyrrhura hypoxantha

Salvadori

Pyrrhura hypoxantha Salvadori 1899, p. 1 (Urucúm, Matto Grosso) Status  Invalid taxon. Putative range  Matto Grosso, Brazil.

Known from four specimens (according to Peters 1937) or three specimens (according to Forshaw 1989), all from western Matto Grosso. Now considered to be an aberrant ­Green-­cheeked Parakeet Pyrrhura molinae sordida (Juniper & Parr 1998).

Bechstein’s Parrot  Myiopsitta buccalis

(Bechstein)

La Perruche à joues grises. Levaillant 1801, p. 188, pl. 67 Psittacus buccalis Bechstein 1811–12, p. 83, pl. 10 Psittacus cinereicollis Vieillot 1817, p. 365 Status  Invalid taxon. Putative range French Guiana.

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Extinct Birds

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.

Blue-backed Parrot  Forpus sp. Blue-backed Parrot. Latham 1822, p. 272 (Brazil) Status  Doubtful taxon. Putative range  Brazil.

Known only from Latham’s description. May represent a Forpus sp., otherwise unidentifiable.

Edwards’s Parakeet  Brotogeris sp. Golden-winged Parrakeet. Edwards 1760, p. 177, pl. 293, fig. 2 (East Indies = error) Status  Known only from Edwards’s description and plate. Putative range  East Indies.

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.

Yellow-breasted Caique  ?Pionites guineensis

(Miller)

Psittacus guineensis Miller 1779, pl. 29 (Guinea) Status  Invalid taxon. Putative range  East Indies.

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.

Cobalt Parrot  Pionus cobaltinus

(Masséna & Souancé)

Psittacus cobaltinus Masséna & Souancé 1854, p. 74. (Colombia) Pionus maximiliani (part) Souancé 1856, p. 155 Pionus cobaltinus Salvadori 1891a, p. 325 Status  Known from two specimens, one of which is a juvenile. Putative range  Colombia.

Probably ­Blue-­headed Parrot P. menstruus reichenowi (Salvadori 1891a).

Dominican Macaw  Amazona atwoodi

Clark

Mackaw. Atwood 1791, p. 29 Ara atwoodi Clark 1908b, p. 310 (Dominica) Status  Doubtful taxon. Putative range  Dominica, West Indies.

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’.

402

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Appendix 2: Doubtful and invalid taxa

Gerini’s Amazon  Amazona gerini

(Latham)

Psittacus Brasiliensis viridis capite albo. Manetti 1767, p. 95, pl. 109 Psittacus gerini Latham 1790, p. 119 Gerini’s Parrot. Latham 1802, p. 93, no. 23 ? White-faced Amazon Parrot. Shaw 1811, p. 500 Status  Known only from Manetti’s plate, which agrees with no known parrot. Putative range  Brazil.

Unidentifiable but Finsch (1868) suggests a variety of ­Yellow-­billed Parrot Amazona collaria.

Red-­and-­blue Amazon  Amazona caeruleocephala

(Linnaeus)

Psittacus versicolor seu erythrocyanus. Aldrovandus 1599, p. 675 Psittacus versicolor. Willughby 1676, p. 75 Psittacus violaceus. Barrère 1745, p. 26 Psittacus Guianensis caeruleus. Brisson 1760, p. 304 Psittacus caeruleocephalus Linnaeus 1766, p. 145 Le crick rouge et bleue. Buffon 1779 p. 226 Red and Blue Parrot. Latham 1781, p. 277 Status  Doubtful taxon. Putative range  Guiana.

Known only from the original description of Aldrovandus, from which all others derive. Provenance in doubt.

Heck’s Parrot  Amazona hecki

(Reichenow)

Androglossa hecki Reichenow 1891b, p. 217 (Colombia?) Status  Doubtful taxon. Putative range  Colombia?

Known only from Reichenow’s description and coloured plate. The sole specimen, a cagebird, seems to have disappeared. Peters (1937), following Salvadori (1891a), synonymised A. hecki with race lilacina of the ­Red-­lored Parrot Amazona autumnalis.

Red-­and-­yellow Amazon  Amazona sp. ? Le Perroquet jaune de Cuba. Brisson 1760, p. 308 ? Perroquet de Cuba. Daubenton 1771–86, no. 336 ? Le Papagay de Paradis. Buffon 1771–86, p. 334 Le Perroquet Jaune écaillé de rouge, en Le Perroquet de Cuba. Levaillant 1805, pl. 137 Status  Doubtful taxon. Putative range  Cuba?

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.

Lilac-bellied Amazon  Amazona sp. Le Perroquet à face bleue. Levaillant 1805, pl. 122 Status  Doubtful taxon. Putative range  Mexico?

Known only from Levaillant’s description. Provenance in doubt.

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Extinct Birds

Paradise Amazon  Amazona paradisi

(Linnaeus)

The Parrot of Paradise. Catesby 1754, p. 10, pl. 10 (the plate is lettered ‘Psitticus Paradisis’) Psittacus luteus Insulae Cubae. Brisson 1760, p. 308 Psittacus paradisi Linnaeus 1758, p. 101 Paradise Parrot. Shaw 1811, p. 501, pl. 75 Status  Doubtful taxon. Putative range  Cuba.

Known only from the plate and description of Catesby, based on a live specimen he saw in Carolina. Identified by Salvadori (1891) as a leucistic specimen of Cuban Parrot Amazona leucocephalus.

Primeval Amazon  ?Amazona prisca

(Sclater)

Chrysotis prisca Sclater 1861b, p. 79 Status  Invalid taxon. Putative range  West Indies.

The evidence for this species is spurious at best, and was considered extremely doubtful by Salvadori (1891a).

Finsch’s Amazon  Amazona chloris

(Finsch)

Neuer Amazonen–Papagei. Sacc 1864, p. 21 Chrysotis chloris Finsch 1868, p. 930 Status  Known only from Finsch’s inadequate description. Putative range  Unknown.

Never identified.

Levaillant’s Black Coucal  Centropus maurus

(Stephens)

Le Coucal negre. Levaillant 1806b, p. 63, pl. 222 Poliophilus maurus Stephens 1815, p. 57 Centropus aethiops Cuvier 1817, p. 426 Corydonix nigerrimus Vieillot 1819, p. 297 Status  Doubtful taxon. Putative range  South Africa.

Known only from Levaillant’s plate and figure, on which the other references are based. Never satisfactorily identified and now unidentifiable.

Cave-­nesting Masked Owl  Tyto novaehollandiae troughtoni

Cayley

Tyto novae–hollandiae troughtoni Cayley, 1931, p. 32, fig. 4 (Ooldea, South Australia) Status  Doubtful taxon. Range  Nullarbor Plain, South Australia.

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.

Commerson’s Owl  Scops commersoni

(Oustalet)

Scops commersoni Oustalet 1896, p. 35, fig. 3 (Mauritius) Status  Invalid taxon. Range  Mauritius, Mascarenes.

Now synonomised with Mascarenotus sauzieri ­(Mourer-­Chauviré et al. 1994). 404

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Appendix 2: Doubtful and invalid taxa

Newton’s Owl  Strix newtoni

(Rothschild)

Strix sp. Newton & Gadow 1893, p. 287 (Mauritius) Strix newtoni Rothschild 1907a, p.79 Status  Invalid taxon. Range  Mauritius, Mascarenes.

Now synonomised with Mascarenotus sauzieri ­(Mourer-­Chauviré et al. 1994).

Vieillot’s Wood Owl  Strix fasciata

Vieillot

Strix fasciata Vieillot 1817, p. 21 (Martinique) Status  Doubtful taxon. Putative range  Martinique, West Indies.

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.

China Owl  Strix orientalis

Shaw

China Owl. Latham 1802, p. 368 (China) Strix sinensis Latham 1801b, supplement p. 16 Strix orientalis Shaw 1809, p. 257 (China) Status  Doubtful taxon. Putative range  China.

Known only from Latham’s description of a specimen, now lost. Not identifiable with any known species (Peters 1940).

Vieillot’s Spotted Owl  Bubo maculosus

(Vieillot)

Strix maculosa Vieillot 1817, p. 44 (Cape of Good Hope) Status  Doubtful taxon. Putative range  South Africa.

Known only from Vieillot’s description. Never identified, but the name was formerly used for the Spotted Eagle Owl Bubo africanus.

La Touche’s Eagle Owl  Bubo bubo jarlandi

La Touche

Bubo bubo jarlandi La Touche 1921, p. 14 (Mountains near Mengtz, Yunnan) Status  Doubtful taxon. Putative range  Yunnan, China.

Known only from the unique type, a female with traces of downy juvenile feathers on the head. Peters (1940) doubted its validity, and Vaurie (1960b) merged this race and another in Bubo bubo kiautschensis.

Wapacuthu Owl  Nyctea wapacuthu

(Gmelin)

Wapacuthu Owl. Pennant 1785, p. 231 Strix wapacuthu Gmelin 1788, p. 290 (based on Pennant) Status  Doubtful taxon. Putative range  Hudson’s Bay, Canada.

Known only from Pennant’s description. McAtee (1963) considered it synonymous with the Snowy Owl Bubo scandiaca. 405

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Extinct Birds

Kaup’s Owl  Pseudoscops macrurus

(Kaup)

Otus macrurus Kaup 1859, p. 232 (Mexico) Status  Doubtful taxon. Putative range Mexico?

Known only from the type, now lost, stated to have been in the Würtzburg Museum. Considered unidentifiable (Peters 1940).

Krusenstern’s Japanese Owl  Strix sp. Japanese Owl. Hartert 1916, p. 94 Status  Doubtful taxon. Putative range  Japan.

Known only from Krusenstern’s (1814) plate. Never satisfactorily identified.

Rufous Owl of Latham  Gisella lathami

(Bonaparte)

Rufous Owl. Latham 1821, p. 375 (no locality) Strix lathami Bonaparte 1830, p. 56 Gisella lathami Bonaparte 1854b, p. 541 (designation of the genus Gisella) Status  Doubtful taxon. Putative range  Unknown.

Known only from Latham’s description. Peters (1940) stated that is was practically unidentifiable.

Bartram’s Owl  ?Strix peregrinator

(Bartram)

Strix peregrinator Bartram 1793, p. 285 (Pennsylvania, Carolina and Florida) Status  Doubtful taxon. Putative range  Eastern United States.

Known only from Bartram’s inadequate description.

Carriker’s Owl  Otus minimus

(Carriker)

Ciccaba minima Carriker 1935, p. 313 (Santa Ana, Rio Coroico, Department La Paz, Bolivia) Status  Doubtful taxon. Putative range  Bolivia.

Known only from the type, an adult male. Bond (1951) and Traylor (1952) considered this bird synonymous with Rufescent Screech Owl Otus ingens.

Stephenson Clarke’s Nightjar  Caprimulgus ludovicianus

Stephenson Clarke

Caprimulgus ludovicianus Stephenson Clarke 1913, p. 108 (S.W. Ethiopia) Status  Invalid taxon. Putative range ­South-­west Ethiopia.

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).

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Appendix 2: Doubtful and invalid taxa

Elliot’s Topaz  Crinis chlorolaema

(Elliot)

Chrysolampis chlorolaema Elliot 1870, p. 346 (Colombia = error for Dominica?) Lampornis calosoma Elliot 1872b, p. 351 Crinis chlorolaema Mulsant & Verreaux 1874, p. 177 Status  Doubtful taxon. Putative range  Colombia or Dominica?

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) considered it to be a hybrid ­Black-­throated Mango Anthracothorax nigricollis x Ruby Topaz Chrysolampis mosquitus.

Lesson’s Amethyst  Calliphlox orthura

(Lesson)

Trochilus orthura Lesson 1832, p. 85, 88, (Cayenne) Ornismya amethystoides Lesson 1832, p. 79, pl. 25 C[alliphlox] Roraimae Boucard 1891, p. 30 Status  Doubtful taxon. Putative range  French Guiana.

Known from the type, an immature male collected by Longuemare, and considered an immature Amethyst Woodstar C. amethystina (Schuchmann 1999).

Sooty Hermit  Phaethornis anthophilus fuliginosus

Simon

Phaethornis fuliginosus Simon 1901a, p. 201 (Colombia?) Status  Doubtful taxon. Putative range  Colombia?

Known only from Bogotá trade skins. Considered a melanistic bird of doubtful origin (Schuchmann 1999).

Porculla Hermit  Phaethornis porcullae

Carriker

Phaethornis porcullae Carriker 1935, p. 345 (Porculla Pass, Department of Lambayeque, Peru) Status  Invalid taxon. Putative range  Peru.

Known only from two females. Considered invalid (SACC 2007).

Schlüter’s Hermit  Phaethornis fumosus

(Schlüter)

Phaetornis fuliginosus Schlüter 1913, p 32 (Bogotá, Colombia) Phaetornis fumosus Schlüter 1915, p. 21 Status  Invalid taxon. Putative range  Bogotá, Colombia.

Known only from the type. Now considered a melanistic specimen of an unknown hermit (Hinkelmann 1999).

Gould’s Hermit  Phaethornis zonura

Gould

Phaëthornis zonura Gould 1860, p. 305 Status  Doubtful taxon. Putative range  Peru.

Known only from the type. Affinities uncertain, generally regarded as conspecific with the ­Grey-­chinned Hermit P. griseogularis (Peters 1945). 407

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Extinct Birds

Heine’s Hermit  Phaethornis apheles

Heine

Phaethornis apheles Heine 1884, p. 235 (Peru) Status  Doubtful taxon. Putative range  Peru.

Known only from the type, an imperfect specimen. Considered a synonym of P. zonura (Peters 1945), which itself is now a subspecies of ­Grey-­chinned Hermit P. griseogularis.

Laugier’s Sabrewing  Campylopterus lazulus

(Vieillot)

Trochilus lazulus Vieillot 1822, p. 557 Status  Doubtful taxon. Putative range  Central America.

Known only from Vieillot’s description. Never identified.

Buckley’s ­Violet-­ear  Colibri buckleyi

(Gould)

Pinarolaema buckleyi Gould 1880, p. 489. (Misqui, Bolivia) Status  Invalid taxon. Putative range  Ecuador.

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).

Elliot’s ­Violet-­ear  Colibri rubrigularis

(Elliot)

Petasophora sp. Elliot 1876b, p. 405 Petasophora rubrigularis Elliot 1878 p. 51 footnote (no locality) Status  Invalid taxon. Putative range  Unknown.

Known only from a specimen in the Elliot Collection. Now considered a hybrid ­Black-­throated Mango Anthracothorax nigricollis x Ruby Topaz Chrysolampis mosquitus (Peters 1945).

Steel-­blue-­tailed Mango  Anthracothorax caeruleicauda

(Floericke)

Lampornis nigricollis caeruleicauda Floericke 1920, p. 3 (Colombia) Status  Doubtful taxon. Putative range  Colombia.

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).

Giglioli’s Topaz  Chrysolampis gigliolii

Oustalet

Chrysolampis Gigliolii Oustalet 1885, p. 108 (Colombia). Status  Invalid taxon. Putative range  Colombia.

Known only from the type. Generally considered to be an artefact (Simon & Hellmayr 1908).

Dusky Coquette  Lophornis melaniae

Floericke

Lophornis melaniae Floericke 1920, p. 2 Status  Invalid taxon. Putative range  Central America.

Known only from Floericke’s account, based on specimens in his collection, now lost. 408

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Appendix 2: Doubtful and invalid taxa

Bearded Coquette  Lophornis insignibarbis

Simon

Lophornis insignibarbis Simon 1890, p. 17 (Colombia) Status  Invalid taxon. Putative range  Colombia.

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).

Elliot’s Sapphire  Chlorostilbon subcaerulea

(Elliot)

Eucephala subcaerulea Elliot 1874a, p. 87 (Brazil?) Status  Doubtful taxon. Putative range  Brazil?

Known only from a unique specimen in the Elliot Collection. Considered an aberrant ­Blue-­chinned Sapphire Chlorostilbon notatus (Schuchmann 1999).

Blue-­breasted Sapphire  Chlorostilbon hypocyanea

(Gould)

Eucephala hypocyanea Gould 1860a, p. 306. (Brazil) Status  Invalid taxon. Putative range  Bahia, Brazil.

Known only from two specimens, both ‘Bahia’ trade skins. Considered an aberrant ­Blue-­chinned Sapphire Chlorostilbon notatus or a hybrid C. notata x ­White-­chinned Sapphire Hylocharis cyanus (Schuchmann 1999).

Brilliant Emerald  Chlorostilbon micans

(Salvin)

Panychlora micans Salvin 1891, p. 375 (no locality) Status  Doubtful taxon. Putative range  Unknown.

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).

Wied’s Emerald  Chlorostilbon wiedi

Boucard

Chlorostilbon wiedi Boucard 1895, p. 120. (Brazil, type a Rio trade skin) Status  Doubtful taxon. Putative range  Brazil.

Known only from the type series, and considered a synonym of the eastern Brazilian race of ­Glittering-­bellied Emerald, Chlorostilbon aureoventris pucherani (Cory 1918; Peters 1945).

Berlepsch’s Emerald  Chlorostilbon inexpectatus

(Berlepsch)

Panychlora inexpectatus Berlepsch 1879, p. 63 (Bogotá) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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).

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Extinct Birds

Cabanis’s Emerald  Chlorostilbon auratus

(Cabanis & Heine)

Panychlora aurata Cabanis & Heine 1860, p. 50 (Peru) Status  Doubtful taxon. Putative range  Peru.

Known only from the type in the Museum Heine. Probably an aberrant ­Short-­tailed Emerald C. poortmani (Schuchmann 1999), which would make the locality of Peru in error.

Iridescent Emerald  Smaragdochrysis iridescens

(Gould)

Calliphlox? iridescens Gould 1860, p. 310 (Nova Friburgo, Rio de Janeiro, Brazil) Smaragdochrysis Gould 1861, pl. 13 and pl. 359 Status  Invalid taxon. Putative range  Rio de Janeiro, Brazil.

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; Graves 1999b).

Natterer’s Emerald  Ptochoptera iolaima

(Reichenbach)

Trochilus iolaimus Natterer 1840 (on authority of Reichenbach) Chlorestes [delta] Riccordia iolaimus Reichenbach 1854, p. 8. (Brazil). Chlorestes iolaimus Reichenbach 1855a, p. 4 (Brazil) Ptochoptera Elliot 1874b, p. 261 Status  Doubtful taxon. Putative range  São Paulo, Brazil.

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 undetermined species.

Lerch’s Woodnymph  Thalurania lerchi

Mulsant & Verreaux

Talurania [sic] Lerchi Mulsant & Verreaux 1872, p. 108 (Colombia) ? Agyrtria tenebrosa Hartert 1899, p. xv (Bogotá, Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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 ­Golden-­tailed Sapphire Chrysuronia oenone (Schuchmann 1999).

Berlioz’s Woodnymph  Augasma cyaneoberillina

Berlioz

Augasma cyaneo–berillina Berlioz 1965, p. 7 (Bahia) Status  Doubtful taxon. Putative range  Bahia, South America.

Known only from two Bahia trade skins. Considered a hybrid, with parents unknown (Meyer de Schauensee 1966, 1970).

Southern Woodnymph  Augasma chlorophana

Simon

Augasma chlorophana Simon 1897, p. 20 (no locality) Status  Doubtful taxon. Putative range  Unknown.

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Appendix 2: Doubtful and invalid taxa

Known only from the unique type. Thought to be the female of the Emerald Woodnymph Augasma smaragdinea (Simon & Hellmayr 1908), itself considered to be a hybrid (see below). Berlioz (in Peters 1945) considered it a hybrid.

Emerald Woodnymph  Augasma smaragdinea

Gould

Augasma smaragdineum Gould 1860, p. 305 (Nova Friburgo, Rio de Janeiro) Eucephala ­smaragdo-­caerulea Gould 1861, pl. 331 Status  Doubtful taxon. Putative range  Rio de Janeiro, Brazil.

Known from four or five males. Status unresolved (Meyer de Schauensee 1970), but considered a hybrid of unknown parentage (Berlioz 1965).

Black-­bellied Woodnymph  Eucephala scapulata

Gould

Eucephala scapulata Gould 1861, p. 166 (‘supposed to be Cayenne’) Status  Doubtful taxon. Putative range  French Guiana.

Known only from the type, an adult male in the Gould Collection. Believed to be a hybrid ­Blue-­chinned Sapphire Chlorostilbon notatus x ­Fork-­tailed Woodnymph Thalurania furcata (Berlioz 1932).

Reeves’s Woodnymph  Eucephala caeruleolavata

(Gould)

Eucephala caeruleo–lavata Gould 1860, p. 306 (São Paulo, Brazil) Status  Doubtful taxon. Putative range  São Paulo, Brazil.

Known only from one unique specimen, a male collected by T. Reeves. Peters (1945) considered it a hybrid, parentage uncertain.

Sapphire-­crowned Woodnymph  Eucephala smaragdocaerulea

Berlioz

Eucephala smaragdocaerulea Berlioz 1951, p. 281 Status  Invalid taxon. Putative range  Uncertain.

Based on a specimen of uncertain affinities in Tring, collected by Whitely in around 1881 and formerly part of the ­Godman-­Salvin Collection. Berlioz (1951) considered that it was probably a hybrid between the ­Violet-­capped Woodnymph Thalurania glaucopis and the ­Blue-­chinned Sapphire Chlorostilbon notatus, as did Meyer de Schauensee (1966).

Bourcier’s Woodnymph  Eucephala chlorocephalus

(Bourcier)

Hylocharis Chlorocephalus Bourcier 1854, p. 457 (Guaranda, Ecuador = error for Brazil?) Status  Doubtful taxon. Putative range  Brazil?

Known only from a unique specimen held at Tring, an adult male. Peters (1945) treated it as a hybrid of undetermined parentage, whereas Meyer de Schauensee (1966) considered it a synonym of Augasma smaragdinea (see above).

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Extinct Birds

Nehrkorn’s Hummingbird  Neolesbia nehrkorni

(Berlepsch)

Cyanolesbia nehrkorni Berlepsch 1887, p. 326 (Bogotá, Colombia) Neolesbia Salvin 1892, p. 115 (in key), p. 145. Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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 ­Purple-­backed Thornbill Ramphomicron microrhynchum.

Flame-­rumped Sapphire  Hylocharis pyropygia

(Salvin & Godman)

Eucephala pyropygia Salvin & Godman 1881, p. 596, pl. 16 (Ecuador = Bahia) Status  Doubtful taxon. Putative range  Ecuador.

Known only from three specimens. Berlioz (1938) considered the bird to be a hybrid ­White-­chinned Sapphire Hylocharis cyanus x ­Glittering-­bellied Emerald Chlorostilbon aureoventris pucherani.

Black-and-blue Carib  Trochilis venustissimus

(Gmelin)

Gros Colibri. du Tertre 1667, p. 263 ? Avicula mexicana cyaneo colore venutissima. Seba 1734, p. 102, fig. 3 (no locality) ? Avicula cyaneo colore venustissima. Klein 1750, p. 107 (based on Seba) Troglodyte adfinis. Möering 1752, no. 102 (based on Seba). Polytmus mexicanus cyaneus. Brisson 1760, p. 681 Black and Blue Hummingbird. Bancroft 1769, p. 167 Le Colibri bleu. Buffon 1771–86, p. 61 Crimson-­headed Blue Humming Bird. Latham 1782, p. 762 (Mexico) Black and Blue Humming Bird. Latham 1782, p. 782 Trochilus venustissimus Gmelin 1788, p. 490. (Mexico). Based on Brisson, du Tertre, Seba and Latham. Trochilus cyanomelas Gmelin 1788, p. 498 Trochilus bancrofti Latham 1790, p. 317 Status  Doubtful taxon. Putative range  Unknown.

Known only from the descriptions of du Tertre and Bancroft, of uncertain origin

Taylor’s Streamertail  Trochilus taylori

(Rothschild)

Aithurus taylori Rothschild 1894a, p. 47 (St Andrews, north of Kingston, Jamaica) Status  Invalid taxon. Putative range  Jamaica.

Known only from the type series in the Rothschild Collection, and considered to be a variety of the ­Red-­billed Streamertail Trochilus polytmus.

Friburgo Whitethroat  Leucochloris malvina

(Reichenbach)

Chlorestes malvina Reichenbach 1854, supplement p. 7 (Brazil) Status  Doubtful taxon. Putative range  Rio de Janeiro, Brazil.

Known only from the unique type collected by Beske. Considered a possible hybrid ­White-­throated Hummingbird Leucochloris albicollis x ­Glittering-­bellied Emerald Chlorostilbon aureoventris pucherani (Berlioz 1938; Schuchmann 1999). 412

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Appendix 2: Doubtful and invalid taxa

Reichenbach’s Whitethroat  Leucippus nigrirostris

Reichenbach

Leucippus nigrirostris Reichenbach 1855, p. 8 Status  Doubtful taxon. Putative range  Unknown.

Known only from Reichenbach’s plate. Placed in the synonymy of the ­White-­bellied Hummingbird Amazilia chionogaster (Salvin 1892).

Blue-­spotted Hummingbird  Amazilia cyaneotincta

(Gounelle)

Polyerata cyaneotincta Gounelle 1909, p. 17 and plate (Bogotá) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Known only from a Bogotá trade skin in the Paris Museum and another in the ­Rousseau-­Decelle collection, Paris. Considered a probable aberrant ­Glittering-­throated Emerald A. fimbriata (Schuchmann 1999).

Small-­billed Azurecrown  Amazilia microrhyncha

(Elliot)

Cyanomyia microrhyncha Elliot 1876a, p. 316 (Honduras?) Status  Invalid taxon. Putative range  Honduras.

Based on inadequate subspecific characters and relegated to synonymy of Azure-crowned Hummingbird Agyrtria c. cyanocephala (Weller 1999a).

Sapphire-­fronted Emerald  Amazilia caeruliceps

(Gould)

Thaumatias caeruliceps Gould 1860, p. 307 (Colombia) Status  Doubtful taxon. Putative range  Colombia.

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.

Dusky Emerald  Amazilia veneta

(Simon)

Agyrtria veneta Simon 1921, p. 112, 328 (Bogotá, Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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).

Brown Emerald  Amazilia aenobrunnea

Chapman

Amazilia aenobrunnea Chapman 1889, p 163 Status  Invalid taxon. Putative range  Bogotá, Colombia.

Known only from the type, which is an artefact, with the body of a ­Ruby-­topaz Hummingbird Chrysolampis mosquitus and the head of a ­Red-­bellied Emerald Chlorostilbon gibsoni (Simon & Hellmayr 1908; Simon 1921).

413

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Extinct Birds

de Oca’s Hummingbird  Amazilia ocai

Gould

Amazilia Ocai Gould 1859, p. 96 (Jalapa, Vera Cruz) Status  Doubtful taxon. Putative range  Xalapa, Mexico.

Known only from the type, an adult male collected by de Oca. Considered to be a hybrid A ­ zure-­crowned Hummingbird Amazilia cyanocephala x Berylline Hummingbird A. beryllina (Berlioz 1932).

Lerdo’s Hummingbird  Amazilia lerdi

(de Oca)

Thaumatias Lerdi de Oca 1875, p. 24 (Paso del Mancho, Vera Cruz) Status  Doubtful taxon. Putative range  Paso del Mancho, Vera Cruz, Mexico.

Known only from de Oca’s description and plate. Generally placed in the synonomy of Amazilia ocai (Peters 1945; see above).

Florence van Rossem’s Hummingbird  Amazilia florenceae

(van Rossem & Hachisuka)

Saucerottia florenceae van Rossem & Hachisuka 1938, 407 (Rancho Santa Barbara, 20 miles ­north-­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).

Salvin’s Azurecrown  Amazilia salvini

(Brewster)

Cyanomyia salvini Brewster 1893, p. 214 (Nacosari, Sonora, Mexico) Status  Invalid taxon. Putative range  Sonora, Mexico.

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).

Neglected Hummingbird  Amazilia neglectus

(Elliot)

? Ornismya bicolor d’Orbigny & Lafresnaye 1838, p. 30 (this name is very doubtfully assigned to this species) Thaumatias neglectus Elliot 1877a, p. 140 (Moxos male and Yungas female, Bolivia). Status  Invalid taxon. Putative range  Bolivia.

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.

Elliot’s Hummingbird  Amazilia lucida

Elliot

Amazilia lucida Elliot 1877b, p. 404 (Colombia) Status  Doubtful taxon. Putative range  Colombia.

Known only from an adult male specimen held in New York. Status and affinities uncertain (Peters 1945; Meyer de Schauensee 1966; Sibley & Monroe 1990). 414

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Appendix 2: Doubtful and invalid taxa

Bangs’s Hummingbird  Amazilia bangsi

(Ridgway)

Amazilis bangsi Ridgway 1910, p. 54 (Volcán de Miravalles, Costa Rica) Status  Invalid taxon. Putative range  Costa Rica.

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).

Elliot’s Hummingbird  Amazilia inculta

(Elliot)

Eriocnemis incultus Elliot 1889, p. 210 (Bogotá) Status  Invalid taxon. Putative range  Bogotá, Colombia.

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 ­Green-­bellied Hummingbird A. viridigaster and it was placed in the synonymy of A. viridigaster by Peters (1945). Weller (2001) considered it to be a melanistic form of the ­Indigo-­capped Hummingbird A. cyanifrons, rather than A. viridigaster.

President Nunez’s Hummingbird  Amazilia nunezi

(Boucard)

Saucerottia nunezi Boucard 1892, p. 81 (Bogotá) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Apparently known from two males. Cory (1918) listed it as a melanistic variety of ­Green-­bellied Hummingbird A. viridigaster, and Peters (1945) also synonymised it with A. viridigaster.

Wiener’s Hummingbird  Amazilia versicolor meracula

(Simon)

Agyrtia nitidifrons meracula Simon 1921, p. 114, 329 (Rio Napo? Locality doubtful) Status  Doubtful taxon. Putative range  Unknown.

Known only from specimens in the Simon collection. Status and affinities uncertain.

Purple Inca  Coeligena purpurea

Gould

Coeligena purpurea Gould 1854, pl. 12 and pl. 256 (Popayán, Colombia) Status  Invalid taxon. Putative range  Popayán, Colombia.

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).

Lilac-­fronted ­Star-­frontlet  Coeligena traviesi

(Mulsant & Verreaux)

Diphlogaena (Helianthea) Traviesi Mulsant & Verreaux 1867, p. 389 (Colombia) Status  Invalid taxon. Putative range  Colombia.

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). 415

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Extinct Birds

Lawrence’s ­Star-­frontlet  Coeligena lawrencei

(Boucard)

Homophania lawrencei Boucard 1892, p. 87 (Bogotá trade skin) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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).

Lowe’s Swordbill  Ensifera caerulescens

(Lowe)

Docimastes ensiferus caerulescens Lowe 1939, p. 73 (South America) Status  Doubtful taxon. Putative range  Unknown.

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.

Hartert’s ­Sun-­angel  Heliangelus dubius

Hartert

Heliangelus dubius Hartert 1897, p. 532 (Bogotá trade skin) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Known only from the type held at Tring, and possibly another in the Simon collection. Considered a probable melanistic ­Amethyst-­throated ­Sun-­angel Heliangelus amethysticollis (Fjeldså & Krabbe 1990; Schuchmann 1999).

Claudia’s ­Sun-­angel  Heliangelus claudia

Hartert

Heliangelus claudia Hartert 1895, vol. 2, p. 484 (Bogotá, Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Known only from Bogotá trade skins. Considered an aberrant ­Amethyst-­throated ­Sun-­angel Heliangelus amethysticollis of race clarisse (Peters 1945, Fjeldså & Krabbe 1990).

Henry’s ­Sun-­angel  Heliangelus henrici

Boucard

Heliangelus henrici Boucard 1891, p. 26 (Ecuador) Status  Doubtful taxon. Putative range  Ecuador.

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 ­Sun-­angel is a specimen of Gorgeted ­Sun-­angel H. strophianus with the central rectrices tinged with bronze green.

Violet ­Sun-­angel (Sarayacu ­­Sun-­­angel)  Heliangelus violicollis

Salvin

Heliangelus violicollis Salvin 1891, p. 376 (Sarayacú, Ecuador) Status  Doubtful taxon. Putative range  Sarayacú, Ecuador.

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. 416

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Appendix 2: Doubtful and invalid taxa

Olive-­throated ­Sun-­angel  Heliangelus squamigularis

Gould

Heliangelus squamigularis Gould 1871, p. 503 (Colombia) Heliotrypha Barrali Mulsant & Verreaux 1872, p. 106 (Banks of the Río Saldana, Antioquia, Colombia) Nodalia barrali Mulsant 1877, p. 100. Status  Invalid taxon. Putative range  Antioquia and Bogotá, Colombia.

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.

Green-­throated ­Sun-­angel  Heliangelus speciosus

(Salvin)

Heliotrypha speciosa Salvin 1891, p. 376. (Colombia) Status  Invalid taxon. Putative range  Bogotá, Colombia.

Known from Bogotá trade skins. Only the type, an adult male, is listed by Salvin (1892). Graves (1990) determined that this is a hybrid ­Amethyst-­throated ­Sun-­angel Heliangelus amethysticollis x ­Copper-­bellied Puffleg Eriocnemis cupreoventris.

Boucard’s ­Sun-­angel Heliangelus simoni

(Boucard)

Heliotrypha simoni Boucard 1892, p. 76 (Bogotá, Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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).

Rothschild’s ­Sun-­angel  Heliangelus rothschildi

Boucard

Heliangelus rothschildi Boucard 1892, p. 77 (Bogotá, Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Known only from the type, an adult male Bogotá trade skin. Considered a probable hybrid Heliangelus sp. x Eriocnemis sp. or ­Purple-­backed Thornbill Ramphomicron microrhynchum (Meyer de Schauensee 1966).

Balen’s ­Sun-­angel  Heliangelus prosantis

(Oberholser)

Aeronympha prosantis Oberholser 1905a, p. 161 (Bogotá, Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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.

Glistening ­Sun-­angel  Heliangelus luminosus

(Elliot)

Iolaema luminosus Elliot 1878, p. 188 (no locality = Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

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). 417

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Extinct Birds

Simon’s ­Sun-­angel  Heliangelus fulvicrissa

Simon

Heliangelus Clarissae [sic] var. fulvicrissa Simon 1921, p. 179 (Bogotá) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Known only from the type, a Bogotá trade skin, which Simon (in Peters 1945) suggested may be a hybrid ­Amethyst-­throated ­Sun-­angel Heliangelus amethysticollis of race clarisse x ­Buff-­tailed Coronet Boissoneaua flavescens.

Söderström’s Puffleg  Eriocnemis soderstromi

Butler

Eriocnemis söderströmi Butler 1926, 7, p. 62 (west side of Mount Pichincha, Ecuador) Status  Invalid taxon. Putative range  Mount Pichincha, Ecuador.

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.

Isaacson’s Puffleg  Eriocnemis isaacsonii

(Parzudaki)

Ornysmia [sic] Isaacsonii Parzudaki 1845, p. 95 (Bogotá, Colombia) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Known only from three Bogotá trade skins, which differ somewhat. Possibly a hybrid Eriocnemis x Coeligena (Schuchmann 1999).

Black Puffleg  Eriocnemis dyselius

Elliot

Eriocnemis dyselius Elliot 1872a, p. 294 Status  Invalid taxon. Putative range  Unknown.

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 ­Copper-­bellied Puffleg Eriocnemis cupreoventris, but did not comment on the Gould specimen.

Simple Puffleg  Eriocnemis simplex

(Gould)

Eripous simplex Gould 1849, p. 96 (Colombia) Eriocnemis simplex Gould 1852, pl. 271 Status  Doubtful taxon. Putative range  Colombia.

Known from three specimens, all from the Gould collection, an adult male, an adult unsexed skin, and a ‘variety’. Peters (1945) placed it in the synonymy of the ­Copper-­bellied Puffleg Eriocnemis cupreoventris.

Berlepsch’s Puffleg  Eriocnemis berlepschi

Hartert

Eriocnemis berlepschi Hartert 1897, p. 531 (Bogotá) Status  Doubtful taxon. Putative range  Bogotá, Colombia.

Known only from the type, considered by Peters (1945) to be a synonym of the Glowing Puffleg Eriocnemis vestitus. 418

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Appendix 2: Doubtful and invalid taxa

Amethyst-­vented Puffleg  Eriocnemis ventralis

Salvin

Eriocnemis ventralis Salvin 1891, p. 378 (Colombia) Status  Doubtful taxon. Putative range  Colombia.

Known only from the type, an adult male Bogotá trade skin. Schuchmann et al. (2000) considered it a probable hybrid Glowing Puffleg E. vestitus x ­Copper-­bellied Puffleg E. cupreoventris.

Ridolfi’s Hummingbird  Eriocnemus ridolfi

Benvenuti

Mellisuga (Eriocnemus) ridolfi E. Benvenuti 1865, p. 205 (‘Nova Granata’) Status  Doubtful taxon. Putative range  New Granada, Colombia.

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.

Purple-­tailed Comet  Zodalia (Lesbia) glyceria

(Gould)

Lesbia glyceria ‘Gould 1854’. Bonaparte 1854c, p. 252 (nomen nudum) Cometes? glyceria Gould 1858, pl. 12 and pl. 176 (near Popayán, Colombia) Status  Invalid taxon. Putative range  Popayán, Colombia.

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.

Orton’s Comet  Lesbia ortoni

Lawrence

Lesbia ortoni Lawrence 1869, p. 269 (Quito Valley, Ecuador) Status  Invalid taxon. Putative range  Ecuador.

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 microrhynchum, and this identification has been supported by Graves (1997a).

Purple-­tailed Thornbill  Metallura purpureicauda

(Hartert)

Chalcostigma purpureicauda Hartert 1898a, p. 28 (Bogotá trade skin) Zodalia thaumasta Oberholser 1902, p. 338 (Illalo, Valley of Chillo, Ecuador). Status  Doubtful taxon. Putative range  Ecuador and Colombia.

Known only from three specimens, a male and a female from Illalo, Ecuador, and a Bogotá trade skin. Considered a probable a hybrid ­Long-­tailed Sylph Aglaiocercus kingi x Black-tailed Trainbearer Lesbia victoriae (Schuchmann 1999).

Boucard’s Metaltail  Metallura griseocyanea

Boucard

Metallura griseo–cyanea Boucard 1893, p. 75 Status  Doubtful taxon. Putative range  Unknown.

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. 419

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Extinct Birds

Violet-­throated Hummingbird  Archilochus (Mellisuga) violajugulum

(Jeffries)

Trochilus violajugulum Jeffries 1888, p. 168 (Santa Barbara, California) Status  Doubtful taxon. Putative range  Santa Barbara, California.

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; Ridgeway 1911).

Underwood’s Hummingbird  Selasphorus underwoodi

Salvin

Selasphorus underwoodi Salvin 1897, p. 38 (Volcán de Irazú, Costa Rica) Status  Invalid taxon. Putative range  Volcán de Irazú, Costa Rica.

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).

Floresi’s Hummingbird  Selasphorus floresii

Gould

Selasphorus floresii Loddiges in Gould 1861, pl. 10, pl. 139 (Bolaños, Jalisco, Mexico) Trochilus rubromitratus Ridgway 1891, p. 114 Status  Doubtful taxon. Putative range  Jalisco, Mexico.

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. Ridgway 1911).

Morcom’s Hummingbird  Atthis morcomi

Ridgway

Atthis morcomi Ridgway 1898, p. 325 (Huachuca Mountains, Arizona) Status  Doubtful taxon. Putative range  Huachuca Mountains, Arizona.

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).

Decorated Woodstar  Acestrura decorata

(Gould)

Calothorax decoratus Gould 1860, p. 309. (‘supposed to be from Antioquia in Colombia’) Status  Invalid taxon. Putative range  Antioquia, Colombia.

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 ­White-­bellied Woodstar Chaetocercus mulsant x Gorgeted Woodstar C. heliodor, which was upheld by Graves (1997b).

Hartert’s Woodstar  Chaetocercus harterti

(Simon)

Polyxemus Harterti Simon 1901b, p. 202 (Western Colombia) Status  Doubtful taxon. Putative range  Ibagüe, western Colombia.

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. 420

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Appendix 2: Doubtful and invalid taxa

Splendid Brilliant  Heliodoxa splendens

(Gould)

Leadbeatera splendens Gould 1861, p. 74 (Venezuela) Status  Doubtful taxon. Putative range  Venezuela.

Known only from Gould’s description. The type appears to be lost. Salvin (1892) synonymised H. splendens with ­Violet-­fronted Brilliant Heliodoxa leadbeateri.

Burton’s Hummingbird  Sephanioides burtoni

(Boucard)

Eustephanus burtoni Boucard 1891, p. 18 (Chile) Status  Doubtful taxon. Putative range  Chile.

Known only from Boucard’s description of a specimen in his collection. Peters (1945) placed it in the synonymy of the ­Green-­backed Firecrown Sephanoides sephanoides.

Molina’s Hummingbird  Sephaniodes galeritus

(Molina)

Trochilus galeritus Molina 1782, p. 247. (Chile) Status  Doubtful taxon. Putative range  Chile.

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.

Penard’s Hummingbird  Glaucis philippinae

(Penard)

Heteroglaucis philippinae T. Penard 1922, p. 27 (Lelydrop, Surinam) Status  Doubtful taxon. Putative range  Surinam.

Known only from the type. Peters (1945) 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. Munroe (1963) carried out a detailed revision of the group and considered it invalid.

Fifth Hummingbird of Marcgrave  ‘Trochilus’ guainumbiquinta  

Marcgrave

Guainumbi quinta. Marcgrave in Piso and Marcgraf 1648, p. 197 Status  Doubtful taxon. Putative range  Brazil.

Known only from Marcgrave’s description. Never satisfactorily identified.

421

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Extinct Birds

Seventh Hummingbird of Marcgrave  ‘Trochilus’ septimus

Marcgrave

Guainumbi septima. Marcgrave in Piso 1648, p. 197. Status  Doubtful taxon. Putative range  Brazil.

Known only from Marcgrave’s description. Never satisfactorily identified.

Ninth Hummingbird of Marcgrave  ‘Trochilus’ nona

Marcgrave

Guainumbi nona. Marcgrave in Piso 1648, p. 197 Status  Doubtful taxon. Putative range  Brazil.

Known only from Marcgrave’s description. Never satisfactorily identified.

Dombey’s Motmot  Baryphthengus dombeyi

Lesson

Le Motmot dombé. Levaillant 1806a, p. 113, pl. 39 (Perú) ? Baryphonus ruficapillus Vieillot 1818, p. 315 ?Prionites dombeyanus Ranzini 1822, p. 158 (based on Vieillot) Momotus dombeyi Lesson 1831, p. 251 Status  Doubtful taxon. Putative range  Peru.

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).

Adanson’s ­Bee-­eater  Merops adansonii

Kuhl

Guepier à longue queue du Senegal. Daubenton 1770–86, no. 314 (Senegal) Le Guepier Adanson. Levaillant 1807, pl. 13, p. 47 (Senegal) Merops badius L. var (beta) Merops Adansonii ‘Vaill. 13’. Kuhl 1820b, p. 6 Status  Doubtful taxon. Putative range  Senegal.

Known only from Kuhl’s description, based on the plates of Daubenton and Levaillant. Considered a probable artefact (Peters 1945)

Red-headed ­Bee-­eater  Merops erythrocephalus

Gmelin

Apiaster indicus erythrocephalus Brisson 1760, p. 563, pl. 44, fig. 3A Le Guepier à tete rouge. Montbeillard 1779, p. 508 Red-headed Bee Eater. Shaw & Nodder 1778 pl. 357 Merops erythrocephalus Gmelin 1788, p. 463 Status  Doubtful taxon. Putative range  Unknown.

Known only from Brisson’s description. Unidentifiable (see Whistler & Kinnear 1935).

Cave’s ­Wood-­hoopoe  Phoeniculus (Rhinopomastus) cavei

(Macdonald)

Scoptelus cavei Macdonald 1946, p. 5 (Boma Hills, ­south-­east Sudan) Status  Doubtful taxon. Putative range  Boma Hills, ­south-­east Sudan.

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. 422

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Appendix 2: Doubtful and invalid taxa

Bertoni’s Puffbird  Nonnula galbuloides

(Bertoni)

Microtrogon galbuloides Bertoni 1901, p. 42 (Paraguay) Status  Doubtful taxon. Putative range  Paraguay.

Known only from Bertoni’s description. Probably based on memory (Peters 1948).

Maroon-backed Araçari  Pteroglossus olallae

Gyldenstolpe

Petroglossus olallae Gyldenstolpe 1942, p. 8 (João Pessôa, Rio Juruá, Brazil) Status  Invalid taxon. Putative range  Rio Juruá, Brazil.

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.

Hauxwell’s Araçari  Pteroglossus didymus

Sclater

Pteroglossus didymus Sclater 1890, p. 403 (Upper Amazonia) Status  Invalid taxon. Putative range  ‘Upper Amazonia’.

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).

Albin’s Toucan  Pteroglossus picatus

(Linnaeus)

Pica brasiliensis. Aldrovandus 1599, p. 801 Pica brasiliensis. Albin 1738, p. 24, pl. 25 Ramphastos picatus Linnaeus 1758, p. 103 (‘in America meridionalis’) Status  Doubtful taxon. Putative range  Unknown.

Known only from the plates and descriptions of Albin and Aldrovandus. Never identified with any known species. Peters (1930) considered it an artefact.

Edwards’s Toucan  Ramphastos piscivorus

Linnaeus

The Toucan or Brasilian Pie. Edwards 1743, pl. 64 Tucana brasiliensis gutture albo. Brisson 1760, p. 413 (based on Edwards) Ramphastos piscivorus Linnaeus 1766, p. 151 Status  Doubtful taxon. Putative range  Brazil.

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.

Pöppig’s Curl-crested Araçari  Pteroglossus poeppigii

Wagler

Pteroglossus poeppigii Wagler 1832, p. 1230 Status  Doubtful taxon. Putative range  Unknown.

Known only from Wagler’s description. This bird has been generally synonymised with the common Curlcrested Araçari Pteroglossus beauharnaesii. 423

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Extinct Birds

Mysterious Toucan  Ramphastos piperivorus

Linnaeus

Ramphastos piperivorus Linnaeus 1758, p. 103 Status  Doubtful taxon. Putative range  Central America.

Known only from Linnaeus’s description. Peters (1930) considered it unidentifiable.

Friedmann’s Honeyguide  Indicator propinquus

Friedmann

Indicator propinquus Friedmann 1943, p. 250. (Donenking, Bafia, Cameroons) Status  Doubtful taxon. Putative range  Cameroon.

Known only from the type. Generally considered to be a young specimen of the Least Honeyguide Indicator exilis (Friedmann 1954).

Natterer’s Piculet  Picumnus fuscus

Pelzeln

Picumnus fuscus Pelzeln 1870, p. 242 (nomen nudum), p. 335 (Rio Guaporé, Mato Grosso, Brazil) Status  Doubtful taxon. Putative range  Mato Grosso, Brazil.

Known only from the type, a young female, perhaps merely the young of some other species, but never identified.

Berlepsch’s Woodpecker  Celeus immaculatus

Berlepsch

Celeus immaculatus Berlepsch 1880, p. 113 (Agua Dulce, Panamá) Status  Doubtful taxon. Putative range  Agua Dulce, Panamá.

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.

Pennant’s Ferruginous Woodpecker  Celeus sp. Woodpecker. Pennant 1785, p. 271 Status  Doubtful taxon. Putative range  South Carolina.

Known only from the description of Pennant, based on a specimen sent to him by Dr. Garden of Charlestown, South Carolina. Unidentified.

Crawfurd’s Woodpecker  Dryocopus crawfurdi

(Gray)

Picus crawfurdi J. E. Gray 1829, p. 513 (no locality, ‘from an Indian drawing’) Status  Doubtful taxon. Putative range  Unknown.

Known only from J. E. Gray’s description from a drawing. Peters (1948) considered it to be an aberrant specimen of ­White-­bellied Woodpecker D. javensis javensis.

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Appendix 2: Doubtful and invalid taxa

Yellow-legged Woodpecker  Picus flavipes

Gmelin

Lesser Black Woodpecker. Albin 1738, p. 22, pl. 23 Le pic noir de la Nouvelle Angleterre. Brisson 1760, p. 24 Yellow-legged Woodpecker. Pennant 1781, p. 275 Picus flavipes Gmelin 1788, p. 438 Status  Doubtful taxon. Putative range  New England, United States.

Known only from Albin’s description and plate, from which the others derive. Unidentifiable and probably a ­dreamed-­up bird.

Hargitt’s Woodpecker  Melanerpes hargitti

Du Bois

Melanerpes sp. inc. Du Bois 1897, p. 783 (no locality) Melanerpes hargitti Du Bois 1899 p. 68, pl. 2, fig. 2 Status  Invalid taxon. Putative range  No locality.

Known only from the type in Brussels. Peters (1948) 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).

Grand Bahama Woodpecker  Melanerpes superciliaris bahamensis

(Cory)

Centurus bahamensis Cory 1892, p. 270 (Grand Bahama Island) Status  Doubtful taxon. Range  Grand Bahama, Bahamas.

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. The population disappeared in the 1950s.

Swainson’s Tyrant  Tyrannula pusilla

(Swainson)

Platyrhynchus pusillus Swainson 1827b, p. 366 (Maritime parts of Mexico) Status  Invalid taxon. Putative range  Mexico.

Known only from Swainson’s description. Unidentifiable (Cory & Hellmayr 1927).

Heine’s Kingbird  Tyrannus apolites

(Cabanis & Heine)

Laphyctes apolites Cabanis & Heine 1859, p. 77 (no locality) Status  Doubtful taxon. Putative range  Unknown.

Known only from the type. It is now considered to be a probable hybrid Tropical Kingbird Tyrannus melancholicus x Varied Flycatcher Empidonomus varius.

Berlepsch’s Tody-tyrant  Todirostrum hypospodium

Berlepsch

Todirostrum hypospodium Berlepsch 1907, p. 354 (‘Bogotá’, Colombia) Status  Invalid taxon. Putative range  Bogotá, Colombia.

Known only from a solitary Bogotá trade skin. Cory & Hellmayr (1927) believed it to be an aberrant specimen of Lawrence’s Tody-tyrant Todirostrum sylvia superciliare, and Traylor (1979) regarded it as a synonym of that form. 425

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Extinct Birds

Philippi’s Spinetail  Sylviorthorhynchus fasciolatus

Philippi

Sylviorthorhynchus fasciolatus F. Philippi 1909, p. 65 (Valdivia) Status  Invalid taxon. Putative range  Valdivia, Chile.

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.

Lillo’s Canastero  Asthenes leptasthenuroides

(Lillo)

Siptornis leptasthenuroides Lillo 1905, p. 52 (Norco, Tucumán, Argentina) Status  Doubtful taxon. Putative range  Tucumán, Argentina.

Known only from the type. Olrog (1963) treated it as a synonym of the Chilean ­Sharp-­billed Canastero Asthenes pyrrholeuca sordida.

Puno Canastero  Asthenes sclateri

(Cabanis)

Synallaxis Sclateri Cabanis 1878, p. 196 (Sierra de Cordova, Argentina) Status  Extant. Range  Sierra de Cordova, Argentina.

A. sclateri was originally known only from Cabanis’s description, based on the type, and one other specimen 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.

Snethlage’s Creeper  Xiphocolaptes franciscanus

Snethlage

Xiphocolaptes franciscanus Snethlage 1927a, p. 8 (Brejo Januaria, left bank of the Rio São Francisco, Minas Gerais) Status  Doubtful taxon. Putative range  East Brazil, west of the Rio São Francisco.

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).

Vila Nova Woodcreeper  Xiphocolaptes villanovae

Lima

Xiphocolaptes albicollis villadenovae (corrected to villanovae on errata slip) Lima 1920, p. 104, fig. 1 (Villa Nova, Bahia, Brazil) Status  Data deficient. Range  Vila Nova, Bahia, Brazil.

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.

Black-­and-­green Sericornis  Sericornis nigroviridis

Miller

Sericornis nigroviridis Miller 1964, p. 2 (Edie Creek, Wau, Moroke district, eastern New Guinea) Status  Doubtful taxon. Putative range  New Guinea.

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Appendix 2: Doubtful and invalid taxa

Known only from the unique type. Regarded as a melanistic specimen of the ­Buff-­faced Scrub Wren Sericornis perspicillatus (Beehler 1978; Mayr et al. 1986).

Charleville Heath Wren  Hylacola tyrannulus

(De Vis)

Sericornis tyrannula De Vis 1905, p. 42 (Charleville, southern Queensland) Status  Doubtful taxon. Putative range  Southern Queensland, Australia.

Known only from the type, now lost. Regarded as an immature of the ­Chestnut-­rumped Heath Wren Calamanthus pyrrhopygius Parker (1984).

Grampian Pied Currawong  Strepera graculina ashbyi

Mathews

Strepera graculina ashbyi Mathews 1913, p.78 (Black Spur, Victoria) Status  Status uncertain. Range  Grampians and ­south-­west Victoria, Australia.

Listed as a morph of S. g. nebulosa of ­south-­eastern Australia (Schodde & Mason 1999). Population has hybridised with S. g. nebulosa over much of its range, with a possible tiny population of ­pure-­bred birds remaining. If it is a valid taxon, it is critically endangered if not already extinct.

Baber’s Cuckoo-shrike  Lalage melanothorax

Sharpe

Lalage melanothorax Sharpe 1879, p. 91 (Madras) Status  Invalid taxon. Putative range  Western India.

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 ­Cuckoo-­shrike Coracina melanoptera (Sharpe 1879).

Maros Cicadabird  Coracina tenuirostris edithae

Stresemann

Coracina tenuirostris edithae Stresemann 1932, p. 109 (Sulawesi) Status  Doubtful taxon. Putative range  South Sulawesi.

Known from a single specimen collected in 1931. Probably a vagrant of the Australian nominate race of Cicadabird.

Natka Shrike  Lanius natka

Gmelin

Natka Shrike. Pennant 1785, p. 239 Lanius Natka Gmelin 1788, p. 309 Status  Doubtful taxon. Putative range  Nootka Sound, Vancouver.

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.

Vera Paz Vireo  Vireo propinqua

(Baird)

Vireosylvia propinqua Baird 1866, p. 345 (in key), p. 348 (Cobán, Vera Paz =Alta Verapaz, Guatemala) Status  Doubtful taxon. Putative range  Alta Verapaz, Guatemala.

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Extinct Birds

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 ­Blue-­headed Vireo V. solitarius.

White-headed Oriole  Oriolus violaceus

Boddaert

Le Cassique de la Louisiane. Buffon 1775, p. 242 Cassique de la Louisiane. Daubenton 1771–86, pl. 646 White-headed Oriole. Latham 1782, p. 422 Oriolus violaceus Boddaert 1783 White-headed Oriole. Pennant 1785, p. 260 Oriolus ludovicianus Gmelin 1789, p. 387 Oriolus leucocephalus Latham 1790, p. 175 Status  Doubtful taxon. Putative range  Louisiana, United States.

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.

Ingram’s Figbird  Sphecotheres stalkeri

Ingram

Sphecotheres stalkeri Ingram 1908, p. 100 (Mt Elliot, near Townsville, Queensland, Australia) Status  Doubtful taxon. Putative range  Queensland, Australia.

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.

Tongatabu Monarch  Pomarea tabuensis

Mathews

Muscicapa atra Forster 1844, p. 171 Pomarea nigra tabuensis Mathews 1929, p. 60 (Tongatabu, Tonga Islands) Status  Invalid taxon. Putative range  Tongatabu, Society Islands.

Known only from Forster’s description. Todd (1984) has shown that it was not a Pomarea, but a Polynesian Triller Lalage maculosa tabuensis.

Stresemann’s Crow Corvus difficilis

Stresemann

Corvus difficilis Stresemann 1943, p. 125 (Malbon, Cloncurry district, ­north-­western Queensland) Status  Invalid taxon. Putative range ­North-­western Queensland, Australia.

Known only from the type. Considered by Vaurie (1962) to be an abnormal specimen of the Australian Raven Corvus coronoides or a hybrid of C. coronoides and the Little Crow C. bennetti.

Brüggemann’s Crow  Corvus fallax

Brüggemann

Corvus fallax Brüggemann 1877, p. 76 (Celebes?) Range  Known only from the type. Putative range  Celebes?

Sharpe (1877) placed C. fallax in the synonymy of the ­Slender-­billed Crow Corvus enca.

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Appendix 2: Doubtful and invalid taxa

Rosenberg’s Crow  Corvus modestus

Brüggemann

Corvus modestus Brüggemann 1877, p. 77 (Celebes?) Status  Doubtful taxon. Putative range  Celebes?

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.

Currie’s Crimson Bird of Paradise ­(Lyre-­tailed King)  Cicinnurus lyogyrus

Currie

Cicinnurus lyogyrus R.P. Currie 1900, p. 497, pl. 17 (New Guinea) Status  Invalid taxon. Putative range  New Guinea.

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).

Lyre-­tailed King Bird of Paradise  Cicinnurus goodfellowi

Ogilvie-Grant

Cicinnurus goodfellowi Ogilvie-Grant 1906, p. 39 (Cyclops Mountains, Humboldt’s Bay, Dutch New Guinea) Putative range  New Guinea.

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).

Mantou’s Riflebird  Craspedophora (Heteroptilornis) mantoui

Oustalet

Craspedophora Mantoui Oustalet 1891, p. 260 (North-west New Guinea) Status  Invalid taxon. Putative range ­North-­west New Guinea.

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).

Bensbach’s Bird of Paradise  Craspedophora bruyni

Büttikofer

Craspedophora bruyni Büttikofer 1895a, p. 161 (Arfak Mountains) Status  Invalid taxon. Putative range ­North-­west New Guinea.

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 ­Twelve-­wired Bird of Paradise Seleucidis melanoleucus (Fuller 1995).

King William III Bird of Paradise  Diphyllodes (Rhipidornis) gulielmitertii

Meyer

Diphyllodes gulielmi III Meyer 1875, p. 31. (Waigeu = error) Status  Invalid taxon. Putative range  New Guinea.

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). 429

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Extinct Birds

Astrapian Sicklebill  Epimachus astrapioides

Rothschild

Epimachus astrapioides Rothschild 1897, p. 22 (Dutch New Guinea) Status  Invalid taxon. Putative range  New Guinea.

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).

Wonderful Bird of Paradise  Janthothorax mirabilis

(Reichenow)

Paradisea mirabilis Reichenow 1901, p. 185 (Madang, Astrolabe Bay) Status  Invalid taxon. Putative range  Astrolabe Bay, New Guinea.

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).

Wilhelmina’s Bird of Paradise  Lamprothorax wilhelminae

Meyer

Lamprothorax wilhelminae Meyer 1894, Ap. 3 (Arfak Mountains) Status  Invalid taxon. Putative range  Arfak Mountains, New Guinea.

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).

Captain Blood’s Bird of Paradise  Paradisaea bloodi

Iredale

Paradisea bloodi Iredale 1948, p. 161 (Miniyip, Mt Hagen)) Status  Invalid taxon. Putative range ­Mount Hagen, central New Guinea.

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).

Frau Reichenow’s Bird of Paradise  Paradisaea maria

Reichenow

Paradisaea maria Reichenow 1894, p. 22 (Finisterre Mountains, New Guinea) Status  Invalid taxon. Putative range  Finisterre Mountains, New Guinea.

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).

Duivenbode’s Bird of Paradise  Paradisaea duivenbodei

Menegaux

Paradisaea duivenbodei Menegaux 1913, p. 49 (Geelvink Bay = error, probably Astrolabe Bay, New Guinea). Status  Invalid taxon. Putative range  New Guinea.

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). 430

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Appendix 2: Doubtful and invalid taxa

Rothschild’s Bird of Paradise  Paradisaea mixta

Rothschild

Paradisea mixta Rothschild 1921a, p. 127 (locality unknown, probably upper Ramu) Status  Invalid taxon. Putative range  New Guinea.

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.

Lupton’s Bird of Paradise  Paradisaea apoda luptoni

Lowe

Paradisea apoda luptoni Lowe 1923, p. 110 (Marauke district) Status  Invalid taxon. Putative range  Marauke district, New Guinea.

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).

Duivenbode’s Six-wired Bird of Paradise  Parotia duivenbodei

Rothschild

Parotia duivenbodei Rothschild 1900, p. 100 (Dutch New Guinea) Status  Invalid taxon. Putative range  New Guinea.

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).

Lesson’s Kioea  Strigiceps leucopogon

(Lesson)

Strigiceps leucopogon Lesson 1840, p. 266 (Australia = error for Hawaiian Islands?) Status  Doubtful taxon. Range  Possibly Australia, but perhaps a Pacific island (Rothschild 1907a).

Unidentifiable. Known only from Lesson’s description, from a type now lost. Generally considered to be identical with the Kioea Chaetoptila angustipluma (see p. 251), but the description does not agree at all, and the provenance is in extreme doubt.

Strömian Titmouse  Parus ignotus

Ström

Parus ignotus Ström 1762, p. 240. Parus ignotus Brünnich 1764, p. 73 (based on Ström) Norway Titmouse. Latham 1781 p. 537 Strömian Titmouse. Pennant 1785, p. 426 Parus ignotus Gmelin, 1789, p. 1006 ? Parus stromei Bonnaterre & Vieillot 1823, p. 504 Status  Doubtful taxon. Putative range  Norway.

Known only from the descriptions of Brünnich, Latham and Pennant, based on Ström. Synonymised under Great Tit Parus major (Gadow 1883).

Zagros Coal Tit  Periparus ater phaeonotus

Blanford

Periparus ater phaeonotus Blandford 1873, p. 88 Status  Doubtful taxon. Putative range  Zagros Mountains, ­south-­west Iran.

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Extinct Birds

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.

Sumatra ­Blue-­wattled Bulbul  Pycnonotus nieuwenhuisii inexspectatus

(Chasen)

Euptilosus nieuwenhuisii inexspectatus Chasen 1939 p. 184 (Upper Kayan River, central Borneo) Status  Doubtful taxon. Putative range  Central Borneo.

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).

Ticehurst’s Warbler  Sylvia deserticola ticehursti

Meinertzhagen

Sylvia ticehursti Meinertzhagen 1939, p. 69 (Tinghir = Tinrhir, Ouarzazate district, Moroccan Sahara) Status  Doubtful taxon. Putative range  Ouarzazate, Morocco.

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).

Fayyum Warbler  Sylvia melanocephala norissae

Nicoll

Sylvia norissae Nicoll 1917, p. 28 (north side of Lake Kar-n, Fayoum, Egypt) Status  Doubtful taxon. Range  Nile Delta region, Egypt.

Not recorded since the 1940s, but probably a ­colour ­morph of Sardinian Warbler Sylvia melanocephala momus (Cabot & Urdiales 2005).

Cuvier’s Kinglet  Regulus cuvieri

Audubon

Regulus cuvieri Audubon 1829, pl. 55 (Fatland Ford, about 10 miles west of Norristown, Pennsylvania) Status  Doubtful taxon. Putative range  Pennsylvania, United States.

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.

Vanderbilt’s Babbler  Malacocichla vanderbilti

Meyer de Schauensee & Ripley

Malacocichla vanderbilti Meyer de Schauensee & Ripley 1940, p. 351, pl. 20 (Koengki, Atjeh, north Sumatra) Status  Invalid taxon. Putative range  Atjeh, north Sumatra.

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).

Amik Gölü Bearded Tit  Panurus biarmicus kosswigi

Kumerloeve

Panurus biarmicus kosswigi Kumerloeve 1958, p.197 (Amik Gölü, southern Turkey) Status  Doubtful taxon, but requires further study. Range  Amik Gölü, southern Turkey and ­north-­west Syria.

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Appendix 2: Doubtful and invalid taxa

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).

Leguat’s Starling  Orphanopsar leguati

(Forbes)

Necropsar leguati Forbes 1898, p. 34, pl. 1 (no locality) Status  Invalid taxon. Putative range  Rodrigues, Mascarenes in error. Specimen comes from Martinique in the West Indies.

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).

Bicoloured Chough  Testudophaga bicolor

Hachisuka

Testudophaga bicolor Hachisuka 1937d, p. 212 (Islet au Mât) Status  Invalid taxon. Putative range  Rodrigues, Mascarene Islands.

The Rodrigues Starling Necropsar rodericanus was described from subfossil remains and from a description by Tafforet (see p. 273), who mentioned the starling’s ability to predate or scavenge giant tortoises or turtles. Hachisuka (1953) compared it with the closely related, but generally frugivorous Réunion Starling Fregilupus varius (see p. 271), 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.

Pacific Shrike-starling  Aplonis pacifica

(Gmelin)

Pacific Shrike. Latham 1781, p. 164 Lanius pacificus Gmelin 1788, p. 306 Status  Doubtful taxon. Known only from Latham’s description, based on a specimen formerly in the Banksian Collection but no longer extant. Range  South Pacific.

The description is considered to be based on Coracias striata = Striated Starling Aplonis striata.

La Touche’s Shortwing  Brachypteryx joannae

(La Touche)

Heteroxenicus joannae La Touche 1922, p. 21 Status  Invalid taxon. Putative range  China.

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).

Sandwich Thrush  Turdus sandwichensis

(Gmelin)

Sandwich Thrush. Latham 1871–85, p. 39 (Sandwich Islands = Hawaiian Islands) Turdus sandwichensis Gmelin 1789, p. 813 Status  Invalid taxon. Putative range  Hawaii.

Known only from Latham’s description. Considered to be a possible immature Hawaii ’Elepaio Chasiempis sandwichensis (Olson 1989). 433

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Extinct Birds

Yakushima Thrush  Turdus celaenops yakushimensis

(Ogawa)

Merula celaenops yakushimensis Ogawa 1905, p. 180 (Yakushima, northern Riukiu Islands) Status  Doubtful taxon. Range  Yakushima, Ryukyu Islands.

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.

Muriel’s Chat  Saxicola dacotiae murielae

Bannerman

Saxicola dacotiae murielae Bannerman 1913b Status  Invalid taxon. Range  Alegranza and Montaña Clara, north of Lanzarote, Canary Islands.

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).

Tonkean ­Henna-­tailed Jungle Flycatcher  Rhinomyias colonus subsolanus

Meise

Rhinomyias colonus subsolanus Meise 1932, p. 80 (East Sulawesi) Status  Doubtful taxon. Range  Eastern Sulawesi, Indonesia.

Known only from the type, now lost. Considered dubious (BirdLife International 2011).

Cyprus Dipper  Cinclus cinclus olympicus

Madarász

Cinclus olympicus Madarász 1903, p. 6 (Trodos Mountains, Cyprus) Status  Doubtful taxon. Range  Mountains of Cyprus.

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.

Painted Sunbird  Cinnyris picta

Hachisuka

Cinnyris picta Hachisuka 1941, p. 52 Status  Invalid taxon. Putative range  Basilan, Philippine Islands.

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).

Daubenton’s Fody  Foudia bruante

(Müller)

Bruante de l’Ile de Bourbon. Daubenton 1771–86, pl. 321, fig. 2 Fringilla bruante P. L. S. Müller 1776b, supplement p. 164 (based on Daubenton) Le Mordré. Buffon 1778, p. 366 Emberiza fuscofulva Boddaert 1783, p. 20 Bourbon Bunting. Latham 1783, p. 210 Emberiza borbonica Gmelin 1789, p. 886 (Réunion) Status  Invalid taxon. Putative range  Réunion, Mascarene Islands.

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Appendix 2: Doubtful and invalid taxa

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).

Black-throated Euphonia  Euphonia vittata

Sclater

Euphonia vittata Sclater 1861b, p. 129 (Brazil) Tanagra catasticta Oberholser 1918, p. 125 Status  Doubtful taxon. Putative range  Brazil.

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.

Fantail Grosbeak  Spermophila laticauda

(Müller)

Grosbec appellé queue en eventail de Virginie. Daubenton 1770–86, no. 380 Loxia laticauda Müller 1776b, supplement, p. 151 Loxia flabellum Boddaert 1783, p. 23 Loxia flabellifera Gmelin 1788, p. 850 Status  Doubtful taxon. Putative range  Virginia, United States.

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.

Yellow-­bellied Grosbeak  Loxia virginica

Gmelin

Yellow-­bellied Grosbeak. Pennant 1785, p. 351 Loxia virginica Gmelin 1788, p. 849 Status  Doubtful taxon. Putative range  Virginia, United States.

Known only from the descriptions of Pennant and Latham, clearly from the same specimen. Unidentifiable, and ‘not an American bird’ (McAtee 1963).

Grey Grosbeak  Loxia grisea

Gmelin

Le Grisalbin. Buffon 1770–86, p. 467 Grosbec de Virginie. Daubenton 1770–86, 393, no. 1 Grey Grosbeak. Pennant 1785, p. 352 Loxia grisea Gmelin 1788, p. 857 Status  Doubtful taxon. Putative range  Virginia, United States.

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).

Linnaeus’s Serin  Loxia butyracea

Linnaeus

Loxia butyracea Linnaeus 1758, p. 174 (India) Status  Invalid taxon. Putative range  India.

Known only from Linnaeus’s description. Unidentifiable. 435

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Extinct Birds

Edwards’s Serin  Serinus butyracea

(Linnaeus)

Chloris indicus. Edwards 1747, p. 84 Fringilla butyracea Linnaeus 1758, p. 181 (Madeira) Status  Doubtful taxon. Putative range  Madeira.

Known only from the descriptions of Edwards and Linnaeus. Never identified.

Perkins’s ’Alauahio  Pareoreomyza perkinsi

(Rothschild)

Oreomyza perkinsi Rothschild 1900, p. 129 (Puulehua, Hawaii) Status  Invalid taxon. Putative range  Hawaii.

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).

Sassi’s Honeycreeper  Sassius simplex

Rothschild & Hartert

Sassius simplex Rothschild & Hartert 1926, p. 51 (no locality) Status  Invalid taxon. Putative range  Unknown.

Known only from the type, of unknown origin. Considered an artefact (Olson & Schifter 1989).

Brewster’s Linnet  Acanthis brewsteri

(Ridgway)

Aegiothus (flavirostris var) brewsteri Ridgway 1872, p. 434 (Waltham, Massachussets) Status  Invalid taxon. Putative range  Massachusetts, United States.

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.

Small-headed Flycatcher  Wilsonia microcephala

(Ridgway)

Muscicapa minuta Wilson 1812, p. 62, pl. 50, fig. 5 (New Jersey) Sylvania microcephala Ridgway 1885b, p. 354 Status  Doubtful taxon. Putative range  New Jersey, United States.

Known only from the descriptions of Wilson (and later Audubon 1829). Chapman (1888) listed it as hypothetical.

Louisiana Warbler  ‘Motacilla ludoviciana’

Gmelin

Le figuier à gorge jaune. Buffon, 1770–86, p. 288 Ficedula ludoviciana Brisson 1760, p. 500, pl. 26, fig.4 Louisiane Warbler. Pennant 1785, p. 407 Motacilla ludoviciana Gmelin 1789, p. 983 Status  Doubtful taxon. Putative range  Louisiana, United States.

Known only from the descriptions of Pennant and others. Considered unidentifiable (McAtee 1963). 436

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Appendix 2: Doubtful and invalid taxa

Orange-bellied Warbler  ‘Motacilla fulva’

Gmelin

Le figuier a gorge jaune. Buffon 1770–86, p. 317 Orange-bellied Warbler. Pennant 1785, p. 410 Motacilla fulva Gmelin 1789, p. 973 Status  Doubtful taxon. Putative range  Louisiana, United States.

Known only from the descriptions of Pennant and others. Considered unidentifiable (McAtee 1963).

Half-collared Warbler  ‘Motacilla semitorquata’

Gmelin

Le figuier a demi collier. Buffon 1770–86, p. 316 Half-collared Warbler. Pennant 1785, p. 410 Motacilla semitorquata Gmelin 1789, p. 972 Status  Doubtful taxon. Putative range  Louisiana, United States.

Known only from the descriptions of Pennant and others. Considered unidentifiable (McAtee 1963).

Bush Warbler  ‘Passer virginianus’

(Brisson)

Little Brown Sparrow? Catesby 1731 , p. 35 Passerculus simpliciter Klein 1750, p. 89 Passer virginianus Brisson 1760, p. 101 Rush Warbler. Latham 1783, p. 420 Bush? Warbler. Pennant 1785, p. 415 Status  Doubtful taxon. Putative range  Carolina and Virginia, United States.

Known only from the description of Catesby, from which are derived those of Pennant and others. Considered unidentifiable (McAtee 1963).

American Nightingale  ‘Motacilla calidris’

(Linnaeus)

American Nightingale. Edwards 1750, p. 121, pl. 121, fig. 2 Motacilla calidris Linnaeus 1758, p. 184 (based on Edwards) Status  Doubtful taxon. Putative range  Jamaica.

Known only from the description of Edwards, and never satisfactorily identified.

Awatcha Warbler  ‘Motacilla awatcha’

Gmelin

Awatcha Warbler. Pennant 1785, p. 422 (possibly based on Pallas’s manuscript) Motacilla awatcha Gmelin 1789, p. 986 Status  Doubtful taxon. Putative range  Kamtchatka.

Known only from Pennant’s description. Unidentifiable.

Long-­billed Warbler  ‘Motacilla camtschatkensis’

Gmelin

Long-­billed Warbler. Pennant 1785, p. 420 (possibly based on Pallas’s manuscript) Motacilla camtschatkensis Gmelin 1789, p. 986 Status  Doubtful taxon. Putative range  Kamtchatka.

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Extinct Birds

Known only from Pennant’s description. Bechstein (1795) considered it to be an earlier name for Acrocephalus palustris. Unidentifiable.

Cincinnati Warbler  Vermivora cincinatiensis

(Langdon)

Helminthophaga cincinatiensis Langdon 1880, p. 119 (Madisonville, Hamilton County, Ohio) Status  Invalid taxon. Putative range  Ohio, United States.

Known only from the type. Considered to represent a hybrid ­Blue-­winged Warbler Vermivora pinus x Kentucky Warbler Oporornis formosus (Ridgway 1880; Graves 1988).

Cowled Yellowthroat  Geothlypis cucullata

(Latham)

Sylvia cucullata Latham 1790, p. 528 (no locality) Cowled Warbler. Latham 1801a, supplement p. 243 Status  Doubtful taxon. Putative range  Unknown.

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.

Carbonated Warbler  Dendroica carbonata

(Audubon)

Sylvia carbonata Audubon 1829, pl. 60 (near Henderson, Kentucky) Status  Doubtful taxon. Putative range  Kentucky, United States.

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).

Blue Mountain Warbler  Dendroica montana

(Wilson)

Sylvia montana Wilson 1812, p. 113, pl. 44, fig. 2 (near the Blue Mountains, Virginia) Status  Doubtful taxon. Putative range  Virginia, United States.

Known only from the plates of Wilson and Audubon. Equated with Black-throated Green Warbler Dendroica virens (Mayr et al. 1986).

Sutton’s Warbler  Dendroica potomac

Haller

Dendroica potomac Haller 1940, p. 50 (12 miles south of Martinsburg, Berkeley County, West Virginia) Status  Invalid taxon. Putative range  West Virginia, United States.

Known from a few specimens; generally regarded as a hybrid Yellow-throated Warbler Dendroica dominica x Northern Parula Parula americana (Dunn & Garnett 1997).

Yellow-­tailed Flycatcher  ?Parula sp. Yellow-­tail Flycatcher. Edwards 1758, pl. 257, p. 101 Muscicapa ruticilla (part) Gmelin 1788, pt. 1, p. 935

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Appendix 2: Doubtful and invalid taxa Status  Doubtful taxon. Putative range  Hispaniola, West Indies.

Known only from Edwards’s and Pennant’s descriptions. Unidentifiable.

Cassin’s Cacique  Cacicus melanurus

(Cassin)

Cassicus melanurus Cassin 1867, p. 66 (Guayquil, Ecuador) Status  Invalid taxon. Putative range  Guayquil, Ecuador.

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.

Villavicencio’s Oriole  Icterus xantholemus

Gil Lletget

Icterus xantholemus Gil Lletget 1918, p. 340 (Ecuador) Status  Doubtful taxon. Putative range  Ecuador.

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).

Maelby Bunting  Emberiza maelbyensis

Sparrman

Emberiza Maelbyensis Sparrman 1786, tab. 21 Status  Doubtful taxon. Putative range  Maelby, Sweden.

Known only from Sparrman’s plate. Unidentifiable.

Louisiana Bunting  Emberiza ludovicia

Linnaeus

Hortulanus Ludovicianus. Brisson 1760, p. 278 L’Ortolan de la Louisiane. Buffon 1770–86, p. 325 Louisiane Bunting. Pennant 1785, p. 363 Emberiza Ludovicia Linnaeus 1766, p. 310 Status  Doubtful taxon. Putative range  Louisiana, United States.

Known from the descriptions of Brisson and Buffon, from which those of Pennant and Latham derive. Unidentifiable, and ‘not an American bird’ (McAtee 1963).

Norton Finch  Calcarius nortoniensis

(Gmelin)

Norton Finch. Latham 1783, p. 274 Fringilla nortoniensis Gmelin 1789, p. 922 Status  Doubtful taxon. Putative range  Norton Sound, Alaska.

Known only from the descriptions of Latham and Pennant. Unidentifiable (McAtee 1963).

Unalaskan Snow Bunting  Plectrophenax aunalaschkensis

(Gmelin)

Aoonalashkan Oriole. Latham 1782, p. 447 Unalaschka Oriole. Pennant 1785, p. 261 Oriolus aunalaschkensis Gmelin 1788, p. 394

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Extinct Birds Status  Doubtful taxon. Putative range  Unalaska, Aleutian Islands.

Known only from the descriptions of Pennant and Latham. Unidentifiable (McAtee 1963).

Hudsonian Snow Bunting  ?Plectrophenax hudsonius

(Gmelin)

Hudsonian White-headed Oriole. Pennant 1785, p. 260 Oriolus hudsonius Gmelin 1788, p. 387 Status  Doubtful taxon. Putative range  Hudson’s Bay, Canada.

Known only from Pennant’s description of a specimen, now lost, in the Leverian Museum. McAtee (1963) considered it a partial albino, and unidentifiable.

Texas Henslow’s Sparrow  Ammodramus henslowii houstonensis

Arnold

Ammodramus henslowii houstonensis Arnold 1983, p. 505 (Houston, Texas) Status  Invalid taxon. Putative range  Texas.

Now considered to fall within the range of variation seen in A. henslowii (Browning 1990).

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).

Gifford’s Finch  Camarhynchus giffordi

(Swarth)

Cactospiza giffordi Swarth 1929, p. 32 (Indefatigable Island, Galápagos) Status  Invalid taxon. Putative range  Indefatigable Island, Galápago Islands.

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).

Beck’s Finch  Camarhynchus conjunctus

Swarth

Camarhynchus conjunctus Swarth 1929, p. 33 (Charles Island, Galápagos) Status  Invalid taxon. Putative range  Charles Island, Galápagos Islands.

Known from two specimens, both adult males collected by Rollo Beck on February 28, 1906. Now considered to be a hybrid Small Tree Finch Camarhynchus parvulus x Warbler Finch Certhidea olivacea (McCarthy 2006).

Swarth’s Finch  Camarhynchus aureus

Swarth

Camarhynchus aureus Swarth 1929, p. 32 (Chatham Island, Galápagos) Status  Invalid taxon. Putative range  Chatham Island, Galápagos Islands.

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Appendix 2: Doubtful and invalid taxa

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).

Humbert’s Cardinal  Paroaria humberti

Angelini

Paroaria humberti Angelini 1901, p. 17 (South America) Status  Invalid taxon. Putative range  South America.

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).

Gould’s Tanager  Tangara gouldi

(Sclater)

Calliste gouldi Sclater 1886, p. 849 ­(south-­eastern Brazil) Status  Invalid taxon. Putative range ­South-­eastern Brazil.

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.

Arnault’s Tanager  Tangara arnaulti

Berlioz

Tangara arnaulti Berlioz 1927, p. 95 (South America) Status  Doubtful taxon. Putative range  South America.

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).

Purple Honeycreeper  Chlorophanes purpurascens

Sclater & Salvin

Chlorophanes purpurascens Sclater & Salvin 1873, p. 157 (Venezuela) Status  Doubtful taxon. Range  Unknown, but probably Trinidad.

Known only from the type, an adult male in Tring of uncertain origin. Considered to be a hybrid ­Red-­legged Honeycreeper Cyanerpes cyaneus x Green Honeycreeper Chlorophanes spiza (Storer 1957).

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Appendix 3 Rediscovered taxa

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 rediscovered 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 rediscovered 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. Species

Last seen

Found

Giant Canada Goose Branta canadensis maxima Mascarene Petrel Pseudobulweria aterrima Chatham Island Taiko Pterodroma magentae Bermuda Petrel Pterodroma cahow Jerdon’s Courser Rhinoptilus bitorquatus White-winged Guan Penelope albipennis Slender-billed Flufftail Sarothrura watersi Snoring Rail Aramidopsis plateni Drummer Rail Habroptila wallacii Takahe Porphyrio hochstetteri Blue-fronted Lorikeet Charmosyna toxopei Rufous-fronted Parakeet Bolborhynchus ferrugineifrons Indigo-winged Parrot Hapalopsittaca fuertesi Lear’s Macaw Anodorhynchus leari Seychelles Scops Owl Otus insularis Honduran Emerald Amazilia luciae Black-breasted Puffleg Eriocnemis nigrivestis Sri Lankan Dollarbird Eurystomus orientalis irisi Juan Fernandez Tit-tyrant Anairetes fernandezianus Royal Cinclodes Cinclodes aricomae White-browed Tit-spinetail Leptasthenura xenothorax Hoary-throated Spinetail Synallaxis kollari Orinoco Soft-tail Thripophaga cherriei Black-hooded Antwren Formicivora erythronotos

1970 1917 1931 1915 1831 1899 c.1870

>1960 1970 1978 1951 1986 1977 1988 1980 1981 1948 >1980 1985 1989 1978 1960 1988 1980 >1980 >1980 1982 1974 1956 1970 1970

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Appendix 3: Rediscovered taxa

Rio Branco Antbird Cercomacra carbonaria Moustached Antpitta Grallaria alleni Grey-winged Cotinga Tijuca condita Yellow-bellied Asity Neodrepanis hypoxantha Red-tailed Newtonia Newtonia fanovanae Everett’s White-eye Zosterops everetti everetti Yellow-throated Serin Serinus flavigula Noisy Scrub-bird Atrichornis clamosus Niceforo’s Wren Thryothorus nicefori Golden-naped Weaver Ploceus aureonucha Bronze-brown Cowbird Molothrus (aeneus) armenti

1831 1911 1942 1980 1989 1961 1989 1986 1969

(Oustalet)

Talegallus Bruijnii Oustalet 1880, p. 906. (Waigeo Island) Status  Last collected in 1932 and thought to be possibly extinct. Rediscovered in 2002. Range  Waigeo Island, ­north-­west of New Guinea.

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 incubation 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.

Manipur Bush-quail  Perdicula manipurensis

Hume

Perdicula manipurensis Hume 1880, 9, p. 467 (Manipur Hills) Status  Last recorded in 1932. Rediscovered in 2006. Range  West Bengal, Assam, Nagaland, Manipur and Meghalaya, North-west India and also Bangladesh.

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.

Madagascar Pochard  Aythya innotata

(Salvadori)

Nyroca innotata Salvadori 1894, p. 2 (Madagascar) Status  Feared extinct after death of a bird in captivity in 1992. Rediscovered in 2006. Range  Central Madagascar.

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.

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Extinct Birds

Beck’s Petrel  Pseudobulweria becki

(Murphy)

Pterodroma becki Murphy 1928, p. 1 (at sea, 3°S, 155°E, east of New Ireland and north of Buka in the Solomons) Status  Known from two specimens collected in 1928 and thought to be probably extinct. Rediscovered in 2007. Range  Solomon Islands and Bismarck Archipelago, western Pacific.

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.

Fiji Petrel  Pseudobulweria macgillivrayi

(Gray)

Thalassidroma (Bulweria) Macgillivrayi G.R. Gray 1859a, p. 56 (Ngau, Fiji Islands) Status  Originally only known with certainty from the type collected in 1855. Recently recorded off Gau Island, Fiji. Range  Gau Island, Fiji.

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.

New Zealand ­Storm-­petrel  Oceanites maorianus

(Mathews)

Pealeornis maorianus Mathews 1932, p.132 (Banks Peninsula, New Zealand) Status  Thought to be extinct since 1850, rediscovered in 2003. Range  North Island, New Zealand.

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 rediscovery 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 ­Storm-­petrel remains mysterious. The location of its distribution and breeding areas are still unknown.

Príncipe Dwarf Olive Ibis   Bostrychia bocagei rothschildi

(Bannerman)

Lampibris rothschildi Bannerman 1919, p. 6 (Infant d’Henrique, Príncipe Island, Gulf of Guinea) Bostrychia olivacea rothschildi Steinbacher 1979, p. 261 (= B. b. rothschildi) Status  Originally known from the type and a few other specimens. Rediscovered in 1991. Range  Príncipe Island, off the coast of West Africa.

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).

Tasman Booby  Sula dactylatra tasmani

van Tets et al. 1988

Sula tasmani van Tets et al. 1988, p. 45. (Kingston, Norfolk Island) Status  Originally known from subfossil remains and early accounts, and thought to have become extinct in 1788. Range  Lord Howe and Norfolk Islands.

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Appendix 3: Rediscovered taxa

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).

Madagascar ­Serpent-­eagle  Eutriorchis astur

Sharpe

Eutriorchis astur Sharpe 1875, p.73, pl. 13 (Madagascar) Status  Last collected in 1930 and thought to be extinct. Rediscovered in 1993. Range  Eastern Madagascar.

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.

Volcano Islands Peregrine Falcon  Falco peregrinus fruitii

Momiyama

Falco peregrinus fruitii Momiyama 1927, p. 71 (in Japanese), 97 (in English) ­(Isino-­mura, San Alessandro Islands = Iwo Jima) Status  Not reliably recorded since 1945 and thought possibly extinct. Survives on Kita Iwo Jima in small numbers. Range  Kazan Rettō or Volcano Islands, south of Honshu, Japan, and possibly the Ogasawara (Bonin) Islands.

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 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.

Austral Rail  Rallus antarcticus

King

Rallus antarcticus King 1828, p. 95 (Straits of Magellan) Status  Last recorded in 1959. Rediscovered in 1998. Range  Central Chile and central Argentina.

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.

Sakalava Rail  Amaurornis olivieri

Grandidier & Berlioz

Amaurornis olivieri Grandidier & Berlioz 1929, p. 83 (Madagascar) Status  Last recorded in 1962. Rediscovered in 1995. Range  Western Madagascar.

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. 445

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Utila Chachalaca  Ortalis vetula deschauenseei

Bond

Ortalis vetula deschauenseei Bond 1936b, p. 356 (Utila Island) Status  Thought to have become extinct since 1953, but rediscovered in 2005. Range  Utila Island, off the Caribbean coast of Honduras.

The Utila Chachalaca is a subspecies of the ­wide-­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.

Chinese Crested Tern  Sterna bernsteini

(Schlegel)

Sterna bernsteini Schlegel 1863, p. 9 (Kaou, eastern coast of Halmahera) Status  Recorded in 1863, 1906, 1937 and 1991; breeding birds rediscovered in 2000. Range  Breeds off the east coast of China. Non-breeding range includes Thailand, the Philippines and Indonesia.

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).

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.

Silvery Pigeon  Columba argentina

Bonaparte

Columba argentina Bonaparte 1855b, p. 36 (Indian Archipelago = Borneo) Status  Last officially recorded in 1931. Rediscovered in 2008. Range  Simeulue and Mentawai Islands; Riau and Lingga Archipelagos; Anambas Island, North Natunas and Karimata Island, Burong Island (Malaysia), mainland Borneo and possibly Sumatra.

The Silvery Pigeon was originally a common species most often observed on small islands, but it disappeared 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.

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Bonin Islands Wood Pigeon Columba janthina nitens

(Stejneger)

Janthoenas nitens Stejneger 1887d, p. 421 (Bonin-Shima, Japan) Status  Last seen in the 1980s, but rediscovered in 1998. Range  Ogasawara or Bonin Islands – Mukojima, Chichijima, Hahajima and Kitanojima – and Iwo Jima in the Volcano Islands, Japan.

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.

Blue-fronted Macaw  Ara glaucogularis

Dabbene

Ara glaucogularis Dabbene 1921, p. 225 (Santa Cruz de la Sierra, Santa Cruz, Bolivia) Status  Known from captive birds only since 1922. Rediscovered in the wild in 1992. Range  Llanos de Mojos, northern Bolivia.

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.

Sumatran Ground Cuckoo  Carpococcyx viridis

Salvadori

Carpococcyx viridis Salvadori 1879, p. 187 (Mount Singgalang, Sumatra) Status  Last collected in 1916. Rediscovered in 1997. Range  Sumatra, Indonesia

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.

Taliabu Masked Owl (Black-brown Owl)  Tyto nigrobrunnea

Neumann

Tyto nigrobrunnea Neumann 1939, p. 89 (Taliaboe, Sula Islands) Status  Originally known from the type, collected in October 1938. Rediscovered in 1991. Range  Taliabu, Sula Archipelago.

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 ­long-­term survival.

Comoro Scops Owl  Otus capnodes

(Gurney)

Scops capnodes Gurney 1889, p. 104 (Anjouan) Status  Originally known from just over 30 specimens, all collected in the mid-1880s by Humblot, and thought to be probably extinct. Rediscovered in June 1992. Range  Anjouan, Comoros Islands.

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The Comoro Scops Owl had apparently not been seen since Humblot collected the type series in the mid1880s. 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 Anjouan must, however, remain a concern.

Flores Scops Owl  Otus alfredi

(Hartert)

Pisorhina alfredi Hartert 1897, p. 527 (Repok Mts, Flores) Otus alfredi Peters 1940, p.89 Status  Last collected in 1896. Rediscovered in 1994. Range  Flores, Indonesia.

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 considered endangered.

Forest Owlet  Heteroglaux blewitti

Hume

Heteroglaux blewitti Hume 1873, p. 468 (Busnah, Phooljan State, India) Status  Known from seven specimens and last recorded in 1884. Rediscovered in 1997. Range  Central India.

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 (Rasmussen & King 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.

Sulawesi Eared Nightjar (Satanic Nightjar)   Eurostopodus diabolicus

Stresemann

Eustopodius diabolicus Stresemann 1931, p. 103 (Kumarsot, base of Klabat Volcano, north Sulawesi) Status  Last recorded in 1931. Rediscovered in 1996. Range  North and central Sulawesi.

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 difficulty 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.

Mexican Woodnymph  Thalurania ridgwayi

Nelson

Thalurania ridgwayi Nelson 1900, p. 262 (San Sebastian, Jalisco, Mexico) Status  Originally known only from the type. Extant but rare and declining. Range  Western Mexico.

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. 448

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Javan Buff-rumped Woodpecker  Meiglyptes tristis tristis

(Horsfield)

Meiglyptes tristis Horsfield 1821, p.177 (west Java) Status  The last authenticated record was in 1880. Rediscovered in 2010. Range  Java.

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 (Piauí Woodpecker)  Celeus obrieni

Short

Celeus spectabilis obrieni Short 1973, p. 465 (Piauí, Brazil) Status  Originally known only from the type, a female collected in 1926, and not seen again. Rediscovered in 2006. Range  Piauí, Brazil.

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.

Golden-crowned Manakin  Lepidothrix vilasboasi 

Sick

Lepidothrix vilasboasi Sick 1959, p. 111 (headwaters of River Cururu-ri, south-west Pará, Brazil) Status  Not seen since 1957. Rediscovered in 2002. Range  Rio Tapajós, Serra do Cachimbo and Rio Jamanxim, near Novo Progresso, south-west Pará, Brazil.

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 situated 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 Goldencrowned Manakin is considered vulnerable, with a declining population (BirdLife International 2011).

Cocha Antshrike  Thamnophilus praecox

Zimmer

Thamnophilus praecox Zimmer 1937, p. l (mouth of Lagarto Cocha, Ecuador) Status  Originally known only from the type, a female, taken along the Ecuador/Perú border on the upper River Napo. Rediscovered in 1991. Range  Along the Río Lagarto in east Napo, and near La Selva on the north and south sides of the Río Napo, east Ecuador.

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).

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White-masked Antbird  Pithys castaneus

Berlioz

Pithys castaneus Berlioz 1938 (Andoas, lower Río Pastaza, Ecuador-Peru) Status  Originally known from a single specimen collected in 1938. Rediscovered in 2001. Range  Northern Peru

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 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 widespread (van Osten et al. 2007).

Brown-banded Antpitta  Grallaria milleri milleri

Chapman

Grallaria milleri Chapman, 1912, p. 147 (Laguneta, near Quindío Pass, Caucam, Colombia) Status  Last recorded in 1942. Rediscovered in 1994. Range  Volcán Ruíz Tolima massif, central Andes, Colombia.

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).

Recurve-billed Bushbird  Clytoctantes alixii

Elliot

Clytoctantes alixii Elliot 1870, p. 242 Status  Last recorded in 1965. Rediscovered in 2004. Range  Norte de Santander, Colombia and Sierra de Perijá, Venezuela.

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 distribution 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.

Stresemann’s Bristlefront  Merulaxis stresemanni

Sick

Merulaxis stresemanni Sick 1960, p.155 (Bahia, Brazil) Status  Originally known from two specimens, the last taken in 1945, and thought possibly extinct. Rediscovered in 1995. Range  Coastal Bahia, east Brazil.

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 ­valley-­bottom forest for logging, cattle ranching and agriculture continues to adversely affect the population. It is considered critically endangered, and the population is now tiny (BirdLife International 2011).

Obscure Berrypecker  Melanocharis arfakiana

(Finsch)

Dicaeum arfakianum Finsch 1900, p. 70 (Arfak Mountains, New Guinea) Status  Originally known from two specimens, the last collected in 1933. Rediscovered in the 1990s. Range  Arfak Mountains and Matsika, ­south-­eastern New Guinea.

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, 450

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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.

Cebu Mottle-breasted Bulbul  Hypsipetes siquijorensis monticola

(Bourns & Worcester)

Iole monticola Bourns & Worcester 1894, p. 25 (Cebu) Status  Thought to have become extinct in 1906; rediscovered in the 1990s. Range  Cebu, Philippines.

The Cebu ­Mottled-­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.

Sooty Babbler  Stachyris herberti

(Baker)

Nigravis herberti Baker 1920, p. 10 (Ban Sao, Thailand; error = Ban Lak Sao, Laos) Stachyris nigriceps nigriceps Baker 1930, p. 52 Status  Last recorded in 1920. Rediscovered in 1994. Range  Central Laos and central Vietnam.

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.

Rusty-throated Wren-babbler  Spelaeornis badeigularis

Ripley

Spelaeornis badeigularis Ripley 1948, p. 103 (Dreyi, Mishmi Hills, north-eastern Assam, India) Status  Last recorded in 1947. Rediscovered in 2004. Range  Mishmi Hills, eastern Arunachal Pradesh, India.

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 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.

Grey-crowned Crocias  Crocias langbianis

Gyldenstolpe

Crocias langbianis Gyldenstolpe 1939, p. 2 (Langbian plateau, Annam) Status  Last recorded in 1938–39. Rediscovered in 1994. Range  Da Lat plateau, southern Vietnam.

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.

Cherry-­throated Tanager  Nemosia rourei

Cabanis

Nemosia rourei Cabanis 1870, p. 459 Status  Originally known only from the type collected in 1870. Rediscovered in 1998. Range  Atlantic forest in Espírito Santo, Brazil.

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The enigmatic ­Cherry-­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 endangered, with a population of probably less than 250 birds (BirdLife International 2011).

Gold-ringed Tanager  Bangsia aureocincta

(Hellmayr)

Buthraupis aureocincta Hellmayr 1910, 3 (Tatamá Mountains, Chocó, Colombia) Bangsia aureocincta Ridgely & Tudor 1989 p. 293 Status  Last recorded in 1946. Rediscovered in 1992. Range  Pacific Andean coast, Colombia.

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.

Cone-­billed Tanager  Conothraupis mesoleuca

(Berlioz)

Rhynchothraupis mesoleuca Berlioz 1939, p. 102 (Juruena, ­north-­east of Cuyaba, Matto Grosso, Central Brazil) Status  Found in 1938 but only seen again in 2003. Range  Mato Grosso, Brazil.

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.

Daitō Bush Warbler  Cettia diphone restricta

(Kuroda)

Horornis cantans restrictus Kuroda, 1923, p. 122. ­(Minami-­daito-­jima, Borodino Islands) Status  Last recorded around 1942. Rediscovered in 2002. Range  Minami ­Daitō-­jima, Daitō (or Borodino) Islands.

The Daitō Bush Warbler was last observed on Minami ­Daitō-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 Daitō Bush Warbler is not uncommon, and the population appears to be stable.

São Tomé Short-tail  Amaurocichla bocagii

Sharpe

Amaurocichla bocagii Sharpe 1892, p. 228, pl. 20 (San Miguel, west coast of São Tomé) Status  Not seen since 1928. Rediscovered in 1990. Range  São Tomé, Gulf of Guinea.

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. 452

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Large-­billed Reed Warbler  Acrocephalus orinus

Oberholser

Phyllopneuste macrorhyncha Hume 1869, p. 357, nomen nudum Acrocephalus macrorhynchus Hume 1871, p. 31 (‘not far from Rampoor’ = Ramphur, Sutlej Valley, Himal Pradesh, India) Acrocephalus orinus Oberholser 1905b, p. 899 Status  Originally only known from the type specimen. Rediscovered in 2006. Range  Ranipur, Himalayas, India.

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 ­Blunt-­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 addition, 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 exhibition 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.).

Long-­legged Thicketbird  Trichocichla rufa rufa

Reichenow

Trichocichla rufa Reichenow 1890, p. 489 (Vitu Levu, Fiji) Status  Originally known from four specimens, the last obtained in 1894 and not seen again until rediscovered in 2002. Range  Vitu Levu, Fiji.

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 observed by BirdLife International researchers during ornithological surveys of the island (Dutson & Masibalavu 2004). The ­Long-­legged Thicketbird appears to be comparatively widespread and the population stable. Unless major changes occur, it is in no immediate threat of extinction.

Lompobatang Flycatcher  Ficedula bonthaina

(Hartert)

Siphia bonthaina Hartert 1896, p. 157 (Bonthain Peak = Mount Lompobattang, Sulawesi) Ficedula bonthaina White & Bruce 1986, p. 357 Status  Last recorded in 1931. Rediscovered in 1995. Range  Lompobatang massif, Sulawesi, Indonesia.

The Lompobatang Flycatcher was considered common when it was first discovered, but numbers subsequently 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 continuing loss of habitat, the long term future of the Lompobatang Flycatcher appears to be bleak.

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Paler Flowerpecker  Dicaeum trigonostigma pallidius

Bourns & Worcester

Dicaeum pallidior Bourns & Worcester 1894, p. 18 (Cebu) Status  Thought extinct since 1910 but rediscovered in 2003. Range  Cebu, Philippine Islands.

The Paler Flowerpecker, a race of the widespread ­Orange-­bellied Flowerpecker Dicaeum trigonostigma, was thought to have become extinct soon after McGregor collected it in 1906. At this time it was the commonest flowerpecker on the island of Cebu (McGregor 1907). Another specimen was reported by Richardson & Baker (1981), collected by J. C. Mounsey at Camp Ialiti, Cebu, on July 26, 1910. This was thought to have been among the last of the birds, but the Paler Flowerpecker had managed to survive in tiny pockets of remnant forest, and was rediscovered in February 2003 (Paguntalan & Jakosalem 2008). The Paler Flowerpecker is considered to be extremely rare.

Four-coloured Flowerpecker  Dicaeum quadricolor

(Tweeddale)

Prionochilus quadricolor Tweeddale 1877, p. 762, pl. 77, fig. 2 (Cebu) Status  Believed to have become extinct in 1906, but rediscovered in the 1990s. Range  Cebu, Philippine Islands.

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 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.

São Tomé Fiscal  Lanius newtoni

Bocage

Lanius newtoni Bocage 1891, p. 79 (São Tomé) Status  Last recorded in 1928 and thought to be possibly extinct. Rediscovered in 1990. Range  São Tomé, in the Gulf of Guinea.

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 ­south-­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 endangered, and the population is declining (Dallimer et al. 2009).

Cerulean Paradise-flycatcher  Eutrichomyias rowleyi

(Meyer)

Zeocephus rowleyi Meyer 1878, p. 163 (Tabukan, Great Sangi =Sangihe) Eutrichomyias rowleyi Meise 1947, p. 135 Status  Known from a single specimen collected in 1873. Rediscovered in 1998. Range  Sangihe Island, Indonesia.

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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 considered 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.

Banggai Crow  Corvus unicolor

(Rothschild & Hartert)

Gazzola unicolor Rothschild & Hartert 1900, p. 29 (Banggai, Sula Islands) Status  Last recorded in the late 19th century. Rediscovered in 2007. Range  Peneng Island, Banggai Archipelago, Indonesia.

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 population 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.

King Island Brown Thornbill  Acanthiza pusilla archibaldi

Mathews

Acanthiza archibaldi Mathews 1910a, p. 501 (King Island) Status  Long thought to be probably extinct, but two were seen in March 2002. Range  King Island, off Tasmania.

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 ­mist-­netted at Loorana in 1971 (McGarvie & Templeton 1974). The thornbill was thought to be possibly extinct until two individuals were located in March 2002 (Donaghey & Lloyd 2003). It is considered critically endangered.

Nendö Shrikebill  Clytorhynchus sanctaecrucis

Mayr

Clytorhynchus nigrogularis sanctaecrucis Mayr 1933c, p.20 (Nendo, Santa Cruz Islands, Solomon Islands) Status  Originally known from two specimens taken in 1927. Rediscovered in 2004. Range  Nendö, Santa Cruz Islands, Solomon Islands.

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.

White-tailed Tityra  Tityra leucura

Pelzeln

Tityra (Erator) leucura Pelzeln 1868, pp. 120, 183 (Salto do Girao = Girau, Rio Madeira, Brazil) Status  Originally known only from the type, collected in October 1829. Rediscovered in 2007. Range  Rio Madeira, Brazil.

The original specimen of the ­White--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 455

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taxon. Its scarcity is due to habitat destruction, and although the size of the population is unknown, it must be extremely small.

Salvin’s Purpletuft  Iodopleura pipra leucopygia

Salvin

Iodopleura leucopygia Salvin 1885, p. 305 (British Guiana) Status  Originally known from two specimens of unknown origin. It was thought to be possibly extinct. Range  Paraíba, Pernambuco, Alagoas and Bahia, ­north-­east Brazil.

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.

St Lucia Forest Thrush  Cichlherminia lherminieri sanctaeluciae

(Sclater)

Margarops ­sanctae-­luciae Sclater 1880, p. 73 (St Lucia, Lesser Antilles) Status  Thought to have become extinct by 1980, but rediscovered in 2007. Range  St Lucia, West Indies.

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.

Javan Large Wren Babbler  Turdinus macrodactyla lepidopleura

(Bonaparte)

Myiothera lepidopleura Bonaparte 1850, p. 257 (Java) Status  Not recorded since the mid-20th century, but rediscovered in 2000. Range  Java, Indonesia.

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 mid20th 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).

Lord Howe ­Silver-­eye  Zosterops tephropleura

Gould

Zosterops lateralis tephropleura Gould 1855, p. 166 (Lord Howe Island) Status  Thought to have become extinct by 1940, but extant and reasonably common. Range  Lord Howe Island.

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. 265). 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).

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Eastern ­Thick-­billed Grasswren  Amytornis textilis modestus

(North)

Amytis modesta North 1902, p. 103 (Meerenie Bluff, Macdonnell Ranges, central Australia) Status  Suffered serious decline in the late 19th and early 20th centuries, and thought possibly extinct. Survives in low densities. Range  South Australia (formerly New South Wales).

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.

Lufira Masked Weaver  Ploceus ruweti

Louette & Benson

Ploceus ruweti Louette & Benson 1982, p. 26 (Lake Lufira, DR Congo) Status  Thought to be possibly extinct, but rediscovered in 2009. Range  Lake Lufira, DR Congo.

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.

Pale-headed Brush-finch  Atlapetes pallidiceps

(Sharpe)

Buarremon pallidiceps Sharpe 1900, p. 2 (Guishapa, Oña, Azuay, Ecuador) Atlapetes pallidiceps Hellmayr 1938, p. 411. Status  Last recorded in 1969. Rediscovered in 1998. Range  Río Jubones basin, in Azuay and Loja, southern Ecuador.

The Pale-headed Brush-finch was not seen from 1969 until its rediscovery in November 1998, when extensive 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.

São Tomé Grosbeak  Neospiza concolor

(Bocage)

Amblyospiza concolor Bocage 1888, p. 229 (São Tomé) Status  Originally known from three ­19th-­century specimens and considered probably extinct. Rediscovered in 1991. Range  São Tomé, Gulf of Guinea.

The São Tomé Grosbeak was first collected in the late 19th century and not seen again, despite several extensive surveys in the first half of the 20th century (Collar & Stuart 1950). 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 population 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.

AUSTRALIA Adelaide, South Australia South Australian Museum Brisbane, Queensland Queensland Museum Hobart, Tasmania Tasmania Museum and Art Gallery Launceston, Tasmania Queen Victoria Museum and Art Gallery Melbourne Museum Victoria Perth, Western Australia Western Australian Museum Sydney Australian Museum MacLeay Museum AUSTRIA Vienna Naturhistorisches Museum BELGIUM Antwerp Natuurwetenschappelijk Museum Brussels Institut Royal des Sciences Naturelles de Belgique Mons Musée d’Histoire Naturelle Tervuren Musée Royal de l’Afrique Central

Ottawa, Ontario Canadian Museum of Nature Quebec Musée de la Province de Québec St. John, New Brunswick The New Brunswick Museum Toronto, Ontario Royal Ontario Museum Winnipeg, Manitoba The Manitoba Museum COLOMBIA Bogotá Museo de la Universidad de la Salle Unidad Investigativa Federico Medem Medellin Museo del Colegio San José Popayan Museo de Historica, Natural de la Universidad del Cauca CUBA Havana Felipe Poey Natural History Museum National Natural History Museum CZECH REPUBLIC Prague Národni Muzeum

BRAZIL São Paulo Museu de Zoologia da Universidade de São Paulo

DENMARK Copenhagen Zoologisk Museum Uppsala Museum of Evolution

BULGARIA Sofia National Museum of Natural History

FIJI Suva Fiji Museum

CANADA Kingston, Ontario Biological Museum

FINLAND Helsinki Finnish Museum of Natural History

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FRANCE Autun Musée d’Histoire Naturelle de Ville d’Autun Besançon Musée d’Histoire Naturelle Bordeaux Musée d’Histoire Naturelle de Bordeaux Caen Musée d’Histoire Naturelle de Caen Cassel Musée d’Histoire Naturelle de Kassel La Rochelle Muséum d’Histoire Naturelle Lyon Musée Guimet d’Histoire Naturelle Metz Musée Municipal Mountauban Musée d’Histoire Naturelle de Mountauban Nancy Muséum–Aquarium de Nancy Nantes Muséum d’Histoire Naturelle de Nantes Paris Muséum National d’Histoire Naturelle Poitiers Musée d’Histoire Naturelle Réunion Muséum d’Histoire Naturelle Rouen Musée d’Histoire Naturelle de Rouen Strasbourg Musée Zoologique de l’Université Louis Pasteur et de la Ville de Strasbourg Toulouse Muséum d’Histoire Naturelle de Toulouse Troyes Musée des Sciences Naturelles Vernon Musée de Vernon GERMANY Augsberg Naturwissenschaftliches Museum Berlin Museum für Naturkunde Bonn Zoologisches Forschungsinstitut und Museum A. Koenig Braunschweig Staatliches Naturhistorisches Museum Bremen Überseemuseum Dresden Staatliches Museum für Tierkunde Frankfurt am Main Forschungsinstitut und Naturmuseum Senckenberg

Görlitz Staatliches Museum für Naturkunde Görlitz Göttingen Zoologisches Museum Halberstadt Museum Heineanum Hamburg Zoologisches Institut und Zoologisches Museum Kassel Naturkunde Museum im Ottoneum Kiel Zoologisches Museum der ChristianAlbrechts-Universität Leipzig Naturkundermuseum Leipzig Munich Zoologische Staatssammlung Stuttgart Staatliches Museum für Naturkunde Wiesbaden Landesmuseum Wiesbaden GUADALOUPE Point a Pitre Musée l’Herminier ITALY Bologna Museo dell’Istituto Nazionale per la Fauna Selvatica Florence Museo Zoologico ‘La Specola’, Sezione del Museo di Storia Naturale Genoa Museo Civico di Storia Naturale ‘Giacomo Doria’ Milan Museo Civico di Storia Naturale de Milan Naples Centro ‘Museo delle Scienze Naturali’ Pisa Museo Storia Naturale e del Territoro Rome Museo Civico di Zoologia di Roma Siena Museo Zoologico dell’ Accadamia dei Fisiocritici Turin Museo Regionale di Scienze Naturali INDONESIA Cibinong, Bogor Museum Zoologicum Bogoriense IRELAND Dublin National Museum of Ireland 459

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JAMAICA Kingston The Institute of Jamaica JAPAN Tokyo Imperial University Ministry of Agriculture and Forestry Yamashina Institute for Ornithology MADAGASCAR Antananarivo, Madagascar Museum of Ethnology and Palaeontology THE NETHERLANDS Amsterdam Zoölogisches Museum, University of Amsterdam Leiden Naturalis, Nationaal Natuurhistorisch Museum Nijmegen Natuurmuseum Nijmegen

PAPUA NEW GUINEA Port Moresby National Museum and Art Gallery THE PHILIPPINES Manila Philippines National Museum, UPLB Museum of Natural History, Los Baños, Luzon POLAND Stettin Museum des Staat Stettin PORTUGAL Lisbon Museo Bocage Madeira Museum of Natural History Funchal QATAR Doha Al-Wabra Wildlife Preservation Center Qatar National Museum

NEW CALEDONIA Nouméa Départment des Sciences Humaines

RODRIGUES Rodrigues François Leguat Museum

NEW ZEALAND Auckland Auckland Institute and Museum Christchurch Canterbury Museum Dunedin Otago Museum Invercargill Southland Museum and Art Gallery Manawatu Te Manawa Museum of Art, Science and History Nelson Nelson Provincial Museum Norfolk Island Norfolk Island Museum Otorohanga Waitomo Caves Museum Wanganui Whanganui Regional Museum Wellington Museum of New Zealand Te Papa Tongarewa

RUSSIA Moscow Darwin Museum St Petersburg Zoologicheskii Institut, Rossiiskoi Akademii Nauk (Russian Academy of Sciences)

NORWAY Bergen Bergens Museum Oslo Zoologisk Museum

SINGAPORE Raffles Museum of Biodiversity Research SOUTH AFRICA Durban, South Africa Durban Natural Science Museum SPAIN Tenerife Museo de Ciencias Naturales de Tenerife Museo Canario Santa Cruz de Tenerife SWEDEN Gothenburg Göteborgs Naturhistoriska Museum Lund Zoological Institute and Musuem of the State University Malmö Malmö Natural History Museum Stockholm Naturhistoriska Riksmuseet

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Uppsala Evolutionsmusset, Zoologi–aektionen, Uppsala Universitet SWITZERLAND Basel Naturhistoriches Museum Geneva Muséum d’Histoire Naturelle Lausanne Musée Zoologique de l’Université Neuchâtel Muséum d’Histoire Naturelle TAHITI Institut de Paléontologie, Muséum National d’Histoire Naturelle  THAILAND Bangkok Thailand Institute for Scientific and Technological Research. UNITED KINGDOM Bath Bath Royal Literary and Scientific Museum Birmingham Birmingham Museum and Art Gallery Bolton Bolton Museums, Art Gallery, and Aquarium Brighton Booth Museum of Natural History Cambridge University Museum of Zoology Cardiff National Museum and Galleries of Wales Edinburgh National Museums of Scotland Royal Museum of Scotland Exeter Royal Albert Memorial Museum and Art Gallery Glasgow Hunterian Museum and Art Gallery Huddersfield Tolson Memorial Museum Leeds Leeds Museum Resource Center Liverpool World Museum London Grant Museum of Zoology and Comparative Anatomy Museum of the Royal College of Surgeons of England The Natural History Museum Newcastle-upon-Tyne The Hancock Museum

Norwich Norwich Castle Museum Nottingham Nottingham Natural History Museum Oxford Oxford University Museum of Natural History Sheffield Weston Park Museum Tring The Natural History Museum Wakefield Wakefield Museum UNITED STATES Albany, New York New York State Museum Alfred, New York Allen Steinheim Museum of Natural History Andover, Massachusetts Phillips Andover Academy Ann Arbor, Michigan Michigan Academy of Science, Arts and Letters Museum of Zoology, University of Michigan Baton Rouge Louisiana State University Museum Berkeley, California University of California Museum of Vertebrate Zoology Bismarck, North Dakota Museum of Natural History of the State University and School of Mines of North Dakota Bloomfield Hills, Michigan Cranbrook Institute of Science Buffalo, New York Buffalo Museum of Science Burlington, Vermont Robert Hull Fleming Museum (University of Vermont) Cambridge, Massachusetts Museum of Comparative Zoology, Harvard University Charleston, South Carolina Charleston Museum Chicago, Illinois Chicago Academy of Sciences Field Museum of Natural History Cleveland, Ohio Cleveland Museum of Natural History Davenport, Iowa Davenport Public Museum Dayton, Ohio Dayton Public Library and Museum Denver, Colorado Denver Museum of Natural History 461

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Fond du Lac, Winsconsin Fond du Lac Public Museum Gainsville, Florida Florida Museum of Natural History Galesburg, Illinois Albert Hurd Museum of Natural History of Knox College Granville, New York Pember Library and Museum Green Bay, Wisconsin Neville Public Museum Grinnell, Iowa Parker Museum of Natural History Hastings, Nebraska Hastings Museum Hawaii Bernice Pauahi Bishop Museum Hawaii Volcanoes National Park St. Louis College Ithaca Cornell University Lawrence, Kansas Dyche Museum of Natural History (University of Kansas) Los Angeles, California Los Angeles County Museum of Natural History Madison, Wisconsin Zoological Museum (University of Wisconsin) Milwaukee, Wisconsin Milwaukee Public Museum Minneapolis, Minnesota Museum of Natural History (University of Minnesota) Newark, New Jersey Newark Museum New Haven, Connecticut Peabody Museum of Natural History (Yale University) New London, Wisconsin New London Public Museum New York American Museum of Natural History Northfield, Minnesota Carleton College Museum Oakland, California Oakland Public Museum Oklahoma Sam Noble Oklahoma Museum of Natural History

Palo Alto, California Zoological Museum of Leland Stanford Junior University Philadelphia Academy of Natural Sciences Pittsburgh, Pennysylvania Carnegie Museum Pittsfield, Massachusetts Berkshire Athanaerum and Museum Princeton, New Jersey Natural Science Museum of Princeton University Providence, Rhode Island Park Museum Raleigh, North Carolina North Carolina State Museum St. Johnsbury, Vermont The Fairbanks Museum of Natural Science St. Paul, Minnesota Hamline University Museum San Diego, California Natural History Museum San Francisco California Academy of Sciences Pacific Museum of Ornithology Santa Barbara, California Santa Barbara Museum of Natural History Seattle, Washington University of Washington Thomas Burke Memorial Washington State Museum Springfield, Illinois Illinois State Museum Springfield, Massachusetts Museum of Natural History Syracuse, New York Natural Science Museum of Syracuse University Trenton, New Jersey New Jersey State Museum University, Alabama Alabama Museum of Natural History Washington, D. C United States National Museum Wilmington, Delaware Delaware Museum of Natural History Worcester, Massachusetts Worcester National History Museum

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531

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Index

Acanthis brewsteri 436 Acanthiza pusilla archibaldi 455 Accipiter alphonsi 370–1 butleri butleri 80–1 efficax 80 ferox 370 quartus 80 Aceros sp. 212 Acestura decorata 420 Acrocephalus caffer garretti 258 caffer musae 258–9 familiaris familiaris 259 longirostris 257–8 (luscinia) astrolabii 256 luscinius yamashinae 257 orinus 453 Adzebill, North Island 37 South Island 37–8 Aegotheles novaezealandiae 203 savesi 352 Aegypius chincou 368 cristatus 368 Aepyornis gracilis 26 hildebranti 26 maximus 24–6 medius 26–7 bruijnii 443 Aerodramus manuoi 204 spodiopygius ingens 205 whiteheadi whiteheadi 353 Agapornis picta 397 Aidemedia chascax 300 lutetiae 301 zanclops 300 Aimophila ruficeps sanctorum 322 ‘Akepa, Maui 301–2 Oahu 301 ‘Akialoa, H ­ oope-­billed 296 Kaua’i (Greater) 293–4 Lesser (Hawaii) 294–5 Maui Nui 294 Oahu 295–6 ‘Alauahio, Lanai 289–90 Molokai 290 Oahu 291 Perkins’s 436 Alcedo euryzona euryzona 210 Alectroenas sp. 161–2 nitidissima 159–60 Alisterus sp. 390 Alopochen kervazoi 46–7 mauritiana 46

532

08 Index.indd 532

‘Amahiki, Greater 292–3 Amaui 281 Amaurocichla bocagii 452 Amaurolimnas concolor concolor 101 Amaurornis olivieri 445 Amazilia aenobrunnea 413 alfaroana 206–7 bangsi 415 caeruliceps 413 cyaneotincta 413 florenceae 414 inculta 415 lerdi 414 lucida 414 microrhyncha 413 neglectus 414 nunezi 415 ocai 414 salvini 414 veneta 413 versicolor meracula 415 Amazon, Aruba 186 Culebra Island 186 Finsch’s 404 Gerini’s 403 Guadeloupe 338–9 Lilac 403 Martinique 339 Paradise 404 Primeval 404 ­Red-­and-­blue 403 ­Red-­and-­yellow 403 Amazona sp. 403 atwoodi 402 barbadensis canifrons 186 caeruleocephala 403 chloris 404 gerini 403 hecki 403 martinicana 339 paradisi 404 prisca 404 violacea 338–9 vittata gracilipes 186 Amethyst, Lesson’s 407 Ammodramus henslowii houstonensis 440 maritimus nigrescens 320–1 Amytornis textilis modestus 457 Anas acuta modesta 50 fuscescens 329 gattair 362

georgica niceforoi 50 gibberifrons remissa 50 latirostra 362 marecula 48–9 peregrina 362 (strepera) couesi 48 theodori 50–1 Anhinga rufa chantrei 78 Anodorhynchus ater 398 glaucus 180–1 purpurascens 399 Anomalopteryx didiformis 31 Anser grandis 362 lacteus 361 Antbird, ­White-­masked 450 Anthracothorax caeruleicauda 408 Antpitta, Antioquia B ­ rown-­banded 226 ­Brown-­banded 450 Ecuadorian 225 Northern Giant 225 Táchira 226 Antrostomus maculosus 351 Antshrike, Cocha 449 Apalis chariessa chariessa 262 Apalis, Northern W ­ hite-­winged 262 Apalopteron familiare familiare 266–7 ‘Apapane, Laysan 311–12 Aphanapteryx bonasia 108–9 Aplonis corvina 276 diluvialis 277–8 fusca fusca 274–5 fusca hulliana 275 mavornata 276–7 pacifica 433 ulietensis 278–9 Aprosmictus insignissimus 390 Aptenodytes chilensis 363 torquata 363 Apteribis brevis 69 glenos 68 Apteryx owenii iredalei 23–4 Aptornis defossor 37–8 otidiformis 37 Apus barbatus glanvillei 353 Aquila sp. 369, 370 alba 369 candidus 370 dicronyx 369 germanicus 369 tigrina 369

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Index Ara sp. 399–400, 400 autochthones 185 erythrocephala 399 erythrura 399 glaucogularis 447 gossei 335–6 guadeloupensis 183–4 martinica 399 tricolor 182–3 tricolor haitius 400 Araçari, Hauxwell’s 423 ­Maroon-­backed 423 Pöppig’s ­Curl-­crested 423 Aramides gutteralis 373 Aratinga sp. 337, 338, 400 chloroptera maugei 185 flaviventris 401 labati 336 leucophthalma nicefori 401 pertinax ssp. 336–7 pertinax griseipecta 347 Archilochus violajugulum 420 Ardea aequinoctialis 366 cristata 366 cyanocephala 365 erythrocephala 366 naevia 365 rubiginosa 366 Ardeotis arabs lynesi 87 Argus, ­Two-­banded 328–9 Argusianus bipunctatus 328–9 Asthenes leptasthenuroides 426 sclateri 426 Astrapia, ­False-­lobed 342 Athene cunicularia amaura 198 cunicularia guadeloupensis 197–8 Atlantisia elpenor 100–1 podarces 99–100 Atlapetes blancae 323 pallidiceps 457 Atthis morcomi 420 Augasma chlorophana 410–11 cyaneoberillina 410 smaragdinea 411 Auk, Great 131–2 Aythya sp. 51 innotata 443 Azurecrown, Salvin’s 414 ­Small-­billed 413 Babbler, B ­ lack-­browed 262–3 Irrawaddy Jerdon’s 263 Javan Large Wren 456 Sooty 451 Vanderbilt’s 432 Bangsia aureocincta 452 Barbet, Botero W ­ hite-­mantled 213 Northern W ­ hite-­mantled 213 Barn Owl, Bahaman 193 Barbudan 193 Hispaniolan 192–3 New Caledonian 192 Puerto Rican 192

Barnardius crommelinae 390 Baryphthengus dombeyi 422 Bee-­eater, Adanson’s 422 ­Red-­headed 422 Bellbird, Chatham Island 231 Bermuteo avivorus 81 Berrypecker, Obscure 450–1 Bird of Paradise, Bensbach’s 343, 429 Captain Blood’s 430 Currie’s Crimson 429 Duivenbode’s 430 Duivenbode’s ­Six-­wired 431 Elliot’s 342 Frau Reichenow’s 430 King William III 429 Lupton’s 431 ­Lyre-­tailed King 429 Rothschild’s 431 Rothschild’s ­Lobe-­billed 343–4 Ruys’s 344 Sharpe’s ­Lobe-­billed 344 Wilhelmina’s 430 Wonderful 430 Bittern, New Zealand Little 70–1 Biziura delautouri 55 Bleeding-­heart, Catanduanes 153 Sulu 152–3 Bobwhite, Key Quest 361 Boobook, Lord Howe 198 Norfolk Island 199 Booby, Tasman 444–5 Ua Huka 75 Bostrychia bocagei rothschildi 444 Bowdleria rufescens 260 Brachygalba lugubris phaenota 341 Brachypteracias langrandi 208–9 Brachypteryx joannae 433 Branta sp. 44–5 canadensis ssp. 361 hutchinsii asiatica 44 hylobadistes 45 Brilliant, Splendid 421 Bristlebird, Western Rufous 231–2 Stresemann’s 450 Broadbill, Guam 244 Brotogeris sp. 402 Brush-­finch, Antioquia 323 ­Pale-­headed 457 Bubo bubo jarlandi 405 maculosus 405 Bulbul, Cebu ­Mottle-­breasted 451 Sumatra ­Blue-­wattled 432 Bullfinch, St Kitts 324–5 Bulweria bifax 62 Bunting, Hudsonian Snow 440 ­Long-­legged 322–3 Louisiana 439 Maelby 439 Townsend’s 346 Unalaskan Snow 439–40 Bush-­quail, Manipur 443 Bushbird, ­Recurve-­billed 450

Bustard, Moroccan 87 Norton Sound 372 Buteo cooperi 370 Buttonquail, New Caledonia 121–2 ­Tawi-­tawi Small 122 Buzzard, Cooper’s 370 Cacicus melanurus 439 Cacique, Cassin’s 439 Cacroenis inornatus 374 Caique, ­Yellow-­breasted 402 Calcarius nortoniensis 439 Calidris cooperi 376 paramelanotos 376 Callaeas cinerea cinerea 235–6 Calliphlox orthura 407 Caloenas canacorum 147 maculata 147–8 Calyptophilus frugivorus abbotti 326 frugivorus frugivorus 325 Calyptura cristata 354–5 Calyptura, Kinglet 354–5 Camarhynchus aureus 440–1 conjunctus 440 giffordi 440 Campephilus bairdii 217–18 imperialis 214–16 principalis 216–17 Camptorhynchus labradorius 53–4 Campylopterus lazulus 408 Canastero, Lillo’s 426 Puno 426 Capellirallus karamu 111 Capito hypoleucus carrikeri 213 hypoleucus hypoleucus 213 Caprimulgus centralasicus 352 ludovicianus 406 prigoginei 351 solala 351 Caracara tellustris 85 Caracara, Bahamas 84 D’Orbigny’s 371 Guadalupe 83–4 Jamaican 85 Puerto Rican 84 Cardinal, Humbert’s 441 Carduelis aurelioi 316 triasi 316 Carib, ­Black-­and-­blue 412 Carpococcyx viridis 447 Carpodacus mexicanus mcgregori 315–16 Celeus sp. 424 immaculatus 424 obrieni 449 Centropus maurus 404 toulou assumptionis 191 Cettia diphone restricta 452 Ceyx fallax sangirensis 210 lepidus mal 352 madagascariensis dilutus 210 Chachalaca, Utila 446 Chaetocercus harterti 420

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Extinct Birds Chaetophila sp. 253 angustipluma 251–2 Chalcopsitta stavorini 384 Charadrius atricapillus 376 Charmosyna sp. 389 amabilis 348 bontii 388–9 diadema 170 samoensis 333–4 Chat, Muriel’s 434 Chaunoproctus ferreirostris 314–15 Chelychelynechen quassus 57 Chenonetta finschi 47–8 Chiffchaff, Lanzarote 255 Chloridops kona 304–5 regiskongi 306 wahi 305 Chlorophanes purpurascens 441 Chlorostilbon auratus 410 bracei 206 elegans 206 hypocyanea 409 inexpectatus 409 micans 409 subcaerulea 409 wiedi 409 Chough, Bicoloured 433 Chrysolampis gigliolii 408 Cicadabird, Maros 427 Cichlherminia lherminieri sanctaeluciae 456 Cicinnurus goodfellowi 429 lyogyrus 429 Cinclosoma punctatum anachoreta 237 Cinclus cinclus olympicus 434 Cinnyris picta 434 Circus dossenus 79 eylesi 80 Ciridops anna 313 tenax 314 Claravis plumbea 383 Clytoctantes alixii 450 Clytorhynchus sanctaecrucis 455 vitiensis powelli 243–4 Cnemiornis calcitrans 43–4 gracilis 44 Coeligena lawrencei 416 purpurea 415 traviesi 415 Coenocorypha barrierensis 125–6 chathamica 127 iredalei 126–7 miratropica 127 Colaptes cafer rufipileus 214 Colibri buckleyi 408 rubrigularis 408 Colinus virginianus insularis 361 Columba sp. 141, 379, 382 argentina 446 australis 382 brasiliensis 379 caerulea 378 cinerea 378

dominicensis 378 dorsocaerulea 378 fusca 379 janthina nitens 447 jouyi 139–40 melancholica 383 mexicana 378–9 palumbus maderensis 138–9 rosea 379 sumatrensis 379 surinamensis 383 versicolor 140–1 vitiensis ssp. 333 vitiensis godmanae 139 Comet, Orton’s 419 ­Purple-­tailed 419 Conothraupis mesoleuca 452 Conure, ‘Conure that never was’ 401 Mona Island 185 Sinú ­Brown-­throated 347 Conuropsis carolinensis carolinensis 186 carolinensis ludovicianus 186–8 Coot, Chatham Island 121 Colenso’s 375 Mascarene 120 New Zealand 121 Coquette, Bearded 409 Dusky 408 Coracina coerulescens altera 238 coerulescens deschauenseei 238 striata cebuensis 238 tenuirostris edithae 427 Cormarant, Pallas’s 76–7 Mascarene Reed 75 Spectacled 76–7 Steller’s White 367 Violet 367 ­White-­rumped 367 ­White-­tailed 367 Corvus antipodum antipodum 245 antipodum pycrofti 245–6 difficilis 428 fallax 428 impluviatus 246 modestus 429 moriorum 245 pumilus 246 unicolor 455 viriosus 247 Cotinga, Kinglet 354–5 Coturnicops noveboracensis goldmani 88 gomerae 42 novaeguineae 360 novaezelandiae novaezelandiae 41–2 Coua berthae 191 Coua (cristata) maxima 349 Coua delalandei 189–90 Coua primaeva 191 Coua, Ancient 191 Bertha’s 191 Delelande’s 189–90 Greater Crested 349 ­Snail-­eating 189–90

Coucal, Assumption Island 191 Levaillant’s Black 404 Crake, Bornean Baillon’s 347 Eastern Hawaiian 104 Iwo Jima 102–3 Keplers’ 106 Kosrae Island 101–2 Laysan 106–7 Mangaia 103 Miller’s 331–2 Molokai 105 Ralphs’ 106 Severns’ 106 St Helena 107–8 Western Hawaiian 103–4 Ziegler’s 105 Crane, Audubon’s Great White 375 Luzon Sarus 375 Marco Polo’s 375 Cranioleuca pyrrhophia rufipennis 224 Craspedophora bruyni 429 duivenbodei 341–2 mantoui 429 Creeper, Snethlage’s 426 Crinis chlorolaema 407 Crocias langbianis 451 Crocias, G ­ rey-­crowned 451 Crombec, Chapin’s 254 Crow, Banggai 455 Brüggemann’s 428 ­Deep-­billed 246 Puerto Rican 246 Robust 247 Rosenberg’s 429 Stresemann’s 428 Crypturellus (erythropus) saltuarius 17–18 Cuckoo, Bahia R ­ ufous-­vented Ground 348 St Helena Dwarf 188–9 Sumatran Ground 447 Cuckoo-­shrike, Baber’s 427 Cebu ­Bar-­bellied 238 Cebu Blackish 238 Marinduque Blackish 238 Curlew, Eskimo 128–9 Currawong, Grampian Pied 427 Cyanoliseus whitleyi 401 Cyanopsitta sp. 398 Cyanoramphus sp. 392 capitatus 393 erythronotus 392 intermedius 392 novaezelandiae erythrotis 170–1 rowleyi 392 subflavescens 171 ulietanus 171–2 verticalis 392 zealandicus 172–3 Cygnus sp. 45–6 davidi 362 immutabilis 362 Cyornis ruckii 357

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Index Daptrius gymnocephalus 371 Darter, Levant 78 Dasyornis broadbenti litoralis 231–2 Dendroica carbonata 438 montana 438 potomac 438 Dendroscansor decurivrostris 223 Diablotin 363 Diaphorapteryx hawkinsi 111–12 Dicaeum quadricolor 454 trigonostigma pallidius 454 Didunculus placopedetes 156 Didus herberti 374 Dinornis novaezealandiae 28–30 robustus 30 Diphyllodes gulielmitertii 429 Dipper, Cyprus 434 Discosura letitiae 205–6 Doctorbird 283–4 Dodo 134–6 Réunion White 377–8 Dove, Amirante Turtle 143 Callao’s ‘Columbicolin laure’ 380 Cebu Amethyst Brown 446 Chamisso’s 382 China 381 Ebon Island Fruit 156–7 Fermin’s 383 Gloger’s 382 Gmelin’s 379 Great Ground 151 Henderson Island Ground 151 Hesler’s 379 Hiva Oa ­Red-­moustached Fruit 158 Immaculate Tambourine 383 Indian Blue 378 Jamaican Blue Ground 383 Levaillant’s 380 Marquesas Cuckoo 143–4 Mauke Fruit 158–9 Mauritius Turtle 142–3 Mexican 378–9 Mexican Blue 378 Negros Fruit 156 New Caledonian Ground 151–2 Norfolk Island Ground 148–9 Nuka Hiva R ­ ed-­moustached Fruit 158 Portugal 382 Puerto Rican quail 147 Rodrigues Turtle 141–2 ­Rose-­coloured 379 San Domingo 378 Seba’s 379 Senegal Cuckoo 380–1 Seychelles Turtle 143 Society Islands Cuckoo 144 Society Islands Ground 149 Solomons W ­ hite-­bibbed Ground 152

­ potted-­necked Turtle 380 S Tanna Ground (Dove of Tanna) 150–1 ­Thick-­billed Ground 152 Tschudi’s 383 ­White-­headed Polynesian Ground 149–50 Drepanis funerea 309–11 pacifica 308–9 Dromaius ater 19–21 baudianus 21–2 diemenensis 22–3 Dryocopus crawfurdi 424 Dryolimnas augusti 99 cuvieri abbotti 98–9 Duck, Amsterdam Island 48–9 Chatham 51 Finsch’s 47–8 Gattair 362 Gaulaund 362 Gmelin’s Wandering 362 Kauai Mole 57, 58 Labrador 53–4 Lappmark 362 New Zealand Musk 55 New Zealand ­Pink-­eared 47 New Zealand ­Stiff-­tailed 55 Newfoundland (Brown) 329 ­Pink-­headed 51–3 Scarlett’s 47 Ducula sp. 381, 382 carola nigrorum 164 david 163 harrisoni 164 lakeba 162–3 Dysmorodrepanis munroi 291–2 Dysmoropelia dekarchiskos 146 Eagle, B ­ lack-­cheeked 369 Conciliating 370 Courland 369 European White 369 Fierce 370 Genoese 370 Haast’s 78–9 Heude’s 368 Louisiana White 370 Macarran 369 Malagasy Crowned 81–2 Maritime 368 Tiger 369 Earthcreeper, Peruvian S ­ cale-­ throated 224 Eclectus infectus 173–4 roratus westermani 334 Ectopistes migratorius 144–6 Elephant Bird, Coastal 27–8 Giant 24–6 Gracile 26 Greater 26–7 Hildebrandt’s 26 Robust 27

Emberiza alcoveri 322–3 ludovicia 439 maelbyensis 439 Emerald, Berlepsch’s 409 Brace’s 206 Brilliant 409 Brown 413 Cabani’s 410 Dusky 413 Iridescent 410 Natterer’s 410 Wied’s 409 Emeus crassus 33 Emu, Kangaroo Island 21–2 King Island 19–21 Tasmanian 22–3 Ensifera caerulescens 416 Eos sp. 385 bornea 384 coccinea 385 goodfellowi 385 histrio challengeri 169 histrio histrio 169 unicolor 385 variegata 385 Epimachus astrapioides 430 ellioti 342 Eremomela turneri kalindei 262 Eremomela, Western Turner’s 262 Eriocnemis berlepschi 418 dyselius 418 godini 353–4 isaacsonii 418 ridolfi 419 simplex 418 soderstromi 418 ventralis 419 Erythromachus leguati 110 Estrilda nigriloris 358 Eucephala caeruleolavata 411 chlorocephalus 411 scapulata 411 smaragdocaerulea 411 Eudyptes sp. 58, 363 Euphonia vittata 435 Euphonia, ­Black-­throated 435 Eurochelidon sirintarae 356 Eurostopodus diabolicus 448 exsul 202 Euryapteryx curtus 31 Eutrichomyias rowleyi 454–5 Eutriorchis astur 445 Euxenura galatea 365 pilla 365 Falco sp. 371 Falco albus 372 desertorum 372 duboisi 85–6 fuscus 371 kurochkini 86 leucocephalus 372 peregrinus fruitii 445

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Extinct Birds rubiginosus 371 sacer 372 stellaris 372 Falcon, Latham’s Brown 371 Rougri 372 Rusty 371 Salt’s 371 Starry 372 Volcano Islands Peregrine 445 White 372 ­White-­headed 372 Fantail, Guam Rufous ­(Fawn-­ breasted) 240–1 Lord Howe (­ Fawn-­breasted) 240 Fernbird, Chatham Islands 260 Ficedula bonthaina 453 Figbird, Ingram’s 428 Finch, Beck’s 440 Bonin Islands 314–15 ­Cone-­billed 307–8 Darwin’s Large Ground 323–4 Gifford’s 440 Greater Koa 302–3 Kiwi S ­ hovel-­billed 299–300 Lesser Koa 303–4 Makawehi 288–9 Maui Nui 289 McGregor’s House 315–16 Norton 439 Oahu Koa 304 Pololei S ­ hovel-­billed 300 ­Ridge-­billed 308 ­Scissor-­billed Koa 304 Sillem’s Mountain 345–6 Swarth’s 440–1 Firecrown, Alejandro Selkirk Island 207–8 Fiscal, São Tomé 454 Flicker, Guadalupe 214 Flowerpecker, F ­ our-­coloured 454 Paler 454 Flycatcher, Grenada Euler’s 226–7 Guam 244 Lompobatang 453 San Cristóbal Vermilion 227 ­Small-­headed 436 Tonkean H ­ enna-­tailed Jungle 434 ­Yellow-­tailed 438–9 Fody, Daubenton’s 434–5 Réunion 287 Forbus sp. 402 Foudia bruante 434–5 delloni 287 Francolin, Amik Gölü Black 41 Francolinus francolinus billypayni 41 Fregilipus varius 271–3 Fulica chathamensis 120–1 newtoni 120 novazealandiae 375 prisca 121 Gadwall, Coues’s 48 Washington Island 48

Gallicolumba eimeensis 383 erythroptera ssp. 149 erythroptera albicollis 149–50 ferruginea 150–1 jobiensis chalconata 152 leonpascoi 151 longitarsus 151–2 luzonica rubiventris 153 menagei 152–3 norfolciensis 148–9 nui 151 salamonis 152 Gallinula hodyeni 117 nesiotis 117–18 Gallinule, Hova 113 Huahine 116 Lord Howe 114–16 Makira 119–20 Marquesas Islands 116–17 New Caledonia 116 New Zealand Flightless 117 Réunion Blue 113–14 Samoan 118–19 Tristan Island 117–18 Gallirallus sp. 96 epulare 94 ernstmayri 94 forbesi 373 gracilitibia 94 huiatua 95 minor 374 modestus 91–2 pacificus 92–3 pendiculentus 93–4 philippensis dieffenbachii 90–1 philippensis macquariensis 90 pisonii 93 ripleyi 95 roletti 94 sharpei 373 steadmani 96 storrsolsoni 95 temptatus 93 vekamatolu 95–6 wakensis 89–90 Gaper, Maui Nui 301 ­Sickle-­billed 300 ­Straight-­billed 300 Garefowl, Black 377 Collared 377 Geoffroyus aureus 389 dorsalis 389 simplex 389 Geopelia sinica 381 Geospiza magnirostris magnirostris 323–4 Geothlypis cucullata 438 rostrata rostrata 318–19 Geotrygon larva 147 Gerygone insularis 232–3 Gerygone, Lord Howe 232–3 Giant, Leguat’s 373 Gisella lathami 406

Glaucis philippinae 421 Goose, Bering Cackling 44 ­Cream-­coloured 361 Giant Hawaiian 44–5 Great 362 Greater Hawaiian 45 Kauai ­Turtle-­jawed 57 North Island 44 Snow’s Canada 361 South Island 43–4 Goshawk, Gracile 80 Powerful 80 Goura hottentotta 380 Grackle, S ­ lender-­billed 319–20 Grallaria chthonia 226 gigantea hylodroma 225 gigantea lehmanni 225 milleri gilesi 226 milleri milleri 450 Grallistrix auceps 195–6 erdmani 196 geleches 197 orion 196 Grasswren, Eastern ­Thick-­billed 457 Grebe, Alaotra 65 Atitlán 65–6 Colombian 66–7 Gadow’s 365 Tota 66–7 Greenfinch, ­Slender-­billed 316 Trías 316 Grosbeak, Fantail 435 Grey 435 King Kong 306 Kona 304–5 Mauka 306 São Tomé 457 Wahi 305 ­Yellow-­bellied 435 Ground-­roller, Ampoza 208–9 Grouse, New Mexico S ­ harp-­tailed 40–1 Grus sp. 375 antigone luzonica 375 polii 375 Guineafowl, Moroccan 38 Gull, Huahine (Society Islands) 130 Gymnocrex plumbeiventris intactus 330 Gymnogyps sp. 367 Haematopus meadewaldoi 122–3 Halcyon sp. 340 gambieri gambieri 209 miyakoensis 340–1 Haliaeetus maritimus 368 niger 368 Harpagornis moorei 78–9 Harrier, Eyles’s 80 Hawaiian 79 Hawk, Bermuda 81 Speckled Partridge 372 Heliangelus claudia 416 dubius 416

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Index fulvicrissa 418 henrici 416 luminosus 417 prosantis 417 rothschildi 417 simoni 417 speciosus 417 squamigularis 417 violicollis 416 zusii 208 Heliodoxa splendens 421 Hemignathus (ellisianus) ellisianus 295–6 (ellisianus) lanaiensis 294 (ellisianus) obscura 294–5 (ellisianus) stejnegeri 293–4 lucidus ssp. 298–9 lucidus affinis 297–8 lucidus hanapepe 296 lucidus lucidus 297 sagittirostris 292–3 upupirostris 296 vorpalis 299 Hemiphaga sp. 165–6 brunnea 381 novaeseelandiae ssp. 165 novaeseelandiae spadicea 164–5 Hen, Heath 38–40 Herbert’s 374 Hermit, Gould’s 407 Heine’s 408 Porculla 407 Schlüter’s 407 Sooty 407 Heron, Ascension Night 73–4 Bermuda Night 74 ­Blue-­headed 365 Bonin Night 72–3 Catesby’s Great Crested 366 Mauritius Night 71 Miller’s 365 Niue Night 73 ­Red-­billed 366 ­Red-­headed 366 Réunion Night 72 Rodrigues Night 72 ­Rusty-­crowned 366 Heteralocha acutirostris 233–5 Heteroglaux blewitti 448 Heteromirafra archeri 355 Heteroptilornis mantoui 429 Hihi 236 Himatione sanguinea freethii 311–12 Hirundo perdita 355–6 Hoa 309–11 Honeycreeper, Laysan 311–12 Purple 441 Sassi’s 436 Honeyguide, Friedmann’s 424 Hookbill, Lanai 291–2 Hoopoe, St Helena Giant 211 Hornbill, Lifou 212 Ticao Tarictic 212

Hovacrex roberti 113 Huia 233–5 Hummingbird, Alfaro’s 206–7 Bangs’s 415 ­Blue-­spotted 413 Burton’s 421 de Oca’s 414 Elegant 206 Elliot’s 414, 415 Fifth of Marcgrave 421 Florence van Rossem’s 414 Floresi’s 420 Lerdo’s 414 Molina’s 421 Morcom’s 420 Neglected 414 Nehrkorn’s 412 Ninth of Marcgrave 422 Penard’s 421 President Nunez’s 415 Ridolfi’s 419 Seventh of Marcgrave 422 Underwood’s 420 ­Violet-­throated 420 Wiener’s 415 Huppe 271–3 Hylacola tyrannulus 427 Hylocharis pyropygia 412 Hypositta perdita 358 Hypothymis coelestis rabori 243 Hypsipetes siquijorensis monticola 451 Ibis, Maui Flightless 69 Molokai Flightless 68 Príncipe Dwarf Olive 444 Réunion 67–8 Icterus leucopteryx bairdi 319 xantholemus 439 Inca, Purple 415 Indicator propinquus 424 Iodopleura pipra leucopygia 456 Ixobrychus minutus novaezelandiae 70–1 Jabiru, Chilean 365 Jacamar, Klages’s 341 Janthothorax bensbachi 343 mirabilis 430 Kagu, Lowland 87–8 Kaka, Chatham Islands 167 ­Long-­billed 383–4 Lord Howe Island 333 Masséna’s 384 Norfolk Island 166–7 Small 384 Kakariki, Intermediate 392 ­Large-­billed 393 Mysterious 392 ­Red-­hooded 393 ­Red-­topped 392 Rowley’s 392 Kakawahie 290

Kama’o 282–3 Kestrel, Cuban 86 Réunion 85–6 Kingbird,Heine’s 425 Kingfisher, Javan ­Blue-­banded 210 Latham’s ­Black-­capped 340 Malaita Variable 352 Mangareva 209 Miyako (Ryukyu) 340–1 Sakaraha Pygmy 210 Sangihe Dwarf 210 Kinglet, Cuvier’s 432 Guadalupe ­Ruby-­crowned 267 Kioea 251–2 Lesson’s 431 ­Narrow-­billed 253 Kite, Cape Verde 371 Kiwi, North Island Little Spotted 23–4 Kokako, South Island 235–6 Kuinia mundyi 373 Lalage leucopyga leucopyga 238–9 melanothorax 427 Lampornis viridipallens connectens 421 Lamprothorax wilhelminae 430 Lanius natka 427 Lanius newtoni 454 Lapwing, Javan 124 Madagascar 125 Lark, Archer’s 355 Larus utunui 130 Laterallus jamaicensis jamaicensis 88–9 Lathrotriccus euleri flaviventris 226–7 Leguatia gigantea 373 Leiothrix (lutea) astleyi 345 Leiothrix, Astley’s 345 Lepidothrix vilasboasi 449 Lesbia glyceria 419 ortoni 419 Leuchochloris malvina 412 Leucippus nigrirostris 413 Leucopeza semperi 318 Leucosticte sillemi 345–6 Lewinia pectoralis clelandii 97–8 pectoralis exsul 98 Linnet, Brewster’s 436 Loborhamphus nobilis 343–4 ptilorhis 344 Loophortyx leucoprosopon 361 Lophopsittacus mauritianus 180–1 Lophornis insignibarbis 409 melaniae 408 Loriculus cervicalis 397 philippensis chrysonotus 168 philippensis siquijorensis 168–9 salvadorii 397 Lorikeet, Aru Islands Rainbow 386 Conquered 170 Kotzebue’s 333–4 Latham’s 389 New Caledonian 170 ­Red-­throated 348

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Extinct Birds Sinoto’s 169–70 Wilson’s 388 Lorius sp. 386, 386–7, 387 amabilis 388 brasiliensis 388 cyanolenus 388 incarnatus 387 mexicanus 387 rex 387 tibialis 388 Lory, Albin’s 388 ­Blue-­tailed 385 Bontius’s 388–9 Challenger R ­ ed-­and-­blue 169 Goodfellow’s 385 Gray’s 386 ­Green-­thighed ­Yellow-­collared 386 Jamrach’s 388 ­Jay-­winged 387 Layard’s Mysterious 385–6 Levaillant’s 385 Little R ­ ed-­winged 387 Mamberiok 384 Meyen’s 388 ­Red-­and-­gold 387 ­Red-­and-­green 384 Sangihe R ­ ed-­and-­blue 169 Stavorini’s 384 Stresemann’s 388 Unicoloured 385 Variegated 385 Verreaux’s 386 Walpole’s 386–7 ­White-­collared 386 Lovebird, Leona 397 Loxia butyracea 435 grisea 435 virginica 435 Loxigilla (portoricensis) grandis 324–5 Loxioides kikuchi 306 Loxops ochraceus 301–2 wolstenholmei 301 Macaw, Albin’s 400 Black 398 ­Blue-­fronted 447 Cuban 182–3 Dominican 402 Glaucous 181–2 Guadeloupe 183–4 Hispaniolan 400 Humboldt’s 398 Jamaican B ­ lue-­and-­yellow 399–400 Jamaican ­Yellow-­headed 335–6 Marcgrave’s 400 ­Orange-­bellied 399 Puerto Rican 185 ­Red-­headed 399 Satin 399 Seaforth’s 400 Violet 399

Macropygia arevarevauupa 144 heana 143–4 macerona 380–1 Malacochichla vanderbilti 432 Malacocincla perspicillata 262–3 Malacorhynchus scarletti 47 Mamijiro Kuina 102–3 Mamo 308–9 Black 309–11 Manakin, ­Golden-­crowned 449 Mango, ­Steel-­blue-­tailed 408 Margarops fusca atlantica 270 Martin, ­White-­eyed River 356 Mascarenotus grucheti 194–5 murivorus 195 sauzieri 194 Mascarinus mascarinus 177–8 Mata 260 Megadyptes waitaha 59 Megalapteryx didinus 34 Megalurulus grosvenori 260–1 rufa cluniei 261 Megapode, Andaman 360 Anderson’s 327 Burnaby’s 360 Consumed 34–5 Kermadec 327–8 Lord Howe Island 360 Noble 35 ­Pile-­builder 35 Megapodius sp. 327–8, 360 alimentum 34–5 amissus 35 andamanensis 360 andersoni 327 burnabyi 360 molistructor 35 Megascops nudipes newtoni 350 Megavitiornis altirostris 35 Meguro 266–7 Meiglyptes tristis tristis 449 Melamprosops phaesoma 312–13 Melanerpes hargitti 425 superciliaris bahamensis 425 Melanocharis arfakiana 450–1 Melanodryas cucullata melvillensis 247 Mellisuga violajugulum 420 Melospiza melodia amaka 440 melodia graminea 321–2 Merganser, Auckland Island 54–5 Mergus australis 54–5 borealis 362 Merops adansonii 422 erythrocephalus 422 Merulaxis stresemanni 450 Metallura griseocyanea 419 purpureicauda 419 Metaltail, Boucard’s 419 Micrathene whitneyi graysoni 197 Microgoura meeki 154–6 Millerbird, Laysan 259 Milvus milvus fasciicauda 371

Moa-­nalo, Maui Nui 56 Oahu Nui 56 Stumbling 56 Moa, ­Broad-­billed 31 Crested 32 Eastern 33 ­Heavy-­footed 32–3 Little Bush 31 North Island Giant 28–30 South Island Giant 30 ­Stout-­legged 31–2 Upland 34 Moho sp. 251 apicalis 248–9 bishopi 250–1 braccatus 248 nobilis 249–50 Monarch, Eiao 243 Hiva Oa 241–2 Maupiti 241 Negros Celestial 243 Nuku Hiva 242 Tongatabu 428 Ua Pou 242 Moorhen, Tristan Island 117–18 Morepork, Lord Howe 198 ‘Motacilla awatcha’ 437 ‘calidris’ 437 ‘camtschatkensis’ 437–8 ‘fulva’ 437 ‘ludoviciana’ 436 ‘semitorquata’ 437 Motmot, Dombey’s 422 Mountain Gem, Salvador 421 Moupinia altirostris altirostris 263 Mullerornis agilis 27–8 rudis 27 Myadestes elisabeth retrusus 280–1 lanaiensis lanaiensis 281–2 lanaiensis rutha 282 myadestinus 282–3 woahensis 281 Myiagra freycineti 244 Myiopsitta buccalis 401–2 Nannococcyx psix 188–9 Natunaornis gigoura 162 Necropsar rodericanus 273–4 Necropsittacus francicus 398 rodericanus 178–9 Nemosia rourei 451–2 Neochmia ruficauda ruficauda 288 Neolesbia nehrkorni 412 Neomorphus geoffroyi maximiliani 348 Neoparadisea 344 Neophron leucocephalus 368 Neospiza concolor 457 Nesillas aldabranus 255–6 Nesoclopeus poeciloptera 89 Nesoenas sp. 142–3 duboisi 141 picturata aldabrana 143

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Index picturata rostrata 143 rodericana 141–2 Nesofregetta fuliginosa moestissima 365 Nesotrochis debooyi 110–11 picapicensis 111 steganinos 111 Nestor sp. 167, 333, 384 esslingi 384 norfolcensis 383–4 productus 166–7 Nightingale, American 437 Nightjar, Cayenne 351 Itombwe (Prigogine’s) 351 Nechisar 351 New Caledonian 202 Stephenson Clarke’s 406 Sulawesi (Satanic) 448 Vaurie’s 352 Ninox novaeseelandiae albaria 198 undulata 199 Nonnula galbuloides 423 Nothoprocta kalinowskii 359 Notiomystis cincta cincta 236 Nukupu’u, Giant ­Scimitar-­billed 299 Hawaii 298–9 Kaua’i 296 Maui 297–8 Oahu 297 Numenius borealis 128–9 Numida melegris sabyi 38 Nyctanassa carcinocatactes 74 Nyctea wapacuthu 405 Nycticorax caledonicus crassirostris 72–3 duboisi 72 kalavikai 73 mauritianus 71 megacephalus 72 olsoni 73–4 ‘O’o, Hawaii 249–50 Kaua’i 248 Maui 251 Molokai (Bishop’s) 250–1 Oahu 248–9 ‘O’u 307 Oceanites maorianus 444 Oceanodroma bicolor 364 macrodactyla 64–5 Oiseaux Bleu 113 Oloma’o, Lanai 281–2 Molokai 282 Ophrysia superciliosa 43 Oriole, Cebu 239 Grand Cayman 319 Villavicencio’s 439 ­White-­headed 428 Oriolus steerii assimilis 239 violaceus 428 Orphanopsar leguati 433 Ortalis vetula deschauenseei 446

Orthiospiza howarthi 306 Ostrich, Levaillant’s Dwarf 359 Syrian 18–19 Otis sp. 372 Otus alfredi 448 capnodes 447–8 manadensis siaoensis 350 minimus 406 Owl, Antigua Burrowing 198 Bartram’s 406 Buru Masked 349 Carriker’s 406 ­Cave-­nesting Masked 404 China 405 Commerson’s 404 Comoro Scops 447–8 Flores Scops 448 Kauai Stilt 195–6 Kaup’s 406 Krusenstern’s Japanese 406 La Touche’s Eagle 405 ­Marie-­Galante Burrowing 197–8 Maui Stilt 196 Mauritian Lizard 194 Molokai Stilt 197 Natuna Bay 350 Newton’s 405 North Island Laughing 200–1 Oahu Stilt 196 Peleng Masked 349 Réunion Lizard 194–5 Rodrigues Lizard 195 Rufous Owl of Latham 406 Sauzier’s 194 Siau Scops 350 Socorro Elf 197 South Island Laughing 199–200 Taliabu Masked 447 Vieillot’s Spotted 405 Vieillot’s Wood 405 Virgin Islands Screech 350 Wapacuthu 405 Owlet-­nightjar, New Zealand 203 New Caledonia 352 Owlet, Forest 448 Oxyura vantetsi 55 Oystercatcher, Canary Islands ­(Meade-­Waldo’s) 122–3 Pachyana chathamica 51 Pachyornis australis 32 elephantopus 32–3 geranoides 31–2 Pachyplichas jagmi 222 yaldwyni 222 Palila, Pila’s 306 Palmcreeper, Hawaii 313 Kaua’i 314 Panurus biarmicus kosswigi 432–3 Papasula abbottii costelloi 75 Paradisaea apoda luptoni 431 bloodi 430 duivenbodei 430

maria 430 mixta 431 Paradise-­flycatcher, Cerulean 454–5 Parakeet, Azara’s 401 Barbados 338 ­Blue-­headed 398 ­Crimson-­banded 396 ­Crimson-­crowned 390 Eastern Carolina 186 Edwards’s ­Long-­tailed 397 Guadeloupe 336 Guaicaramo W ­ hite-­eyed 401 Intermediate 398 Jonquil 398 Lord Howe 171 Macquarie Island 170–1 Marcgrave’s 400 Martinique 336–7 Newton’s 175–6 Ober’s 337 Réunion R ­ ing-­necked 176–7 Rodrigues R ­ ing-­necked 175–6 Seychelles 174–5 Sinú Painted 185 Western (Louisiana) Carolina 186–8 Whitley’s 401 ­Yellow-­sided 401 Pareoreomyza perkinsi 436 Pareudiastes pacifica 118–19 silvestris 119–20 Paroaria humberti 441 Paroreomyza flammea 290 maculata 291 montana montana 289–90 Parotia duivenbodei 431 Parrakeet, B ­ lack-­necked 393 ­Blue-­faced 395 Cayenne 395 Crested ­Red-­and-­green 394 Edwards’s 402 Japanese 394 Lineated 393 Nooffy 395 Pale 393 ­Short-­tailed 395 Sulphur 391 Parrot, Alexander’s 397 Amboina 390 Andai 389 Bastard King 390 Bechstein’s 401–2 Bechstein’s Horned 392 ­Black-­fronted 172–3 ­Blue-­backed 402 ­Broad-­billed 180–1 Cebu Hanging 168 Chlorotic 391 Cobalt 402 Crested Mexican 387 ­Crimson-­vented 391 David Livingstone’s 396 Dubious 396

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Extinct Birds Golden 389 ­Green-­crested 396 Heck’s 403 Hecquard’s 396 Heuglin’s 396 Kuhl’s 392 Lady Tavistock’s 390 Marcgrave’s 394 Mascarene 177–8 Mauritian 398 Moseley’s 333 Obscure 335 Pacific Islands Eclectus 173–4 Paradise 173 Peregrine 393 Raven 180–1 ­Red-­naped 397 ­Red-­winged 394 Réunion 179–80 Rodrigues 178–9 ­Rose-­fronted 334–5 ­Ruby-­capped 396 ­Rufous-­tailed 389 Salvadori’s Hanging 397 Scopoli’s 389 Siquijor Hanging 168–9 Society 171–2 Society Islands 172 Tanna 334 Taraba 395 Thirioux’s Grey 174 Westerman’s Eclectus 334 Partridge, Italian 361 Parula sp. 438–9 Parus ignotus 431 Paryphephorus duivenbodei 341–2 ‘Passer virginianus’ 437 Pauraque, Cuban 202–3 Jamaican 202 Pelecanus (conspicillatus) novaezealandiae 366 sharpei 366 Pelican, New Zealand 366 Sharpe’s 366 Penelopides panini ticaensis 212 Penguin, Chatham Islands 58 Commerson’s 363 Hunter Island 57–8 Larsen’s 363 Molina’s 363 Waitaha 59 Perdicula manipurensis 443 Perdix perdix italica 361 Periparus ater phaeonotus 431–2 Petrel, Agile 364 Beck’s 444 Fiji 444 Greater St Helena 62 Guadalupe 64–5 Hindwood’s 363 Jamaican 59–60 Kuril 364 Lesser Hawaiian 61

Lesser St Helena 62 Mount Pitt 60–1 Mourning 364 Sailing 364 Samoan 365 Sordid 364 ­Two-­coloured 364 Pezophaps solitaria 137–8 Phaethornis anthophilus fuliginosus 407 apheles 408 fumosus 407 porcullae 407 zonura 407 Phalacrocorax (africanus) nanus 75 albidus 367 leuconotus 367 leucurus 367 novaezelandiae 367 perspicillatus 76–7 violaceus 367 Phapitreron amethystina frontalis 446 Phasianus sp. 361 superbus 361 Pheasant, Florida 361 Superb 361 Phodilus badius arixuthus 350 Phoeniculus cavei 422 Phylloscopus canariensis exsul 255 Piculet, Natterer’s 424 Picumnus fuscus 424 Picus flavipes 425 Pigeon, Bonin Islands 140–1 Bonin Islands Wood 447 Bougainville’s 382 Brown 381 Burnes’s 380 Choiseul Ground 154–6 Earl’s 382 Henderson Island Imperial 164 ­Hook-­billed 382 Hottentot 380 Indian 381 Kanaka 147 King’s White 381 Lakeba Imperial 162–3 Lefèbvre’s 381 Liverpool 147–8 Lord Howe 139 Lord Howe Island 165–6 Madeira Wood 138–9 Mauritius Blue (Pigeon Hollandais) 159–60 Mauritius Wood 141 Negros Spotted Imperial 164 Norfolk Island 164–5 Passenger 144–6 Providence Blue 161–2 ­Purple-­breasted 383 Raoul Island 165 Réunion Blue 161 Réunion Pink 141 Rodrigues Blue 161 Ryukyu 139–40

Siamese 379 Silvery 446 Sonnerat’s 378 Spotted Green 147–8 St Helena 146 Tongan T ­ ooth-­billed 156 ‘Uvea Imperial 163 Vava’u 333 Viti Levu Giant 162 Pinguinus sp. 377 impennis 131–3 Pintail, Nicéforo’s 50 Tristram’s 50 Pionites guineensis 402 Pionus cobaltinus 402 Piopio, North Island 230 South Island 228–9 Stephens Island 229–30 Pipilo erythropthalmus consobrinus 321 Pithys castaneus 450 Pitta anerythra nigrifrons 223–4 anerythra pallida 223 Pitta, Bougainville B ­ lack-­faced 223 Choiseul B ­ lack-­faced 223–4 Platycercus sp. 391 cornutus 392 erythropeplus 390 ignitus 391 leverianus 391 masterianus 391 paradiseus 391 Plectrophenax aunalaschkensis 439–40 hudsonius 440 Ploceus ruweti 457 Plover, B ­ lack-­crowned 376 Ross’s Shore 375 Po’ouli 312–13 Pochard, Madagascar 443 Réunion 51 Podiceps andinus 66–7 gadowi 365 Podilymbus gigas 65–6 Pogoniulus makawai 354 ‘Poicephalus’ sp. 396 Poicephalus kintampoensis 397 rubricapillus 396 Polyborus creightoni 84 latebrosus 84 lutosus 83–4 Polystictus pectoralis bogotensis 227 Pomarea fluxa 243 mendozae 241–2 mira 242 nukuhivae 242 pomarea 241 tabuensis 428 Poorwill, Cuban 202–3 Jamaican 202 Popelairia letitiae 205–6 Porphyrio albus 114–16 caerulescens 113–14 kukwiedei 116 mantelli 114

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Index mcnabi 116 paepae 116–17 Porzana astrictocarpus 107–8 cinerea brevipes 102–3 circoleps 375 keplerorum 106 menehune 105 monasa 101–2 nigra 331–2 palmeri 106–7 pusilla mira 347 ralphorum 106 rua 103 sandwichensis millsi 104 sandwichensis sandwichensis 103–4 severnsi 106 (tabuensis) tenebrosa 331 ziegleri 105 Procellaria curilica 364 Prosobonia cancellata 332 ellisi 332 leucoptera 129–30 Prosopeia sp. 334 Psammoenas burnesii 380 Psephotus pulcherrimus 173 Pseudastrapia lobata 342 Pseudobulweria becki 444 macgillivrayi 444 rupinarum 62 Pseudoscops macrurus 406 Psittachus multicolor 386 Psittacula sp. 334–5 bensoni 174 eques 176–7 exsul 175–6 indica 398 intermedia 398 narcissus 398 rufirostris 397 wardi 174–5 Psittacus erythrochlorus 394 erythroleucus 394 obscurus 335 paraguanus 394 sosove 395 vaillanti 394 ‘Psittacus’ sp. 395, 396 dubius 396 japonicus 394 lineatus 393 magnirostris 393 nigricollis 393 pallidus 393 peregrinus 393 taraba 395 Psittirostra psittacea 307 Psittrichas sp. 333 Ptaiochen pau 56 Pterocles exustus floweri 133 Pterodroma agilis 364 atrata 364 caribbaea 59–60 diabolica 363

hindwoodi 363 jugabilis 61 (solandri?) 60–1 sordida 364 velificans 364 Pteroglossus didymus 423 olallae 423 picatus 423 poeppigii 423 Ptilinopus arcanus 156 mercierii mercierii 158 mercierii tristrami 158 porphyraceus marshallianus 156–7 rarotongensis ‘byronensis’ 158–9 Ptochoptera iolaima 410 Puffbird, Bertoni’s 423 Puffinus olsoni 63–4 spelaeus 63 Puffleg, ­Amethyst-­vented 419 Berlepsch’s 418 Black 418 Isaacson’s 418 Simple 418 Söderström’s 418 ­Turquoise-­throated 353–4 Purpletuft, Salvin’s 456 Pyconotus nieuwenhuisii inexspectatus 432 Pyrocephalus rubinus dubius 227 Pyrrhura beryllina 401 hypoxantha 401 picta subandina 185 Quail, Canary Islands 42 Himalayan Mountain 43 New Guinea 360 New Zealand 41–2 Reichenow’s 361 Quiscalus palustris 319–20 Quail-­thrush, Mount Lofty Spotted 237 Rail, Aguiguan 93 Ascension 100–1 Assumption W ­ hite-­throated 98–9 Austral 445 ­Blue-­bellied 374 Bruner’s 374 Chatham Islands 91–2 Cleland’s 97–8 Dark 331 De Booy’s 110–11 Dieffenbach’s 90–1 Eua 95–6 Fiji B ­ ar-­winged 89 Flores 98 Gilbert 374 Goldman’s Yellow 88 Grey 110 Haiti 111 Hawkins’s 111–12 Huahine 95 Jamaican Black 88–9

Jamaican Wood 101 Laysan 106–7 Leguat’s 110 Macquarie Island 90 Mangaia 95 Mauritius Red 108–9 Miller’s 331–2 Mysterious 375 New Caledonian Wood 96–7 New Ireland 94 Niue 95 Norfolk Island 96 Notable 373 Nuka Hiva 94 Peter Mundy’s 373 Picapica 111 ­Red-­throated Wood 373 Réunion 99 Ripley’s 95 Rodrigues 110 Rota 93 Sakalava 445 Sharpe’s 373 ­Short-­winged 374 Solomon Islands B ­ are-­eyed 330 St Helena 99–100 Tabuai 96 Tahiti R ­ ed-­billed 92–3 Tahuata 94 Tinian 93–4 Ua Huka 94 Viti Levu 112 Wake Island 89–90 Rallus antarcticus 445 ecaudatus 374 ecaudotus 374 Ramphastos piperivorus 424 piscivorus 423 Raphus cucullatus 134–6 solitarius 377 Raven, Chatham Island 245 North Island 245 South Island 245–6 Regulus calendula obscurus 267 cuvieri 432 Rhagorhina auricularis 382 Rhea nana 360 Rhea, Dwarf 360 Rhinopomastus cavei 422 Rhipidornis gulielmitertii 429 Rhipidura cervina 240 rufifrons uraniae 240–1 Rhodacanthis flaviceps 303–4 forfex 304 litotes 304 palmeri 302–3 Rhodonessa caryophyllacea 51–3 Rhynochetus orarius 87–8 Riflebird, Duivenbode’s 341–2 Mantou’s 429 Robin, Tiwi Islands Hooded 247 Rosella, Fiery 391 Masters’s 391

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Extinct Birds Mrs Bligh’s 390 Mysterious 390 Paradise 391 Sabrewing, Laugier’s 408 Saggitarius philippensis 368 Salpinctes obsoletus exsul 268 Sandgrouse, Fayum 133 Sandpiper, Cooper’s 376 Cox’s 376 Ellis’s 332 Kiritimati 332 Sakhalin 376 Uniform 376 Waved 376 ­White-­winged 129–30 Sapphire, ­Blue-­breasted 409 Elliot’s 409 ­Flame-­rumped 412 Sarcorhamphus sacer 329–30 Sassius simplex 436 Saucerottia alfaroana 206–7 Saxicola dacotiae murielae 434 Sceloglaux albifacies albifacies 199–200 albifacies rufifacies 200–1 Scolopax anthonyi 127–8 Scops commersoni 404 Scrubfowl, New Caledonian Giant 36 Viti Levu 35 Secretary Bird, Sonnerat’s 368 Seedeater, Hooded 358 Selasphorus floresii 420 underwoodi 420 Sephanioides burtoni 421 galeritus 421 Sephanoides fernandensis leyboldi 207–8 Sericornis nigroviridis 426–7 Sericornis, B ­ lack-­and-­green 426–7 Serin, Edwards’s 436 Linnaeus’s 435 Serinus butyracea 436 Serpent-­eagle, Madagascar 445 Shag, Forbes’s 367 Shearwater, Lava 63–4 Scarlett’s 63 Sheldgoose, Mauritius 46 Réunion 46–7 Shelduck, Crested 47 Short-­tail, São Tomé 452 Shortwing, La Touche 433 Shrike, Natka 427 Shrike-­starling, Pacific 433 Shrikebill, Manu’a 243–4 Nendö 455 Sicklebill, Astrapian 430 Silver-­eye, Lord Howe 456 Robust 265 Siphonorhis americana 202 daiquiri 202–3 Smaragdichysis iridescens 410 Snipe, Forbes’s 127 Little Barrier Island 125–6

North Island 125–6 South Island 126–7 Viti Levu 127 Snipe-­rail, New Zealand 111 Solitaire, Green 292–3 Isle of Pines 280–1 Réunion 67–8 Rodrigues 137–8 Sparrow, Amak Island Song 440 Dusky Seaside 320–1 Santa Barbara Song 321–2 Texas Henslow’s 440 Todos Santos ­Rufous-­sided 322 Sparrowhawk, Car Nicobar 80–1 Mauritius 370–1 Spelaeornis badeigularis 451 Spermophila laticauda 435 Sphecotheres stalkeri 428 Spinetail, Northern S ­ tripe-­crowned 224 Philippi’s 426 Spiza townsendi 346 Sporophila melanops 358 Stachyris herberti 451 Star Finch, Eastern 288 Star-­frontlet, Lawrence’s 416 ­Lilac-­fronted 415 Starling, Hoopoe 271–3 Huahine 277–8 Kosrae 276 Leguat’s 433 Lord Howe 275 Mauke (Mysterious) 276–7 Norfolk Island 274–5 Réunion Crested 271–3 Rodrigues 273–4 Stephanoaetus mahery 81–2 Sterna bernsteini 446 camtschatica 377 naevia 377 Stitchbird, North Island 236 Stork, Galatea 365 Storm-­petrel, New Zealand 444 Streamertail, Taylor’s 412 Strepera graculina ashbyi 427 Streptopelia sp. 380 lusitanicus 382 Strigiceps leucopogon 431 Strix sp. 406 fasciata 405 newtoni 405 orientalis 405 peregrinator 406 Struthio bidactylus 359 (camelus) syriacus 18–19 Stubtail, Babar 254 Sula dactylatra tasmani 444–5 Sun-­angel, Balen’s 417 Bogotá 208 Boucard’s 417 Claudia’s 416 Glistening 417 ­Green-­throated 417

Hartert’s 416 Henry’s 416 ­Olive-­throated 417 Rothschild’s 417 Simon’s 418 Violet 416 Sunbird, Painted 434 Swallow, Jamaican Golden 253 Red Sea 355–6 Swan, Chatham Islands Black 45–6 David’s 362 Polish 362 Swift, Sierra Leone Black 353 Swiftlet, Luzon Whitehead’s 353 Mangaia 204 Vanuata W ­ hite-­rumped 205 Swordbill, Lowe’s 416 Sylvia deserticola ticehursti 432 melanocephala norissae 432 Sylvietta leucophrys chapini 254 Sylviornis neocaledoniae 36 Sylviorthorhynchus fasciolatus 426 Tachuri, Bogotá Bearded 227 Tachybaptus rufolavatus 65 Tachycineta euchrysea euchrysea 253 Tadorna cristata 47 Takahe, North Island 114 Talpanas lippa 57, 58 Tanager, Arnault’s 441 ­Cherry-­throated 451–2 ­Cone-­billed 452 ­Gold-­ringed 452 Gonâve Eastern Chat 326 Gould’s 441 Samaná Eastern Chat 325 Tangara arnaulti 441 gouldi 441 Tanygnathus sp. 390 Tanygnathus heterurus 389 Tasidyptes hunteri 57–8 Teal, Mascarene 50–1 Rennell Island 50 Telespiza persecutrix 288–9 ypsilon 289 Tern, Chinese Crested 446 ­Cloven-­footed 377 Kamchatkan 377 Testudophaga bicolor 433 Thalurania belli 339–40 lerchi 410 ridgwayi 448 Thambetochen chauliodous 56 xanion 56 Thamnophilus praecox 449 Thicketbird, Bismarck 260–1 ­Long-­legged 453 Vanua Levu L ­ ong-­legged 261 Thinornis rossii 375 Thornbill, King Island Brown 455 ­Purple-­tailed 419 Coppery 205–6

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Index Thrasher, Barbados S ­ caly-­breasted 270 Cozumel 270–1 Threskiornis solitarius 67–8 Thrush, Bay 278–9 Cauca B ­ lack-­hooded 286 Choiseul R ­ usset-­tailed 280 Grand Cayman 285–6 Kibale B ­ lack-­eared 280 Kittlitz’s 279 Lanai 281–2 Large Kaua’i 282–3 Lifou 284–5 Lord Howe 283–4 Maré 285 Molokai 282 Norfolk Island 284 Oahu 281 Sandwich 433 St Lucia Forest 456 Yakushima 434 Thryomanes bewickii brevicauda 269 bewickii leucophrys 268 Tinamou, Huila Black 17 Kalinowski’s 359 Magdalena 17–18 Pernambuco Solitary 359 Tinamus osgoodi hershkovitski 17 solitarius pernambucensis 359 Tinkerbird, W ­ hite-­chested 354 Tit, Amik Gölü Bearded 432–3 Zagros Coal 431–2 Titmouse, Strömian 431 Tityra leucura 455–6 Tityra, ­White-­tailed 455–6 Toatoa 230 Todiramphus miyakoensis 340–1 Todirostrum hypospodium 425 Tody-­tyrant, Berlepsch’s 425 Topaz, Elliot’s 407 Giglioli’s 408 Toucan, Albin’s 423 Edwards’s 423 Mysterious 424 Towhee, Guadalupe 321 Toxostoma guttatum 270–1 Treron asiatica 381 curvirostra 382 phasianella 381 Trichocichla rufa rufa 453 Trichoglossus sp. 385–6 coccineifrons 386 haematodus brooki 386 multicolor 386 verreauxius 386 Tricholimnas conditicius 374 lafresnayanus 96–7 Triller, Norfolk Island 238–9 Tringa meleagris 376 undata 376 uniformis 376 Trochilis venustissimus 412

Trochilus taylori 412 ‘Trochilus’ guainumbiquinta 421 nona 422 septimus 422 Troglodytes aedon martinicensis 269–70 troglodytes orii 345 Turdinus macrodactyla lepidopleura 456 Turdus celaenops yakushimensis 433 olivater caucae 286 poliocephalus mareensis 285 poliocephalus poliocephalus 284 poliocephalus pritzbueri 284–5 poliocephalus vinitinctus 283–4 ravidus 285–6 sandwichensis 433 ulietensis 278–9 Turkey, Bruijn’s Brush 443 Turnagra capensis capensis 228–9 capensis minor 229–30 turnagra 230 Turnix sylvatica suluensis 122 varius novaecaledoniae 121–2 Turtur virgo 383 Tympanuchus cupido cupido 38–40 phasianellus hueyi 40–1 Tyrannula apolites 425 pusilla 425 Tyrant, Swainson’s 425 Tyto cavatica 192 letocarti 192 neddi 193 nigrobrunnea 447 novaehollandiae troughtoni 404 ostologa 192–3 pollens 193 rosenbergii pelengensis 349 sororcula cayelii 349 ‘Ula-­’ai-­Hawane 313 Upucerthia dumetaria peruana 224 Upupa antaios 211 Urosphena subulata advena 254 Vanellus macropterus 124 madagascariensis 125 Vanga, ­Short-­toed Nuthatch 358 Vangulifer mirandus 299–300 neophasis 300 Vermivora bachmanii 317–18 cincinatiensis 438 Verrulia carunculata 380 Vini cyanescens 388 sinotoi 169–70 vidivici 170 Violet-­ear, Buckley’s 408 Elliot’s 408 Vireo propinqua 427–8 Vireo, Vera Paz 427–8 Vittirallus watlingi 112 Vulture, ­Ash-­coloured 368 Chincou 368

Hare 368 Painted 329–30 ­White-­winged 367 Warbler, Aldabra Brush 255–6 Astrolabe 256 Awatcha 437 Bachman’s 317–18 Blue Mountain 438 Bush 437 Carbonated 438 Cincinnati 438 Daito Bush 452 Fayyum 432 ­Half-­collared 437 Huahine 258 ­Large-­billed Reed 453 ­Long-­billed 437–8 Louisiana 436 Moorea 257–8 ­Orange-­bellied 437 Pagan Nightingale 257 Raiatea 258–9 Semper’s 318 Sutton’s 438 Ticehurst’s 432 Waxbill, B ­ lack-­lored 358 Weaver, Lufira Masked 457 White-­eye, ­Chestnut-­flanked 264 Guam Bridled 264–5 Lord Howe 265 Mukojima 266–7 Sangihe 356–7 ­White-­breasted (Norfolk Island) 266 Whitethroat, Friburgo 412 Reichenbach’s 413 Wilsonia microcephala 436 Wood-­hoopoe, Cave’s 422 Woodcock, Puerto Rico 127–8 Woodcreeper, Vila Nova 426 Woodhen, Little 374 Woodnymph, Bell’s 339–40 Berlioz’s 410 ­Black-­bellied 411 Bourcier’s 411 Emerald 411 Lerch’s 410 Mexican 448 Reeve’s 411 ­Sapphire-­crowned 411 Southern 410–11 Woodpecker, Berlepsch’s 424 Crawfurd’s 424 Cuban ­Ivory-­billed 217–18 Grand Bahama 425 Hargitt’s 425 Imperial 214–16 ­Ivory-­billed 216–17 Javan ­Buff-­rumped 449 Kaempfer’s (Piauí) 449 Pennant’s Ferruginous 424 ­Yellow-­legged 425

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Extinct Birds Woodstar, Decorated 420 Hartert’s 420 Wren-­babbler, R ­ usty-­throated 451 Wren, Charleville Heath 427 Colombian Whiskered 357 Daito 345 Guadalupe 269 ­Long-­billed 223 Martinique House 269–70 North Island Bush 219 North Island S ­ tout-­legged 222 San Benedicto Rock 268 San Clemente 268

South Island Bush 220–1 South Island ­Stout-­legged 222 Stead’s Bush 219–20 Stephens Island 221 Xenicus longipes longipes 220–1 longipes stokesii 219 longipes variabilis 219–20 lyalli 221 Xestospiza conica 307–8 fastigialis 308 Xiphocolaptes franciscanus 426 villanovae 426

Yellowthroat, New Providence 318–19 Zodalia glyceria 419 Zoothera cameronensis kibalensis 280 heinei choiseuli 280 terrestris 279 Zosterops albogularis 266 conspicillatus conspicillatus 264–5 (mayottensis) semiflava 264 nehrkorni 356–7 strenua 265 tephropleura 456

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