HBW 7 - Foreword on extinct birds by Errol Fuller

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  The subject of recent extinctions is one that seems, increasingly, to be a part of popular culture. Today the words Great Auk (Pinguinus impennis), moa (Dinornithidae), Thylacine or Tasmanian Wolf (Thylacinus cynocephalus), Quagga (Equus quagga) and, of course, Dodo (Raphus cucullatus) are names with which many people are familiar. Yet, while the subject has always held an intrinsic fascination, it is one that comparatively few have cared to pursue - until recently. Now, the increasing vigour of the conservation movement, combined with the new vogue for popular science, has brought with it a fresh awareness of, and enthusiasm for, the subject.

  It is not difficult to see why. First, there is the mystery - and mystery exerts a great influence over the human psyche. Yet, paradoxically, recently extinct creatures are accessible in a way that animals known only from the fossil record are not. They sometimes seem almost touchable but, of course, this is only an illusion. In reality, they remain as elusive as those species that passed away aeons ago.

  Recent extinctions have a major advantage over those from the deep past: they offer the possibility that we may learn something significant from their example. Perhaps if we listen to these woeful stories we may be able to find a way to prevent similar tragedies from occurring in the future. Or, perhaps not!

  Then, also, there is the tantalizing possibility of rediscovery. While there may be no chance of finding a Tyrannosaurus lurking is some jungle fastness, who can say with certainty that Pink-headed Ducks (Rhodonessa caryophyllacea) do not still survive in an Indian wilderness? There have been numerous examples of rediscovery and, doubtless, there will be others.

  It has become traditional to regard 1600 as an approximate cut-off date from which to determine recently extinct bird species. Lionel Walter Rothschild's pioneering work of 1907 began this trend, since which time the date of 1600 has been adopted by several other writers, including the present one. The date is not quite as arbitrarily chosen as it may seem. The year 1600 represents a date heralding a period at which relatively reliable records began to accumulate; before this time it is generally impossible to make realistic sense of the few records that exist. The date also largely eliminates fossil birds or most birds known only from skeletal material. Such a subject is a vast and complex one and totally beyond the scope of a work such as this. For various technical and practical reasons it is impossible to put an exact figure on the number of bird species that have become extinct during the last 400 years. Obviously, any number may have passed away without leaving any trace at all. Concerning those of which we have certain knowledge, most authors would agree that there are something between 80 and 100, depending on how one interprets evidence. Sadly, this number is likely to increase dramatically during the next few years.

  At many practical levels the subject of recent extinctions is a difficult one, for how can these be accurately and consistently defined? First comes the very complex question, “How do you know it is extinct?” In many cases - the Dodo, for instance - it is obvious: a very large, conspicuous bird restricted to a relatively small island could certainly not survive unnoticed. In others the issue is rather more difficult to resolve. From the ranks of the world's bird species it is possible to select a considerable number that might be extinct. But on the other hand, they might not be!

  One well-known definition states that any creature not seen for a period of 50 years may be regarded as extinct. Unfortunately, however, such a definition - understandable and persuasive though it may be - is unworkable from any realistic point of view. Patterns of decline, remoteness of terrain occupied, recent changes to the environment and a whole variety of other elements may be relevant in this or that particular instance. There are a number of bird species that went missing for upwards of 50 years and then returned from seeming oblivion, notably the Forest Owlet (Athene blewitti) and the New Caledonian Owlet-nightjar (Aegotheles savesi) “absent” for 113 years and 118 years respectively, and even more recently Bruijn's Brush-turkey (Aepypodius bruijnii), rediscovered on the remote and almost inaccessible New Guinean island of Waigeo. Yet there are others that have been seen in the last decade or two that could now categorically be considered extinct.

  Rarity is a peculiar thing. Some species can stay rare for decades, or even centuries, without actually dropping into the doleful condition of extinction. Others become extinct in much more dramatic style.

  The celebrated case of the Takahe (Porphyrio mantelli) of New Zealand provides a perfect example. Known from just four specimens taken during the nineteenth century, the species “vanished” for a period of 50 years before being rediscovered during 1948. Since this time the species has limped on in extreme rarity and managed to survive into the twenty-first century. This might be compared with the famous downfall of the Passenger Pigeon (Ectopistes migratorius), which at the start of the nineteenth century was possibly amongst the most numerous birds on Earth. In the space of two or three decades numbers plummeted from many, many millions to just a handful of individuals, and during 1914 the last individual of all died in her cage at the Cincinnati Zoo.

  What does all this mean? Simply that, while there may be guidelines to govern the nature of extinction, there are no hard and fast universal rules. Each case must be judged on its own merits.

  This introduction to the subject features only those birds that can claim without doubt the title of extinct. Other, more controversial, cases are reviewed in brief.

  Another great difficulty that surrounds extinct birds concerns the definition of a species. As so many of these creatures are poorly known it is by no means certain that some of them had actually passed the boundary at which they could be regarded as full species. This might particularly be the case where a form is known only from skeletal remains, but even with more substantial evidence there may still be controversy over definitions, as indeed is apparent with the differing taxonomic views regarding even common and well-known taxa. One authority might define an extinct form as simply a subspecies, while others may see it as a full and legitimate species. The Norfolk Kaka (Nestor (meridionalis) productus) and the Guadalupe Caracara (Polyborus (plancus) lutosus) are good examples. Both hover rather uneasily at the junction where full species meet subspecies, and either can be fairly defined in either way.

  Then there is the matter of mystery species and others that might be termed “hypothetical”. A surprisingly large number of species have been described from just a single specimen with no other similar bird ever turning up. Sharpe's Rail (Gallirallus sharpei), for instance, was described from a single specimen that was purchased by the Leiden Museum during 1865 and came from an unknown source. Since that time no comparable specimen has come to light. Is the bird a freak or is it an otherwise unknown - but legitimate and presumably extinct - species? Probably it is the latter, but no-one knows for sure.

  “Hypothetical” birds come into a related but slightly different category. These are forms that have been described from early travellers' tales or descriptions. The desire to name new species seems so great that many ornithologists (who should have known better) have succumbed to temptation and named birds on evidence that is totally unsatisfactory and against the rules of scientific nomenclature. Perhaps the most famous case of this kind is that of the White Dodo (Raphus solitarius) of Reunion. There is a very, very small amount of evidence that such a bird may once have existed but absolutely no hard proof of any kind. Many other “hypothetical” bird species have been proposed but, while their stories may be fascinating, none of these have any real place in a serious list of extinct birds.

  It may be worth briefly remarking on another category of disappearing species, namely those classified by BirdLife International as “Extinct in the Wild”, at present numbering just three species, the Alagoas Curassow (Mitu mitu), the Guam Rail (Gallirallus owstoni) and the Socorro Dove (Zenaida graysoni). Sadly, it is very likely that within just a few years quite a number of other species may join this group. The prime candidate must be Spix's Macaw (Cyanopsitta spixii), with a wild population of only a single bird from 1990 until it disappeared in 2001, but with at least 60 birds (probably all closely related) currently held by aviculturalists. By definition, all such species survive in captivity, and the chances of their eventual reintroduction to their natural range should not be discounted, so they remain outwith the strict scope of this foreword.

  All these considerations mean that no-one can come up with a definitive extinct list that is totally satisfactory to everyone else. Even the birds featured here are at some variance with the list given in the present author's recent (2001) book Extinct Birds. In order to ensure continuity and conformity with HBW, certain eliminations and additions have been made. Most notably, several species covered in Extinct Birds have been excluded from this foreword because they have already received full treatment in the main body of HBW, with individual species account, illustration and map; most of these were, perhaps optimistically, listed as “Almost certainly Extinct”. These variations should be regarded in the spirit in which such determinations must necessarily be made. No decisions on what is extinct or what is not, what represents a subspecies and what does not, should be regarded as graven in stone. All such things change according to current knowledge and attitude. The information given here represents a good and honest guess at the truth - but it should never be considered as an absolute truth. 

Reasons for recent extinctions are, of course, many and various but most are due to human intervention. Hunting and shooting, though the most obvious kind of destructive processes, are not - at the species level - necessarily the most devastating. Destroying habitats and/or altering them have caused immense damage. The problems caused by complete destruction of habitat are obvious, but alteration, in the form of altering land use or introducing alien species, is similarly critical and both of these are among the main factors that destroy bird populations and ultimately lead to extinctions.

  The geography of recently extinct birds is a particularly revealing one. By and large such extinctions come from islands rather than from the major land masses. Although there are a number of spectacular exceptions - and this number is likely to increase in the coming decades - it is on oceanic islands and island groups that the major casualties have occurred. Thus, the Hawaiian Islands, the Mascarenes, New Zealand and the islands south of Japan are all among the great theatres of extinction.

  Clearly, if a species' habitat is demolished or radically altered, sooner or later that species will inevitably vanish along with the environment. With many island birds being endemics, it must follow that a species lost from its island home is a species lost entirely. Many, many island forms are already gone and many more are today under threat; there is little doubt that true island insularity hugely increases any species' vulnerability.

  At a less dramatic level, local populations of otherwise flourishing species are often wiped out when a marsh is drained or a forest chopped down. If the species is plentiful elsewhere, its loss in one particular place may well pass unnoticed. Even subtle interference can sometimes have the same result and such subtle interventions can take many forms. The results of the random (or sometimes highly organized) introduction of predators to predator-free places are obvious but the introduction of animals with no predatory intent can cause equal chaos. Competition for limited territory or an ecological niche is one obvious area of concern. Equally insidious are the possibility of introduced disease, the spread of alien plants pushing out the natural flora, or the introduction of poisons.

  There is a tendency to think that the creation of reserves might provide a convenient solution. Unfortunately this setting aside of portions of land does not necessarily work. When environments become fragmented, it is often not possible to foresee the ultimate consequences of such fragmentation. 

The birds featured here are listed in an order conforming to the general arrangement of HBW. A short preface for each major bird group that is currently affected by extinction is included to facilitate some discussion of controversial or problematical forms.

  Concerning many extinct species virtually nothing is known; about others there is a considerable body of information. If basic ornithological information is not given in any particular case, it may be assumed that this is because such information remains unknown.

   A computation has recently been made (for BirdLife International's book Threatened Birds of the World, 2000) that one in eight of the world's bird species (around 1200 species) are at risk of extinction during the next 100 years. No-one should assume, therefore, that extinct species are irrelevant. They are not things that are gone and best forgotten. Rather, they should be seen as something to illuminate the future. And that future will be much drabber, should we not look at these pictures from history and learn from them.

 
RATITES

The celebrated nineteenth century ornithologist Alfred Newton defined the ratites as:

That division of the Class Aves whose sternum developing no “keel” resembles a raft or flat-bottomed boat.

  This is, then, a rather loose assemblage of creatures but from it can be selected a number of the largest birds known to us. Among living ratites are the Ostrich (Struthionidae), the rheas (Rheidae), the cassowaries (Casuariidae) and the Emu (Dromaiidae), and the much smaller and different-looking kiwis (Apterygidae). Among extinct groups are the elephantbirds (Aepyornithidae) and the moas (Dinornithidae).

  The origins of these rather un-birdlike birds are unclear and how each of the distinct groups relate to one another - if at all - is a matter of some controversy.

  The major areas of mystery concerning the ratites can be summed up in two questions. Did the various ratite orders evolve separately or are they all derived from a common ancestral stock? Was this ancestral stock - or these stocks - made up from true flying birds, or were these prehistoric creatures that never actually acquired the power of flight?

  As far as the second question is concerned, it can be said that the extinct moas show no vestige of a wing. In simplistic terms they are the most perfect of bipeds, their forelimbs having vanished without trace. Notwithstanding this curious fact, it seems most likely that the ancestors of the birds now called ratites could all fly perfectly well and that they gradually lost the power in much the same way, and for much the same reasons, as other flightless birds.

  The question of common ancestry is, perhaps, rather more contentious. Although most of the ratites show some overall similarity in general appearance, this may be entirely due to the factor known as convergent evolution. In other words, these creatures may have grown to look alike simply because they share similarities in their lifestyles.

  Traditionally, the five extant ratite families have been awarded four or five separate orders, but in recent years there has been an increasing tendency to lump them all in a single order, Struthioniformes, with the different groupings being reduced to four suborders, the families Casuariidae and Dromaiidae being placed in the same suborder. How this affects the extinct Aepyornithidae and Dinornithidae, each traditionally placed in its own separate order too, is not always discussed, but the general consensus of opinion is that they too are probably most usefully placed in this all-enveloping Struthioniformes.

  Ratites have certainly been very successful in many parts of the world and Africa, Asia, South America and Australia have all been occupied. So too have smaller land masses: New Zealand, Madagascar, New Guinea and many of the islands that make up Indonesia. Yet despite this evident success, the ratites have been in general decline during the age of man. There is, for instance, no doubt that until comparatively recently Ostriches occurred in many places where they can no longer be found. They have vanished from many parts of Africa, from Asia, and from Arabia, where the last individuals may have died during the 1940's or, perhaps, during the 1960's. There is evidence that Ostriches once lived in parts of southern Russia. Whether the birds that inhabited all these places belonged to just a single species or whether there were, in fact, several different kinds, is difficult to determine.

  Similarly, it is difficult to determine exactly which ratite species have become extinct since 1600. For the purposes of this work four species have been selected, an elephantbird and three moas, although in each of these cases there is doubt over the actual date of extinction, and some might consider that these birds were gone before 1600. Each of the four is known only from skeletal remains, skin or egg fragments, and in some cases complete eggs. Curiously, the eggs of moas are excessively rare whereas their bones are relatively common; the reverse is the case with the elephantbirds and their eggs turn up far more frequently than their bones.

  Two other forms are sometimes listed as recently extinct species. These are dwarf emus from Kangaroo and King Islands, off the south coast of Australia, known from skeletal remains and a single stuffed specimen now in Paris. Although there is no doubt that these two island forms are now extinct, both having vanished during the early years of the nineteenth century, there is uncertainty over whether either had passed the point at which it could be considered specifically distinct from their extant relative on the mainland, the Emu (Dromaius novaehollandiae).

  There has been a certain amount of sloppiness over the way in which scientific names for these forms have been changed. The Kangaroo Island birds are now usually listed as Dromaius novaehollandiae baudinianus (formerly D. n. diemenianus) and those from King Island as D. n. ater (previously minor). There is, in fact, no certainty over which specimens came from which island, and, as a result, the switch to these names from older ones may not be justified. The controversy over this is still raging (see Greenway, 1958, Jouanin, 1959, Fuller, 2000a) so no hard and fast determination can be made.

  Another rather distinct form was the Tasmanian Emu (Dromaius novaehollandiae diemenensis) but, like D. n. baudinianus and D. n. ater, it had probably not evolved into a species that could be regarded as separate from the Emu of the Australian mainland. This Tasmanian population became extinct during the middle years of the nineteenth century.

AEPYORNITHIDAE

Great Elephantbird Aepyornis maximus

The Great Elephantbird of Madagascar holds the distinction of having laid the largest egg known to man, in fact the largest of single-celled objects. This extraordinary egg is far bigger than that laid by any known dinosaur or any other giant reptile and is thought to have given rise to Arab tales of the “roc”.

  Nothing is known of the bird's habits and even its exact appearance in life is unknown; it is assumed that it looked something like an enormous Emu. Standing some 3 m high, this was an incredibly ponderous and heavy bird and in terms of sheer weight it is the largest bird ever known to have lived.

  There appear to have been several related species occupying the island of Madagascar and this particular one - the largest - seems to have been the last to become extinct. The testimony of the first French Governor of Madagascar, Étienne de Flacourt, seems to indicate that individuals still survived in the 1650's. He mentioned a large bird that laid an egg similar to that of an Ostrich. Sadly for Monsieur de Flacourt, he was killed by Algerian pirates on his way back to France without further elaborating on his story.

  The gigantic eggs of the bird, as big as an American football or a rugby ball, are still found from time to time in Madagascar and fragments of them are common at certain localities. Although virtually complete skeletons exist, bones seem to be much, much rarer than eggs.

DINORNITHIDAE

Slender Moa Dinornis torosus

Because the original fauna of New Zealand contained no mammals, these islands were often called “The Land of Birds”. Such a title has long been inappropriate, as introduced mammals now rule the land in much the same way as they do elsewhere - but for aeons birds represented the dominant lifeform. Quite why mammals were originally excluded from this corner of the planet is one of the mysteries of prehistory, but excluded they certainly were. In the land now known as New Zealand, birds adapted to fill many of the niches more familiarly occupied by mammals.

  The most spectacular of these creatures were the birds now known as moas. There were a number of species and argument rages today over just how many. Some authorities put the figure as low as 10, others identify 30 or 40; such controversy is typical of the kind of difficulty experienced when ornithologists try to make sense of osteological material and have little else to go on. In the case of the moas there are some skin and feather fragments as well as a very few eggs and egg pieces, but these remains are tantalizing and reveal little of significance.

  The largest of the moas, Dinornis giganteus, was the tallest known bird ever and it raised its head, upon a long serpentine neck, to the extraordinary height of around 4 m. Although not as hugely ponderous as the Madagascan Great Elephantbird, this was a considerably taller creature and presumably it must have been an awe-inspiring sight.

  The evidence suggests that this particular species was probably gone by 1600 but it seems probable that a smaller - but still gigantic - relative, Dinornis torosus, survived after this date.

  There is, in fact, much controversy over the date at which the last of the moas became extinct. Some argue that this was at a comparatively antique time, others suggest the extinction was much more recent.

  A fragment of bone from Dinornis torosus seems to have been the first moa relic to come to the attention of ornithologists. This was in 1839 when the celebrated comparative anatomist Richard Owen stuck his neck out and made his famous announcement that there had lived in New Zealand - or even perhaps at that time still lived - a gigantic bird similar to an Ostrich. Owen based this deduction on a very small fragment of bone that had been brought to him for identification by a Dr John Rule. Despite the unpleasant things that are often said of him, Owen was certainly not lacking in courage: had he been proved wrong, his reputation, and indeed his whole career, would have been in tatters. However, only a short time after his sensational announcement, solid proof, in the form of skeletons, came to light and proved his deductions to have been accurate.

  Bones of D. torosus have been found at widely scattered localities on New Zealand's South Island. Their comparative rareness in swamp deposits suggests that the species may have preferred hill country. Some of the tops of the skulls of this species show deep pits in the surface and this may indicate that birds carried crests.

Greater Broad-billed Moa Euryapteryx gravis

There were several moas belonging to the genus Euryapteryx and these are distinguished from other moas by their broad, round-tipped beaks. Species have been described from both the South Island of New Zealand and the North. Euryapteryx gravis seems to have been the most widespread of these, and it has left its remains scattered throughout New Zealand. Although it was rather smaller than birds belonging to the genus Dinornis, it was nonetheless a very large and bulky creature. Its bones are said to be those that are most commonly found in Polynesian kitchen middens, so there is little doubt that this bird was often eaten by man.

  At what date this supply of meat ran out is not known and it is possible that the species was extinct before 1600. On the other hand, birds of this species may have been the creatures referred to by ancient Maoris when they recalled moa hunts that took place during the last half of the eighteenth century. These intriguing stories have long fascinated researchers and they centre around the reminiscences of aged warriors who told their tales of long ago during the middle years of the nineteenth century. Whether these old men were simply enjoying being the centre of attention or whether there was some substance in their stories is a matter of speculation.

Lesser Megalapteryx Megalapteryx didinus

Although gigantic by bird standards, this creature was diminutive in comparison to other moas, standing little more than a metre tall. It may have been restricted only to the South Island.

  It seems quite likely that the species survived to a later date than any other, and it may have been extant during the early nineteenth century; there are even those who hope that it survives still in some remote mountain fastness of New Zealand's fiordland. Its small size and the wild terrain it occupied may have combined to enable it to survive when the lines of its larger relatives failed.

  There are several relics of this species that have survived in a remarkably good state of preservation. Dried heads and necks with ligaments and flesh attached have been found and so too have feathers and mummified feet. Such finds come from the province of Otago and they make it clear that the species was feathered right down to the toes. The feathers show an open structure, lacking barbicles and presenting a rather hair-like appearance. They are greyish-brown in colour, some with a rufous tinge and some tipped with white.

  Sir George Grey, at one stage Prime Minister of New Zealand, was once told at Preservation Inlet, South Island, of the recent capture and killing of a small moa out of a drove of six or seven. This incident is supposed to have happened around 1868. Did it really occur, or is it just a story?

 
PODICIPEDIFORMES

This group of birds may or may not contain extinct species but it certainly contains several that are under grave threat of extinction. The most seriously threatened is the Atitlan Grebe (Podilymbus gigas), and it now seems highly likely that this form is actually extinct. In Volume 1 of HBW, the species was listed as Almost Certainly Extinct; several years later the “almost” can probably be removed. However, since the genes of this creature may still exist, there may be an argument for retaining it on the list of living creatures for a while longer.

  Affected by habitat degradation, the introduction of an alien, predatory fish and the arrival of a closely related species with which it hybridized, the species - in its purest sense - seems no longer to be extant.

  The Atitlan Grebe was always restricted to a single lake in Guatemala, where it adapted to the prevailing conditions by becoming flightless and isolating itself from its close relative, the widespread Pied-billed Grebe (Podilymbus podiceps). In essence this was simply an outsize version of the Pied-billed, and it was probably always vulnerable to the threat of hybridization with that species. This seems to be the factor that finally led to the loss of Lake Atitlan's grebe, an event that probably took place around 1990. The possibility that a few pure-bred examples of the species still exist cannot be completely excluded, of course, although it seems very doubtful. For more on this species and the cases of the probably extinct Rusty Grebe (Tachybaptus rufolavatus) and Colombian Grebe (Podiceps andinus), see HBW 1, pages 185-194.

 
PROCELLARIIFORMES

This group, containing the albatrosses, shearwaters and petrels, includes many threatened species but only one seems to be extinct.

HYDROBATIDAE

Guadalupe Storm-petrel Oceanodroma macrodactyla

The island of Guadalupe lies off the coast of Baja California, some 320 km south-west of San Diego. It should not be confused with Guadeloupe in the West Indies. Guadalupe of the Pacific was once a home to several forms that are now extinct; these were mainly subspecies of more widespread birds.

  A distinct species of storm-petrel once lived here, a species that nested in burrows carved out of the soft soil beneath trees on the steep north-eastern ridges of this small island. The population was probably never large but there is no reason to suppose that it was anything but thriving until cats and goats were introduced to the island. Cats are known to have infested the nesting areas and, presumably, it was this agency that brought about the destruction of the petrels. Apart from the fact that breeding began in March and continued until May, with a single white egg being laid, nothing of significance is known of the species. It seems to have become extinct during the years immediately preceding World War I.

 
PELECANIFORMES

One member of this order can confidently be listed as extinct, but there is one other rather mysterious form that may be. This is a bird known as Kenyon's Shag (Phalacrocorax kenyoni). It is identified from just three specimens from Amchitka Island, in the Aleutian Archipelago of the north Pacific, collected during the late 1950's and preserved as skeletons. The external appearance of this bird is unknown. So too is its relationship to other cormorants, and it has even been suggested that these may merely have been outsize individuals of the extant Red-faced Cormorant (Phalacrocorax urile), a species that is still common in this same area. It is not even known if the form kenyoni still survives, so this is truly a mystery bird.

PHALACROCORACIDAE

Spectacled Cormorant Phalacrocorax perspicillatus

The spectacular Spectacled Cormorant is a particularly striking member of the list of extinct birds. Discovered on Bering Island during 1741 by the celebrated naturalist George Wilhelm Steller, the species was exterminated in little more than a century. At the time of its discovery this large cormorant was common, although its range was very limited. In addition to Bering Island, it was also found on the nearby Commander Islands, and it probably lived on other islands of the Aleutian Archipelago in the north Pacific. Its general unwariness, its suitability for the pot and its small wings that rendered it almost flightless, combined to make this bird an obvious target for man.

  There are very few specimens in existence and nothing of significance is known of the bird in life.

 
CICONIIFORMES

There are several seriously threatened birds in this order but only one full species is known to have become extinct during the last 400 years.

ARDEIDAE

Rodrigues Night-heron Nycticorax megacephalus

During 1691 a small band of Huguenots, fleeing from religious persecution in France, ended up marooned for around two years on the Mascarene island of Rodrigues, far out in the Indian Ocean. Their leader was one François Leguat, and this gentleman made notes on the things he saw during his stay on the island, notes that included detailed observations of the birds he encountered. Years later, after his return to Europe, Leguat's notes were published in book form (simultaneously, in both English and French) and his book was called A New Voyage to the East Indies (1708). This splendid little volume provides primary source material for a number of extinct species, including the present one.

  Leguat describes a powerfully built and pugnacious heron that had almost lost the power of flight but had developed strong legs that enabled it to move very quickly.

  In their loneliness (there were no women with the small band!), Leguat and his followers found some small solace in the companionship of some endemic geckos, which they shook down from the trees. These little creatures, which were themselves to become extinct around the time of World War I, were surprisingly tame and the Huguenots often allowed them to feed from their own table. So bold and aggressive were the herons, however, that Leguat and his friends had great difficulty in protecting their little pets from the marauding birds.

  Despite their bullying manner, the night-herons died out at an unknown date during the eighteenth century. There is a fleeting mention of them in a document that dates from around 1725 but nothing more recent is recorded of the living bird.

  During the 1870's bones were found on Rodrigues that confirm the earlier written reports and suggest that this species was derived from a night-heron, but nothing more is known of it.

 
ANSERIFORMES

The waterfowl - the swans, geese and ducks - have always been enormously attractive to man. Like parrots, they are comparatively easy to domesticate and they have been bred in captivity, or semi-captivity, for many centuries. Ducks, geese and swans have been used for food, sport and also simply for decorative purposes.

   Many kinds have been seriously affected by changing patterns of land use, habitat alteration and wetland drainage. Many species have declined drastically and four are now regarded as extinct and discussed below, although there is still some hope that two of them might conceivably still hang on in remote parts of their ranges.

  One other species, the Chatham Swan (Cygnus sumnerensis), sometimes occurs on lists of recently extinct birds but there is no real evidence to show when this creature died out. Known only from bones, it may well have become extinct long before 1600. Another species that is sometimes listed as recently extinct is Sarkidiornis mauritiana from Mauritius but, like the Chatham Swan, there is little to suggest that this creature lived on into recent historical times, and again it is known only from bones.

  Were it not for the fact that many kinds of waterfowl can be successfully bred in captivity, there would have been several additional extinctions, and some very notable conservation success stories concern waterfowl. Perhaps the most famous is that of the Néné or Hawaiian Goose (Branta sandvicensis). The population of this striking goose was long in decline and by the early 1950's it had plummeted to around 30 individuals. With numbers at this low level the prospects of the species' survival looked bleak. At various times during the nineteenth century and early years of the twentieth (and also in several places) it had proved possible to breed Nénés under captive conditions and so one last effort was made to save the species at the famous waterfowl reserve at Slimbridge, England. The operation was an enormous success and numbers steadily increased. There are now hundreds of Nénés in existence and captive colonies can be seen in many parts of the world. The species has also been reintroduced to its native Hawaiian Islands where around a thousand individuals probably survive. There is no reason to suppose that its future is anything but secure.

  Another Hawaiian species or subspecies that survives thanks largely to captive breeding is the Laysan Duck (Anas (platyrhynchos) laysanensis). It is recorded that numbers once dropped as low as seven individuals and during the 1930's the tiny surviving population was alleged to contain just a single female. Somehow this form survived, and a captive breeding programme was started in 1958. Many birds have been successfully bred in captivity and populations of Laysan Ducks now exist in a number of waterfowl collections. Unfortunately, some of these populations may no longer be pure.

  A form that has caused considerable confusion is the one known as the Marianas Duck (“Anas oustaleti”) and there is no general agreement about the true nature of this creature. It comes from the Pacific islands of Guam, Tinian and Saipan and there are two distinct colour morphs, one of which resembles a Mallard (Anas platyrhynchos), the other looking rather like a Pacific Black Duck (Anas superciliosa). Because of this resemblance and also because of the extreme isolation of the islands on which it occurs, it seems likely that the population is the result of hybridization. Probably, individuals of the two commoner species arrived on the islands, naturally or with human assistance, and, because of the unusual circumstances, interbred. A viable hybrid population was established and some authorities consider that the colony represents the evolution of a species through the mechanism of hybridization. Unfortunately, this evolutionary experiment - assuming that this is what it is - seems to be coming to an end. The isolated colony is now very close to extinction; indeed, perhaps it is already gone.

ANATIDAE

Crested Shelduck Tadorna cristata

Only a very few specimens of this striking duck are in existence, and virtually nothing is known of it in life. Even the limits of its original range are a matter of speculation; there is an assumption that it bred in eastern Siberia and migrated to Korea and Japan, but this may be incorrect. It is not even certain that it is extinct, although hopes for its continuing survival are probably forlorn.

  The most recently obtained specimen was taken in 1916 but a number of much later sightings are claimed, including a reported observation in southern China as late as 1990. Such unsubstantiated sightings should be taken with caution, but as long as tracts of potentially suitable habitat remain in unsurveyed areas, hope lives on.

  Although the earliest known specimen was taken near Vladivostock during 1877, the Crested Shelduck was not recognized as a species until 1917 when the Japanese ornithologist Nagamichi Kuroda formally described it. Prior to this the form had been regarded as a hybrid between the Ruddy Shelduck (Tadorna ferruginea) and the Falcated Duck (Anas falcata).

   Despite the fact that this is now such a mysterious bird, it seems that it was well known historically to both the Japanese and the Chinese and it is said to be depicted on a number of antique artefacts.

Pink-headed Duck Rhodonessa caryophyllacea

There is a certain amount of controversy over whether or not the Pink-headed Duck of north-eastern India still survives. Probably it does not, but there are those who believe it might and every few years or so an expedition goes in search of it. Thus far none of these expeditions has proved successful.

  This was a shy and wary bird that sometimes emitted a wheezy whistle and was always difficult to flush. Its rather secretive nature is one of the factors that encourages the optimists. Against this it might be said that individuals were found without too much difficulty during the nineteenth century and early years of the twentieth, yet none has been seen in the wild since the 1930's.

  On the water - where the birds spent most of the hot day - individuals showed a rather peculiar, stiff-necked posture and a strange triangular patch was evident on the top of the head. Despite some reports to the contrary, they seem to have been capable divers and were probably omnivorous feeders. Small groups sometimes gathered, particularly during the cooler months, and typically these groups consisted of six to eight birds, although sometimes the groups were considerably larger. The birds were rarely seen in the air and descriptions of their mode of flight are mildly contradictory: one report suggests that it was rapid and powerful, while another maintains it was light and easy.

  Pink-headed Ducks bred in April and May and they built circular nests from dry grass and feathers. The eggs - white or faintly yellowish in colour - were almost spherical in shape and the clutch numbered between five and ten. The species was recorded from Assam, Manipur, Bengal, Bihar, Myanmar and Orissa.

  Curiously, the last Pink-headed Duck of all may have died in England, thousands of miles from its native home. A number of captive individuals were kept in a private collection at Foxwarren Park, Surrey, and a rumour suggests that a male bird may have survived until 1945.

Labrador Duck Camptorhynchus labradorius

The Labrador Duck is one of the most mysterious of North American birds. It was always uncommon, shy and wary and for these reasons seems to have been little affected by shooting.

   Reasons for its extinction are unknown but one school of thought suggests that it was a very specialized feeder unable to cope with the vast changes to the environment that came about as a result of European settlement. This idea is, perhaps, supported by the slightly aberrant development of the bill which shows a curious softness around the edges.

  The species occupied the eastern seaboard of North America where it seems to have ranged from Labrador, in which area it may have spent the summer months, down to Chesapeake Bay. Nesting grounds were never reliably located; they may have been in Labrador or on islands in the Gulf of St Lawrence. Several eggs exist that are attributed to the species, but all are of doubtful provenance.

  Labrador Ducks seem to have declined rapidly after 1840 and the last sighting of the species dates from the 1870's. A specimen was apparently taken from Long Island waters during 1875 and another individual was allegedly shot near Elmira, New York, during 1878.

Auckland Merganser Mergus australis

The Auckland Islands lie some 320 km south of New Zealand and this rather forlorn group was once home to a species of merganser that, while not entirely flightless, showed a marked reduction in wing size compared with its relatives. As often happens when birds develop in evolutionary backwaters (i.e. areas without mammalian predators), the adults begin to resemble the immature stages of their relatives and this is just what was happening in the case of this merganser. Similarly, and typically, its chicks were showing none of the distinctive patterning shown in related species.

  The Auckland Merganser was first discovered in 1840 during a French exploratory visit to New Zealand waters. Two French corvettes, L'Astrolabe and La Zelée, arrived at the rarely visited and remote Auckland Islands, and some of the crew collected specimens. A Lieutenant Charles Jacquinot obtained a merganser and published a description of it, jointly, with ship's surgeon, Jacques Hombron. No further examples were spotted for 30 years or so, but then the species was seen on a number of occasions until the year 1902, after which time it vanished.

  The most extraordinary aspect relating to this species lies, perhaps, in its range. Five kinds of merganser exist north of the equator, a sixth inhabits parts of Brazil but none - other than this one - occurs anywhere near the Auckland Islands. Bones found in Polynesian kitchen middens indicate that the species once occurred on the main islands of New Zealand.

  The reasons for its extinction are unknown, although some reports suggest that individuals were prone to hide among rocks when pressed, rather than take to the water and dive like their European counterparts. Such a defensive strategy obviously did not work well against man and the predatory mammals he introduced.

 
FALCONIFORMES

The birds of prey are, of course, well known as being among the most persecuted of the world's birds, yet, despite the fact that many are seriously threatened, none can be categorically listed as recently extinct.

  One form that might qualify as an extinct species is the Guadalupe Caracara (Polyborus plancus lutosus). There is no doubt that the form is extinct, as no individual has been seen since the year 1900, but what is in doubt is the bird's taxonomic status. This is one of those creatures that hovers close to the border of what defines a full species and what defines a subspecies. Some authorities (including the present writer, 2001) have regarded it as a species in its own right, while others relegate it - with equal justice - to subspecific rank, considering it simply an island subspecies of the Crested Caracara (Polyborus plancus).

  Irrespective of the taxonomic view taken, this bird is one of those that disappeared as a direct result of human persecution. Exasperatingly, the last individuals on record were a flock of eleven birds, of which the scientific collector Rollo Beck shot nine. There were no further records (see HBW 2, page 242)

 
GALLIFORMES

In addition to the single species from this order that is definitely extinct, another may well be, and a third provides one of ornithology's little mysteries.

  The species that may well be extinct is the Himalayan Quail (Ophrysia superciliosa), last reliably recorded during the nineteenth century. Inhabiting the undergrowth of steep Himalayan slopes at around 1650-2100 m, this quail has repeatedly been the object of speculation with regard to its possible continued existence, but, despite a number of claimed sightings, no convincing proof has emerged. Nevertheless, the difficulty of the terrain it inhabits means that the optimists continue to believe in its possible survival (see HBW 2, page 513).

  The mystery bird is the Double-banded Argus (Argus (argus) bipunctatus). This form is known from a single feather found at an unknown location at some time before 1871. This feather, although similar to those of the two known species of argus pheasant, is markedly different to that carried by either. Does it come from a freak? Does it come from a distinct species? Is that species extinct? No-one knows.

PHASIANIDAE

New Zealand Quail Coturnix novaezelandiae

It is difficult to account for the disappearance of the New Zealand Quail. A closely related, possibly conspecific, Australian species, the Stubble Quail (Coturnix pectoralis), flourishes in its homeland, yet the New Zealand birds failed to survive the coming of Europeans.

  The species seems to have been common on the grass-covered downs of New Zealand's South Island until the mid-nineteenth century, but then, suddenly, its numbers plunged and within two decades it was all but gone. The factors responsible for this decline are unknown. Introduced mammalian predators, overhunting (presumably the birds provided the kind of “sport” that colonists were used to in their old homes), the burning off of land, or an unknown - and introduced - avian disease are all reasons that have been speculatively put forward.

  The latest specimens were taken during the late 1860's and individuals were, apparently, seen up to the mid-1870's. After this they vanished.

 
GRUIFORMES

This large and varied order includes the rails, a group of birds which has suffered a large number of extinctions, a fact that can be accounted for by the tendency of birds of this kind to disperse at random. Rails are particularly prone to colonize oceanic islands, diversify, and then lose the power of flight. Such a tendency makes them particularly vulnerable when their safe, island paradises become safe no longer, when they are invaded by mammalian predators.

  Eleven extinct species are recognized here, although differing taxonomic views can lead to rather different lists. To these eleven may be added the Bar-winged Rail (Nesoclopeus poecilopterus), a Fijian species not reliably recorded since 1890, and the Kosrae Crake (Porzana monasa), from the Caroline Islands, not recorded since at least the mid-nineteenth century; both have already been covered by HBW 3, pages 161 and 189. Another supposed member of this latter genus, Miller's or the Tahiti Crake (Porzana nigra), is often listed as an extinct species but in reality this should be considered a hypothetical form (see Walters, 1988, and Fuller, 2000a). Also deserving of mention is the Mascarene Coot (Fulica newtoni), not included in the list herein, as there is no hard evidence that it survived until 1600, although there are seventeenth century reports of it from both Reunion and Mauritius. The difficult case of Sharpe's Rail (Gallirallus sharpei) has already been mentioned (see page 12).

RALLIDAE

Mauritian Red Rail Aphanapteryx bonasia

Scattered among the seventeenth century written accounts and illustrations that relate to the celebrated Dodo are descriptions and pictures of a flightless bird of a rather different kind. These pictures show a creature that in overall shape and appearance looks something like a kiwi. A long, down-curved beak, rather hair-like plumage and stout legs all add to this impression. The bird in question bore no genuine relationship to the kiwis, however. It was a rail and, like the Dodo, it came from the island of Mauritius.

  Like its more famous fellow, the Mauritian Red Rail or Red Hen failed to survive the coming of Europeans by more than a few decades. The Dutch arrived on Mauritius, with devastating effect, during 1598, and it is highly doubtful if there were any red “hens” left a century later.

  Peter Mundy, an English traveller who penned fascinating reminiscences of his exploits, described the species as:

A Mauritius henne, a Fowle as bigge as our English hennes...of which we got only one. It hath a long, Crooked sharpe pointed bill. Feathered all over, butte on their wings they are soe Few and smalle that they cannot with them raise themselves From the ground...They bee very good Meat, and are also Cloven footed, soe that they can Neyther Fly nor Swymme.

  Mundy was recalling events of 1638 and the fact that he and his companions got only a single individual surely indicates that the species was already rare. Earlier accounts suggest that the birds gathered in flocks and were fatally attracted to the colour red. By means of a piece of red material they could be induced to approach and then, of course, they were caught.

Leguat's Rail Aphanapteryx leguati

The Huguenot refugee François Leguat discovered a relative of the Mauritian Red Rail during his two-year sojourn on the island of Rodrigues:

Our Wood-hens are fat all the year round and of a most delicate taste. Their colour is always of a bright grey, and there is very little difference in the plumage between the two sexes. They hide their nests so well that we could not find them 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 them too heavy for it. If you offer them anything that's red, they are so angry they will fly at you and catch it out of your hand, and in the heat of the combat we had an opportunity to take them with ease.

  It is curious to note how this last piece of information conforms to the reported behaviour of the Mauritius Red Rail.

  An anonymous manuscript - alleged to have been written by a marooned sailor named Tafforet - dating from around 1725, described a creature that could not fly, was armed with a heavy beak, made a continual whistling, fed on tortoise eggs (Rodrigues was once home to a now extinct species of giant tortoise) and was a powerful runner.

  Since no subsequent visitor to Rodrigues mentions any such creature it may be assumed that these birds became extinct soon after 1725.

  In 1874 bones were found in a Rodrigues cave and these appear to correspond with the early accounts; they constituted the material on which Alphonse Milne-Edwards based his formal description of this species. Additional skeletal finds have subsequently been made.

Wake Rail Gallirallus wakensis

The Wake Rail has acquired the grim celebrity of having been eaten out of existence by hungry Japanese soldiers during World War II. Unable to fly, these rails could scuttle about their island home quickly, but despite their agility any efforts to escape would have been no match for the concerted efforts of a few peckish men. Before World War II the rails were plentiful but by the time the Japanese garrison left Wake Island in 1945 there were none left.

  Remote Wake Island is situated far out in the Pacific, hundreds of kilometres from the Hawaiian group. Despite its extreme isolation, rails something like today's Buff-banded Rail (Gallirallus philippensis) managed to colonize it at some period in the distant past, and from this ancestral stock the form developed that has become known as the Wake Rail. Over generations the size became smaller, the plumage lost its brightness and the wings became rudimentary. As Wake is small, low and scrub-covered, the rails probably had to make use of any food source that came their way; molluscs, insects and other invertebrates are likely to have provided the basic diet.

  Other than the fact that breeding was observed to take place in July and August, and that individuals made the low chattering and clucking that is typical for a rail, nothing of substance is known of this species.

Tahiti Rail Gallirallus pacificus

The Tahitian Red-billed or Tahiti Rail is known only from a painting that survives today in the collection of the Natural History Museum, London. This painting was produced by Georg Forster, one of the naturalists who sailed with Captain James Cook on his second epic voyage around the world during the 1770's. Forster's credentials are impeccable, so there need be no doubt that he saw just such a bird as that depicted.

  The individual that served as a model for his picture was found on Tahiti, where it was known as tevea, oomnaa or eboonaa. No-one knows why it became extinct - or when - but it seems likely that this was connected with the introduction of cats and rats to the island.

Dieffenbach's Rail Gallirallus dieffenbachii

The remote Chatham Islands lie way to the south of New Zealand and here, in isolation, two species seem to have developed from an ancestral stock that resembled the Buff-banded Rail. The less evolved of these is Dieffenbach's Rail, and this form seems to have developed from a comparatively recent invasion of the ancestral birds. Indeed there is some doubt as to whether or not this form had actually passed beyond the stage at which it could be regarded as just a subspecies. Apart from some minor plumage differences, it sports a sharply downturned beak and this is clearly an adaptation to the conditions prevailing on the island.

  Nothing is known of the bird in life. It is known from just a single specimen collected during the 1840's and now in the collection of the Natural History Museum, London.

Chatham Rail Gallirallus modestus

This species seems to provide a fairly typical example of the way in which rails develop on isolated islands. It is assumed that this is another form that has evolved from an ancestral stock resembling the Buff-banded Rail, but in this case the evolution was rather more advanced than in others.

  Following the arrival of this ancestral stock, the birds diversified over aeons, assumed a neotenous appearance (adults began to resemble juveniles), lost the power of flight, and their plumage began to assume a rather hair-like appearance. All these developments enabled the birds to make maximum use of the rather limited resources available on the Chatham Islands, by cutting down on what were, for them, unnecessary and costly forms of energy expenditure, but, of course, they rendered them horribly vulnerable if, and when, the situation changed. The coming of man and his introduction of mammals, either intentional or not, caused just such a change.

  The species seems to have been on several islands of the Chatham group - Chatham Island itself, Pitt Island and Mangere - but it is from Mangere that the majority of surviving specimens come, and this seems to have been the last stronghold. From here the birds vanished around 1900. By this time much of the land had been burned off, sheep were grazing and cats had overrun the island.

  All that is known of the species is that it nested in burrows or in hollow trees, ate insects and laid a creamy-white egg.

Ascension Rail Atlantisea elpenor

The evidence for the former existence of a small rail on Ascension Island is of two kinds. First, bones from a rail have been found on the island and, second, there exists a seventeenth century written account of just such a creature. This account was written by the much travelled Englishman Peter Mundy, the same man who also penned an account of the extinct rail of Mauritius. Mundy's descriptions are always reliable and in this case he also provided a crude but explicit drawing of the birds he saw. He wrote:

Alsoe halfe a dozen of a strange kind of fowle, much bigger than our Sterlings...colour grey 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...It was more than ordinary dainety meat, relishing like a roasting pigge.

  Nothing more is known of the species. Why and when it became extinct are also matters completely shrouded in mystery.

Laysan Crake Porzana palmeri

Laysan is a tiny coral island around 1280 km to the north-east of Hawaii, and one of its few inhabitants was a small species of rail. Because it is so remote, the island was infrequently visited but, nevertheless, there are several good accounts of the bird in life. These consistently tell of how bold the tiny creatures were, hunting for flies between the feet of men, or entering huts and other buildings in search of food. All kinds of insects and their larvae were eaten and the rails would often squabble with Laysan Finches (Telespyza cantans) over the broken eggs of noddies and terns (Sternidae). Scraps were taken from the carcasses of dead seabirds, and vegetable matter was also consumed. In short, no food resource was neglected.

  The Laysan Crake was flightless, but birds could move with incredible swiftness and agility when attacked by frigatebirds (Fregata) and other larger seabirds. The call was described as a warbling and chattering, and there was a peculiar evening chorus that began soon after darkness fell and lasted for just a few seconds. The birds all started their chorus in unison and then all fell silent together. The sound produced was said to be like a handful of marbles being thrown onto a glass roof and then descending in a series of bounds.

  Breeding occurred between April and July. A nest was made of grass and leaves, and this material was arched over to make a roof with an entrance at the side.

  During the 1890's these little birds were common but then rabbits and guineapigs were introduced to the island, and the habitat deteriorated. By 1925 the vegetation was gone, and Laysan was a desert. During this year two rails were seen, but by 1936 there were certainly none left. The species was not quite extinct, however. During 1891 a few individuals had been transferred to Eastern Island in the Midway Atoll, and here the small population flourished. By the start of World War II the birds were common. Then, alas, a US Navy landing craft drifted ashore bringing with it an accidental invasion of rats. Within two years the last Laysan Crakes were gone.

Hawaiian Crake Porzana sandwichensis

A small species of rail once inhabited the main island of Hawaii and perhaps some other Hawaiian islands. A handful of specimens exist in the museums of the world but these have in themselves caused some controversy. Two of them are rather paler in colour than the rest, giving rise to the idea that there may have been two species rather than one. This claimed second species is herein tentatively treated as a separate race, millsi. On the other hand the general consensus of informed opinion is that the paler individuals may simply be immatures.

  Apart from the probability that these birds were extinct by the end of the nineteenth century, nothing is known of them.

Samoan Moorhen Gallinula pacifica

There seem to be eleven specimens of this species in the world's museums, the first collected during 1869 and the last perhaps a mere five years later. Since then the species has never more been located.

  Little is known of it other than that it inhabited the Samoan island of Savaii. It was called the puna'e by the local inhabitants, it was probably flightless (or almost so), and it had exceptionally large eyes, from which fact it has been deduced that it was nocturnal or crepuscular in habit.

  The reasons for its extinction are unknown but, presumably, introduced predators will have played some part.

White Gallinule Porphyrio albus

When British ships belonging to what was known as the First Fleet were dispatched under Governor Phillip to Australia during the late 1780's, their task was to found a penal colony to which British felons could be transported. Much of interest surrounds this rather unpleasant endeavour and one of the by-products of the epic journey was a great upsurge in knowledge concerning Australian natural history.

  One of the creatures that was brought to attention as a result of the expedition was a rather strange and albinistic swamphen that occurred on Lord Howe Island, a small and isolated piece of land lying some 480 km off the eastern coast of Australia. These birds were white, or mostly so, and considerably larger and heavier than ordinary swamphens and they had comparatively formidable bills. In fact, they seem to have been evolving independently into an entity that was reminiscent of New Zealand's Takahe (Porphyrio mantelli).

  Unfortunately for the birds, men who visited the island were easily able to kill them with sticks and this fact alone makes it highly likely that they were flightless. Just two specimens of the White Gallinule (or Lord Howe Swamphen) are in existence today, one in Vienna, the other in Liverpool.

  Once man began to visit their island home regularly, the species' days were numbered, and it seems to have survived for only a few decades after the coming of the First Fleet.

 
CHARADRIIFORMES

The Charadriiformes comprise a diverse group containing waders, gulls and auks. In addition to the three species discussed below, there are two others that are often listed as extinct, and may well be so, though herein the possibility of their survival is given the benefit of the doubt. These are the Javanese Wattled Lapwing (Vanellus macropterus), last definitely recorded in 1939, and the Eskimo Curlew (Numenius borealis) for which sightings are occasionally claimed right up to the present time, but which, if extant, could probably only number a handful of individuals (see HBW 3, pages 418 and 503).

SCOLOPACIDAE

White-winged Sandpiper Prosobonia leucoptera

Ellis's Sandpiper Prosobonia ellisi

Whether there was one extinct species of sandpiper occupying the Pacific islands of Tahiti and Moorea or whether there were, in fact, two remains something of an enigma. The naturalists who actually saw the birds in life and handled fresh specimens were convinced that there was only one, but more recent commentators have produced reasonable arguments that there were two: a form from Tahiti (leucoptera) with a more sandy-coloured breast; and another from Moorea (ellisi), with more reddish underparts. No definitive solution to this problem is possible.

  Although there were once several specimens in existence, only one (an example now in Leiden) survives today. This is one of the more sandy-coloured individuals. The species was only ever seen by naturalists who sailed with Cook and one of them, Georg Forster, painted a picture of a Moorea bird showing the more reddish breast. It is on the basis of this painting that the form ellisi is named.

  Clearly the birds - whether there was once species or two - became extinct at an early date. Other than the fact that Forster found it close to small brooks, nothing is known of it in life.

ALCIDAE

Great Auk Pinguinus impennis

The Great Auk is one of the true stars of extinction. Its story rises and falls like a Greek tragedy. This was a creature that had evolved perfectly to take advantage of a particular ecological niche but its very adaptation rendered it totally incapable of withstanding the depredations of man. It is probably true to say that the species was doomed from the prehistoric day when man first invented the boat. Use of boats enabled men to visit lonely islands and skerries, to round capes and headlands and reach otherwise inaccessible stretches of coast, and thus to pursue Great Auks to their final refuges.

  The weakness in the Great Auk's lifestyle lay in the fact that it had lost the power of flight, and when it waddled clumsily ashore to breed in large colonies on islands of the North Atlantic it was totally vulnerable to a predator as ruthless and inventive as man. In addition, its large size made it a natural target. For ten months of each year Great Auks lived at sea, and it was only the necessity of breeding that brought them ashore for a few brief and doom-laden weeks.

  First, Great Auks were driven back from all their most accessible breeding haunts. Then they were steadily forced back into even more remote areas. By the start of the eighteenth century they bred only on a few far-flung islands, each of which was difficult to land upon. The greatest of the breeding colonies appears to have been at Funk Island, off the coast of Newfoundland, and here the birds assembled in vast numbers during the months of May and June. Unfortunately for the auks the island lay near to the end of a long stretch of open water on the sea route from Europe to America. Hungry sailors feasted on the defenceless creatures and discovered that not only did they provide a handy food source, but also their feathers and the oil from their bodies were very useful commodities. By the late eighteenth century this vast colony had been wiped out and for all practical purposes the Great Auk bred only on a few islands off the coast of Iceland. On one of these in particular, the species seemed safe. This was an island known as the Geirfuglasker (which loosely translated means “Great Auk island”). The birds were safe here simply because the prevailing currents were so fierce and unpredictable that the island was almost impossible to land upon. While the species was wiped out elsewhere, the stronghold on the Geirfuglasker still held out. Then, catastrophe befell. A submarine volcanic explosion during the winter of 1830 caused the Geirfuglasker to sink beneath the surface of the waves never to reappear. When the few surviving Great Auks returned to their breeding rock and found it gone, they were forced to choose another site. This they did and they selected the infamous island of Eldey, a great rock that held one major disadvantage: although difficult to land upon, such a feat was by no means impossible. And land man did. In the first raid upon the island 24 birds were caught. A year or so later 13 were obtained. Each successive raid brought a dwindling haul until in June of 1844 just two individuals - a male and a female - were killed. None was ever seen again.

  The story of the killing of these supposedly “last” two birds has acquired almost legendary status and it occurs in many variants. All of these variants derive from the notes of two Victorian ornithologists, John Wolley and Alfred Newton, who recorded the tale after conducting extensive interviews with the Icelanders who actually participated in the event. Their notes show that on either 2nd or 3rd June an eight-oared fishing boat left the Icelandic mainland and made for the island of Eldey. Here, three of the crew struggled ashore and spotted two Great Auks among hundreds of smaller seabirds. They pursued the pair, caught them both and strangled them. The men returned to the boat and with great difficulty managed to scramble back into it. Once safely back on the mainland the leader of the raiding party took the road to the Icelandic capital Reykjavik, where he hoped to sell the birds. On the road it seems that he met, by chance, a trader with an interest in Great Auks - and the dead birds were sold on the spot. Curiously, the skins themselves vanished and no-one knows for certain what became of them (there is, however, some evidence to show that they might be the specimens now in Los Angeles and Brussels), although the internal organs of these two birds, preserved in spirits are now at the Zoological Museum in Copenhagen.

  After its extinction, the surviving relics of the Great Auk - in the form of eggs and stuffed birds - became highly sought after. Through the second half of the nineteenth century, and afterwards, such items changed hands for large sums of money. In 1890 a stuffed bird would have cost around £400 (the price of four or five ordinary-sized houses) and an egg only slightly less.

  Although the species has acquired a sort of cult status, surprisingly little is known of it in life. We know it was flightless, that it lived in and around the waters of the North Atlantic and must therefore have preyed on fish and other oceanic creatures. We know it laid a large pyriform egg, that it came ashore - helplessly - to breed for a few weeks each year, and that it spent the rest of its time at sea. The rest is speculation.

 
COLUMBIFORMES

The pigeons as a whole have been badly affected by extinction. The reasons for this are probably those that apply broadly in the cases of other badly affected groups, the rails and the parrots. Each of these groups shows a marked tendency to disperse to oceanic islands where the species that evolve show a terrible vulnerability to changing conditions.

  In addition to the two members of the extinct family Raphidae, and the seven species of Columbidae covered below, two other pigeon species may well be extinct but are not included here as there is a faint possibility that they may still survive. These are the Red-moustached Fruit-dove (Ptilinopus mercierii) from the Marquesas Islands in the Pacific and the Choiseul Pigeon (Microgoura meeki) of the Solomon Islands; the last definite records of these two were, respectively, some 80 and 100 years ago. Both have already been covered (see HBW 4, pages 187 and 216).

RAPHIDAE

Rodrigues Solitaire Pezophaps solitaria

  The island of Rodrigues, far out in the Indian Ocean, is one of the Mascarene group and here a dodo-like species evolved from an ancestral pigeon stock. This species has become known as the Rodrigues Solitaire.

  The Huguenot refugee François Leguat, whose name occurs so regularly in the stories of extinct birds, took a particular liking to this species and it is from his account that most information concerning it is derived.

  Leguat was marooned on Rodrigues during 1691 and stayed for around two years. He and his small band of followers became entranced by the strange bird they found there and Leguat penned a number of observations of the Solitaire. One of these constitutes the first recorded observation of territorial behaviour in birds:

All the while they are sitting...or bringing up their young one, which is not able to provide for its self in several Months, they will not suffer any other Bird of their Species to come within 200 yards round of the Place; but what is very singular, is that Males will never drive away the Females; only when he perceives one he makes a noise with his wings to call [his] Female, and she drives the unwelcome Stranger away, not leaving it till 'tis without her Bounds. The Female do's the same as the Males, whom she leaves to the Male.

  The only other early authority for the birds of Rodrigues is an anonymous author, usually thought to have been a marooned sailor by the name of Tafforet, who wrote a document that has come to be known as the Relation de l'Île Rodrigue. A translation of the part of this document that relates to the Solitaire reads:

The Solitary...weighs...40 or 50 pounds. They have a very big head, with a...frontlet, as if of black velvet. Their feathers are neither feathers nor fur...of a light grey colour, with a little black on their backs. Strutting proudly about...they preen their plumage...and keep themselves very clean...[They] run with quickness among the rocks, where a man, however agile, can hardly catch them...They do not fly at all, having no feathers to their wings but they flap them and make a great noise...I have never seen but one little one alone with them, and if anyone tried to approach it, they would bite him severely. These birds live on seeds and leaves of trees.

  Confirmation of the one-time existence of this creature comes from bones found in caves on Rodrigues. Complete skeletons have been assembled and these show that there was an extraordinary divergence in the size of the sexes with the males being much bigger. The bones also confirm some anatomical details that were mentioned by Leguat.

  The remarks in the Relation indicate that the birds were still plentiful during the 1720's, and the document is thought to date from around 1725. However, at some time soon afterwards, they seem to have declined rapidly. During 1761 the Abbé Pingré arrived on the island with the intention of viewing the Transit of Venus. He arrived in time to see the heavenly wonder but not, it seems, in time to see the Solitaire. If the species still existed at this time it was so rare that it stood at the very brink of extinction.

  Solitaires seem to have been tasty birds and it can only be assumed that settlers who arrived on Rodrigues during the eighteenth century ate them out of existence.

Dodo Raphus cucullatus

The Dodo is probably the most powerful of all icons of extinction. “Dead as a Dodo” is an expression familiar to almost everybody in the English-speaking world. The name is unforgettable and so too is the bird's appearance, an appearance made memorable through the paintings of Roelandt Savery and their many derivatives, chiefly, perhaps, John Tenniel's illustrations for Lewis Carroll's Alice's Adventures in Wonderland. On the Dodo's home island of Mauritius the image of the bird is everywhere - on crockery, T-shirts, sweets and all kinds of ornamental souvenir ware.

  More has been written about the Dodo than about any other extinct bird, but the reader should beware. Dodo literature is studded with inaccuracies and ludicrous theories, and despite the flood of words almost nothing is known of the bird in life.

  Dutch mariners arrived on the Dodo's home island of Mauritius during September of 1598. We have no way of knowing how the species was faring before this but we do know that the arrival of the Dutch meant doom for the Dodo. Within 40 years of Dutch arrival the Dodo was virtually extinct, unable to withstand the joint assaults of man and the mammals he introduced - pigs, rats and monkeys, creatures that may not have threatened the adult birds but certainly threatened the young and the eggs. The existing seventeenth century reports become very thin on the ground after 1640 and even more so do original pictures. Indeed, after 1638 all Dodo images are derivations of earlier ones. The Dodo may have been in existence for several more decades but as a species it was a spent force from around this time.

  There is no doubt that hungry sailors visiting Mauritius played a large part in the extermination of the Dodo and a very considerable proportion of the sparse contemporary literature concerns the edibility of the bird. Apparently, the flesh was rather tough and not particularly good to eat so whenever possible the men turned their attention to smaller, tastier birds. The Dodo held one big advantage for them, however. Being large and flightless, it was easy to locate and catch.

  Although it interacted with man for such a short period of time, this exotic creature quickly gained some celebrity. A few living individuals reached Europe; several ornithologists have tried to calculate the number, but the evidence is flimsy and we have no real idea of how many there were. Also, the Dutch seem to have taken live birds to India and Indonesia. It has even been suggested that Dodos were taken to Japan but there is no proper proof of this.

  The actual natural history of the Dodo is shrouded in mystery. How it used its curious beak, what it ate, even what parts of Mauritius it inhabited are all matters of debate. Almost certainly the species lived on fruits, nuts and other vegetable matter that it found on the forest floor but the records seem to indicate that it occurred only in certain areas. Woods close to the shore were probably a favoured habitat. It certainly seems unlikely that the birds inhabited the entire island. Had this been the case, the extinction could not have happened so rapidly.

  Despite the early date of disappearance, the Dodo has left a number of physical relics behind. First, there are bones. With just a few exceptions these come from a small area of swamp known as the Mare aux Songes, close to the southern coast of Mauritius. From this material several complete skeletons have been assembled and these confirm the impression (gained from seventeenth century pictures and writings) that this was a large and heavy bird. It does seem, however, that it may not have been quite so bulky and clumsy as previously supposed and probably adopted a rather upright stance.

  The most famous Dodo relics of all are a head and a foot that were once part of an entire stuffed bird. The body of this specimen was destroyed long ago but the head and foot were preserved and they are kept today at the University Museum of Zoology, Oxford.

  Many people leave museums convinced that they have seen a stuffed Dodo. They have not, for there are none in existence. What they have seen are carefully constructed models made from the feathers of other birds.

  The best description of the Dodo in life comes from the pen of an English courtier named Thomas Herbert who visited Mauritius in 1628 during a round trip to Persia:

Her body is round and fat...her visage darts forth melancholy, as sensible of Nature's injurie in framing so great a body to be guided with complementall wings, so small and impotent, that they serve only to prove her bird. The halfe of her head is naked seeming couered with a fine vaile, her bill is crooked downwards, in midst is the trill [nostril], from which part to the end tis of a light green, mixed with pale yellow tincture; her eyes are small and like to Diamonds, round and rowling, her clothing downy feathers, her train three plumes, short and inproportionable, her legs suiting to her body, her pounces sharp, her appetite strong and greedy.

  The actual date of extinction is a matter of some doubt. Although the Dodo was on its last legs as a functioning species by 1640, there is little doubt that individuals lingered for some time afterwards. The last record has traditionally been regarded as a mention made by one Benjamin Harry who was writing in 1681. Following a visit to Mauritius, Harry compiled a list of the island's products and Dodos were included in this list. It is by no means certain, however, that Harry actually saw these birds. He may merely have been compiling a list of items that he knew had come from the island. Anthony Cheke (1987), who has conducted much research on the ornithological history of the Mascarenes, believes that the last Dodo observation was made by a certain Volquard Iversen and relates to the year 1662. Iversen was marooned on Mauritius for several months and claimed to have encountered Dodos on a small offshore island to which he waded at low tide. Even this record may not stand up to scrutiny, however. Some evidence suggests that the name Dodo was transferred to the Mauritius Red Rail during the second half of the seventeenth century, and Iversen's record may perhaps result from confusion over terminology.

COLUMBIDAE

Bonin Woodpigeon Columba versicolor

The avifaunas of several islands to the south of Japan have been badly depleted, and one of these island groups, the Bonins (Ogasawara), has lost several distinct forms.

  One of these was the Bonin Pigeon, a large, dark bird that was discovered in 1827 during the exploratory voyage of HMS Blossom. It seems to have survived until around 1890, the last known specimen being collected during 1889. The bird's general unwariness and the implication that it showed no fear of man doubtless contributed to its downfall. Very little is known of its habits but it seems likely that it required well-wooded territory where it probably fed upon fruit, seeds and buds. The species is known to have inhabited Peel Island (Chichijima) and Nkondo-shima and it probably lived on other islands of the Bonin group.

  There seem to be only three specimens of the species in existence. One belongs to the Natural History Museum, London, a second is in St Petersburg and the third is in Frankfurt.

Ryukyu Woodpigeon Columba jouyi

This species, closely related to the last, occurred on the Ryukyu and Daito Islands, another of the island groups to the south of Japan. It was last recorded on the Daito Islands during 1936. Although certainly extinct on Okinawa where it was last seen in 1904, the species may possibly cling to existence on some of the smaller islands. Like its relative it appears to have required heavily forested country.

  Nothing is known of the bird in life and reasons for its disappearance are unclear.

Passenger Pigeon Ectopistes migratorius

The celebrated Passenger Pigeon has, perhaps, the most extraordinary story of any extinct bird. It may once have been the most numerous bird on Earth, and at the start of the nineteenth century vast flocks of this species blackened American skies. Yet during the course of 100 years the tremendous numbers dwindled until just a handful of birds remained. The last individual of all, a female named Martha, died in her cage at the Cincinnati Zoological Gardens at 1:00 p.m. on 1st September 1914.

  Early descriptions of the massed flights of the Passenger Pigeon are remarkable. In 1759 a certain Peter Kalm wrote: 

In the spring of 1749...there came from the north an incredible number of these pigeons to Pennsylvania and New Jersey. Their number, while in flight, extended 3 or 4 English miles in length, and more than one such mile in breadth, and they flew so closely together that the sky and the sun were obscured by them, the daylight becoming sensibly diminished by their shadows.

The big as well as the little trees...sometimes covering a distance of 7 English miles, became so filled with them that hardly a twig or a branch could be seen which they did not cover...when they alighted on the trees their weight was so heavy that not only big limbs and branches were broken straight off, but less firmly rooted trees broke down completely under the load. The ground below the trees where they had spent the night was entirely covered with their dung, which lay in great heaps.

  During the 1830's John James Audubon penned his famous account:

The air was literally filled with pigeons, the light of noonday was obscured as by an eclipse; the dung fell in spots not unlike melting flakes of snow...pigeons were still passing in undiminished numbers and continued to do so for three days in succession.

  These vast and spectacular flights rendered the Passenger Pigeon vulnerable in a quite remarkable way. So easy was it to shoot the birds that one simply had to point a rifle skywards and repeatedly pull the trigger. Hunting competitions were organized and for one of these it is known that a haul of more than 30,000 dead birds was necessary to claim a prize. With many hands turned against it there is little wonder that Passenger Pigeon numbers began to fall drastically.

  However, it is simply not possible to shoot such a numerous species out of existence. It seems that there were additional factors and that these factors centered around the species' need to operate in such vast groups. The birds moved around nomadically, looking for rich crops of beechmast, acorns or chestnuts, and when an exceptionally good site was located the birds were attracted in huge numbers. The forests were steadily depleted, however, and so good crops became fewer and farther between. To compound this, so many thousands of pigeons were shot that their scouting capacity dropped, making it even more difficult for them to locate adequate food supplies. Once numbers had fallen below a certain level, even though that level may have been incredibly high, the Passenger Pigeon was doomed as a species. Clearly this was a creature that could only survive in huge numbers. The critical figure may have been a million individuals, it may have been twenty million; we shall never know. But at some point during the mid-nineteenth century, the figure was passed and from that moment onwards the species was spiralling downwards to extinction. The decline became apparent during the 1870's. At the start of this decade the flocks seemed as healthy as ever, but by its end the ranks were noticeably thinned and by the end of the century the species was virtually gone from the wild, leaving only a few individuals in captivity. It seems that the decline was accelerated too by an outbreak of Newcastle disease, a paramyxoviral disease that attacks the digestive tract and nervous system, and is known to affect domestic poultry as well as many species of wild bird.

  The Passenger Pigeon was quite unlike most other pigeons. Its body was beautifully streamlined, with a small head and long, pointed wings and tail. This body design allowed the bird to fly with great swiftness and agility. When wheeling, dropping or rising in the air, the beautiful, yet subtly marked, plumage would flash spectacularly in the sun.

  Naturally, such enormous numbers had a marked effect on the vegetation. The birds ate acorns and nuts of all kinds as well as fruit, grain, insects and other invertebrates.

  When they nested they did so in colossal colonies and sometimes these extended for many kilometres. Nesting sites of 65 km were recorded and it is estimated that 16 km by 5 km were characteristic. The nest was flimsily made from twigs, and one white egg was laid. The peak breeding period occurred in April and May, although breeding could take place at any time between March and September. Both parents helped with the incubation, and the single chick was cared for by the adults until it was around two weeks old. Then, suddenly, the old birds would depart leaving the fat chick abandoned and crying in the nest. After a while the baby bird would drop to the ground and a day or two later it would take to the air and leave.

Liverpool Pigeon Caloenas maculata

Two hundred years ago there were two specimens of this species in existence. Now there is just one. No-one knows where it came from or when it was collected, although this was certainly at some time during the last half of the eighteenth century. The specimen is now in the collection of the Liverpool Museum, and it has been there since 1851, when it was bequeathed to the people of Liverpool by Edward Stanley the 13th Earl of Derby.

  Stanley was one of the great collectors of the nineteenth century, a period that is rightly celebrated for the collecting zeal that inspired so many rich men. Not only did he collect preserved birds and other animals, he also collected live ones, and the menagerie he put together in the grounds of his home, Knowsley Hall, became famous throughout Europe. The Earl collected hundreds of pictures and albums of pictures relating to natural history and commissioned the famous artist Edward Lear to paint portraits of the animals in his menagerie. He assembled a magnificent library and acquired two copies of what is now the world's most valuable illustrated book, Audubon's Birds of America (a copy of this work recently changed hands for £8,000,000), one of which he promptly cut up to make a scrapbook featuring pictures of “birds of the world”!

  Considering its antiquity, his specimen of the Liverpool Pigeon is in a remarkably fresh state of preservation. In appearance, the bird shows affinity to the widespread Nicobar Pigeon (Caloenas nicobarica), but notwithstanding the similarity it is clearly distinct. For reasons that are not entirely clear, the specimen has been overlooked by many commentators and there has been a tendency among those who do know of its existence to regard it as a freak. Few of these have actually seen it, however.

  It is assumed that the specimen was brought back from an expedition to the South Pacific and that the species inhabited a Pacific island - but this is just a guess. Obviously, nothing is known of the living bird.

Tanna Ground-dove Gallicolumba ferruginea

The Tanna Ground-dove is known today from just a single, rather crude, painting by Georg Forster that was produced during Captain Cook's second voyage around the world. This painting is in the Forster portfolio at the Natural History Museum, London, and in the margin the following words are inscribed:

Tanna, female, 17th August 1774.

  Nothing more is known of the species. A specimen of it did once exist although this, like the species itself, vanished long ago. Concerning the events of 17th August Georg Forster's father, Johann Reinhold, made this spare record:

I went ashore, we shot a new pigeon and got a few plants.

  The island of Tanna is one of the islands of Vanuatu, and for many years it had an ugly reputation on account of the ferocity of the native inhabitants. For this reason it was seldom visited during the nineteenth century, and at some time during that period the native pigeon seems to have passed away.

Mauritius Blue-pigeon Alectroenas nitidissima

When the first Dutch mariners landed on Mauritius, they were hungry after weeks at sea on meager rations. Naturally, they caught Dodos and ate them. Soon, however, they tired of the rather tough meat on these easily caught creatures and turned their attentions to smaller, tastier birds. Among those that they mentioned particularly were small pigeons and these seemed to provide a welcome alternative to the cloying flesh of the Dodos. A remarkable pair of illustrations drawn in Mauritius and dating from 1601 show a dead individual of the species. It is in the journal, or log, of one of the first Dutch ships to visit Mauritius and, although anonymous, the pictures are brilliantly and expressively rendered. This journal is now kept at the Rijksarchiv in the Hague and it relates to the voyage of the ship Gelderland.

  Despite the existence of the journal, almost two hundred years were to pass before the species was formally described and brought to zoological attention. In 1782 Pierre Sonnerat mentioned a specimen of the bird that he had collected in 1774, during his extensive travels to the East. However, as he did not adhere to the Linnaean principle of scientific naming, it was left to Giovanni Antonio Scopoli some four years later to give it its scientific name.

  This striking pigeon, with strange wax-like, elongated head and neck feathers, is now represented in the world's museums by just three specimens, one in Mauritius itself, one in Paris and one in Edinburgh. The living bird, with its peculiar headdress and beautiful colouring, must have been a remarkable sight, contrasting markedly with the three dingy, antiquated stuffed examples that survive. These specimens date from the last part of the eighteenth century and the first decades of the nineteenth, the most recent having been collected in 1826. Soon after this date the species seems to have become extinct. Given the early disappearance of so many Mauritian birds it is perhaps surprising that this pigeon lasted so long.

  Julien Desjardins, a specimen dealer who spent many years on Mauritius and who received the 1826 specimen, provided the only information of the bird in life. He described how it lived alone near riverbanks and fed on fruit and freshwater molluscs.

Rodrigues Pigeon Alectroenas rodericana

This species is known from bones discovered on Rodrigues during the 1870's and described by Alphonse Milne-Edwards, the celebrated expert on fossil birds at the Paris museum. Milne-Edwards described the species primarily on the evidence of a sternum, and chose to assign it to the genus Columba, whereas later researchers have shown an inclination to place it in the genus Alectroenas, alongside the extinct pigeon of Mauritius Alectroenas nitidissima. Either designation may be correct.

   This is another of those rather mysterious birds that seem to have been observed in life by the Huguenot refugee François Leguat. He wrote:

The pigeons here are somewhat less than our own [European pigeons] and all of a slate colour, fat and good. 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...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.

  Presumably this description can be correlated with the evidence of the bones. Leguat's account clearly indicates two factors that contributed to the species' extinction - tameness and the depredations of rats.

 
PSITTACIFORMES

A great many parrots are threatened with extinction or are in serious decline. Partly this is due to the vulnerability of island forms, for many parrots are inhabitants of small islands, but partly, and paradoxically, it is due to their great popularity. For centuries man has been unable to resist the temptation to interfere with populations, move them about and keep individuals as pets.

  Nine extinct species are dealt with herein. Two others have already been considered “Almost certainly Extinct” in HBW, and so are not covered here, although elsewhere they have been listed by the present author as extinct (see Fuller, 2000a). These are the Paradise Parrot (Psephotus pulcherrimus) and the Glaucous Macaw (Anodorhynchus glaucus). There is much negative evidence to show that both are extinct, but perhaps there remains a very small chance that they survive (HBW 4, pages 382 and 419).

  The names of a number of what can only be termed “hypothetical” parrot species sometimes occur on lists of the extinct. Among these are several amazons and macaws alleged to come from the West Indies that were described by nineteenth and twentieth century ornithologists on the basis of written reports made by travellers in earlier centuries. The problems with these reports are twofold. First, they are usually vague; second, parrots have been moved around so much by humans that a sighting of an unexpected parrot on one island need not signify a new species. These names cannot, therefore, be accepted as valid.

PSITTACIDAE

Norfolk Kaka Nestor productus

This is one of these forms that can be interpreted either as a subspecies or as a full species in its own right. In this particular case interpretation is a subjective commodity and each of the interpretations has its merits.

  The Kaka (Nestor meridionalis) is a relatively well-known representative of the New Zealand avifauna, widely distributed, although nowhere plentiful, on both main islands and on a number of smaller offshore islands. It is a fairly stocky parrot of medium size with striking but quite variable plumage.

  Until around the middle of the nineteenth century a closely related bird occurred on Norfolk Island and its close neighbour Phillip Island, both of which lie in the Tasman Sea almost half way between New Zealand and Australia. The plumage of the Norfolk and Phillip Island birds is somewhat different to those from New Zealand, but it seems quite likely that the two could have interbred, had they come into contact. However, the two forms have acquired a fairly separate identity in ornithological literature and if museum specimens are compared it is quite easy to tell them apart - even without reference to locality data.

  All birds on Norfolk Island were badly affected by the establishment of penal settlements for criminals transported from Britain. These were founded during the last decade or so of the eighteenth century and they were used during the first few decades of the nineteenth. There is no doubt that convicts and early settlers entirely disrupted the peace and tranquility of the island and the large, brightly coloured parrots would have made a very tempting target. Norfolk Kakas were probably caught primarily for food but there is no doubt that they were also taken as pets. The famous ornithological writer John Gould saw one in Sydney during his visit in the late 1830's and the last individual of all may well have been a bird that died in its cage in London in 1851, or soon after.

  Almost nothing is known of the bird in the wild. Apparently it nested in holes in trees and laid up to four eggs. It frequented the rocks and the tree tops, was very tame, as might be expected in an island form, and was seen feeding on blossoms. The population seems to have been prone to a strange deformity of the beak.

  As far as is known these parrots were first extirpated on Norfolk Island, and held out for a little longer on Phillip.

Raiatea Parakeet Cyanoramphus ulietanus

This is another of the species known only from the records and specimens brought back by those who sailed with Cook on his three exploratory voyages around the world.

  There is some doubt over which particular expedition encountered this bird, but it seems likely that the encounter took place when naturalists of the third voyage called at Raiatea (then called Ulietea) in the Society Islands of the South Pacific, during November 1777. At least two specimens were collected and these are now in the Natural History Museum, London and the Naturhistorisches Museum, Vienna.

  Nothing is known of the species, nor can any realistic reasons be advanced for its extinction which, presumably, took place soon after Cook's last voyage, for the birds were never encountered again. Perhaps they were unable to withstand the attacks of introduced mammals or maybe man's meddling with parrots was a factor.

Black-fronted Parakeet Cyanoramphus zealandicus

Like the Raiatea Parakeet, this was another inhabitant of the Society Islands, but as far as is known this particular species was restricted to Tahiti.

  Both species are closely related to the well known Cyanoramphus parakeets of New Zealand but they are physically separated from these by 3220 km of open sea. Probably, they had quite similar lifestyles although this cannot be confirmed as there are no records concerning the two extinct species.

  Again like the Raiatea Parakeet, this species was first brought to light by naturalists who sailed with Cook, although in this particular case it was also recorded by a few later visitors.

  A painting by Sydney Parkinson, made during Cook's first voyage, is kept at the Natural History Museum, London, and there are also a handful of specimens in existence - one in London, two in Liverpool, one in Paris and one in Perpignan.

  The last record of the species dates from 1844, but nothing is recorded of the bird in life.

Newton's Parakeet Psittacula exsul

  A greyish-blue parakeet belonging to the familiar ring-necked group once lived on the Mascarene island of Rodrigues. Curiously, it survived for much longer than most of the endemic bird species of the island. Certainly, individuals were still alive during the 1870's, although it is clear that all were gone soon after that.

  Little is known of the species but it seems that the two early chroniclers of the birds of Rodrigues, François Leguat and the anonymous author who is usually considered to be a certain Monsieur Tafforet, both saw this species and made observations of it. Certainly, both authors mention parakeets. One complication of these early reports is that Leguat mentioned seeing green as well as blue parakeets. There may have been different colour morphs of the species or the two colours might be explained by the occasional arrival of storm-blown individuals belonging to the more familiar Psittacula species. Possibly, such individuals could interbreed with the endemic birds. In addition to these complicating factors, there is skeletal evidence to show that more than one parakeet species once inhabited the island.

  Leguat and his followers saw birds - presumably Newton's Parakeets - feeding on the nuts of an olive-like tree, and they taught some of these parakeets to speak, surely an indication of just how tame such birds were. Apparently they became bi-lingual; they could speak in both French and Flemish! When Leguat and his little band of followers fled the island they took a parakeet with them on their voyage to Mauritius.

  Although the species was obviously common during the period of Leguat's visit (1691), it was rare by the nineteenth century. Just two specimens are known and both are in the collection of the University Museum of Zoology, Cambridge. Both were collected during the 1870's, the last on 14th August 1875.

  The reasons for extinction are unknown, but it is likely that the bird's original tameness and the infestation of the island by mammals were primary causes.

Mascarene Parrot Mascarinus mascarinus

Like the preceding species, just two specimens of this striking, middle-sized parrot exist. One is in Paris, the other in Vienna. Both probably date from the last years of the eighteenth century or the early years of the nineteenth. A record of an individual surviving in a European aviary until 1834 (see Hahn, 1841) is now considered to be false.

  Very little is known of the species other than the fact that its home was the Mascarene island of Reunion. Why it vanished and when is not known but it can be assumed that it became extinct during the first decades of the nineteenth century.

Broad-billed Parrot Lophopsittacus mauritianus

Pictures drawn by Dutch explorers visiting the island of Mauritius during the early seventeenth century clearly show a rather strange and aberrant parrot, and these pictures can be correlated with bones found on the island. The bones, like the pictures and a few contemporary written descriptions, indicate the former existence of a large-headed parrot.

  Apart from the fact that such a bird once existed, little can be meaningfully said of it. It certainly still lived during the early seventeenth century, but it probably became extinct not long afterwards. It may have lasted until the last decades of the seventeenth century or it may have been lost well before this.

Rodrigues Parrot Necropsittacus rodericanus

A parrot comparable in size to a large cockatoo with an outsize beak once inhabited the island of Rodrigues. An almost complete skull of this creature has been found and this relic can be matched with an account in the anonymous document - written around 1725 - known as the Relation de l'Île Rodrigue. This account describes a long-tailed parrot with a large head. The writer suggests that the species was green in colour and lived on islets to the south of the main island. Probably it had been penned back to these refuges on account of the incursions of rats. To find water the parrots had to fly across to the main island of Rodrigues itself, surely an indication that the islets would not have been the birds' first and original choice as a home. Individuals were seen eating small black seeds from a tree that smelled of lemons.

  Probably the species became extinct during the late eighteenth century.

Cuban Macaw Ara tricolor

This distinctive and beautiful macaw, around a third smaller in size than its largest relatives, was an inhabitant of the island of Cuba.

  The most recent record of a living bird comes from La Vega, close to the Zapata Swamp, where an individual was killed during 1864. It is thought that a few birds survived for a decade or two after this date, but there is no actual proof of this. Presumably, the species' decline was due to human interference: the taking of birds for food or their capture as pets.

  Although it has a very distinctive identity, almost nothing is known of the species in life. It is assumed that its habits were similar to those of its more familiar, extant relatives.

  Attempts have been made to create artificial Cuban Macaws by selectively breeding their near relatives. Some parrot fanciers have claimed that a degree of success has been achieved but the word of many parrot breeders is notoriously unreliable - and in any case such created creatures would have little, if any, real connection with the actual species.

Carolina Parakeet Conuropsis carolinensis

The Carolina Parakeet, like its compatriot the Passenger Pigeon, declined from vast numbers to just a few individuals within the period of a century. This was once considered a pest species, with communal feeding habits, that ranged across the southern and eastern USA. It ruined orchards, wrecked corn fields, destroyed grain stocks and thereby aroused the anger of man. The species seems to have had few defensive skills. When individuals were shot, their companions would fly squawking above the dead or wounded and eventually they would settle among their downed fellows. Naturally, they then made easy targets themselves.

  A parrot having the capability of inhabiting the USA is in itself something of a peculiarity, and there are surprising records of this species flying over snow-covered fields. In the years of its abundance the species liked forested lowland and showed a preference for land close to water. It lived mostly among buttonwood, cypress or sycamore and roosted in hollow stumps into which individuals would crowd and cluster together. At dawn the birds would flock to the top branches of the trees and then they stayed quiet for much of the day. Late afternoon and early evening would see great bursts of activity. When the birds decided to eat they would fly swiftly down to the chosen feeding area and arrive in a blaze of colour.

  As far as breeding is concerned, there are conflicting descriptions. Some records tell of several parakeets laying their eggs together in tree holes, while others indicate that fragile nests were made from twigs and that these were sited in the forks of branches.

  Two races of the species are commonly recognized, the nominate form and a western subspecies named ludovicianus. These two races are rather poorly distinguished, however.

  Through much of the nineteenth century the Carolina Parakeet was a particularly common bird. Even as late as the 1880's it could be found in some numbers. Yet soon after this the species could hardly be found in the wild at all.

  As the century drew to its close, a few individuals still survived in captivity, most particularly a group in the same Cincinnati Zoo that provided a home to the last Passenger Pigeon. The very last Carolina Parakeets were a pair named Lady Jane and Incas, and by 1917 these two birds had been cage-mates for something like 32 years. Then, Lady Jane died, leaving Incas the sole representative of the species. He survived, alone, for just a few months until February 1918, when he died in his cage surrounded by his keepers. They were in unanimous agreement: their bird had died of grief. His little body was frozen in a block of ice and sent for preservation to the Smithsonian Institution in Washington but, curiously, it never arrived or, if it did, it was stolen. Perhaps this hardly matters; there are several hundred specimens of this species in the world's museums.

  There are several alleged late records dating from the 1920's and 1930's that supposedly relate to Carolina Parakeets still surviving in the wild, but these are probably bogus.

 
CUCULIFORMES

Delalande's or the Snail-eating Coua (Coua delalandei), a large, ground-dwelling cuckoo-like bird from Madagascar is almost certainly extinct, and has already been covered thus in HBW (see HBW 4, page 581). It has not been definitely recorded since 1834 but partly because rumours of its continuing existence circulated during the 1920's, there has been reluctance in some quarters to add the species to the ranks of the extinct.

STRIGIFORMES

The Laughing Owl (Sceloglaux albifacies) of New Zealand is usually considered to be extinct but for the purposes of HBW the optimistic view has been taken that there is still a remote chance of its survival (see HBW 5, page 239). It has not been reliably recorded since the early decades of the twentieth century, however, and more recent reports are probably false.

  It seems clear that at least two now-extinct owl species occupied the Mascarenes during recent historical times, but only one of these has left definitive specimen material behind, and is discussed below. The other has come to be known as Otus commersoni. A fairly detailed description of this creature was made by the naturalist/dealer Julien Desjardins, who saw a dead specimen during 1836, and there is a drawing of what seems to be the same species made by one Philippe Sanguin de Jossigny around 1770. This was a largish owl with ear tufts that inhabited the island of Mauritius. Since there is no remaining specimen material of any kind, it is impossible to assign this form to any particular owl group. Other claimed owl species from the Mascarenes were named as Bubo leguati, Strix sauzieri and Strix newtoni, but these forms must be considered even more hypothetical.

STRIGIDAE

Rodrigues Little Owl Athene murivora

The anonymous author of the Relation de l'Île Rodrigue (c. 1725) mentions a small owl that preyed on little birds and lizards and could be heard calling in fine weather.

  This brief account is often correlated with some fairly fragmentary skeletal material that was found on the island around a century and a half later. This correlation is, of course, a little tenuous but it is by no means unreasonable.

  The skeletal remains were assigned to the genus Athene by the celebrated French comparative anatomist Alphonse Milne-Edwards who, during the last decades of the nineteenth century was the leading expert in fossil and sub-fossil birds. Unfortunately, his suggestion that the remains could be assigned to Athene may not be correct. First, the remains by no means represent a complete specimen. Second, the Athene species with which Milne-Edwards compared the Rodrigues material has more recently been reassigned to the genus Ninox. This highlights the difficulty of making judgements when assessing incomplete skeletal material. What the evidence of the bones does show is that this species had legs of particular strength and length for an owl of this rather small stature. It can be deduced, therefore, that it was more terrestrial than most owls.

  The reasons for its extinction are unknown, as too is the time of this event.

 
CAPRIMULGIFORMES

The Jamaican Poorwill (Siphonorhis americana), not definitely recorded since the 1860's, is often treated as extinct, but the secretive nature of nightjars, in conjunction with recent observations of “unidentifiable” nightjars in remoter parts of the island, give some cause for hope to persist (see HBW 5, page 344). Optimists take heart from the recent rediscovery of the New Caledonian Owlet-nightjar (Aegotheles savesi), not definitely recorded since 1880 (or perhaps 1915) until rediscovered in 1998.

APODIFORMES

Although the hummingbirds (Trochilidae) make up a rather homogeneous family, their classification is a very complex matter. Just a small part of the problem is that there are a great many taxa described from single or few specimens of doubtful provenance, a number of which have subsequently been reclassified as hybrids or aberrant individuals. One of the problems with describing a species from just a single individual is that it can be so easy to misunderstand the true nature of that example.

  One such case is that of a form recently described as the Bogota Sunangel (Heliangelus zusii), on the basis of a specimen that was purchased in Bogotá, Colombia, in 1909 and is now in Philadelphia. Many hummingbird specialists are not inclined to accept it as a valid species, reckoning it probably to be a hybrid, though there is much dispute as to its claimed parentage! Another poorly known form sometimes listed as a threatened species, but recently reported to be probably of hybrid origin, is the Tachira Emerald (Amazilia (Polyerata) distans). In such cases, taxonomic difficulties override the debate as to whether or not such forms are extinct. One apparently extinct form has, however, received somewhat more widespread acceptance as a valid species.

TROCHILIDAE

Brace's Emerald Chlorostilbon bracei

Although there is indeed fairly general acceptance of Brace's Emerald as a valid species, the same doubts hang about it as in the aforementioned cases, and the present author considers it to be a rather poorly established species. However, a number of hummingbird experts have spoken in its favour and in line with the designations of these specialists it is included here.

  In fact, there is very little that can be meaningfully said of it (see HBW 5, page 533). The single known specimen was collected in July 1877 and was apparently found on New Providence in the Bahamas. No such bird had ever been seen there before, and there is a reasonable possibility that it may have been a wind-blown vagrant. There has been an attempt to correlate fossil remains found on the island with the preserved skin, but any attempt to make proper sense of tiny hummingbird bones amongst the mass of valid taxa described is really perhaps asking too much.

 
CORACIIFORMES

The large and diverse order Coraciiformes includes the kingfishers (Alcedinidae), and in this family is a very poorly known form that may constitute an extinct species. This is a bird rather similar to the Micronesian Kingfisher (Todiramphus cinnamominus) and known as the Ryukyu Kingfisher (Todiramphus (c.) miyakoensis) after the islands to the south of Japan that it presumably inhabited. Unfortunately, it is one of those creatures known only from a single specimen, taken in 1887, and this extreme limitation makes interpretation difficult, especially as the bill colour is not known. It is probably either an extinct species or an extinct subspecies; either interpretation has its merits. However, there is also some doubt as to whether it might have been brought to the islands by chance vagrancy or through human intervention, or indeed that it might be a mislabelled specimen, so any systematist is in a difficult position in trying to decide. An interesting, if rather sad, related case is that of the nominate race of the Micronesian Kingfisher, endemic to the island of Guam: it is now probably extinct in the wild there, although it still survives in captivity.

PICIFORMES

There are two species in this order that are probably extinct, although optimists still express hope for their survival. These are the Imperial Woodpecker (Campephilus imperialis) and its celebrated relative the Ivory-billed Woodpecker (Campephilus principalis). Both are dealt with elsewhere in this volume (see page 534).

PASSERIFORMES

  As might be expected, an order as vast as the Passeriformes contains quite a number of extinct species. In all, 26 are covered herein. In addition there are two species that are often regarded as extinct and that in all probability are. These are the Bush Wren (Xenicus longipes) and the Aldabra Brush-warbler (Nesillas aldabrana). Despite relatively recent records of each, in 1972 and 1983 respectively, there is only a remote chance that either survives. Both species will be fully covered in HBW in due course.

  Several other species hover around the very brink of extinction. Gurney's Pitta (Pitta gurneyi) is one such species and, despite great conservation efforts over the last 15 years, it could be gone by the time this volume is published. So too could the Po'o'uli (Melamprosops phaeosoma), a species that at the time of writing seems to have only three surviving representatives, all of which are males! Another Hawaiian bird for which there is very little hope is the Molokai Creeper (Paroreomyza flammea), last recorded in 1963 and probably now extinct.

  On the brighter side, the Four-coloured Flowerpecker (Dicaeum quadricolor), once confidently considered extinct, was recently rediscovered, but its hold on survival is very tenuous and it could vanish at any time. Similarly, the Sao Tome Grosbeak (Neospiza concolor), not recorded since 1888 until rediscovered in 1991, is presumed to survive with only a tiny population that urgently needs safeguarding.

ACANTHISITTIDAE

Stephens Wren Xenicus lyalli

Although so tiny in size the Stephens Wren was an altogether remarkable creature. It may have had the smallest natural range of any known bird. It may have been the only flightless passerine. It may have been the only creature discovered and then exterminated by a single animal - a lighthouse keeper's cat.

  During 1894 this single domesticated predator brought to its owner a series of tiny corpses. These grisly little events occurred on the small island of Stephens lying in the Cook Straits, the channel that separates New Zealand's North Island from the South. The lighthouse keeper in question - a Mr Lyall, after whom the species is scientifically named - was something of an amateur ornithologist. He preserved the specimens and, realizing that the birds might be rather unusual, passed them on to a dealer. Soon the majority of them were shipped to Europe where most were bought by Walter Rothschild, the celebrated natural-history collector, who was then busy assembling his wonderful museum at Tring in Hertfordshire, England. Thus, Xenicus lyalli became known to science. By the time its existence was broadcast to the world, via the ornithological journal Ibis, the species was already extinct. The cat had stopped bringing in dead specimens and the birds were never seen again. It seems likely that forest clearance for the construction of the lighthouse, in 1894, may also have made a significant contribution to the bird's demise.

  The only human observation of Stephens Wrens was made by the lighthouse keeper himself. He saw the birds twice, both times in the evening. Disturbed from holes among the rocks, they ran fast in the dusk, like mice. They never tried to take to the air and this suggestion of flightlessness is borne out by the poorly developed wings which indicate weak flight at best.

  Skeletal remains of what seems to have been a flightless wren have been found on the New Zealand mainland. Some writers believe that this proves that Stephens Wrens were once widespread in New Zealand and that the birds discovered during 1894 were simply a relict population, but it seems far more likely that the skeletal remains come from a similar but quite separate creature. There are a number of objections to the case for the Stephens Wren being associated with the mainland skeletal remains. First, what material were the bones compared with? Second, how did a wren that was flightless, or almost so, manage to reach Stephens Island?

  Sir Walter Buller (1905), the great chronicler of New Zealand birds, quoted from a correspondent of The Canterbury Press who had written the following:

And we certainly think that it would be as well if the Marine Department, in sending lighthouse keepers to isolated islands where interesting specimens of native birds are known or believed to exist, were to see that they are not allowed to take any cats with them, even if mouse-traps have to be furnished at the cost of the state.

TURDIDAE

Bay Thrush Turdus ulietensis

This is another of those species known today only from a painting by Georg Forster. This painting, now in the Forster portfolio at the Natural History Museum, London, was produced on 1st June 1774 at Raiatea in the South Pacific, and according to an inscription it shows a female.

  Although no actual specimen is in existence, there was one once, and presumably this was the bird from which the picture was painted. The ornithological writer John Latham saw it in the collection of Sir Joseph Banks and wrote a description of it for his book General Synopsis of Birds (1781-1785), giving it the name “Bay Thrush”. Latham was a pioneering ornithologist, but headstrong, and in his early works he simply gave common English names to the birds he described, deliberately ignoring the fast-gaining Linnaean system of scientific naming. By the time he realized his mistake, he was too late: the lapse had left the field open for Johann Friedrich Gmelin, who in the meantime had produced his own book Systema Naturae (1788-1789), just a few years after Latham's. Gmelin attached scientific names to many of the birds that Latham had originally described and, because of this, it is Gmelin's name that is associated with so many of Latham's original descriptions.

  Naturally, a species represented in such a flimsy manner as is the Bay Thrush is bound to be surrounded by mystery - and this the Bay Thrush most certainly is. Most importantly, it is by no means sure that the species was actually a thrush. Georg Forster and his father Johann Reinhold were efficient naturalists who sailed with Cook, but when they saw this creature they were rather uncertain of its affinities and described it as a thrush with some degree of reluctance. A particularly un-thrushlike feature was a notch in the bill.

  Whatever its precise relationships, this bird seems to have been a valid and distinct species. The reasons for its extinction are entirely unknown but this event must have happened soon after the visit of Cook and the Forsters for no other naturalist ever mentioned seeing the species.

  The Forsters themselves gave a few fleeting details about the bird in life. They said it had a soft, fluting voice and lived among thickets in valleys. Apart from these bare facts, nothing is known of the species.

Grand Cayman Thrush Turdus ravidus

The Grand Cayman Thrush became extinct towards the middle of the twentieth century, but considering the comparative lateness of this date very little is known of it. One of the few things on record is a description of its song, which was apparently rather weak and hesitant, more, perhaps, a subdued warbling than a melodic triumph.

  As the name indicates, the species came only from the island of Grand Cayman, in the West Indies, where it inhabited dense woodland. It was closely related to a widespread West Indian species, the Red-legged Thrush (Turdus plumbeus).

  The Grand Cayman birds were large and beautiful grey thrushes that first came to public attention during 1886, when C. B. Cory described the species. He found it to be common on the island, but its decline was to be rapid. By the outbreak of World War I, it was rare and could only be found in remote areas of woodland. No specimens were taken after this time and the last recorded sighting occurred before the start of World War II. Destruction of habitat was probably a key factor in the species' disappearance, as it seemed to need areas of dense woodland.

   The last stronghold appears to have been at the eastern end of the island. Severe hurricanes during the years 1932 and 1944 are thought to have played some role in the final destruction of the species.

Kittlitz's Thrush Zoothera terrestris

The name of Baron Friedrich Heinrich von Kittlitz is associated with the stories of several extinct birds. During the late 1820's he explored a number of Pacific islands during a round-the-world expedition on a Russian corvette called the Senjawin.

  In June 1829 Kittlitz landed on Peel Island (Chichi-jima), the largest island in the Bonin group (Ogasawara-shoto). Kittlitz was one of only a very few Europeans to see the Bonins in something approaching their pristine state. Within a few years of his visit, the original ecosystem had been shattered. Just two years after he left, a group of American, British and Polynesian colonists arrived and settled. Then whalers began to call and started to use the Bonins as a suitable place to replenish their stores and careen their vessels. Rats arrived and the devastation of the endemic birdlife was horrendous.

  Kittlitz made his landing place at a spot called Port Lloyd that had been visited and named just a year before by a British expedition under the command of a Captain Beechey in HMS Blossom. Surprisingly, Kittlitz found a seemingly common bird that the British had entirely overlooked, the handsome ground thrush that now bears his name. During his short stay, Kittlitz saw this bird quite often but no later naturalist ever did. He sailed away, apparently taking four specimens with him, and the bird was never seen again. His specimens are today divided between the museums of Frankfurt, St Petersburg, Leiden and Vienna.

  Whether Kittlitz's Thrush occupied other islands in the Bonin group is not known.

ZOSTEROPIDAE

Robust White-eye Zosterops strenuus

So impressed was he by the comparatively large size of this species, for a white-eye, that the famous nineteenth century ornithologist and writer John Gould felt inclined to give this bird the common name of “Robust Zosterops”, and this same sensation is perhaps recalled somewhat in the scientific name that he gave it. Gould had developed a considerable interest in the avifauna of Australia and published his celebrated book The Birds of Australia between the years 1840 and 1848. It was later, while he was undertaking research for a Supplement to this work, that he became aware of this particular species.

  The species was endemic to the island of Lord Howe, a tiny piece of land, 11 km long by 1·6 km wide, that lies far out in the Tasman Sea between Australia and New Zealand. During the mid-nineteenth century, when the bird was first identified, it was found to be common, and the population stayed healthy for the next 60 years or so. Then in 1918, an incident occurred that proved fatal for many of the birds of Lord Howe. A ship, the SS Makambo, was accidentally grounded at a site known as Ned's Beach. Black rats (Rattus rattus) left the stricken vessel and poured ashore - and the consequences were disasterous. In a very short space of time the white-eyes were gone. A search made just ten years later could find no trace of them.

  There are a few records concerning the habits of the species. When it was common it was regarded as something of a pest. It was destructive to fruit and crops, and was even said to suck the eggs of other bird species! Its own eggs were laid during November and December. They were blue and, typically, the clutch consisted of two or three. The nest was cup-shaped and made from rootlets and grasses with a soft lining of disintegrated leaves and other suitable materials.

  Curiously, another kind of white-eye lived on tiny Lord Howe. This was a race of the widespread Silvereye (Zosterops lateralis), and to distinguish the larger birds belonging to the species Z. strenuus local people gave them the name “big grinnels”.

MELIPHAGIDAE

Kioea Chaetoptila angustipluma

In terms of extinct birds the Hawaiian Islands are one of the world's black spots. The losses at species and subspecies level have been enormous and there is no sign that this process is letting up. There are several species and races that are unlikely to see out the next decade, such as the Po'o'uli, mentioned above.

  Many Hawaiian species are known only from skeletal material, and it is not known when such creatures died out. Perhaps it was comparatively recently, perhaps it was some way back in time. Other species are known just from small series of specimens taken by ornithological collectors during the last decade of the nineteenth century, at which time there was a great spate of scientific interest in the Hawaiian Islands. What can be said with certainty is that the coming of man entirely shattered the ecosystems of the islands. It is clear that the Polynesian arrival had a devastating effect. Although the coming of Europeans, centuries later, probably increased the pace of extinction, the real damage was already done, and the endemic avifauna consisted largely of scattered, fast-diminishing populations.

  One such badly affected species was the Kioea, a large and handsome honeyeater. The species was first noticed during the visit to Hawaii of Charles Pickering and Titian Ramsay Peale as part of The United States Exploring Expedition of 1838-1842. At the time Peale wrote of it:

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 that are in flower.

  Clearly Pickering and Peale experienced no great difficulty in finding individuals, but the birds were rapidly becoming rarer. In the period leading up to 1859, during which year the last known specimen was taken, a local shopkeeper with an interest in natural history, J. Mills, killed two or three individuals on wooded slopes below the crater of the volcano Kilauea. After Mills's last record, Kioeas were never seen again. Clearly, and notwithstanding the fact that Pickering and Peale located birds reasonably easily, the species was approaching extinction even as it was being discovered.

  Four specimens survive today and there may never have been more. Of these, one is in Honolulu, one in New York, one in Washington and one in Cambridge, England.

  Apart from the brief account given by Peale, there is almost nothing on record concerning the species' habits. Although Peale described the birds as inclined to be musical, the call was described as a loud “chuck”, but this is the sum of additional knowledge.

  The species is said to have been an inhabitant of the high plateau between the mountains and the edge of the forest but whether or not this kind of area was representative of the original range cannot be said. Although the Kioea was only ever located on the island of Hawaii, fossil evidence shows that the species was also once present on Oahu and Maui, and it was also claimed to have occurred on Molokai.

Hawaii 'O'o Moho nobilis

The ancient kings and princes of Hawaii chose this unfortunate creature to be their “royal” bird. As is so often the case when monarchs choose, being the “chosen” one does not necessarily confer safety - and it definitely did not in this case. The honour merely meant that 'O'os were expected to provide plumes for the famous robes, capes and helmets that are today so prized by ethnologists, but which were once an integral part of the whole culture of pre-European Hawaii. The downfall of the 'O'o lay in the beautiful tufts of yellow feathers that grew below the wings and on the lower abdomen. These were ripped from the living bird and then woven into a bed of coarse netting; gradually, after the unwilling input of many hundreds of individuals, the cloak began to take shape.

  After the feathers were stolen from the bird, those who had trapped it were supposed to let it go. Whether or not they did is something of an open question. 'O'os fried in their own fat were, apparently, a great delicacy and it is hard to imagine hungry Hawaiians passing up the chance of a tasty morsel, particularly as any chance of being caught defying the law would presumably be remote. Even if the plumes were carefully removed and the bird released there is no guarantee that it would be capable of surviving, for the shock alone might be sufficient to cause death, but perhaps it could.

  The species was first scientifically discovered by naturalists who sailed with Cook on his third voyage around the world. Cook himself did not survive this expedition and was killed at Kealakakua Bay, Hawaii, during February 1779. The voyage itself was completed, however, and when the ships eventually arrived in England specimens of Moho nobilis were among the treasures brought back. John Latham in his General Synopsis of Birds (1781-1785) described the “Yellow-tufted Bee-eater” from these specimens.

  Although the species must once have been common on Hawaii, by the time of the coming of Europeans it was in decline and no European observer described it as anything but shy and wary. No wonder! It was seen to take nectar from flowers but it also tucked into bananas, as well as insects and their larvae. In flight it showed a dipping action and the wings moved so rapidly that they made a continuous buzzing sound. These birds frequented the topmost branches of the trees and, perhaps as a result of this preference, the nest and eggs were never found, although an experienced ornithologist, R. C. L. Perkins, did see young individuals that were recently out of the nest.

  The call was harsh and clear, and from it the name 'O'o is derived. Since there is often confusion, it might be added that the pronunciation of this name should be in the form of two o's separated by a glottal stop, “oh-oh”, rather than “oooww”.

  The records indicate that the species was once widespread over the whole island of Hawaii, as it was evidently able to live at quite high altitudes.

  Hunting for plumes obviously depleted numbers, but other factors probably also played a part in bringing about its extinction. Perhaps some kind of avian disease carried to Hawaii by an introduced species was partly responsible or maybe it was just the general breakdown of the ecosystem that was responsible for the species' disappearance.

  The last genuine report of the Hawaii 'O'o probably concerns an individual heard singing on the slopes of the volcano Mauna Loa around the year 1934.

Oahu 'O'o Moho apicalis

The main islands of Hawaii each had their own distinctive species of 'O'o. All are closely related but their respective island isolations led to certain clear differences.

  The Oahu 'O'o was distinguished chiefly by its strikingly marked black and white tail. Like its relative on Hawaii it sported yellow flank plumes and undertail-coverts.

  This was probably the first of the 'o'os to become extinct. A certain Herr Deppe, about whom virtually nothing is known, collected a series of specimens during 1837 from the hills behind Honolulu. Never again was the species encountered or, if it was, no-one ever mentioned it. Specimens do exist for which there is no provenance noted, so it is possible that a later collector found the species. Three that are known to have been taken by Deppe are in the museums of Berlin, Vienna and New York, but in addition to these there is a second specimen in New York, two examples in London, one in Paris, and one in Cambridge, Massachusetts.

  Nothing is known of the living bird.

Molokai 'O'o Moho bishopi

The 'o'o that inhabited the island of Molokai was discovered rather later than the others but it was known to science as an extant species for only a very short period. Towards the end of the nineteenth century there was a sudden surge of interest in the birds of Hawaii, and several teams of collectors ranged over the islands in search of specimens. It is a curious fact that a number of Hawaiian birds are known solely from this period. Clearly the species in question were all at the point of extinction when they were discovered, and were it not for the collecting zeal that inspired these Hawaiian expeditions during the 1880's and 1890's, their existence and passing would probably have gone entirely unnoticed by the world of ornithology. Two beautifully illustrated books resulted from the collections made at this time, Aves Hawaiiensis (1890-1899) by S. B. Wilson and A. H. Evans and The Avifauna of Laysan and the Neighbouring Islands (1893-1900) by Walter Rothschild, and it is from the words and pictures contained in these books that almost all knowledge of certain lost species is derived.

  The Molokai 'O'o provides just such a case. Described in 1893 by Walter Rothschild from specimens procured by men in his pay, it was gone by the early years of the twentieth century, the last definite record coming from the year 1904. A few later sightings were claimed but these may or may not be reliable.

  Curiously, and remarkably, the species was reported from the nearby island of Maui during 1981 but this claimed sighting is almost certainly a misidentification. Nevertheless, there is some evidence to show that the species did once inhabit Maui. A certain H. W. Henshaw, who was a very reliable observer, reported seeing an individual during June, 1901.

  Virtually nothing is known of the species in life. It differed from the two previous species by showing bright yellow cheek tufts and by carrying a tail with much less white on it.

Kauai 'O'o Moho braccatus

The fourth species 'o'o clung to survival for much longer than the others. It was also the most divergent of the four. Whereas the other three are very similar looking creatures that show their close affinity quite clearly, the Kauai 'O'o was comparatively aberrant. By no means so flashy in appearance, it was an altogether more sober-looking bird. Clearly, it was taking a rather different evolutionary path on its home island which is farther west and a little more remote than the other 'o'o islands.

  It was probably this very remoteness that enabled the species to survive longer than its relatives, and, although it was extremely rare from the beginning of the twentieth century, a few individuals held out until the 1980's. Their stronghold was a wild place known as the Alaka'i Swamp, a wet montane plateau broken up by ravines.

  This area acquired something of a worldwide celebrity as a natural refuge for endangered birds and several rare species and races clung to existence here and nowhere else. Although it provided a sanctuary of sorts for many years, the Alaka'i Swamp was not in itself enough to save the Kauai 'O'o. The small colony that had established itself there dwindled slowly until by 1981 it seemed that just a pair survived. The female of this pair disappeared during 1983 as a result of the devastation caused by Hurricane Iwa. The male continued to be seen until 1985. A rather evocative photograph of this individual exists. It was taken by Hawaiian bird expert H. Douglas Pratt and shows the bird sheltering forlornly in the branches of a tree. The flute-like call of this bird may have been heard during 1987 but the bird itself was never seen again.

  Although the Kauai 'O'o managed to linger on until comparatively recently, its main decline was remarkably rapid. George Munro, a man who spent much of a very long life studying Hawaiian birds, recorded that in 1891 the species was common and occurred from sea-level right up to the mountain tops. In 1899 he left the island and did not return for 29 years, but when he did finally go back, he failed to see or hear a single individual.

CALLAEATIDAE

Huia Heteralocha acutirostris

Perhaps the most celebrated of extinct passerines is the Huia (pronounced hoo-ee-ah). This strange, funereal-looking creature fascinated all of those who came into contact with it. First, it caught the imagination of the Maoris, who accorded it a special place in the natural order of things. Among the great treasures of ethnology are items known as waka-huias. These are intricately carved wooden boxes that the Maoris made expressly to hold the black tail feathers with white tips that characterized the Huia. To this war-like people the feathers themselves held an almost magical significance. They were worn in battle, given as tokens of friendship or respect, and mourned over in times of grief.

  After the European invasion of New Zealand, the birdlife came under intense scrutiny and no species fascinated ornithologists more than the Huia. This was probably largely due to a unique peculiarity of the Huia's anatomy. The beak of the male differs markedly from that of the female. At first sight this might not seem of any particular importance but no other known bird species shows a comparable difference: although plumages may vary vastly between the sexes, beaks are always fairly similar. In the case of the Huia, however, the beak of the male is powerful and rather crow-like whereas that of the female is long, slender and down-curved. This difference was so unexpected that when specimens first reached ornithologists they were thought to constitute two separate but related species, the female being given the name acutirostris and the male crassirostris.

  What was the reason for this divergence? The answer to this question has to be that pairs co-operated in feeding. If two birds could tap different food sources then the territory that they needed to occupy would be smaller. Perhaps the great wonder is why such a useful-seeming adaptation is not commoner. It could, of course, be argued that the very fact of the Huia's extinction indicates that the strategy does not really work.

  The male Huia could use its chunky beak to break up rotting branches or trunks, leaving the female free to come along behind and use her long, slender bill to probe into the distressed material for insects and their larvae. A favourite food item was, apparently, the huhu, a large plump grub of the nocturnal beetle Prionoplus reticularis; another was the weta (Hemideina megacephala).

  As might be expected in pairs of birds that are so dependant on each other, extreme distress was shown by any individual that lost its mate. There are several accounts that detail the behaviour of birds whose mate had been shot. Instead of flying off at the sound of the gunfire, the survivor would stay and search for its downed fellow. Doubtless, this kind of unwariness was one of the prime reasons for the species' extinction. Probably it fell prey to four-legged mammalian predators as easily as it fell prey to man.

  The range of the Huia was always rather restricted. It seems to have occurred only on New Zealand's North Island - although there are one or two indications that it may have been present at the very north of the South Island. Even on the North Island only certain areas seem to have suited it. The famous chronicler of New Zealand birds Sir Walter Buller commented on the fact that in winter Huias needed to descend from any mountainous territory occupied, in order to avoid the extremes of cold. Probably it was this liking for warmth that prevented Huias from successfully inhabiting the rather cooler South Island.

  Largely thanks to the works of Buller, a body of information exists on the lifestyle of this bird. The call was as the name; Huia is simply a phonetic rendering of it. It was described as soft and fluting and altogether haunting. The species laid 2-4 greyish eggs that were marked with brown and purplish spots and blotches, and breeding occurred during the Southern Hemisphere summer, chiefly around the month of November.

  There is no doubt that the species' general demeanour rendered it ripe for extinction, but other factors probably played a part. Chief among these is the fact that large areas of the North Island became cultivated or were given over to grazing, so the woodland that formed the Huia's natural habitat became ever more fragmented. The last fully accepted sighting of living Huias dates from the year 1907, but it is almost certain that a few birds lingered on after this date.

  The best description of the bird in life comes from the pen of Buller (1887-88):

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 depths 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.

TURNAGRIDAE

Piopio Turnagra capensis

The Piopio, or New Zealand Thrush, may have been two species rather than one. Two quite distinct kinds existed, and they are usually regarded as races of the same species: nominate capensis of the South Island; and race tanagra of the North Island. There are good reasons for supposing that they should be treated as separate species. First, the plumages are very different. Second, the structure of the beak is not quite the same, being considerably heavier and stouter in North Island birds. Third, each had a clearly definable and isolated geographical range, so they had no scope for interbreeding. It is perhaps the extreme rarity of specimens from the North Island that has prevented this idea from being investigated further. A third suggested race, minor, identified from specimens taken from tiny Stephens Island, a short distance offshore from the South Island in the Cook Straits, is only doubtfully distinct from the South Island form.

  Just as the question of species is somewhat doubtful, so too is the bird's place in any systematic list. There is no general agreement as to where this creature's relationships lie and it has been associated with various passerine groups. Certainly it is not a thrush. This term was used simply as a comforter for early settlers in New Zealand who were anxious to see in the new land anything that could remind them of their old homes. Thus, there are New Zealand robins, New Zealand crows, New Zealand wrens and New Zealand thrushes, none of which have any real connection with the species that inhabited the “old” country, but which simply bore a vague, superficial resemblance to them. Systematists have aligned the Piopio (a Maori word that is derived from the bird's call) with the whistlers and the bowerbirds. The bowerbird hypothesis is, perhaps, the more likely, but the Piopio could easily belong close to another group altogether.

  In the early days of European settlement Piopios were common birds, at least on South Island. The species was able to occupy many differing types of environment from sea-level up to the higher alpine country; the preference was for wooded country close to water. Its fate was almost certainly sealed by the fact that it was so tame and confiding. Individuals would hop around doors and windows in the hope of picking up scraps of food, or drop unwarily onto the forest floor. This left them horribly vulnerable to the attacks of dogs and cats, and the species steadily succumbed. Although much later dates are sometimes given, it seems that the last reliable record of the North Island Piopio comes from the year 1902. A few South Island birds survived for rather longer but by the early years of the twentieth century this form too was realistically finished. The most recent record which has any claim to credibility comes from Lake Hauroko in 1949.

  Piopios lived on a fairly wide variety of different kinds of food. They would eat insects, worms, fruit, buds, seeds, leaves and any kitchen scraps that they could get hold of. A cup-shaped nest was built in the trees from twigs and mosses, with a lining of soft grasses or down. Two white eggs, spotted and blotched with brown, were laid in December.

  The musicality of the species was mentioned by Buller (1887-88). Describing a captive individual, he wrote:

It was when I obtained a caged Piopio that I first became acquainted with its superior vocal powers...He often astonished me with the power and variety of his notes. Commencing sometimes with the loud strains of the Thrush, he would suddenly change his song to a low flute-note of exquisite sweetness; and then abruptly stopping, would give vent to a loud rasping cry, as if mimicking a pair of Australian Magpies confined in the same aviary. During the early morning he emitted at intervals a short flute-note, and when alarmed or startled uttered a sharp repeated whistle.

STURNIDAE

Kosrae Starling Aplonis corvina

Three starlings of the genus Aplonis have vanished. The species from Kosrae Island, one of the Caroline group in the Pacific, is known from just a series of specimens collected by F. H. von Kittlitz in December 1827. Five skins exist today, three in St Petersburg and two in Leiden; there may be another in Frankfurt but it has not proved possible to verify this rumour. Kittlitz took both adult and immature individuals and expressed the opinion that he would not have believed these to belong to the same species had he not shot an intermediate example.

  His observations reveal that the species lived on small mammals, lizards and insects. Its call was loud, and individuals were solitary in their habits. He recorded that it seemed rare at the time of his visit, and doubtless it was well on the way to extinction, as no later naturalist ever managed to locate it.

  Kosrae had become a popular resort for visiting whalers, who used the island to repair their vessels and generally recuperate after their exertions. Rats were introduced and these animals quickly overran the island. They had a greatly detrimental effect on the bird populations, and doubtless their presence was a major factor in the starling's extinction.

Mysterious Starling Aplonis mavornata

This species is known from just a single skin in the collection of the Natural History Museum, London. For many years this skin was a complete mystery and its origin was entirely unknown, but thanks to some comprehensive detective work conducted by Storrs Olson of the Smithsonian Institution in Washington, its origin is now relatively clear.

  The bird was collected by Andrew Bloxam, a naturalist who served aboard HMS Blonde when it sailed to Honolulu carrying the bodies of Liholiho and Kamamalu, King and Queen of Hawaii, who had died of measles during a trip to London. While in the Pacific the Blonde called at the Cook Islands and on the afternoon of 9th August 1825 Bloxam went ashore on the island of Mauke. Here, during a visit of no more than two hours, he shot a pigeon, a kingfisher and a starling. Nothing more is known of his unique starling. Bloxam simply stated that his bird was killed “hopping about a tree”.

Norfolk Starling Aplonis fusca

The third extinct member of the genus Aplonis inhabited the islands of Norfolk and Lord Howe. Both these Tasman Sea islands have lost several of their endemic birds and the starling vanished during the first half of the twentieth century. The species has been divided into two races, the nominate from Norfolk Island and race hulliana from Lord Howe.

  Although the reasons for extinction on Norfolk Island are unclear, it is quite apparent why the birds vanished from Lord Howe. They were among the creatures doomed by the grounding of a ship, the SS Makambo, in 1918 on a stretch of shore known as Ned's Beach. Rats escaped from the vessel and quickly infested the previously rat-free island. Within just a few years the starlings were gone.

  Little is on record concerning the species. In the days of its abundance it was something of a pest, often feeding on fruit and crops. The nest was loosely built in a tree hollow from twigs and grasses. The birds laid 3-5 bluish eggs, speckled and blotched with red. Lord Howe birds went locally by the name cudgimaruk, a name derived from the call.

Reunion Starling Fregilupus varius

This large and rather beautiful species, also known as the Bourbon Crested Starling, was characterized by an extraordinary lace-like crest. It was an inhabitant of the Mascarene island of Reunion (formerly called Bourbon). It was known locally by the name huppe, which is also the French name for the Hoopoe (Upupa epops). It is perhaps not surprising, then, that the starling was originally placed in the Hoopoe's genus, Upupa.

  Like so many island birds, this was a tame and confiding species. Islanders described how easily individuals could be knocked down with sticks. Despite this, the Reunion Starling survived for much longer than many other Mascarene bird species. It seems to have been relatively common right through into the early years of the nineteenth century, although by the middle years of that century it had disappeared altogether.

  Nothing, apart from its appearance, is known of this species. Indeed, its taxonomic placement is also disputed, as some authors consider it to be a helmet-shrike (Prionopidae).

Rodrigues Starling Necropsar rodericanus

The skeletal remains of a starling were found on the island of Rodrigues during the 1870's. Although there may be no connection, these bones have been associated with a brief account in the document known as the Relation de l'Île Rodrigue, which was probably written by a marooned sailor named Tafforet during 1725. A translation reads:

A little bird is found which is not common...One sees it on the Islet de Mat, 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, and 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 some turtles dead of hunger, which they...tear out of their shells. These birds are a little larger than a blackbird, and have white plumage, part of the wings and tail black, the beak yellow, as well as the feet, and make a wonderful warbling.

  Whether these words should really be matched with the bones cannot be said with any degree of certainty.

FRINGILLIDAE

Bonin Grosbeak Chaunoproctus ferreorostris

A large, spectacular grosbeak-like bird once lived on the Bonin Islands to the south of Japan. It is known from nothing more than two series of skins that were collected during the 1820's, skins that are in themselves a little puzzling. Some are rather larger than others, giving rise to the supposition that individuals may have been collected from different islands and that each island may have supported its own population.

  All the known specimens derive from two expeditions. The first of these was the voyage of HMS Blossom, which vessel called at Peel Island (Chichi-jima), one of the Bonins, during June 1827. The tameness of the birds led the leader of the expedition, Captain Beechey, to assume that there had been no permanent human presence on the island. When the Blossom sailed away several grosbeak specimens were aboard.

  Just months later F. H. von Kittlitz landed on Peel during the voyage of the Russian corvette Senjawin. Whether Kittlitz took birds from elsewhere is not known. His brief notes provide the only record of the bird in life. He noticed them on the forest floor, singly or in pairs, and described the call as a soft, pure and high piping.

  No naturalist ever saw the grosbeak again. By the 1850's, when Peel Island was searched by members of an American naval expedition, there was no sign of it.

DREPANIDIDAE

Ula-ai-Hawane Ciridops anna

Although it was once well known to the natives of Hawaii, by the time that Europeans began their ornithological exploration of the island, this species has virtually disappeared.

  The striking red, black and silver Ula-ai-Hawane is known from just five specimens, two in New York (one of which is either an immature or a female, and shows a greenish plumage), one in Cambridge (Massachusetts), one in Honolulu and one in Tring. The first of these was taken by an amateur naturalist and shopkeeper by the name of Mills about the year 1859. The last was collected from Mount Kohala in 1892. Probably the species became extinct soon after this date, although the experienced and reliable Hawaiian ornithologist George C. Munro reckoned he might have glimpsed an individual as late as 1937.

  Quite clearly, this is one of those Hawaiian species that stood at the brink of extinction even as it was being discovered to science. Native Hawaiian knowledge of the species indicated that it was once widely distributed on the island of Hawaii, although it was only ever seen close to the hawane palm.

Koa Finch Rhodacanthis palmeri

The Koa Finch is something of a mystery. Was it one species or was it two? Were the “Greater” and “Lesser” Koa Finches both members of the same species?

  During 1891 Henry Palmer was busy collecting specimens on the Hawaiian Islands for Walter Rothschild. The enormously wealthy scion of the famous banking family was at this time frantically acquiring material for his museum at Tring, Hertfordshire, England. Nothing interested him more than rare, extinct and curious birds, and he was later to write his famous book Extinct Birds, 1907. Palmer had been commissioned to sail to Hawaii in search of whatever he could find. He linked up with George C. Munro, and together the two men combed Hawaii.

  One of their “finds” was a series of large “finches” that they took from the koa forests in the Kona district of Hawaii. First of all it should be made clear that these were not finches at all; they were Hawaiian honeycreepers that had evolved a finch-like form, in order to exploit a vacant ecological niche. The original collectors, Palmer and Munro, were quite clear in their minds about the nature of the birds they had discovered. They felt that these creatures belonged to a single, rather variable, species, and it is worth noting that the two forms were collected on the very same trees. They then dispatched the specimens of their new bird to Rothschild.

  When the preserved specimens reached Rothschild in England, he formed a different conclusion to that of his collectors, and he delightedly separated his new possessions into two groups; nothing pleased Rothschild more than to be able to name a new form! The larger individuals, with more orange heads, he assigned to a new species that he called, in honour of his collector, Rhodacanthis palmeri. To cover the smaller birds with heads that were rather more yellow he proposed the name Rhodacanthis flaviceps. Rothschild identified eight individuals that he felt belonged to this second species, two males and six females. After Palmer's original collecting success, no further specimens that could be assigned to flaviceps were ever taken or seen. However, the larger birds were observed again, and captured. Over a period of five years a few individuals were encountered. Then these birds vanished too. There is no record later than 1896.

  Palmer's notes were bequeathed by Rothschild to the Natural History Museum, London, but, unfortunately, many of them were destroyed by tidy-minded museum workers, so we will never know exactly what was in his mind.

  The argument for or against two species is a fairly basic one. Does one agree with the experienced field ornithologists who actually saw the living bird, or does one take notice instead of undoubted differences that were noticed by cabinet naturalists? It is possible that the smaller, yellower birds were immatures or first-year birds, and this idea might tidily explain why they were never encountered after 1891. There were no more immatures because the species was standing at the edge of extinction and no longer breeding properly. The last sighting of the species actually involved an observation of some very young birds but, probably, these did not live long.

  The big problem with the evidence of the specimens is that is difficult to correlate it with the birds in life. Although the most extreme examples of each form appear to be rather different, there are areas of overlap between some of the other specimens. The real truth is probably unresolvable, and is simply a matter of interpretation.

  Unfortunately, knowledge of the living bird in no way matches the amount that has been written concerning the supposed status of the species - or pair of species. The best description of the bird in life was given in 1903 by R. C. L. Perkins, a man who left behind some of the most evocative writings on the extinct birds of the Hawaiian Islands:

Although spending most of its time in the tops of the loftiest koa trees, Rhodacanthis occasionally visits the lesser trees...chiefly for the sake of the caterpillars that feed upon them...Its chief food, however, is the green pod of the koa tree, which it swallows in large-sized pieces and its blue bill is often stained with the green juice and fragments of the pods...The song...consists of four, five or even six whistled notes, of which the latter ones are much prolonged...Although the notes are not loud, they are very clear, and...easily imitated...Were it not for this fact Rhodacanthis, when keeping to the leafy crowns of tall koa trees...would be most difficult to get sight of...The green plumaged young...are fed partly on fragments of koa pods.

Kona Grosbeak Chloridops kona

The Hawaiian honeycreepers form one of the most striking illustrations of adaptive radiation, but, unfortunately, so many of them are extinct that the example is now historical rather than living.

  At some point in prehistory an ancestral honeycreeper stock somehow arrived at the Hawaiian Islands and found a situation that was ripe for exploitation. Due to the remoteness of the islands there were many ecological niches that were not successfully occupied. The ancestors of the honeycreepers adapted accordingly and lifestyles were adopted that would in more normal circumstances be the prerogative of other kinds of birds. Thus, there are honeycreepers that came to resemble finches, some that developed almost parrot-like bills, still others that acquired long, slender, down-curved beaks useful for probing, and yet others with more general all-purpose beaks. This fascinating diversity has been horribly diminished by the process of extinction.

  The Kona Grosbeak was a perfect example of the way in which the ancestral honeycreepers adapted. It had evolved a short and incredibly powerful bill that enabled it to exploit food types that more normally developed birds could not. Indeed, the sounds of hard seeds and nuts being split by the awesome bill of this species betrayed its presence to field collectors.

  This is another bird that was best described by R. C. L. Perkins (1903). He wrote:

It is a dull, sluggish bird, and very silent - its whole existence may be summed up in the words “to eat”. Its food consists of seeds of the aaka, and as these are very minute, its whole time seems to be taken up in cracking the extremely hard shells...Its beak is nearly always dirty, with a brown substance adherent to it, which must be derived from the sandal nuts.

  The species was discovered on the slopes of the famous volcano Mauna Loa in the Kona district of Hawaii during 1887. It was seen again several times during the next few years, but the last sighting occurred in 1894 and it is assumed that these birds became extinct soon after that date.

Greater Amakihi Hemignathus sagittirostris

The Greater Amakihi, a rather non-descript little honeycreeper coloured olive green, was discovered by the world of ornithology during 1892 when Rothschild's collector Henry Palmer took four specimens. In December 1895 a few individuals were collected, and the species was located again during 1900. The following year it was found once more, since which time it has never been seen again.

  By the time of its ornithological discovery the species was restricted to a very small part of the island of Hawaii, the dripping rainforest to either side of the Wailuku River at elevations of 300-900 m.

  Whenever it was seen, this species was observed creeping along the branches of ohia trees or in the foliage. Individuals ate insects and their grubs and other invertebrates. They were also seen to feed on the nectar of the ohia flower.

  The habitat that this species occupied is long gone and the area is now covered with sugar cane.

Akialoa Hemignathus obscurus

Because it is so poorly known, the honeycreeper family, in particular its extinct members, arouses a certain amount of controversy. It is classified and reclassified over and over again, and drastic revisions of the family are made from time to time. Unfortunately, these revisions are, of necessity, made from specimen material rather than from any real recourse to the living birds. For this reason these revisions should be viewed with a certain degree of caution. They tend to be prone to the whims and prejudices of individuals, and also subject to the dictates of fashion.

  In no case is this better exemplified than in the instance of the Akialoa. Perhaps the Akialoa constitutes one species, perhaps two, perhaps even four. It has been divided into two, a “Greater” and a “Lesser”, and into four with each inhabited Hawaiian island being held to have its own species. The specimen material is very difficult to interpret, however. The forms from Lanai (lanaiensis) and O'ahu (ellisianus) are known only from very, very few specimens, so there is no real series on which to base conclusions. The other two forms, nominate obscurus from Hawaii and procerus from Kauia, are known from rather more skins, but even in these cases comparisons are clouded by various factors. The main problem is that the Akialoa was a generally variable bird and individuals of each kind show some overlap in terms of size and colour. It is possible that individuals became brighter in colour when they were breeding, although it should be said that there is no real evidence of this. Bill length is held to be a significant character, but this too is variable: there is evidence to show that towards the end of its existence as a species, the Akialoa's beak tended to be rather shorter than in earlier years. Perhaps this was because the birds were generally in poor health, and collectors who took specimens during the 1890's noticed that individuals were often covered in sores, tumours and swellings, particularly on the head and feet; or maybe the new circumstances on the islands were inhibiting proper growth.

  The most sensible way forward is, perhaps, to regard the Akialoa as a single species with four races. Bearing in mind the paucity of specimen material available, no other course seems meaningful.

  However they may be interpreted, the birds that are known as Akialoas were among the most intriguing of the honeycreepers. The extraordinarily delicate beak is in complete contrast to the shorter, stumpy bills of the Koa Finch and the Kona Grosbeak. The lower mandible was often considerably shorter than the upper, and with this strange device individuals were able to suck nectar from the flowers of ohias and lobelias, and they could also probe into cavities for insects and their larvae. R. C. L. Perkins noticed them clinging to tree trunks like “true” creepers when they were feeding.

  While it is known that there were Akialoas on Hawaii, O'ahu, Lanai and Kauai, it is quite possible that birds were present on other islands. It is surprising to find, for instance, that they were never reported from Molokai and Maui. Perhaps they vanished from these islands at a comparatively early date. It is likely that the Akialoas from Oahu became extinct around 1840, although there are claimed records, made by experienced ornithologists, of possible sightings made during the late 1930's. The Lanai and Hawaii populations disappeared around 1900, but Kauai birds held out for much longer. Their stronghold was the same Alaka'i Swamp that was home to the last Kauai 'O'os. A few individuals were still in existence during the early 1960's but the last record of them seems to date from 1967.

  The obvious frailty of the Akialoa reflects the fragility of the honeycreepers as a whole. In addition to all the other factors that depleted them, the danger that the mosquito posed cannot be overstated. Once this insect arrived in the Hawaiian Islands, birds that lived at lower altitudes were seriously affected by the bites of these creatures. Even those species living at altitudes that were unattractive to mosquitoes could be vulnerable. Sometimes they would descend to lower levels to ride out the effects of hurricanes, and then the bites could prove fatal. Captive birds that have been brought down from altitude have been observed to die within minutes of sustaining a bite.

Hawaii Mamo Drepanis pacifica

The Hawaii Mamo suffered in the same way and for the same reason as the Hawaii 'O'o. The rich yellow feathers of the uppertail- and undertail-coverts attracted the attention of the kings and princes of Hawaii, and these feathers were used in the making of ceremonial cloaks and other artifacts. The most famous cloak of all, that of Kamehameha I was made from the feathers of an extraordinary number of birds. An estimate has been made that 80,000 individuals were used before the garment could be brought to completion.

  The Hawaii Mamo was one of the first Hawaiian birds to come to the attention of Europeans and specimens were brought back from the Pacific to England by the naturalists who participated in James Cook's third voyage. Surprisingly little is known of it, however. The birds fed on nectar, particularly from the flowers of arborescent lobelias, but they are thought also to have eaten insects. The call was described as a single, long, rather mournful and haunting note.

  The species was only ever noticed on the island of Hawaii and it was last seen during 1898.

Black Mamo Drepanis funerea

A bird very closely related to the Hawaii Mamo once lived on the island of Molokai. Here it was discovered by R. C. L. Perkins at an altitude of 1525 metres during June 1893. What is known of the species comes largely from Perkins's account.

  Individuals were only ever seen low down in the underbrush. Dangerously, for a creature that spent its time close to the ground, the birds were rather tame, and Perkins found it easy to observe them at close quarters. He believed that they fed exclusively on nectar, and he wrote:

I saw three adult males of this bird in one low bush passing from flower to flower and spending only a few seconds over each...Even those flowers which were at a height of no more than a foot from the ground were carefully explored. The crown of the head of each of these birds was plentifully encrusted...with the sticky white or purplish-white pollen of the lobelias and gave them a singular appearance...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.

  Following Perkins's discovery of the species, it was found on several more occasions, but was last seen in June 1907. There is no real chance that it could have survived as the habitat in which it lived has been virtually destroyed.

PARULIDAE

Bachman's Warbler Vermivora bachmanii

This tiny species divided its time between the south-eastern USA and Cuba, where it wintered. It was first identified by the Reverend John Bachman, a close personal friend of the famous painter and writer John James Audubon. Bachman was a resident of Charleston, and in July 1833 he found, in a local swamp, a small yellow and black bird that he did not recognize. He sent it on to Audubon for formal identification and the famous ornithologist quickly realized that it was something entirely new. Audubon, quite naturally, gave the species his friend's name.

  For a period of more than 50 years after that nothing further was heard of the new creature. Then, in 1886, a hunter by the name of Charles Galbraith shot a bird he had never seen before, just north of New Orleans. The next year he shot six more of the same birds, and during the following year he killed no less than 31. At this point he decided to have his mysterious specimens identified. They proved to be Bachman's Warblers.

  A year later, in March 1889, on a single day 21 warblers of the species struck a lighthouse on the Florida Keys. Just three years later a hunter killed 50 individuals on Florida's Suwannee River.

  Through the first half of the twentieth century it proved possible to locate Bachman's Warblers from time to time, but then the observations proved much less frequent. By the 1980's the species was probably extinct, although, given its tendency to disappear, there is a faint chance that it might still survive.

  Although Bachman's Warblers were always surrounded by a certain amount of mystery, their nests were found on a number of occasions. These were built low down in dense patches of bramble, situated along forested river courses.

  Reasons for the species' extinction are unclear. The effect of hurricanes, changing land use, and the fragmentation of suitable territory have all been proposed as factors. Another suggestion is that the species was coming into increasing contact with the parasitic Brown-headed Cowbird (Molothrus ater), due to extensive forest clearance, and that the encroachments of this bird proved irresistible.

 
Hypothetical Species and Mystery Birds

No account of the world's recently extinct birds can be quite complete without some mention of “hypothetical” extinct species and “mystery” birds. The names of such creatures frequently litter the lists of extinct species and create endless confusion.

  Hypothetical species can be defined as birds that have been scientifically named, mainly by nineteenth and twentieth century ornithologists, on the basis of early traveller's tales or antique paintings, and of which no specimen material exists. The problems associated with such names are self-evident. First, they contravene the rules of zoological nomenclature by being too vague. Second, there is no actual proof that the described creatures ever existed, or that they were what they seemed to be. The names may be founded upon mistakes, misunderstandings or even plain old-fashioned lies.

  Mystery birds are seemingly distinct kinds that are known from just a single specimen, or a very small number of specimens, that, for a variety of reasons, may not be exactly what they seem to be.

  Although the two categories are obviously related, it is easiest to discuss them separately.

  It is a surprising fact that so many hypothetical species have been named (by ornithologists who should have known better!), and that they have gained a significant place in ornithological literature - but such is the case.

  Most are forms resurrected from the accounts of travellers from the seventeenth or eighteenth centuries, and given formal, albeit invalid, scientific identities by more modern writers. The problems with such accounts are several. First, they may have been - and usually were - written by men with no real background in natural history. Second, they may have been based on hearsay. Third, the accounts may be flimsy and superficial in the extreme. Fourth, there was often a tendency among early travel writers to fill up the pages of their books with accounts of creatures that were entirely, or partly, fanciful. The fourth category obviously speaks for itself and needs no further discussion. Similarly, the problems with the second and third categories are perfectly apparent. A more complex set of problems is posed by the first. When men have no knowledge of natural history (or even when they do), they are likely to misinterpret what they see. A description of a new creature may be entirely truthful in spirit, yet still be absolutely misleading in effect. Perhaps the observer saw something that was new to him but was otherwise well known. Perhaps he casually mentioned a crow when the bird he had seen was actually a starling, and the words get in the way of the truth. Sometimes, the description may be entirely inaccurate.

  Early paintings of birds that cannot be precisely aligned with any known species are also problematical. To take a picture, even when painted by an artist of talent, at face value, is often unrealistic. The problem here is one of intention. There is often no way of knowing whether an artist was trying to produce a true likeness, whether his intention was simply decorative, or whether it lay somewhere between the two. There is also the matter of technique. Many of the pictures on which names have been based were drawn by men with little or no skill. Yet, nuances of detail on such pictures have been taken absolutely seriously by some ornithologists when, quite clearly, such details may simply be the result of inadequate drawing ability. These pictures may be charming and curious, and they may show something of genuine interest. On the other hand they may not! When scientific names are proposed solely on the basis of such pictures these names should be regarded as invalid.

  The inclusion of several species known today only from the paintings of Georg Forster, who was by no means an efficient technician, may seem to contravene these ideas, but it should be borne in mind that Forster's pictures were originally backed up by specimen material. Unfortunately, in the cases in question, that material has not survived. 

The best known of all hypothetical species is the so-called White Dodo of Reunion. The evidence that such a creature existed is flimsy in the extreme yet many ill-judged words have been written about it (including some by the present author), and it was given the technical name Raphus solitarius. Two accounts from the early seventeenth century suggest that there may have been a creature that looked something like a White Dodo on Reunion, but these are far too vague to enable the building of any kind of sensible hypothesis. Another strand of evidence concerns a well-known series of seventeenth century Dutch paintings by Pieter Holsteyn (later copied by another painter named Pieter Witthoos) that show a White Dodo. Unfortunately, however, there is absolutely nothing to link these pictures to Reunion. Nor is there anything to show that the model for the pictures was anything other than an albinistic Dodo of the more ordinary kind. Similarly, because we know nothing of the intention of the original artist, it is quite fair to suppose that he made his Dodo white simply because he wanted to. What can be said with some certainty is this: had a Dodo-like bird evolved independently on the island of Reunion, it would have looked radically different from its relative on Mauritius, rather than looking just like a regular Dodo that had been bleached. It would necessarily have developed in a different direction, in much the same way as did the dodo-like solitaire of the island of Rodrigues. This creature, while showing many dodoesque features, has a number of entirely independent ones - and this is only to be expected of a flightless creature evolving independently on a separate island. It is quite possible that a relative of the Dodo did actually live on Reunion, but there is not one scrap of hard evidence to show this.

  Another hypothetical species that has acquired almost equal celebrity has come to be known as Leguat's Giant (Leguatia gigantea). This is a bird that the Huguenot refugee François Leguat claimed to have seen on Mauritius during the 1690's. His description makes it clear that this was a slender, very tall creature. As Leguat was generally a very truthful recorder it may be assumed that he really did see something, but just what that something was is difficult to say. The likelihood is that he may have seen a flamingo and that he was entirely unfamiliar with such a creature.

  The list of hypothetical species is fairly endless, and by and large such mythical birds are properly beyond the scope of this work. 

The problems associated with mystery birds are equally difficult to resolve. If a bird is known from just a single specimen or just a very small number of specimens, there can be all kinds of difficulty involved in assessing the correct position. This is especially so if the bird was never observed properly in the wild. The possibility that such birds may be simply freaks or perhaps hybrids between better-known species cannot be discounted. Naturally, if birds turn up just once and then many years pass without anything similar being found, there is always the possibility that the species - if species it truly is - has become extinct.

  There are, in fact, a surprisingly large number of birds known from only very limited specimen material. The family of birds-of-paradise (Paradisaeidae), for instance, contains more than 20 mysterious forms known only from excessively rare and isolated specimens. These are generally considered to be hybrids between some of the better-known kinds, but some may not be.

  Ultimately, such decisions are often based on individual prejudices, as the actual hard evidence is not sufficient to build a proper determination.

  The case of the Lanai Hookbill (Dysmorodrepanis munroi), a honeycreeper from the Hawaiian island of Lanai, provides a good example. It may be a “good” species and, if it is, it is almost certainly extinct. On the other hand, it may be a freak. Just a single example of this creature is known and this was collected during 1913 by George C. Munro, whose name so often occurs in accounts of Hawaiian birds. The specimen shows a very distinct but rather peculiar arrangement of the beak and experts interpret this in two contrasting ways: some argue that the beak's peculiar nature is the sure mark of a freak; others suggest that it is a clear and distinct indication of a “good” species.

  In some respects similar is the case of Townsend's Bunting (Spiza townsendi). This is a form described by the celebrated John James Audubon from a unique specimen taken in Pennsylvania. No similar bird has ever come to light, but, since the specimen shows some relationship to the Dickcissel (Spiza americana), the form is usually relegated to the status of freak or hybrid.

  Ross's Plover (Thinornis rossi), a strange shorebird from New Zealand waters was described from a single specimen taken in 1840, during the exploratory voyage of the British vessels Erebus and Terror. Usually, it is regarded as an aberrant, perhaps young, individual of the Shore Plover (Thinornis novaeseelandiae), itself a very rare bird. It may or may not be. Similarly, Cooper's Sandpiper (Calidris cooperi), collected during the nineteenth century on Long Island, New York, might be a “good” species, although nowadays the general consensus is that it is a hybrid of the Curlew Sandpiper (Calidris ferruginea) and the Sharp-tailed Sandpiper (Calidris acuminata).

  The big problem with all these forms (and there are a considerable number of them) is that we shall simply never know what they really were.

  To these forms can be added many that are simply known from fragmentary bone material. For the purposes of a work like this there is little point in pursuing such insufficient yet tantalizing material. Such a pursuit leads to endless list-making, argument and counter-argument. Those recently extinct creatures that we know from clear-cut specimen material are tantalizing and mysterious enough in themselves and, most regrettably, it seems inevitable that, unless conservation action is given full institutional backing right away, there will soon be many more well-known, and not so well-known, names to add to the ranks of the extinct. 

  Errol Fuller 
 
 

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