Authorities.—A. Milne-Edwards,Recherches anatomiques et paléontologiques pour servir à l’histoire des oiseaux fossiles de la France(Paris, 1867-1868); F.P. Moreno and A. Mercerat,Catalogo de los Pajaros fosiles de la Republica Argentina. Anales Mus. La Plata, 1891, 21 pls.; O.C. Marsh,Odontornithes: A monograph of the Extinct Toothed Birds of North America(New Haven, Conn., 1880); R. Lydekker, article “Fossil Birds,” in A. Newton’sDictionary of Birds(London, 1893); Cat. Foss. Birds, Brit. Museum, 1891; K. v. Zittel,Handbuch der Palaontologie, i. 3 (1887-1890); C.W. Andrews, “On the Extinct Birds of Patagonia,”Tr. Zool. Soc. xv., 1899, pp. 55-86, pls. 14-17.
Authorities.—A. Milne-Edwards,Recherches anatomiques et paléontologiques pour servir à l’histoire des oiseaux fossiles de la France(Paris, 1867-1868); F.P. Moreno and A. Mercerat,Catalogo de los Pajaros fosiles de la Republica Argentina. Anales Mus. La Plata, 1891, 21 pls.; O.C. Marsh,Odontornithes: A monograph of the Extinct Toothed Birds of North America(New Haven, Conn., 1880); R. Lydekker, article “Fossil Birds,” in A. Newton’sDictionary of Birds(London, 1893); Cat. Foss. Birds, Brit. Museum, 1891; K. v. Zittel,Handbuch der Palaontologie, i. 3 (1887-1890); C.W. Andrews, “On the Extinct Birds of Patagonia,”Tr. Zool. Soc. xv., 1899, pp. 55-86, pls. 14-17.
C. Geographical Distribution
The study of the extinct organisms of any country leads to a proper appreciation of its existing flora and fauna; while, on the other hand, a due consideration of the plants and animals which may predominate within its bounds cannot fail to throw more or less light on the changes it has in the course of ages undergone. That is to say, the distribution of forms in time is a subject so much connected with the distribution of forms in space, that the one can hardly be separated from the other. Granting this is a general truth, it must yet be acknowledged as a special fact, that in fossil birds we have as yet but scanty means of arriving at any precise results which will justify bold generalization in the matter of avine distribution. Remains of extinct birds are, compared with those of other classes of vertebrates, exceedingly scarce, and these have been found in very few, widely separated countries. The great problems involved in the study of geographical distribution must therefore be based mainly upon the other classes, both vertebrate and invertebrate, which, moreover, enjoy less great facilities of locomotion than the birds.
Yet it so happens that the great zoogeographical regions of the world, now more or less generally accepted, have been based upon the distribution of birds. The whole subject was properly introduced by Treviranus,1who in his large philosophical work devotes considerable space to the “geographical distribution of animals.” Next we have to mention F. Tiedemann,2the Heidelberg anatomist, who has been generally ignored, although he surpassed many a recent zoogeographer by the wide view he took of the problem; in fact he was the first to connect distribution with environmental or bionomic factors;e.g. the remark on p. 481 of his work that “the countries of the East Indian flora have no kinds of birds in common with America which are vegetable feeders.” L.K. Schmarda3divided the land into twenty-one realms, characterizing these mainly by their birds. P.L. Sclater4was the first to divide the world into a few great “regions,” the Palaearctic, Ethiopian, Indian and Australian forming one group, the “Old World” (Palaeogaea); and the Nearctic and Neotropical forming a second, the New World (Neogaea). Birds being of all animals most particularly adapted for extended and rapid locomotion, it became necessary for him to eliminate from his consideration those groups, be they small or large, which are of more or less universal occurrence, and to ground his results on what was at that time commonly known as the order Insessores or Passeres, comprehending the orders now differentiated as Passeriformes, Coraciiformes and Cuculiformes, in other words the mass of arboreal birds. His six main divisions—practically adopted by A.R. Wallace5in his epoch-making work—are excellent, taken separately. They express the main complexes of land with their dependencies in well-chosen terms; for instance the “Neotropical region” stands short for South and Central America with the Antilles.
But these six divisions of Sclater and Wallace are not all equivalent, only some are of primary importance; they require co- and sub-ordination. This most important advance was made by T.H. Huxley.6Some of the “regions” have now to be called subregions,e.g. the Nearctic and the Palaearctic. The reduction of the Oriental to a subregion, with consequent “provincial” rank of its main subdivisions, will probably be objected to, but these are matters of taste and prejudice. Above all it should be borne in mind that nearly all the last subdivisions or provinces are of very little real value and most of them are inapplicable to other classes of animals.
Besides some occasional references in the text, only a few more of the general works dealing with the distribution of birds can here be mentioned. Especial attention has to be drawn to the article “Geographical Distribution,” in Newton’sDictionary of Birds. See also A. Heilprin,The Geographical and Zoological Distribution of Animals(New York, 1887); W. Marshall and A. Reichenow, two maps with much detail, although badly arranged, in Berghaus’Physikalischer Atlas, pt. vi. (Atlas d. Thierverbreitung), (Gotha, 1887); A. Reichenow, “Die Begrenzung zoogeographischer Regionen vom ornithologischen Standpunkte,”Zoolog. Jahrb.iii., 1888, pp. 671-704, pl. xxvi.; E.L. Trouessart,La Géographie zoologique. (Paris, 1890).The scheme adopted in the following account stands as follows:—(A)Austrogaeaor I. Australian RegionNew Zealandsubregion.Australian”Papuan”(B)Neogaeaor II. Neotropical RegionAntillean”Columbian”Patagonian”(C)ArctogaeaIII. Holarctic RegionNearctic”Palaearctic”IV Palaeotropical RegionEthiopian”Oriental”
Besides some occasional references in the text, only a few more of the general works dealing with the distribution of birds can here be mentioned. Especial attention has to be drawn to the article “Geographical Distribution,” in Newton’sDictionary of Birds. See also A. Heilprin,The Geographical and Zoological Distribution of Animals(New York, 1887); W. Marshall and A. Reichenow, two maps with much detail, although badly arranged, in Berghaus’Physikalischer Atlas, pt. vi. (Atlas d. Thierverbreitung), (Gotha, 1887); A. Reichenow, “Die Begrenzung zoogeographischer Regionen vom ornithologischen Standpunkte,”Zoolog. Jahrb.iii., 1888, pp. 671-704, pl. xxvi.; E.L. Trouessart,La Géographie zoologique. (Paris, 1890).
The scheme adopted in the following account stands as follows:—
In the following account the characterization of the various regions and subregions has to a very great extent been adopted from Newton’s article in hisDictionary of Birds, and from the chapter on distribution in the article on “Birds” in theEncyclopaedia Britannica, 9th edition. This applies especiallyto those instances in which the members of families, genera and species are mentioned. The families are those which are enumerated in Garow’s classification. The numbers of genera and species of birds are, of course, a matter of personal inclination. If we take a moderate computation the number of recent species may be taken at 10,000-11,000.7Dr R.B. Sharpe increases their number to about 15,000 in theNew Hand-List of Birds, published by the British Museum. In the first two volumes fossil birds, occasionally based upon a fragmentary bone only, are also included.
(A)Austrogaea, the Australian region in the wider sense, with the Papuan, Australian and New Zealand subregions, including also Polynesia. We may here quote Newton (Encyclopaedia Britannica, 9th ed., “Birds,” p. 738) on the remarkable differences between this region and the rest of the Old World:—“The prevalent zoological features of any Region are of two kinds—negative and positive. It is therefore just as much the business of the zoogeographer, who wishes to arrive at the truth, to ascertain what groups of animals are wanting in any particular locality (altogether independently of its extent) as to determine those which are forthcoming there. Of course, in the former case it would be absurd to regard as a physical feature of any great value the absence from a district of groups which do not occur except in its immediate neighbourhood; but when we find that certain groups, though abounding in some part of the vicinity, either suddenly cease from appearing or appear only in very reduced numbers, and occasionally in abnormal forms, the fact obviously has an important bearing. Now, mere geographical considerations, taken from the situation and configuration of the islands of the so-called Indian or Malay Archipelago, would indicate that they extended in an unbroken series from the shores of the Strait of Malacca to the southern coast of New Guinea, which confronts that of north Australia in Torres Strait, or even farther to the eastward. Indeed, the very name Australasia, often applied to this part of the world, would induce the belief that all the countless islands, be they large or small—and some of them are among the largest on the globe—were but a southern prolongation of the mainland of Asia. But so far from this being the case a very definite barrier is interposed. A strait, some 15 m. or so in width, and separating the two fertile but otherwise insignificant islands of Bali and Lombok, makes such a frontier as can hardly be shown to exist elsewhere. The former of these two islands belongs to the Indian Region, the latter to the Australian, and between them there is absolutely no true transition—that is, no species are common to both which cannot be easily accounted for by the various accidents and migrations that in the course of time must have tended to mingle the productions of islands so close to one another. The faunas of the two are as absolutely distinct as those of South America and Africa, and it is only because they are separated by a narrow strait instead of the broad Atlantic that they have become so slightly connected by the interchange of a few species and genera.“Now, first, of the forms of birds which are prevalent throughout the Indian Region, but are entirely wanting in the Australian, we have at once the bulbuls (Ixidae), very characteristic of most parts of Africa and Asia, including the sub-groupPhyllornithinae, which is peculiar to the Indian Region; the widely-spread families of barbets (Megalaeminae) and vultures (Vulturidae); and the pheasants (Phasianidae), which attain so great a development in various parts of the Asiatic continent and islands that there must their home be regarded as fixed. Some naturalists would add the finches (Fringillidae), rightly if we assume that thePloceidaeor weavers constitute a separate family. Then, of forms which are but weakly represented, we have the otherwise abundant thrushes (Turdidae), and, above all, the woodpeckers (Picidae), of which only very few species, out of 400, just cross the boundary and occur in Lombok, Celebes or the Moluccas, but are unknown elsewhere in the region.”But the Australian region is also remarkable for its ornithic singularity. All the existing Ratitae (with the exception of the ostriches of Africa and South America, belonging to the generaStruthioandRhea, and comprising at most but five species) are found in Austrogaea and nowhere else. Of the Passeres the honey-suckers (Meliphagidae) are most characteristic, and, abounding in genera and species, extend to almost every part of the region, yet only one species ofPtilotisoversteps its limits, crossing the sea from Lombok to Bali. Other peculiar families are much more confined. But the positive characteristics of the region as a whole are not its peculiar forms alone; there are at least four families which, being feebly represented elsewhere, here attain the maximum of development. Such are the thick-headed shrikes (Pachycephalidae), the caterpillar-eaters (Campephagidae), the flower-peckers (Dicaeidae), and the swallow-flycatchers (Artamidae). Besides these, three or perhaps four groups, though widely distributed throughout the world, arrive in the Australian region at their culmination, presenting an abundance of most varied forms. These are the weaver-birds (Ploceidae), and the moreporks (Podargidae), but especially the kingfishers (Alcedinidae) and the pigeons (Columbidae), the species belonging to the two last obtaining in this region a degree of prominence and beauty which is elsewhere unequalled.The boundaries of the subregions are not well defined.Fig.22.—Extinct Phillip-Island Parrot (Nestor productus). (From specimen in the British Museum. Reduced.)TheNew Zealand Subregion, considered by Professors Newton and Huxley and various other zoogeographers as deserving the rank of a region, is, and to all appearance has long been, more isolated than any other portion of the globe. Besides the three larger islands numerous satellites belong to the subregion, as Lord Howe, Norfolk and Kermadec islands, with the Chatham, Auckland and Macquarie groups. The main affinities of the avifauna are, of course, Australian. The most extraordinary feature is unquestionably the former existence of the gigantic Dinornithes or moas (q.v.) and, another family of Ratitae, the weird-looking kiwis or Apteryges, which are totally unlike any other existing birds. Of other peculiar genera it will suffice to mention only the more remarkable.Rallidaepresent the very noteworthy woodhens,Ocydromus, and the takahe,Notornis, which is almost extinct. The widely-spread plovers,Charadriidae, have two not less singular generic developments,Thinornis, and the extraordinary wrybill,Anarhynchus. There is an owl, type of the genusSceloglaux. Of parrots,Stringops, the kakapo or owl-parrot, is certainly peculiar, whileNestorconstitutes a peculiar subfamily of the brush-tongued parrots orTrichoglossidae.XenicusandAcanthosittaform a little family of truly mesomyodean Passeres Clamatores. Of theMeliphagidaethe generaProsthemadera,PogonornisandAnthornisare peculiar. The starlings,Sturnidae, are represented byCallaeas,Creadionand the very abnormalHeterolocha. The gallinaceous birds are represented by a quail,Coturnix novae zealandiae, now exterminated. A large flightless goose,Cnemiornis, allied to the AustralianCereopsis, and the gigantic rapaciousHarpagornis, have died out recently, with the moas. In all, there is a wonderful amount of specialization, though perhaps in a very straight line from generalized forms; but the affinity to Australian or Polynesian types is in many cases clearly traceable, and it cannot be supposed but that these last are of cognate origin with those of New Zealand. A very long period of isolation must have been required to produce the differences so manifestly to be observed, but a few forms seem at rare intervals to have immigrated, and this immigration would appear to be kept up to our own day, as shown by the instance ofZosterops lateralis, which is said to have lately made its first appearance, and to have established itself in the country, as well as by the fact of two cuckoos,the widely-rangingEudynamis taitensisandChrysococcyx lucidus, which are annual visitors.Polynesiaforms, of course, part of Austrogaea. Its extent is so vast that it necessarily contains some peculiar, outlying forms, so to say forgotten, which in their long-continued isolation have specialized themselves. For instance, the kagu (Rhinochetus) of New Caledonia, a queerly specialized form with Gruine affinities pointing only to South America. The toothbilled pigeon (Didunculus) is restricted to Samoa. Most interesting is the avifauna of the Sandwich islands; entirely devoid of Psittaci and of Coraciiformes, these islands show an extraordinary development of its peculiar familyDrepanidae, which are probably of South or Central American descent.Acrulocercusis a Meliphagine, and a peculiar genus. There are a raven (Corvus), a coot (Fulica), the well-known Sandwich island goose (Bernicla sandvicensis), now very commonly domesticated in Europe; and some flycatchers and thrushlike birds.TheAustralian Subregioncomprises Australia and Tasmania. In the north it is influenced, of course, by its proximity to Papuasia, whence there is a considerable admixture of genera which do not proceed beyond the tropics, and of theseCasuariusis a striking example. The Cape York peninsula practically belongs to Papuasia. As a whole, Australia is rich in parrots, of which it has several very peculiar forms, but Picarians in old-fashioned parlance, of all sorts—certain kingfishers excepted—are few in number, and the pigeons are also comparatively scarce, no doubt because of the many arboreal predaceous marsupials. The continent, however, possesses the two important genera of thePseudoscines, namely the lyre-birds (Menura) and the scrub-birds (Atrichia). Among the more curious forms of other land-birds may be especially mentioned theMegapodiidae,LipoaandTalegallus, the railTribonyxandPedionomus, which represents the otherwise palaeotropicalTurnicesin Australia. The presence of bustards (Eupodotis) is a curious example of interrupted distribution, since none other of theOtididaeare found nearer than India. The Ratitae are represented by two species of emeu (Dromaeus), besides the cassowary of Cape York peninsula, and the extinctDromornisandGenyorniswith its enormous skull.ThePapuan Subregion, chiefly New Guinea with its dependencies, the Timor group of islands, the Moluccas and Celebes. On the whole its avifauna presents some very remarkable features. Its most distinctive characteristic is the presence of the birds of paradise, which are almost peculiar to it; for, granting that the bower-birds,Chlamydoderaand others, of Australia, belong to the same family, they are far less highly specialized than the beautiful and extraordinary forms which are found, within very restricted limits, in the various islands of the subregion. Another chief feature is the extraordinary development of the cassowaries, the richness and specialization of the kingfishers, parrots, pigeons, honey-suckers and some remarkable flycatchers. It has several marked deficiencies compared with Australia, among which are the babblers (Timeliidae), weaver birds (Ploceidae), thePlatycercinaeamong parrots, diurnal birds of prey and the emeus. As a whole, the birds of Papua are remarkable for their brilliance of plumage, or their metallic colouring. The birds of paradise, the racquet-tailed kingfishers,Tanysiptera, the largest and smallest of parrots,CalyptdrhynchiisandNasiterna, and the great crowned pigeons,Goura, are very characteristic; and so are the various Megapodes.(B)Neogaea, or theNeotropical region.—Excepting towards the north, where, in Mexico, it meets, and inosculates with the Nearctic subregion, the boundaries of the Neotropical region are simple enough to trace, comprehending as it does the whole of South America and all Central America; besides including the Falkland islands to the south-east and the Galapagos under the equator to the west, as well as the Antilles or West India islands up to the Florida channel.Owing to the comparatively scanty number of harmful mammalian types, the birds play a considerable part in this large region, and some authorities consider its avifauna the richest in the world. The entire number of species amounts to about 3600. Of these 2000, or a good deal more than half, belong to the order Passeriformes. But the characteristic nature of the avifauna is more clearly brought out when we learn that of the 2000 species just mentioned only about 1070 belong to the higher suborder of Oscines, that means to say, nearly one-half belong to the lower suborder Clamatores. This is a state of things which exists nowhere else; for except in Australia, where a few indigenous and peculiar low non-Oscines are found, and in the Nearctic country, whither one family of Clamatores, viz. theTyrannidae, has evidently been led by the geographical continuity of its soil with that of the Neotropical region, such forms do not occur elsewhere. Accordingly their disproportionate prevalence in South America points unerringly to the lower rank of the avifauna of the region as a whole, and therefore to the propriety of putting it next in order to that of the Australian region, the general fauna of which is admittedly the lowest in the world. Huxley has urged with his wonted perspicuity the alliance of these two regions asNotogaea, basing his opinion, besides other weighty evidence, in great measure on the evidence afforded by the two main sections of the Galli, viz. thePeristeropodesand theAlectoropodes, the former composed of the familiesMegapodiidae, almost wholly Australian, and theCracidae, entirely Neotropical. (Cf.P.Z.S., 1868. pp. 294-319.)Leaving, however, this matter as in some degree hypothetical, we have as genera, families, or perhaps even larger groups, a great many very remarkable forms which are characteristic of, or peculiar to, the Neotropical region in part, if not as a whole. Of families we find twenty-three, or maybe more, absolutely restricted thereto, besides at least eight which, being peculiar to the New World, extend their range into the Nearctic region, but are there so feebly developed that their origin may be safely ascribed to the southern portion of America. First in point of importance comes the extraordinarily beautiful family of humming-birds (Trochilidae), with nearly 150 genera (of which only three occur in the Nearctic region) and more than 400 species. Then the tyrants (Tyrannidae), with more than seventy genera (ten of which range into the northern region), and over 300 species. To these follow the tanagers (Tanagndae), with upwards of forty genera (only one of which crosses the border), and about 300 species; the piculules (Dendrocolaptidae), with as many genera, and over 200 species; the ant-thrushes, (Formicariidae), with more than thirty genera, and nearly 200 species; together with other groups which, if not so large as those just named, are yet just as well defined, and possibly more significant, namely, the tapaculos (Pteroptochidae), the toucans (Rhamphastidae), the jacamars (Galbulidae), the motmots (Monotidae), the todies (Todidae), the trumpeters (Psophiidae), and the screamers (Palamedeidae); besides such isolated forms as the seriema (Cariama), and the sun-bittern (Eurypyga).The nature of the South American avifauna will perhaps become still more evident if we arrange the characteristic members as follows:—1. Birds which are restricted to, probably indigenous of the region:Rhea;PalamedeaandChauna, the screamers;Tinami;Psophia, Dicholophus, Eurypyga, Heliornisof the Gruiform assembly;ThinocorysandAttagis; Cracidae; Opisthocomus; of parrotsAraandConuruswith their allies;Monotidae, incl.Todus; Steatornis; GalbulinaeandBucconinae; Rhamphastidae; Formicariidae, Pteroptochidae, and of theTyrannidaetheCotinginae.2. Birds which are indigenous, but extend far into North America:Cathartae, Trochilidae, Tyrannidae.3. Birds which are originally immigrants from North America:Podicipedidae, with the flightlessCentropelmaon Lake Titicaca;Ceryle, the only genus of kingfishers in the New World; all theOscines.More or less cosmopolitan groups like herons,Falconidae, Anseres, Columbae, &c., and circumtropical families likeParridae, Trogonidae, Capitonidae, are to be excluded from these lists as indifferent. The differences between the Neotropical avifauna and that of North America are fundamental and prove the independence or superior value of the Neotropical region as one of the principal realms.It is difficult to subdivide the Neotropical region into subregions; the best suggestion is that of Newton:Antillean, with the exception of the islands of Trinidad and Tobago, as well as those which lie on the northern coast of South America;Patagonian, including Chile and part of Peru;Columbian, comprising the rest of the continent and also Central America.TheAntillean Subregionis in many respects one of the most suggestive and interesting, comparatively small though it be. For narrow as are the channels between Cuba and the opposite coast of Central America, between the Bahamas and Florida, and between Grenada and Tobago, the fauna of the Antillean chain, instead of being a mixture of that of the almost contiguous countries, differs much from all, and exhibits in some groups a degree of speciality which may be not unfitly compared with that of oceanic islands. Except such as are of coral formation, the Antilles are hilly, not to say mountainous, their summits rising in places to an elevation of 8000 ft., and nearly all, prior to their occupation by Europeans, were covered with luxuriant forest, which, assisting in the collection and condensation of the clouds brought by the trade winds, ensured its own vitality by precipitating frequent and long-continued rains upon the fertile soil. Under such conditions we might expect to find an extremely plentiful animal population, one as rich as that which inhabits the same latitudes in Central America, not many degrees farther to the west; but no instance perhaps can be cited which shows more strikingly the difference between a continental and an insular fauna, since, making every allowance for the ravages of cultivation by civilized man, the contrary is the case, and possibly no area of land so highly favoured by nature is so poorly furnished with the higher forms of animal life. Here, as over so large a portion of the Australian region, we find birds constituting the supreme class—the scarcity of mammals being accounted for in some measure as a normal effect of insularity.There is one peculiar subfamily,Todinae, represented by only four species ofTodus. We note the absence ofRatitae, Tinami, Cracidae, Rhamphastidae, and any of those gruiform genera which are so characteristic of the continent. There is no family of birds common to the Nearctic area and the Antillean subregion without occurring also in other parts of the Neotropical region, a fact which proves its affinity to the latter.ThePatagonian Subregion, most extratropical, is naturally devoid of a good many typically tropical birds, or these are but poorly represented, for instanceCaerebidae, Mniotiltidae, Tanagridae, Vireonidae. On the other hand some of the most characteristicfeatures of the whole region are here well represented,e.g.Rhea, Tinami, Chauna, Dicholophus, Attagis, Pteroptochidae, and indeed therein we find some of the best evidence of the antiquity of its population, both recent and extinct (cf. the numerous fossils of the Santa Cruz formation), and also the nearest resemblance to the fauna of Austrogaea.(C)Arctogaeais Huxley’s well-chosen term for all the rest of the world (including the Nearctic, Palaearctic, Indian and Ethiopian regions of P.L. Sclater) in opposition to Notogaea. Faunistically, although not geographically, the Nearctic and Palaearctic areas must form the two subdivisions of one great unit, for which the “Holarctic region” is now the generally accepted term.TheHolarctic Region, comprising North America and the extratropical mass of land of the Old World, may from an ornithological point of view be characterized by the Colymbi,Alcidae, Gallidaeor Alectoropodous Galli, and the Oscines, which have here reached their highest development; while Ratitae, Tinami, Psittaci, and non-Oscine Passeres (with the exception ofTyrannidaeextending into North America andConurus carolinensis) are absent.Nearctic Subregion.—The close affinity of North America with the Palaearctic avifauna becomes at once apparent if we exclude those groups of birds which we have good reason to believe have their original home in the Neotropical region, notably numerousTyrannidae, humming-birds and the turkey-buzzards.The following groups may be mentioned as characteristic and typically American, and, since we consider them as comparatively recent immigrants into the Neotropical region, as originally peculiar to the Nearctic area:Mniotiltidae, Vireonidae, Icteridae, Meleagrisand variousTetraoninae. Restricted to and peculiar to the sub-region is only the little Oscine family ofChamaeidae, restricted to the coast district of California. “More than one-third of the genera of Nearctic birds are common also to the Palaearctic subregion. If we take the number of Nearctic species at 700, which is perhaps an exaggeration, and that of the Palaearctic at 850, we find that, exclusive stragglers, there are about 120 common to the two areas. Nearly 20 more are properly Palaearctic, but occasionally occur in America, and about 50 are Nearctic, which from time to time stray to Europe or Asia. This, however, is by no means the only point of resemblance. Of many genera, the so-called species found in the New World are represented in the Old by forms so like them that often none but an expert can distinguish them, and of such representative ‘species’ about 80 might be enumerated” (Newton,Dict. Birds, p. 335).Of the many attempts to subdivide the Nearctic subregion, the same authority favours that of Dr S.F. Baird, who distinguishes betweenCanadian, Alleghanian, Middleor Missourian,CalifornianandAlaskanprovinces. Dr Hart Merriam takes the broad point of view “that the whole of extratropical North America consists of but two primary life regions, aBorealregion, which is circumpolar, and aSonoranor Mexican tableland region which is unique.” The first of these supports Newton’s contention of the essential unity of the Nearctic and Palaearctic areas. In any case the various Nearctic subdivisions completely merge into each other, just as is to be expected from the physical configuration and other bionomic conditions of the Nort American continent.ThePalaearctic Subregionis, broadly speaking, Europe and Asia, with the exception of India and China. The propriety of comprehending this enormous tract in one zoological “region” was first shown by Dr P.L. Sclater, and as regards the distribution of most classes of animals there have been few to doubt that it is an extremely natural one. Not indeed altogether so homogeneous as the Nearctic area, it presents, however, even at its extreme points, no very striking difference between the bulk of its birds. Though Japan is far removed from western Europe, and though a few generic forms and still fewer families inhabit the one without also frequenting the other, yet there is a most astonishing similarity in a large portion of their respective birds. In some cases the closest examination has failed to detect any distinction that may be called specific between the members of their avifauna; but in most it is possible to discover just sufficient difference to warrant a separation of the subjects. Nevertheless, it is clear that in Japan we have, as it were, a repetition of some of our most familiar species—the redbreast and the hedge-sparrow, for example—slightly modified in plumage or otherwise, so as to furnish instances of the most accurate representation,e.g.Cyanopica cookiof Portugal and Spain, andC. cyanaof Amoorland and Japan.Like the Nearctic the Palaearctic subregion seems to possess but one single peculiar family of land birds, thePanuridae, represented by the beautiful species known to Englishmen as the bearded tit-mouse,Panurus biarmicus. The entire number of Palaearctic families are, according to Newton, 67, and of the genera 323. Of these 128 are common to the Nearctic subregion. Species of 51 more seem to occur as true natives within the Ethiopian and Indian regions, and besides these 18 appear to be common to the Ethiopian without being found in the Indian, and no fewer than 71 to the Indian without occurring in the Ethiopian. To compare the Palaearctic genera with those of the Australian and Neotropical regions would be simply a waste of time, for the points of resemblance are extremely few, and such as they are they lead to nothing. It will therefore be seen from the above that next to the Nearctic are the Palaearctic has a much greater affinity to any other, a fact which might be expected from geographical considerations.Having shown this much we have next to deal with the peculiarities of the vast Palaearctic subregion. At the lowest computation 37 genera seem to be peculiar to it, though it is certain that species of several are regularly wont to wander beyond its limits in winter seeking a southern climate. Of the peculiar genera only a few examples may be mentioned:Eurynorhynchus, the spoon-billed sandpiper of Siberia;Syrrhaptes, the sandgrouse of central Asia;Musicicapaof Europe.We distinguish between aSiberian, Mongolian, MediterraneanandEuropean province, none of which can be well defined. The islands of the Canaries, Madeira and the Azores belong to the Mediterranean province, and offer some peculiarities of great interest. The Azores have been monographed by F.D. Godman (Nat. Hist. of the Azores or Western Islands, London, 1870). There is a general tendency among these insular birds to vary more or less from their continental representatives, and this is especially shown by the former having always darker plumage and stronger bills and legs. In one instance the variation is so excessive that it fully justifies the establishment of a specific distinction. This is the case of the bullfinch of the more western of these islands (Pyrrhula murina), the male of which, instead of the ruddy breast of its well-known congener (P. vulgaris), has that part of a sober mouse-colour. A similar sombre hue distinguishes the peculiar chaffinch of the Canary Islands (Fringilla teydea), but to these islands as well as the Azores and Madeiras there belongs in common another chaffinch (F. tintillon) which, though very nearly allied to that of Mauritania (F. spodogenia) is perfectly recognizable, and not found elsewhere. Madeira has also its peculiar golden-crested wren (Regulus maderensis), and its peculiar pigeon (Columba trocaz), while two allied forms of the latter (C. laurivoraandC. bollii) are found only in the Canaries. Further on this subject we must not go; we can only state that Godman has shown good reason for declaring that the avifauna of all these islands is the effect of colonization extending over a long period of years, and going on now.Palaeotropical Region.—Much can be said in favour of combining the mostly tropical portion of the great mass of land of the Old World (excluding, of course, Austrogaea or the Australian region) into one region, for which Oscar Drude’s well-chosen term “palaeotropical” has been adopted (cf. Bronn’sThierreich, System Part.p. 296, 1893). This region naturally comprises the African and Indian areas, conformably to be called subregions.Both subregions possess, besides others, the following characteristic birds: Ratitae, viz.Struthioin Africa and Arabia, fossil also in the Sivalik Hills, andAepyornithidaein Madagascar;Pittidae, BucerotinaeandUpupinae, of whichUpupaitself in India, Madagascar and Africa;Coraciidae; Pycnonotidaeor bulbuls;Trogonidae, of which the Asiatic genera are the less specialized in opposition to the Neotropical forms;Vulturidae; Leptoptilus, AnastomusandCiconiaamong the storks;Pteroclidae; Treroninaeamong pigeons. Of other families which, however, extend their range more or less far into the Australian realm, may be mentionedOtididae, the bustards;Meropidaeor bee-eaters;Muscicapidaeor flycatchers;Sturnidaeor starlings.TheEthiopian Subregioncomprises the whole of Africa and Madagascar, except the Barbary States, but including Arabia; in the north-east the subregion melts into the Palaearctic between its limits still farther to the eastwards, through Beluchistan and even beyond the Indus.So large a portion of the Ethiopian subregion lies between the tropics that no surprise need be expressed at the richness of its fauna relatively to that of the last two subregions we have considered. Between fifty and sixty so-called families of land birds alone are found within its limits, and of them at least nine are peculiar; the typical genera of which areBuphaga, Euryceros, Philepitta, Musophaga, Irrisor, Leptosoma, Colius, Serpentarius, Struthio, Aepyornis. It is singular that only the first three of them belong to the orderPasseriformes, a proportion which is not maintained in any other tropical region. The number of peculiar genera, besides those just mentioned, is too great for them to be named here; some of the most remarkable on the continent are:Balaeniceps, the whale-headed heron;Balaearica, the crowned crane;Podica, finfoot;Numidaand allied genera of guinea fowls.The natural division of the subregion is that into an African and a Madagascar province. Subdivision of the continental portion is beset with great difficulties, and none of the numerous attempts have proved long-lived. The forest-clad basin of the Congo, with the coastal districts of the bay of Guinea, seem to form one domain in opposition to the rest.The Malagasy province comprises, besides Madagascar, the Mascarene, Comoro and Seyehelle islands. It may be safely deemed the most peculiar area of the earth’s surface, while from the richness and multifariousness of its animal, and especially of its ornithic population, New Zealand cannot be compared with it. In A. Grandidier’s magnificentHistoire physique, naturelle et politique de Madagascar, vol. xii. (Paris, 1875-1884), are enumerated 238 species as belonging to the island, of which 129 are peculiar to it, and among those are no fewer than 35 peculiar genera.Eurycerosof theOscines, andPhilepittaof the Clamatores, are remarkable enough to form the types of Passeriform families, andMesiteshalf-way between Galli and Gruiformes is of prime importance. The PasserineFalculia, with its recently extinguished alliesFregilupusandNecropsarof the Mascarenes; the CoraciineBrachypteracias,AtelornisandGeobiastes, are very abundant, whileHeliodilusis an owl belonging to that subfamily which is otherwise represented only by the widely-spread barn owl,Strix flammea. Lastly must be noted the extinct tall Ratite species ofAepyorniswith its several fancy genera. But, as Newton charmingly puts it (Dict Birds, p. 353), the avifauna of Madagascar is not entirely composed of such singularities as these. We have homely genera, even among the truePasseres, occurring there—such asAlauda, Acrocephalus, MotacillaandPratincola, while theCisticola madagascariensisis only distinguishable from the well-known fan-tailed warbler,C. schoenicolaof Europe, Africa and India by its rather darker coloration. But there are also species, though not Passerine, which are absolutely identical with those of Britain, the barn owl, common quail, pigmy rail, and little grebe or dabchick, all of them common and apparently resident in the island. Mauritius had the dodo (q.v.),LophopsittacusandAphanapteryx. Rodriguez had the solitaire,NecropsittacusandNecropsar. Bourbon or Réunion hadFregilupus.Fig.23.—Extinct Starling of Reunion (Fregilupus varius), adapted from figures by Daubenton, Levaillant and others. Reduced.Some of the Malagasy avifauna is certainly ancient, aboriginal, and even points to India; other forms indicate clearly their African origin; while, lastly, such strikingly characteristic Indo-African birds as hornbills are unaccountably absent.TheOriental Subregioncomprises all the countries and numerous islands between the Palaearctic and Australian areas; it possesses upwards of seventy families, of which, however, only one is peculiar, but this family, theEurylaemidaeor broadbills, is of great importance since it represents all theSubclamatores. Of the many characteristic birds may be mentionedPycnonotidaeor bulbuls, of which thePhyllornithinaeare peculiar,Campephagidaeor cuckoo shrikes,Dicruridaeor drongos,Nectariniidaeor sunbirds; pheasants, together withPavoandGallus. Some of the similarities to the Ethiopian and the great differences from the Australian avifauna have already been pointed out. Naturally no line whatever can be drawn between the Oriental and the Palaearctic subregions, and many otherwise essentially Indo-Malayan families extend far into the Australian realm, far across Wallace’s line, whilst the reverse takes place to a much more moderate extent. Certainly the Oriental area, in spite of its considerable size, cannot possibly claim the standing of a primary region. It is a continuation of the great Arctogaea into the tropics.Following H.J. Elwes we subdivide the whole subregion into a Himalo-Chinese, Indian and Malayan province. These divisions had the approval of W.T. Blanford, who proposed the terms Cis- and Trans-gangetic for the two first. The Himalo-Chinese or Trans-gangetic province shows the characteristics of its avifauna also far away to the eastward in Formosa, Hainan and Cochin China, and again in a lesser degree to the southward in the mountains of Malacca and Sumatra. Indo-China is especially rich inEurylaemidae, China proper and the Himalayas in pheasants.The Indian or Cisgangetic province is the least rich of the three so far as peculiar genera are concerned.The Malayan province comprising the Malay islands, besides the Malay peninsula, and the very remarkable Philippines, possess an extraordinary number of peculiar and interesting genera.The influence of the Australian realm is indicated by a Megapode in Celebes, another in Borneo and Labuan, and a third in the Nicobar islands (which, however, like the Andamans, belong to the Indian province), but there are no cockatoos, these keeping strictly to the other side of Wallace’s line, whence we started on this survey of the world’s avifauna.
(A)Austrogaea, the Australian region in the wider sense, with the Papuan, Australian and New Zealand subregions, including also Polynesia. We may here quote Newton (Encyclopaedia Britannica, 9th ed., “Birds,” p. 738) on the remarkable differences between this region and the rest of the Old World:—“The prevalent zoological features of any Region are of two kinds—negative and positive. It is therefore just as much the business of the zoogeographer, who wishes to arrive at the truth, to ascertain what groups of animals are wanting in any particular locality (altogether independently of its extent) as to determine those which are forthcoming there. Of course, in the former case it would be absurd to regard as a physical feature of any great value the absence from a district of groups which do not occur except in its immediate neighbourhood; but when we find that certain groups, though abounding in some part of the vicinity, either suddenly cease from appearing or appear only in very reduced numbers, and occasionally in abnormal forms, the fact obviously has an important bearing. Now, mere geographical considerations, taken from the situation and configuration of the islands of the so-called Indian or Malay Archipelago, would indicate that they extended in an unbroken series from the shores of the Strait of Malacca to the southern coast of New Guinea, which confronts that of north Australia in Torres Strait, or even farther to the eastward. Indeed, the very name Australasia, often applied to this part of the world, would induce the belief that all the countless islands, be they large or small—and some of them are among the largest on the globe—were but a southern prolongation of the mainland of Asia. But so far from this being the case a very definite barrier is interposed. A strait, some 15 m. or so in width, and separating the two fertile but otherwise insignificant islands of Bali and Lombok, makes such a frontier as can hardly be shown to exist elsewhere. The former of these two islands belongs to the Indian Region, the latter to the Australian, and between them there is absolutely no true transition—that is, no species are common to both which cannot be easily accounted for by the various accidents and migrations that in the course of time must have tended to mingle the productions of islands so close to one another. The faunas of the two are as absolutely distinct as those of South America and Africa, and it is only because they are separated by a narrow strait instead of the broad Atlantic that they have become so slightly connected by the interchange of a few species and genera.
“Now, first, of the forms of birds which are prevalent throughout the Indian Region, but are entirely wanting in the Australian, we have at once the bulbuls (Ixidae), very characteristic of most parts of Africa and Asia, including the sub-groupPhyllornithinae, which is peculiar to the Indian Region; the widely-spread families of barbets (Megalaeminae) and vultures (Vulturidae); and the pheasants (Phasianidae), which attain so great a development in various parts of the Asiatic continent and islands that there must their home be regarded as fixed. Some naturalists would add the finches (Fringillidae), rightly if we assume that thePloceidaeor weavers constitute a separate family. Then, of forms which are but weakly represented, we have the otherwise abundant thrushes (Turdidae), and, above all, the woodpeckers (Picidae), of which only very few species, out of 400, just cross the boundary and occur in Lombok, Celebes or the Moluccas, but are unknown elsewhere in the region.”
But the Australian region is also remarkable for its ornithic singularity. All the existing Ratitae (with the exception of the ostriches of Africa and South America, belonging to the generaStruthioandRhea, and comprising at most but five species) are found in Austrogaea and nowhere else. Of the Passeres the honey-suckers (Meliphagidae) are most characteristic, and, abounding in genera and species, extend to almost every part of the region, yet only one species ofPtilotisoversteps its limits, crossing the sea from Lombok to Bali. Other peculiar families are much more confined. But the positive characteristics of the region as a whole are not its peculiar forms alone; there are at least four families which, being feebly represented elsewhere, here attain the maximum of development. Such are the thick-headed shrikes (Pachycephalidae), the caterpillar-eaters (Campephagidae), the flower-peckers (Dicaeidae), and the swallow-flycatchers (Artamidae). Besides these, three or perhaps four groups, though widely distributed throughout the world, arrive in the Australian region at their culmination, presenting an abundance of most varied forms. These are the weaver-birds (Ploceidae), and the moreporks (Podargidae), but especially the kingfishers (Alcedinidae) and the pigeons (Columbidae), the species belonging to the two last obtaining in this region a degree of prominence and beauty which is elsewhere unequalled.
The boundaries of the subregions are not well defined.
TheNew Zealand Subregion, considered by Professors Newton and Huxley and various other zoogeographers as deserving the rank of a region, is, and to all appearance has long been, more isolated than any other portion of the globe. Besides the three larger islands numerous satellites belong to the subregion, as Lord Howe, Norfolk and Kermadec islands, with the Chatham, Auckland and Macquarie groups. The main affinities of the avifauna are, of course, Australian. The most extraordinary feature is unquestionably the former existence of the gigantic Dinornithes or moas (q.v.) and, another family of Ratitae, the weird-looking kiwis or Apteryges, which are totally unlike any other existing birds. Of other peculiar genera it will suffice to mention only the more remarkable.Rallidaepresent the very noteworthy woodhens,Ocydromus, and the takahe,Notornis, which is almost extinct. The widely-spread plovers,Charadriidae, have two not less singular generic developments,Thinornis, and the extraordinary wrybill,Anarhynchus. There is an owl, type of the genusSceloglaux. Of parrots,Stringops, the kakapo or owl-parrot, is certainly peculiar, whileNestorconstitutes a peculiar subfamily of the brush-tongued parrots orTrichoglossidae.XenicusandAcanthosittaform a little family of truly mesomyodean Passeres Clamatores. Of theMeliphagidaethe generaProsthemadera,PogonornisandAnthornisare peculiar. The starlings,Sturnidae, are represented byCallaeas,Creadionand the very abnormalHeterolocha. The gallinaceous birds are represented by a quail,Coturnix novae zealandiae, now exterminated. A large flightless goose,Cnemiornis, allied to the AustralianCereopsis, and the gigantic rapaciousHarpagornis, have died out recently, with the moas. In all, there is a wonderful amount of specialization, though perhaps in a very straight line from generalized forms; but the affinity to Australian or Polynesian types is in many cases clearly traceable, and it cannot be supposed but that these last are of cognate origin with those of New Zealand. A very long period of isolation must have been required to produce the differences so manifestly to be observed, but a few forms seem at rare intervals to have immigrated, and this immigration would appear to be kept up to our own day, as shown by the instance ofZosterops lateralis, which is said to have lately made its first appearance, and to have established itself in the country, as well as by the fact of two cuckoos,the widely-rangingEudynamis taitensisandChrysococcyx lucidus, which are annual visitors.
Polynesiaforms, of course, part of Austrogaea. Its extent is so vast that it necessarily contains some peculiar, outlying forms, so to say forgotten, which in their long-continued isolation have specialized themselves. For instance, the kagu (Rhinochetus) of New Caledonia, a queerly specialized form with Gruine affinities pointing only to South America. The toothbilled pigeon (Didunculus) is restricted to Samoa. Most interesting is the avifauna of the Sandwich islands; entirely devoid of Psittaci and of Coraciiformes, these islands show an extraordinary development of its peculiar familyDrepanidae, which are probably of South or Central American descent.Acrulocercusis a Meliphagine, and a peculiar genus. There are a raven (Corvus), a coot (Fulica), the well-known Sandwich island goose (Bernicla sandvicensis), now very commonly domesticated in Europe; and some flycatchers and thrushlike birds.
TheAustralian Subregioncomprises Australia and Tasmania. In the north it is influenced, of course, by its proximity to Papuasia, whence there is a considerable admixture of genera which do not proceed beyond the tropics, and of theseCasuariusis a striking example. The Cape York peninsula practically belongs to Papuasia. As a whole, Australia is rich in parrots, of which it has several very peculiar forms, but Picarians in old-fashioned parlance, of all sorts—certain kingfishers excepted—are few in number, and the pigeons are also comparatively scarce, no doubt because of the many arboreal predaceous marsupials. The continent, however, possesses the two important genera of thePseudoscines, namely the lyre-birds (Menura) and the scrub-birds (Atrichia). Among the more curious forms of other land-birds may be especially mentioned theMegapodiidae,LipoaandTalegallus, the railTribonyxandPedionomus, which represents the otherwise palaeotropicalTurnicesin Australia. The presence of bustards (Eupodotis) is a curious example of interrupted distribution, since none other of theOtididaeare found nearer than India. The Ratitae are represented by two species of emeu (Dromaeus), besides the cassowary of Cape York peninsula, and the extinctDromornisandGenyorniswith its enormous skull.
ThePapuan Subregion, chiefly New Guinea with its dependencies, the Timor group of islands, the Moluccas and Celebes. On the whole its avifauna presents some very remarkable features. Its most distinctive characteristic is the presence of the birds of paradise, which are almost peculiar to it; for, granting that the bower-birds,Chlamydoderaand others, of Australia, belong to the same family, they are far less highly specialized than the beautiful and extraordinary forms which are found, within very restricted limits, in the various islands of the subregion. Another chief feature is the extraordinary development of the cassowaries, the richness and specialization of the kingfishers, parrots, pigeons, honey-suckers and some remarkable flycatchers. It has several marked deficiencies compared with Australia, among which are the babblers (Timeliidae), weaver birds (Ploceidae), thePlatycercinaeamong parrots, diurnal birds of prey and the emeus. As a whole, the birds of Papua are remarkable for their brilliance of plumage, or their metallic colouring. The birds of paradise, the racquet-tailed kingfishers,Tanysiptera, the largest and smallest of parrots,CalyptdrhynchiisandNasiterna, and the great crowned pigeons,Goura, are very characteristic; and so are the various Megapodes.
(B)Neogaea, or theNeotropical region.—Excepting towards the north, where, in Mexico, it meets, and inosculates with the Nearctic subregion, the boundaries of the Neotropical region are simple enough to trace, comprehending as it does the whole of South America and all Central America; besides including the Falkland islands to the south-east and the Galapagos under the equator to the west, as well as the Antilles or West India islands up to the Florida channel.
Owing to the comparatively scanty number of harmful mammalian types, the birds play a considerable part in this large region, and some authorities consider its avifauna the richest in the world. The entire number of species amounts to about 3600. Of these 2000, or a good deal more than half, belong to the order Passeriformes. But the characteristic nature of the avifauna is more clearly brought out when we learn that of the 2000 species just mentioned only about 1070 belong to the higher suborder of Oscines, that means to say, nearly one-half belong to the lower suborder Clamatores. This is a state of things which exists nowhere else; for except in Australia, where a few indigenous and peculiar low non-Oscines are found, and in the Nearctic country, whither one family of Clamatores, viz. theTyrannidae, has evidently been led by the geographical continuity of its soil with that of the Neotropical region, such forms do not occur elsewhere. Accordingly their disproportionate prevalence in South America points unerringly to the lower rank of the avifauna of the region as a whole, and therefore to the propriety of putting it next in order to that of the Australian region, the general fauna of which is admittedly the lowest in the world. Huxley has urged with his wonted perspicuity the alliance of these two regions asNotogaea, basing his opinion, besides other weighty evidence, in great measure on the evidence afforded by the two main sections of the Galli, viz. thePeristeropodesand theAlectoropodes, the former composed of the familiesMegapodiidae, almost wholly Australian, and theCracidae, entirely Neotropical. (Cf.P.Z.S., 1868. pp. 294-319.)
Leaving, however, this matter as in some degree hypothetical, we have as genera, families, or perhaps even larger groups, a great many very remarkable forms which are characteristic of, or peculiar to, the Neotropical region in part, if not as a whole. Of families we find twenty-three, or maybe more, absolutely restricted thereto, besides at least eight which, being peculiar to the New World, extend their range into the Nearctic region, but are there so feebly developed that their origin may be safely ascribed to the southern portion of America. First in point of importance comes the extraordinarily beautiful family of humming-birds (Trochilidae), with nearly 150 genera (of which only three occur in the Nearctic region) and more than 400 species. Then the tyrants (Tyrannidae), with more than seventy genera (ten of which range into the northern region), and over 300 species. To these follow the tanagers (Tanagndae), with upwards of forty genera (only one of which crosses the border), and about 300 species; the piculules (Dendrocolaptidae), with as many genera, and over 200 species; the ant-thrushes, (Formicariidae), with more than thirty genera, and nearly 200 species; together with other groups which, if not so large as those just named, are yet just as well defined, and possibly more significant, namely, the tapaculos (Pteroptochidae), the toucans (Rhamphastidae), the jacamars (Galbulidae), the motmots (Monotidae), the todies (Todidae), the trumpeters (Psophiidae), and the screamers (Palamedeidae); besides such isolated forms as the seriema (Cariama), and the sun-bittern (Eurypyga).
The nature of the South American avifauna will perhaps become still more evident if we arrange the characteristic members as follows:—
1. Birds which are restricted to, probably indigenous of the region:Rhea;PalamedeaandChauna, the screamers;Tinami;Psophia, Dicholophus, Eurypyga, Heliornisof the Gruiform assembly;ThinocorysandAttagis; Cracidae; Opisthocomus; of parrotsAraandConuruswith their allies;Monotidae, incl.Todus; Steatornis; GalbulinaeandBucconinae; Rhamphastidae; Formicariidae, Pteroptochidae, and of theTyrannidaetheCotinginae.
2. Birds which are indigenous, but extend far into North America:Cathartae, Trochilidae, Tyrannidae.
3. Birds which are originally immigrants from North America:Podicipedidae, with the flightlessCentropelmaon Lake Titicaca;Ceryle, the only genus of kingfishers in the New World; all theOscines.
More or less cosmopolitan groups like herons,Falconidae, Anseres, Columbae, &c., and circumtropical families likeParridae, Trogonidae, Capitonidae, are to be excluded from these lists as indifferent. The differences between the Neotropical avifauna and that of North America are fundamental and prove the independence or superior value of the Neotropical region as one of the principal realms.
It is difficult to subdivide the Neotropical region into subregions; the best suggestion is that of Newton:Antillean, with the exception of the islands of Trinidad and Tobago, as well as those which lie on the northern coast of South America;Patagonian, including Chile and part of Peru;Columbian, comprising the rest of the continent and also Central America.
TheAntillean Subregionis in many respects one of the most suggestive and interesting, comparatively small though it be. For narrow as are the channels between Cuba and the opposite coast of Central America, between the Bahamas and Florida, and between Grenada and Tobago, the fauna of the Antillean chain, instead of being a mixture of that of the almost contiguous countries, differs much from all, and exhibits in some groups a degree of speciality which may be not unfitly compared with that of oceanic islands. Except such as are of coral formation, the Antilles are hilly, not to say mountainous, their summits rising in places to an elevation of 8000 ft., and nearly all, prior to their occupation by Europeans, were covered with luxuriant forest, which, assisting in the collection and condensation of the clouds brought by the trade winds, ensured its own vitality by precipitating frequent and long-continued rains upon the fertile soil. Under such conditions we might expect to find an extremely plentiful animal population, one as rich as that which inhabits the same latitudes in Central America, not many degrees farther to the west; but no instance perhaps can be cited which shows more strikingly the difference between a continental and an insular fauna, since, making every allowance for the ravages of cultivation by civilized man, the contrary is the case, and possibly no area of land so highly favoured by nature is so poorly furnished with the higher forms of animal life. Here, as over so large a portion of the Australian region, we find birds constituting the supreme class—the scarcity of mammals being accounted for in some measure as a normal effect of insularity.
There is one peculiar subfamily,Todinae, represented by only four species ofTodus. We note the absence ofRatitae, Tinami, Cracidae, Rhamphastidae, and any of those gruiform genera which are so characteristic of the continent. There is no family of birds common to the Nearctic area and the Antillean subregion without occurring also in other parts of the Neotropical region, a fact which proves its affinity to the latter.
ThePatagonian Subregion, most extratropical, is naturally devoid of a good many typically tropical birds, or these are but poorly represented, for instanceCaerebidae, Mniotiltidae, Tanagridae, Vireonidae. On the other hand some of the most characteristicfeatures of the whole region are here well represented,e.g.Rhea, Tinami, Chauna, Dicholophus, Attagis, Pteroptochidae, and indeed therein we find some of the best evidence of the antiquity of its population, both recent and extinct (cf. the numerous fossils of the Santa Cruz formation), and also the nearest resemblance to the fauna of Austrogaea.
(C)Arctogaeais Huxley’s well-chosen term for all the rest of the world (including the Nearctic, Palaearctic, Indian and Ethiopian regions of P.L. Sclater) in opposition to Notogaea. Faunistically, although not geographically, the Nearctic and Palaearctic areas must form the two subdivisions of one great unit, for which the “Holarctic region” is now the generally accepted term.
TheHolarctic Region, comprising North America and the extratropical mass of land of the Old World, may from an ornithological point of view be characterized by the Colymbi,Alcidae, Gallidaeor Alectoropodous Galli, and the Oscines, which have here reached their highest development; while Ratitae, Tinami, Psittaci, and non-Oscine Passeres (with the exception ofTyrannidaeextending into North America andConurus carolinensis) are absent.
Nearctic Subregion.—The close affinity of North America with the Palaearctic avifauna becomes at once apparent if we exclude those groups of birds which we have good reason to believe have their original home in the Neotropical region, notably numerousTyrannidae, humming-birds and the turkey-buzzards.
The following groups may be mentioned as characteristic and typically American, and, since we consider them as comparatively recent immigrants into the Neotropical region, as originally peculiar to the Nearctic area:Mniotiltidae, Vireonidae, Icteridae, Meleagrisand variousTetraoninae. Restricted to and peculiar to the sub-region is only the little Oscine family ofChamaeidae, restricted to the coast district of California. “More than one-third of the genera of Nearctic birds are common also to the Palaearctic subregion. If we take the number of Nearctic species at 700, which is perhaps an exaggeration, and that of the Palaearctic at 850, we find that, exclusive stragglers, there are about 120 common to the two areas. Nearly 20 more are properly Palaearctic, but occasionally occur in America, and about 50 are Nearctic, which from time to time stray to Europe or Asia. This, however, is by no means the only point of resemblance. Of many genera, the so-called species found in the New World are represented in the Old by forms so like them that often none but an expert can distinguish them, and of such representative ‘species’ about 80 might be enumerated” (Newton,Dict. Birds, p. 335).
Of the many attempts to subdivide the Nearctic subregion, the same authority favours that of Dr S.F. Baird, who distinguishes betweenCanadian, Alleghanian, Middleor Missourian,CalifornianandAlaskanprovinces. Dr Hart Merriam takes the broad point of view “that the whole of extratropical North America consists of but two primary life regions, aBorealregion, which is circumpolar, and aSonoranor Mexican tableland region which is unique.” The first of these supports Newton’s contention of the essential unity of the Nearctic and Palaearctic areas. In any case the various Nearctic subdivisions completely merge into each other, just as is to be expected from the physical configuration and other bionomic conditions of the Nort American continent.
ThePalaearctic Subregionis, broadly speaking, Europe and Asia, with the exception of India and China. The propriety of comprehending this enormous tract in one zoological “region” was first shown by Dr P.L. Sclater, and as regards the distribution of most classes of animals there have been few to doubt that it is an extremely natural one. Not indeed altogether so homogeneous as the Nearctic area, it presents, however, even at its extreme points, no very striking difference between the bulk of its birds. Though Japan is far removed from western Europe, and though a few generic forms and still fewer families inhabit the one without also frequenting the other, yet there is a most astonishing similarity in a large portion of their respective birds. In some cases the closest examination has failed to detect any distinction that may be called specific between the members of their avifauna; but in most it is possible to discover just sufficient difference to warrant a separation of the subjects. Nevertheless, it is clear that in Japan we have, as it were, a repetition of some of our most familiar species—the redbreast and the hedge-sparrow, for example—slightly modified in plumage or otherwise, so as to furnish instances of the most accurate representation,e.g.Cyanopica cookiof Portugal and Spain, andC. cyanaof Amoorland and Japan.
Like the Nearctic the Palaearctic subregion seems to possess but one single peculiar family of land birds, thePanuridae, represented by the beautiful species known to Englishmen as the bearded tit-mouse,Panurus biarmicus. The entire number of Palaearctic families are, according to Newton, 67, and of the genera 323. Of these 128 are common to the Nearctic subregion. Species of 51 more seem to occur as true natives within the Ethiopian and Indian regions, and besides these 18 appear to be common to the Ethiopian without being found in the Indian, and no fewer than 71 to the Indian without occurring in the Ethiopian. To compare the Palaearctic genera with those of the Australian and Neotropical regions would be simply a waste of time, for the points of resemblance are extremely few, and such as they are they lead to nothing. It will therefore be seen from the above that next to the Nearctic are the Palaearctic has a much greater affinity to any other, a fact which might be expected from geographical considerations.
Having shown this much we have next to deal with the peculiarities of the vast Palaearctic subregion. At the lowest computation 37 genera seem to be peculiar to it, though it is certain that species of several are regularly wont to wander beyond its limits in winter seeking a southern climate. Of the peculiar genera only a few examples may be mentioned:Eurynorhynchus, the spoon-billed sandpiper of Siberia;Syrrhaptes, the sandgrouse of central Asia;Musicicapaof Europe.
We distinguish between aSiberian, Mongolian, MediterraneanandEuropean province, none of which can be well defined. The islands of the Canaries, Madeira and the Azores belong to the Mediterranean province, and offer some peculiarities of great interest. The Azores have been monographed by F.D. Godman (Nat. Hist. of the Azores or Western Islands, London, 1870). There is a general tendency among these insular birds to vary more or less from their continental representatives, and this is especially shown by the former having always darker plumage and stronger bills and legs. In one instance the variation is so excessive that it fully justifies the establishment of a specific distinction. This is the case of the bullfinch of the more western of these islands (Pyrrhula murina), the male of which, instead of the ruddy breast of its well-known congener (P. vulgaris), has that part of a sober mouse-colour. A similar sombre hue distinguishes the peculiar chaffinch of the Canary Islands (Fringilla teydea), but to these islands as well as the Azores and Madeiras there belongs in common another chaffinch (F. tintillon) which, though very nearly allied to that of Mauritania (F. spodogenia) is perfectly recognizable, and not found elsewhere. Madeira has also its peculiar golden-crested wren (Regulus maderensis), and its peculiar pigeon (Columba trocaz), while two allied forms of the latter (C. laurivoraandC. bollii) are found only in the Canaries. Further on this subject we must not go; we can only state that Godman has shown good reason for declaring that the avifauna of all these islands is the effect of colonization extending over a long period of years, and going on now.
Palaeotropical Region.—Much can be said in favour of combining the mostly tropical portion of the great mass of land of the Old World (excluding, of course, Austrogaea or the Australian region) into one region, for which Oscar Drude’s well-chosen term “palaeotropical” has been adopted (cf. Bronn’sThierreich, System Part.p. 296, 1893). This region naturally comprises the African and Indian areas, conformably to be called subregions.
Both subregions possess, besides others, the following characteristic birds: Ratitae, viz.Struthioin Africa and Arabia, fossil also in the Sivalik Hills, andAepyornithidaein Madagascar;Pittidae, BucerotinaeandUpupinae, of whichUpupaitself in India, Madagascar and Africa;Coraciidae; Pycnonotidaeor bulbuls;Trogonidae, of which the Asiatic genera are the less specialized in opposition to the Neotropical forms;Vulturidae; Leptoptilus, AnastomusandCiconiaamong the storks;Pteroclidae; Treroninaeamong pigeons. Of other families which, however, extend their range more or less far into the Australian realm, may be mentionedOtididae, the bustards;Meropidaeor bee-eaters;Muscicapidaeor flycatchers;Sturnidaeor starlings.
TheEthiopian Subregioncomprises the whole of Africa and Madagascar, except the Barbary States, but including Arabia; in the north-east the subregion melts into the Palaearctic between its limits still farther to the eastwards, through Beluchistan and even beyond the Indus.
So large a portion of the Ethiopian subregion lies between the tropics that no surprise need be expressed at the richness of its fauna relatively to that of the last two subregions we have considered. Between fifty and sixty so-called families of land birds alone are found within its limits, and of them at least nine are peculiar; the typical genera of which areBuphaga, Euryceros, Philepitta, Musophaga, Irrisor, Leptosoma, Colius, Serpentarius, Struthio, Aepyornis. It is singular that only the first three of them belong to the orderPasseriformes, a proportion which is not maintained in any other tropical region. The number of peculiar genera, besides those just mentioned, is too great for them to be named here; some of the most remarkable on the continent are:Balaeniceps, the whale-headed heron;Balaearica, the crowned crane;Podica, finfoot;Numidaand allied genera of guinea fowls.
The natural division of the subregion is that into an African and a Madagascar province. Subdivision of the continental portion is beset with great difficulties, and none of the numerous attempts have proved long-lived. The forest-clad basin of the Congo, with the coastal districts of the bay of Guinea, seem to form one domain in opposition to the rest.
The Malagasy province comprises, besides Madagascar, the Mascarene, Comoro and Seyehelle islands. It may be safely deemed the most peculiar area of the earth’s surface, while from the richness and multifariousness of its animal, and especially of its ornithic population, New Zealand cannot be compared with it. In A. Grandidier’s magnificentHistoire physique, naturelle et politique de Madagascar, vol. xii. (Paris, 1875-1884), are enumerated 238 species as belonging to the island, of which 129 are peculiar to it, and among those are no fewer than 35 peculiar genera.Eurycerosof theOscines, andPhilepittaof the Clamatores, are remarkable enough to form the types of Passeriform families, andMesiteshalf-way between Galli and Gruiformes is of prime importance. The PasserineFalculia, with its recently extinguished alliesFregilupusandNecropsarof the Mascarenes; the CoraciineBrachypteracias,AtelornisandGeobiastes, are very abundant, whileHeliodilusis an owl belonging to that subfamily which is otherwise represented only by the widely-spread barn owl,Strix flammea. Lastly must be noted the extinct tall Ratite species ofAepyorniswith its several fancy genera. But, as Newton charmingly puts it (Dict Birds, p. 353), the avifauna of Madagascar is not entirely composed of such singularities as these. We have homely genera, even among the truePasseres, occurring there—such asAlauda, Acrocephalus, MotacillaandPratincola, while theCisticola madagascariensisis only distinguishable from the well-known fan-tailed warbler,C. schoenicolaof Europe, Africa and India by its rather darker coloration. But there are also species, though not Passerine, which are absolutely identical with those of Britain, the barn owl, common quail, pigmy rail, and little grebe or dabchick, all of them common and apparently resident in the island. Mauritius had the dodo (q.v.),LophopsittacusandAphanapteryx. Rodriguez had the solitaire,NecropsittacusandNecropsar. Bourbon or Réunion hadFregilupus.
Some of the Malagasy avifauna is certainly ancient, aboriginal, and even points to India; other forms indicate clearly their African origin; while, lastly, such strikingly characteristic Indo-African birds as hornbills are unaccountably absent.
TheOriental Subregioncomprises all the countries and numerous islands between the Palaearctic and Australian areas; it possesses upwards of seventy families, of which, however, only one is peculiar, but this family, theEurylaemidaeor broadbills, is of great importance since it represents all theSubclamatores. Of the many characteristic birds may be mentionedPycnonotidaeor bulbuls, of which thePhyllornithinaeare peculiar,Campephagidaeor cuckoo shrikes,Dicruridaeor drongos,Nectariniidaeor sunbirds; pheasants, together withPavoandGallus. Some of the similarities to the Ethiopian and the great differences from the Australian avifauna have already been pointed out. Naturally no line whatever can be drawn between the Oriental and the Palaearctic subregions, and many otherwise essentially Indo-Malayan families extend far into the Australian realm, far across Wallace’s line, whilst the reverse takes place to a much more moderate extent. Certainly the Oriental area, in spite of its considerable size, cannot possibly claim the standing of a primary region. It is a continuation of the great Arctogaea into the tropics.
Following H.J. Elwes we subdivide the whole subregion into a Himalo-Chinese, Indian and Malayan province. These divisions had the approval of W.T. Blanford, who proposed the terms Cis- and Trans-gangetic for the two first. The Himalo-Chinese or Trans-gangetic province shows the characteristics of its avifauna also far away to the eastward in Formosa, Hainan and Cochin China, and again in a lesser degree to the southward in the mountains of Malacca and Sumatra. Indo-China is especially rich inEurylaemidae, China proper and the Himalayas in pheasants.
The Indian or Cisgangetic province is the least rich of the three so far as peculiar genera are concerned.
The Malayan province comprising the Malay islands, besides the Malay peninsula, and the very remarkable Philippines, possess an extraordinary number of peculiar and interesting genera.
The influence of the Australian realm is indicated by a Megapode in Celebes, another in Borneo and Labuan, and a third in the Nicobar islands (which, however, like the Andamans, belong to the Indian province), but there are no cockatoos, these keeping strictly to the other side of Wallace’s line, whence we started on this survey of the world’s avifauna.
D. Classification of Birds
Fürbringer’s great work, published in the year 1888 by theNatura Artis MagistraSociety of Amsterdam, enabled Gadow not only to continue for the next five years the same lines of morphological research, but also further to investigate those questions which were still left in abeyance or seemed to require renewed study. The resulting “classification is based on the examination, mostly autoptic, of a far greater number of characters than any that had preceded it; moreover, they were chosen in a different way, discernment being exercised in sifting and weighing them, so as to determine, so far as possible, the relative value of each, according as that value may vary in different groups, and not to produce a mere mechanical ‘key’ after the fashion become of late years so common” (Newton’sDictionary of Birds, Introduction, p. 103). It is not the quantity but the quality of the anatomical and bionomic characters which determines their taxonomic value, and a few fundamental characters are better indications of the affinities of given groups of birds than a great number of agreements if these can be shown to be cases of isomorphism or heterophyletic, convergent analogy. Nature possesses three great educational or developmental schools—terrestrial, aquatic and aerial life. Each of these affords animal, vegetable or mixed diet. Animal diet implies the greatest variety with regard to locality and the modes of procuring the food. Each of these schools impresses its pupils, in the case of the birds, with its own stamp, but there are many combinations, since in the course of phyletic development many a group of birds has exchanged one school for another. Originally terrestrial groups have taken to an entirely aquatic life, andvice versâ; others, originally endowed with the power of flight, have become, or are transforming themselves into, absolutely cursorial forms; some members of one group live entirely on seeds, while others have become fierce fishers, and so forth. Only by the most careful inquiry into their history can their relationship or pedigree be unravelled. A statement may now be given of Gadow’s classification of birds, in which the extinct forms have been intercalated so far as possible. The few characters assigned to the various groups are sufficiently diagnostic when taken together, although they are not always those upon which the classification has been established:—
Class AVES
I. Sub-classArchaeornithes.—The three fingers and their metacarpals remain separate, each with a claw. Well-developed remiges. Both jaws with alveolar teeth. Amphicoelous. Caudal vertebrae more than thirteen, without a pygostyle, but with about twelve pairs of rectrices.Archaeopteryx, A. lithographica, s. macroura, two specimens from the upper Oolite of Solenhofen, Bavaria.
II. Sub-classNeornithes.—Metacarpals fused. Second finger the longest. Not more than thirteen caudal vertebrae.
I. DivisionRatitae.—Terrestrial, flightless. Without sternal keel. Quadrate bone with single proximal knob. Without pygostyle. Coracoid and scapula fused. Compound rhamphotheca. Adult without apteria. With copulatory organ. A collective polyphyletic or heterogeneous group, originally cosmopolitan; with certainty existing since the Miocene.1. OrderStruthiones.—With pubic symphysis. Two toes only, third and fourth.Struthio, ostrich, Pliocene of Samos and of north-west India, now Africa and Arabia.2. OrderRheae.—With long ischiadic symphysis. Three toes.Mesembriornis, Miocene or Pliocene of Argentina. Rhea, South America.3. OrderCasuarii.—Three toes. Aftershaft as long as the other half.CasuariusandDromaeus, Australian.Hypselornis, Pliocene of Sivalik Hills.4. OrderApteryges.—Four toes. Bill long and slender.Apteryx, New Zealand.5. OrderDinornithes.—Three or four toes. Bill short. Anterior limbs extremely reduced.Dinornis, numerous species, recently extinct, New Zealand.6. OrderAepyornithes.—Aepyornis, recently extinct, Madagascar.To the Ratitae belong possibly also the imperfectly knownDiatryma, Eocene of New Mexico,GastornisandDasornis, Eocene of Europe,Genyornis, Pleistocene of Australia.II. DivisionOdontolcae.—Marine, flightless, without sternal keel. Upper and lower jaws with teeth in furrows. Cretaceous epoch.Enaliornis, England, vertebrae chiefly biconcave;Hesperornis, North America, vertebrae heterocoelous.III. DivisionCarinatae.—With keeled sternum.1. OrderIchthyornithes.—Power of flight well developed. Vertebrae still amphicoelous. With small pygostyle. Incisura ischiadica. With alveolar teeth. Cretaceous of Kansas.Ichthyornis, Apatornis.2. OrderColymbiformes.—Plantigrade, nidifugous, aquatic. All toes webbed, fourth largest, hallux short; metatarsus laterally compressed; tibia with high, pyramidal crest. Bill straight, pointed, with simple sheath.Sub-order 1.Colymbi, Divers. Front toes completely webbed. Holarctic.Colymbus.Sub-order 2.Podicipedes, Grebes. Toes lobated. Cosmopolitan.3. OrderSphenisciformes.—Nidicolous, marine. Flightless, wings transformed into rowing paddles.Sphenisci, penguins. Antarctic and southern temperate coasts. Since the Eocene.4. OrderProcellariiformes.—Well flying, pelagic, nidicolous. Hallux absent or vestigial. Rhamphotheca compound. Cosmopolitan.Tubinares, petrels and albatrosses.5. OrderCiconiiformes.—Swimmers or waders. Desmognathous, without basipterygoid processes; with one pair of sterno-tracheal muscles.Sub-order 1.Steganopodes.—Well flying, aquatic, nidicolous; with all the four toes webbed together. Rhamphotheca compound; cosmopolitan.Phaëthon, tropic-bird;Sula, gannet;Phalacrocorax, cormorant andPlotus, snake-bird;Fregata, frigate-bird; Pelecanus. Here alsoPelagornis, Miocene of France;Argillornisand probablyOdontopteryxfrom the London Clay.Sub-order 2.Ardeae.—Piscivorous, nidicolous, waders; with complicated hypotarsus and with long cervical apteria.Ardeidae, cosmopolitan; includingCancroma, Neotropical,Balaeniceps, Scopidae, Ethiopian.Proherodius, Eocene of England.Sub-order 3.Ciconiae.—Zoophagous, nidicolous, waders; with simple hypotarsus and without cervical apteria. Cosmopolitan.Ciconiidae, storks.Ibidae, ibises and spoonbills.Propelargus, Oligocene.Sub-order 4.Phoenicopteri.—Flamingos. Nidifugous, waders; with simple hypotarsus and without cervical apteria. Front toes completely webbed; hallux very short or absent; feed chiefly on small aquatic invertebrates.Phoenicopterus, cosmopolitan. OligoceneElornisand, allied,Palaelodus.6. OrderAnseriformes.—Desmognathous, nidifugous; with two pairs of sterno-tracheal muscles, with complete basipterygoid processes and with a penis.Sub-order 1.Palamedeae.—Screamers. Ribs without uncinate processes. Hypotarsus simple. Neotropical.Chauna, Palamedea.Sub-order 2.Anseres.—FamilyAnatidae. Hypotarsus complex.Anser, Anas, Cygnus, since Miocene.Cnemiornis, Pleistocene, New Zealand, flightless.7. OrderFalconiformes.—Birds of prey. Carnivorous, desmognathous, nidicolous, without functional caeca. Terrestrial, aerial.Sub-order 1.Cathartae.—American vultures. With nares perviae.Cathartes, turkey buzzards,Sarcorhamphus gryphus, condorGypagus papa, king vulture.Sub-order 2.Accipitres.—With nares imperviae.Serpentariidae, secretary-bird, Ethiopian; Miocene, France.Vulturidae, Old World vultures, excluding Australia.Falconidae, cosmopolitan, since the Eocene.Harpagornis, Pleistocene, New Zealand;Lithornis, Eocene, England.Pandionidae, ospreys or fish hawks, cosmopolitan.8. OrderTinamiformes.—Nidifugous, with incisura ischiadica, without pygostyle. Herbivorous, terrestrial, neotropical.Crypturi, tinamous.9. OrderGalliformes.—Schizognathous, herbivorous, terrestrial. With ten functional remiges. With strong spinae sterni.Sub-order 1.Mesites.—Without basipterygoid processes, and with large spina interna.Mesites, Madagascar.Sub-order 2.Turnices.—Hemipodes or button-quails. Nidifugous; vomer large; sternum without processus obliqui. Hallux absent or vestigial. Old World.Turnix, Pedionomus.Sub-order 3.Galli.—With large spina communis, and with large processus obliqui. Hallux functional.Megapodiidae, Australian region.Cracidae, curassows and guans, neotropical.Gallidae, cosmopolitan.Sub-order 4.Opisthocomi.—Arboreal, with long spina externa; without basipterygoid processes.Opisthocomushoatzin, Guiana, Venezuela and Amazon countries.10. OrderGruiformes.Legs of the wading type. Without basipterygoid processes. Without spina interna. Nidifugous. Essentially Schizognathous.Rallidae, cosmopolitan, since Oligocene.Rallus, Fulica, Ocydromus, &c.,Gallinula nesiotis, Tristan d’Acunha, flightless.Notornis, New Zealand, flightless, nearly extinct.Aptornis, New Zealand, flightless, extinct.Aphanapteryx(Mauritius) =Erythromachus(Rodriguez) =Diaphorapteryx(Chatham Island), flightless and recently extinct.Gypsornis, upper Eocene, France.Gruidae, cranes, cosmopolitan, alliedPhororhacos, Tertiary of Argentina.Dicholophidae, cariamas, neotropical.Otididae, bustards, Old World.Rhinochetidae, kagus, New Caledonia.Eurypygidae, sun-bittern, neotropical.Heliornithidae, finfoots, tropical.11. OrderCharadriiformes.—Schizognathous. With eleven remiges, of which the terminal very short. Aquinto-cubital. Spinae sterni short, separate.Sub-order 1.Limicolae.—Nidifugous, without spina interna sterni. Hypotarsus complicated.Charadriidae, plovers.Chionididae, sheath-bill.Glareolidae, wading swallows and coursers.Thinocorythidae, seed-snipes.Oedicnemididae, thick-knees.Parridae.Sub-order 2.Lari.—Aquatic, vomer complete. Without basipterygoid processes. Front toes webbed; hallux small or absent. Large supraorbital glands. Since Miocene.Laridae, gulls, cosmopolitan.Alcidae, auks, northern half of periarctic region.Sub-order 3.Pterocles.—Sand-grouse. Nidifugous. Vomer vestigial. With large crop and caeca. Hallux vestigial or absent since Oligocene. Africa to India, and Siberia.PteroclesandSyrrhaptes.Sub-order 4.Columbae.—Pigeons. Nidicolous. Vomer vestigial. With large crop, vestigial caeca.Columbidae, cosmopolitan, since Miocene.Dididae, flightless, recently extinct.Didus, dodo, Mauritius.Pezophaps, solitaire, Rodriguez.12. OrderCuculiformes.—Desmognathous, nidicolous; zygodactylous, or with the outer toe reversible.Sub-order 1.Cuculi.—Cuckoos. Quinto-cubital.Cuculidae, cosmopolitan.Musophagidae, plantain-eaters and touracos, Ethiopian since Miocene.Sub-order 2.Psittaci.—Parrots. Zygodactylous; aquinto-cubital. Cosmopolitan, chiefly tropical.Trichoglossidae, lories, Austro-Malayan.Nestor, New Zealand.Cyclopsittacus, Eos, Lorius, &c.Psittacidae, tongue smooth, incl.Stringops.13. OrderCoraciiformes.—Nidicolous. Nares imperviae, holorhinal. Downs restricted to the apteria or absent. Thirteen to fifteen cervical vertebrae. Mostly desmognathous. Deep plantar tendons connected with each other.Sub-order 1.Coraciae.—Either (1) with long spina externa sterni,Coraciidae, rollers, Old World.Momotidae, neotropical, motmots and todies.Alcedinidae, kingfishers, cosmopolitan or (2) with long spina communis.Meropidae, bee-eaters, Old World.Upupidae, Upupinae, hoopoes: palaearctic and palaeotropical.Bucerotinae, hornbills, palaeotropical;Irrisorinae, woodhoopoes, Ethiopian.Sub-order 2.Striges.—Owls. Outer toe reversible. Schizognathous. Long caeca. Flexor tendons normal. Hypotarsus simple. Cosmopolitan.Sub-order. 3.Caprimulgi.—Nightjars. Nocturnal. With gaping mouth. Ten remiges and ten rectrices. Spinae sterni vestigial. Caeca functional.Steatornithidae, Steatornis, oil-bird or guacharo, South America.Podargidae, Australasian,Caprimulgidae, cosmopolitan.Sub-order 4.Cypseli.—Tenth terminal remex the longest. With short spinae sterni. Without caeca.Cypselidae, swifts, cosmopolitan.Trochilidae, humming-birds, American.Sub-order 5.Colii.—Mouse-birds. First and fourth toes reversible. Ethiopian.Sub-order 6.Trogones.—Trogons. Heterodactyle, first and second toes directed forwards, third and fourth backwards. Tropical.Trogon gallicus, Miocene of France.Sub-order 7.Pici.—Zygodactylous. Tendon of the flexor hallucis longus muscle sending a strong vinculum to that of the flexor profundus muscle, the tendon of which goes to the third toe only.Galbulidae, puff-birds and jacamars, neotropical.Capitonidae, barbets, tropical.Rhamphastidae, toucans, neotropical.Picidae, woodpeckers, cosmopolitan, excepting Madagascar and Australian region.14. OrderPasseriformes.—Nidicolous. Aegithognathous, without basipterygoid processes. Spina externa sterni large, spina interna absent. Quinto-cubital, toes normal. Apparently since the upper Eocene.Sub-order 1.Passeres Anisomyodae.—Syrinx muscles entirely lateral or attached to the dorsal or ventral corners of the bronchial semi-rings, (1)Subclamatores. Deep plantar tendons connected by a vinculum.Eurylaemidae, broadbills, Indian and Indo-Malayan. (2)Clamatores. Deep flexor tendons not connected.Pittidae, palaeotropical.Xenicidae, New Zealand.Tyrannidae, American,Formicariidae,Pteroptochidae, neotropical.Sub-order 2.Passeres Diacromyodae.—Syrinx muscles of either side attached to the dorsal and ventral corners of the rings. Hallux strong, with a large claw, (1)SuboscineswithMenura, lyre-bird, andAtrichia, scrub-bird, in Australia. (2)Oscines, the true singing-birds, with more than 5000 recent species, are mostly divided into some thirty “families,” few of which can be defined.
I. DivisionRatitae.—Terrestrial, flightless. Without sternal keel. Quadrate bone with single proximal knob. Without pygostyle. Coracoid and scapula fused. Compound rhamphotheca. Adult without apteria. With copulatory organ. A collective polyphyletic or heterogeneous group, originally cosmopolitan; with certainty existing since the Miocene.
1. OrderStruthiones.—With pubic symphysis. Two toes only, third and fourth.Struthio, ostrich, Pliocene of Samos and of north-west India, now Africa and Arabia.2. OrderRheae.—With long ischiadic symphysis. Three toes.Mesembriornis, Miocene or Pliocene of Argentina. Rhea, South America.3. OrderCasuarii.—Three toes. Aftershaft as long as the other half.CasuariusandDromaeus, Australian.Hypselornis, Pliocene of Sivalik Hills.4. OrderApteryges.—Four toes. Bill long and slender.Apteryx, New Zealand.5. OrderDinornithes.—Three or four toes. Bill short. Anterior limbs extremely reduced.Dinornis, numerous species, recently extinct, New Zealand.6. OrderAepyornithes.—Aepyornis, recently extinct, Madagascar.To the Ratitae belong possibly also the imperfectly knownDiatryma, Eocene of New Mexico,GastornisandDasornis, Eocene of Europe,Genyornis, Pleistocene of Australia.
1. OrderStruthiones.—With pubic symphysis. Two toes only, third and fourth.Struthio, ostrich, Pliocene of Samos and of north-west India, now Africa and Arabia.
2. OrderRheae.—With long ischiadic symphysis. Three toes.Mesembriornis, Miocene or Pliocene of Argentina. Rhea, South America.
3. OrderCasuarii.—Three toes. Aftershaft as long as the other half.CasuariusandDromaeus, Australian.Hypselornis, Pliocene of Sivalik Hills.
4. OrderApteryges.—Four toes. Bill long and slender.Apteryx, New Zealand.
5. OrderDinornithes.—Three or four toes. Bill short. Anterior limbs extremely reduced.Dinornis, numerous species, recently extinct, New Zealand.
6. OrderAepyornithes.—Aepyornis, recently extinct, Madagascar.
To the Ratitae belong possibly also the imperfectly knownDiatryma, Eocene of New Mexico,GastornisandDasornis, Eocene of Europe,Genyornis, Pleistocene of Australia.
II. DivisionOdontolcae.—Marine, flightless, without sternal keel. Upper and lower jaws with teeth in furrows. Cretaceous epoch.Enaliornis, England, vertebrae chiefly biconcave;Hesperornis, North America, vertebrae heterocoelous.
III. DivisionCarinatae.—With keeled sternum.
1. OrderIchthyornithes.—Power of flight well developed. Vertebrae still amphicoelous. With small pygostyle. Incisura ischiadica. With alveolar teeth. Cretaceous of Kansas.Ichthyornis, Apatornis.2. OrderColymbiformes.—Plantigrade, nidifugous, aquatic. All toes webbed, fourth largest, hallux short; metatarsus laterally compressed; tibia with high, pyramidal crest. Bill straight, pointed, with simple sheath.Sub-order 1.Colymbi, Divers. Front toes completely webbed. Holarctic.Colymbus.Sub-order 2.Podicipedes, Grebes. Toes lobated. Cosmopolitan.3. OrderSphenisciformes.—Nidicolous, marine. Flightless, wings transformed into rowing paddles.Sphenisci, penguins. Antarctic and southern temperate coasts. Since the Eocene.4. OrderProcellariiformes.—Well flying, pelagic, nidicolous. Hallux absent or vestigial. Rhamphotheca compound. Cosmopolitan.Tubinares, petrels and albatrosses.5. OrderCiconiiformes.—Swimmers or waders. Desmognathous, without basipterygoid processes; with one pair of sterno-tracheal muscles.Sub-order 1.Steganopodes.—Well flying, aquatic, nidicolous; with all the four toes webbed together. Rhamphotheca compound; cosmopolitan.Phaëthon, tropic-bird;Sula, gannet;Phalacrocorax, cormorant andPlotus, snake-bird;Fregata, frigate-bird; Pelecanus. Here alsoPelagornis, Miocene of France;Argillornisand probablyOdontopteryxfrom the London Clay.Sub-order 2.Ardeae.—Piscivorous, nidicolous, waders; with complicated hypotarsus and with long cervical apteria.Ardeidae, cosmopolitan; includingCancroma, Neotropical,Balaeniceps, Scopidae, Ethiopian.Proherodius, Eocene of England.Sub-order 3.Ciconiae.—Zoophagous, nidicolous, waders; with simple hypotarsus and without cervical apteria. Cosmopolitan.Ciconiidae, storks.Ibidae, ibises and spoonbills.Propelargus, Oligocene.Sub-order 4.Phoenicopteri.—Flamingos. Nidifugous, waders; with simple hypotarsus and without cervical apteria. Front toes completely webbed; hallux very short or absent; feed chiefly on small aquatic invertebrates.Phoenicopterus, cosmopolitan. OligoceneElornisand, allied,Palaelodus.6. OrderAnseriformes.—Desmognathous, nidifugous; with two pairs of sterno-tracheal muscles, with complete basipterygoid processes and with a penis.Sub-order 1.Palamedeae.—Screamers. Ribs without uncinate processes. Hypotarsus simple. Neotropical.Chauna, Palamedea.Sub-order 2.Anseres.—FamilyAnatidae. Hypotarsus complex.Anser, Anas, Cygnus, since Miocene.Cnemiornis, Pleistocene, New Zealand, flightless.7. OrderFalconiformes.—Birds of prey. Carnivorous, desmognathous, nidicolous, without functional caeca. Terrestrial, aerial.Sub-order 1.Cathartae.—American vultures. With nares perviae.Cathartes, turkey buzzards,Sarcorhamphus gryphus, condorGypagus papa, king vulture.Sub-order 2.Accipitres.—With nares imperviae.Serpentariidae, secretary-bird, Ethiopian; Miocene, France.Vulturidae, Old World vultures, excluding Australia.Falconidae, cosmopolitan, since the Eocene.Harpagornis, Pleistocene, New Zealand;Lithornis, Eocene, England.Pandionidae, ospreys or fish hawks, cosmopolitan.8. OrderTinamiformes.—Nidifugous, with incisura ischiadica, without pygostyle. Herbivorous, terrestrial, neotropical.Crypturi, tinamous.9. OrderGalliformes.—Schizognathous, herbivorous, terrestrial. With ten functional remiges. With strong spinae sterni.Sub-order 1.Mesites.—Without basipterygoid processes, and with large spina interna.Mesites, Madagascar.Sub-order 2.Turnices.—Hemipodes or button-quails. Nidifugous; vomer large; sternum without processus obliqui. Hallux absent or vestigial. Old World.Turnix, Pedionomus.Sub-order 3.Galli.—With large spina communis, and with large processus obliqui. Hallux functional.Megapodiidae, Australian region.Cracidae, curassows and guans, neotropical.Gallidae, cosmopolitan.Sub-order 4.Opisthocomi.—Arboreal, with long spina externa; without basipterygoid processes.Opisthocomushoatzin, Guiana, Venezuela and Amazon countries.10. OrderGruiformes.Legs of the wading type. Without basipterygoid processes. Without spina interna. Nidifugous. Essentially Schizognathous.Rallidae, cosmopolitan, since Oligocene.Rallus, Fulica, Ocydromus, &c.,Gallinula nesiotis, Tristan d’Acunha, flightless.Notornis, New Zealand, flightless, nearly extinct.Aptornis, New Zealand, flightless, extinct.Aphanapteryx(Mauritius) =Erythromachus(Rodriguez) =Diaphorapteryx(Chatham Island), flightless and recently extinct.Gypsornis, upper Eocene, France.Gruidae, cranes, cosmopolitan, alliedPhororhacos, Tertiary of Argentina.Dicholophidae, cariamas, neotropical.Otididae, bustards, Old World.Rhinochetidae, kagus, New Caledonia.Eurypygidae, sun-bittern, neotropical.Heliornithidae, finfoots, tropical.11. OrderCharadriiformes.—Schizognathous. With eleven remiges, of which the terminal very short. Aquinto-cubital. Spinae sterni short, separate.Sub-order 1.Limicolae.—Nidifugous, without spina interna sterni. Hypotarsus complicated.Charadriidae, plovers.Chionididae, sheath-bill.Glareolidae, wading swallows and coursers.Thinocorythidae, seed-snipes.Oedicnemididae, thick-knees.Parridae.Sub-order 2.Lari.—Aquatic, vomer complete. Without basipterygoid processes. Front toes webbed; hallux small or absent. Large supraorbital glands. Since Miocene.Laridae, gulls, cosmopolitan.Alcidae, auks, northern half of periarctic region.Sub-order 3.Pterocles.—Sand-grouse. Nidifugous. Vomer vestigial. With large crop and caeca. Hallux vestigial or absent since Oligocene. Africa to India, and Siberia.PteroclesandSyrrhaptes.Sub-order 4.Columbae.—Pigeons. Nidicolous. Vomer vestigial. With large crop, vestigial caeca.Columbidae, cosmopolitan, since Miocene.Dididae, flightless, recently extinct.Didus, dodo, Mauritius.Pezophaps, solitaire, Rodriguez.12. OrderCuculiformes.—Desmognathous, nidicolous; zygodactylous, or with the outer toe reversible.Sub-order 1.Cuculi.—Cuckoos. Quinto-cubital.Cuculidae, cosmopolitan.Musophagidae, plantain-eaters and touracos, Ethiopian since Miocene.Sub-order 2.Psittaci.—Parrots. Zygodactylous; aquinto-cubital. Cosmopolitan, chiefly tropical.Trichoglossidae, lories, Austro-Malayan.Nestor, New Zealand.Cyclopsittacus, Eos, Lorius, &c.Psittacidae, tongue smooth, incl.Stringops.13. OrderCoraciiformes.—Nidicolous. Nares imperviae, holorhinal. Downs restricted to the apteria or absent. Thirteen to fifteen cervical vertebrae. Mostly desmognathous. Deep plantar tendons connected with each other.Sub-order 1.Coraciae.—Either (1) with long spina externa sterni,Coraciidae, rollers, Old World.Momotidae, neotropical, motmots and todies.Alcedinidae, kingfishers, cosmopolitan or (2) with long spina communis.Meropidae, bee-eaters, Old World.Upupidae, Upupinae, hoopoes: palaearctic and palaeotropical.Bucerotinae, hornbills, palaeotropical;Irrisorinae, woodhoopoes, Ethiopian.Sub-order 2.Striges.—Owls. Outer toe reversible. Schizognathous. Long caeca. Flexor tendons normal. Hypotarsus simple. Cosmopolitan.Sub-order. 3.Caprimulgi.—Nightjars. Nocturnal. With gaping mouth. Ten remiges and ten rectrices. Spinae sterni vestigial. Caeca functional.Steatornithidae, Steatornis, oil-bird or guacharo, South America.Podargidae, Australasian,Caprimulgidae, cosmopolitan.Sub-order 4.Cypseli.—Tenth terminal remex the longest. With short spinae sterni. Without caeca.Cypselidae, swifts, cosmopolitan.Trochilidae, humming-birds, American.Sub-order 5.Colii.—Mouse-birds. First and fourth toes reversible. Ethiopian.Sub-order 6.Trogones.—Trogons. Heterodactyle, first and second toes directed forwards, third and fourth backwards. Tropical.Trogon gallicus, Miocene of France.Sub-order 7.Pici.—Zygodactylous. Tendon of the flexor hallucis longus muscle sending a strong vinculum to that of the flexor profundus muscle, the tendon of which goes to the third toe only.Galbulidae, puff-birds and jacamars, neotropical.Capitonidae, barbets, tropical.Rhamphastidae, toucans, neotropical.Picidae, woodpeckers, cosmopolitan, excepting Madagascar and Australian region.14. OrderPasseriformes.—Nidicolous. Aegithognathous, without basipterygoid processes. Spina externa sterni large, spina interna absent. Quinto-cubital, toes normal. Apparently since the upper Eocene.Sub-order 1.Passeres Anisomyodae.—Syrinx muscles entirely lateral or attached to the dorsal or ventral corners of the bronchial semi-rings, (1)Subclamatores. Deep plantar tendons connected by a vinculum.Eurylaemidae, broadbills, Indian and Indo-Malayan. (2)Clamatores. Deep flexor tendons not connected.Pittidae, palaeotropical.Xenicidae, New Zealand.Tyrannidae, American,Formicariidae,Pteroptochidae, neotropical.Sub-order 2.Passeres Diacromyodae.—Syrinx muscles of either side attached to the dorsal and ventral corners of the rings. Hallux strong, with a large claw, (1)SuboscineswithMenura, lyre-bird, andAtrichia, scrub-bird, in Australia. (2)Oscines, the true singing-birds, with more than 5000 recent species, are mostly divided into some thirty “families,” few of which can be defined.
1. OrderIchthyornithes.—Power of flight well developed. Vertebrae still amphicoelous. With small pygostyle. Incisura ischiadica. With alveolar teeth. Cretaceous of Kansas.Ichthyornis, Apatornis.
2. OrderColymbiformes.—Plantigrade, nidifugous, aquatic. All toes webbed, fourth largest, hallux short; metatarsus laterally compressed; tibia with high, pyramidal crest. Bill straight, pointed, with simple sheath.
Sub-order 1.Colymbi, Divers. Front toes completely webbed. Holarctic.Colymbus.
Sub-order 2.Podicipedes, Grebes. Toes lobated. Cosmopolitan.
3. OrderSphenisciformes.—Nidicolous, marine. Flightless, wings transformed into rowing paddles.Sphenisci, penguins. Antarctic and southern temperate coasts. Since the Eocene.
4. OrderProcellariiformes.—Well flying, pelagic, nidicolous. Hallux absent or vestigial. Rhamphotheca compound. Cosmopolitan.Tubinares, petrels and albatrosses.
5. OrderCiconiiformes.—Swimmers or waders. Desmognathous, without basipterygoid processes; with one pair of sterno-tracheal muscles.
Sub-order 1.Steganopodes.—Well flying, aquatic, nidicolous; with all the four toes webbed together. Rhamphotheca compound; cosmopolitan.Phaëthon, tropic-bird;Sula, gannet;Phalacrocorax, cormorant andPlotus, snake-bird;Fregata, frigate-bird; Pelecanus. Here alsoPelagornis, Miocene of France;Argillornisand probablyOdontopteryxfrom the London Clay.
Sub-order 2.Ardeae.—Piscivorous, nidicolous, waders; with complicated hypotarsus and with long cervical apteria.Ardeidae, cosmopolitan; includingCancroma, Neotropical,Balaeniceps, Scopidae, Ethiopian.Proherodius, Eocene of England.
Sub-order 3.Ciconiae.—Zoophagous, nidicolous, waders; with simple hypotarsus and without cervical apteria. Cosmopolitan.Ciconiidae, storks.Ibidae, ibises and spoonbills.Propelargus, Oligocene.
Sub-order 4.Phoenicopteri.—Flamingos. Nidifugous, waders; with simple hypotarsus and without cervical apteria. Front toes completely webbed; hallux very short or absent; feed chiefly on small aquatic invertebrates.Phoenicopterus, cosmopolitan. OligoceneElornisand, allied,Palaelodus.
6. OrderAnseriformes.—Desmognathous, nidifugous; with two pairs of sterno-tracheal muscles, with complete basipterygoid processes and with a penis.
Sub-order 1.Palamedeae.—Screamers. Ribs without uncinate processes. Hypotarsus simple. Neotropical.Chauna, Palamedea.
Sub-order 2.Anseres.—FamilyAnatidae. Hypotarsus complex.Anser, Anas, Cygnus, since Miocene.Cnemiornis, Pleistocene, New Zealand, flightless.
7. OrderFalconiformes.—Birds of prey. Carnivorous, desmognathous, nidicolous, without functional caeca. Terrestrial, aerial.
Sub-order 1.Cathartae.—American vultures. With nares perviae.Cathartes, turkey buzzards,Sarcorhamphus gryphus, condorGypagus papa, king vulture.
Sub-order 2.Accipitres.—With nares imperviae.Serpentariidae, secretary-bird, Ethiopian; Miocene, France.Vulturidae, Old World vultures, excluding Australia.Falconidae, cosmopolitan, since the Eocene.Harpagornis, Pleistocene, New Zealand;Lithornis, Eocene, England.Pandionidae, ospreys or fish hawks, cosmopolitan.
8. OrderTinamiformes.—Nidifugous, with incisura ischiadica, without pygostyle. Herbivorous, terrestrial, neotropical.Crypturi, tinamous.
9. OrderGalliformes.—Schizognathous, herbivorous, terrestrial. With ten functional remiges. With strong spinae sterni.
Sub-order 1.Mesites.—Without basipterygoid processes, and with large spina interna.Mesites, Madagascar.
Sub-order 2.Turnices.—Hemipodes or button-quails. Nidifugous; vomer large; sternum without processus obliqui. Hallux absent or vestigial. Old World.Turnix, Pedionomus.
Sub-order 3.Galli.—With large spina communis, and with large processus obliqui. Hallux functional.Megapodiidae, Australian region.Cracidae, curassows and guans, neotropical.Gallidae, cosmopolitan.
Sub-order 4.Opisthocomi.—Arboreal, with long spina externa; without basipterygoid processes.Opisthocomushoatzin, Guiana, Venezuela and Amazon countries.
10. OrderGruiformes.Legs of the wading type. Without basipterygoid processes. Without spina interna. Nidifugous. Essentially Schizognathous.Rallidae, cosmopolitan, since Oligocene.Rallus, Fulica, Ocydromus, &c.,Gallinula nesiotis, Tristan d’Acunha, flightless.Notornis, New Zealand, flightless, nearly extinct.Aptornis, New Zealand, flightless, extinct.Aphanapteryx(Mauritius) =Erythromachus(Rodriguez) =Diaphorapteryx(Chatham Island), flightless and recently extinct.Gypsornis, upper Eocene, France.Gruidae, cranes, cosmopolitan, alliedPhororhacos, Tertiary of Argentina.Dicholophidae, cariamas, neotropical.Otididae, bustards, Old World.Rhinochetidae, kagus, New Caledonia.Eurypygidae, sun-bittern, neotropical.Heliornithidae, finfoots, tropical.
11. OrderCharadriiformes.—Schizognathous. With eleven remiges, of which the terminal very short. Aquinto-cubital. Spinae sterni short, separate.
Sub-order 1.Limicolae.—Nidifugous, without spina interna sterni. Hypotarsus complicated.Charadriidae, plovers.Chionididae, sheath-bill.Glareolidae, wading swallows and coursers.Thinocorythidae, seed-snipes.Oedicnemididae, thick-knees.Parridae.
Sub-order 2.Lari.—Aquatic, vomer complete. Without basipterygoid processes. Front toes webbed; hallux small or absent. Large supraorbital glands. Since Miocene.Laridae, gulls, cosmopolitan.Alcidae, auks, northern half of periarctic region.
Sub-order 3.Pterocles.—Sand-grouse. Nidifugous. Vomer vestigial. With large crop and caeca. Hallux vestigial or absent since Oligocene. Africa to India, and Siberia.PteroclesandSyrrhaptes.
Sub-order 4.Columbae.—Pigeons. Nidicolous. Vomer vestigial. With large crop, vestigial caeca.Columbidae, cosmopolitan, since Miocene.Dididae, flightless, recently extinct.Didus, dodo, Mauritius.Pezophaps, solitaire, Rodriguez.
12. OrderCuculiformes.—Desmognathous, nidicolous; zygodactylous, or with the outer toe reversible.
Sub-order 1.Cuculi.—Cuckoos. Quinto-cubital.Cuculidae, cosmopolitan.Musophagidae, plantain-eaters and touracos, Ethiopian since Miocene.
Sub-order 2.Psittaci.—Parrots. Zygodactylous; aquinto-cubital. Cosmopolitan, chiefly tropical.Trichoglossidae, lories, Austro-Malayan.Nestor, New Zealand.Cyclopsittacus, Eos, Lorius, &c.Psittacidae, tongue smooth, incl.Stringops.
13. OrderCoraciiformes.—Nidicolous. Nares imperviae, holorhinal. Downs restricted to the apteria or absent. Thirteen to fifteen cervical vertebrae. Mostly desmognathous. Deep plantar tendons connected with each other.
Sub-order 1.Coraciae.—Either (1) with long spina externa sterni,Coraciidae, rollers, Old World.Momotidae, neotropical, motmots and todies.Alcedinidae, kingfishers, cosmopolitan or (2) with long spina communis.Meropidae, bee-eaters, Old World.Upupidae, Upupinae, hoopoes: palaearctic and palaeotropical.Bucerotinae, hornbills, palaeotropical;Irrisorinae, woodhoopoes, Ethiopian.
Sub-order 2.Striges.—Owls. Outer toe reversible. Schizognathous. Long caeca. Flexor tendons normal. Hypotarsus simple. Cosmopolitan.
Sub-order. 3.Caprimulgi.—Nightjars. Nocturnal. With gaping mouth. Ten remiges and ten rectrices. Spinae sterni vestigial. Caeca functional.Steatornithidae, Steatornis, oil-bird or guacharo, South America.Podargidae, Australasian,Caprimulgidae, cosmopolitan.
Sub-order 4.Cypseli.—Tenth terminal remex the longest. With short spinae sterni. Without caeca.Cypselidae, swifts, cosmopolitan.Trochilidae, humming-birds, American.
Sub-order 5.Colii.—Mouse-birds. First and fourth toes reversible. Ethiopian.
Sub-order 6.Trogones.—Trogons. Heterodactyle, first and second toes directed forwards, third and fourth backwards. Tropical.Trogon gallicus, Miocene of France.
Sub-order 7.Pici.—Zygodactylous. Tendon of the flexor hallucis longus muscle sending a strong vinculum to that of the flexor profundus muscle, the tendon of which goes to the third toe only.Galbulidae, puff-birds and jacamars, neotropical.Capitonidae, barbets, tropical.Rhamphastidae, toucans, neotropical.Picidae, woodpeckers, cosmopolitan, excepting Madagascar and Australian region.
14. OrderPasseriformes.—Nidicolous. Aegithognathous, without basipterygoid processes. Spina externa sterni large, spina interna absent. Quinto-cubital, toes normal. Apparently since the upper Eocene.
Sub-order 1.Passeres Anisomyodae.—Syrinx muscles entirely lateral or attached to the dorsal or ventral corners of the bronchial semi-rings, (1)Subclamatores. Deep plantar tendons connected by a vinculum.Eurylaemidae, broadbills, Indian and Indo-Malayan. (2)Clamatores. Deep flexor tendons not connected.Pittidae, palaeotropical.Xenicidae, New Zealand.Tyrannidae, American,Formicariidae,Pteroptochidae, neotropical.
Sub-order 2.Passeres Diacromyodae.—Syrinx muscles of either side attached to the dorsal and ventral corners of the rings. Hallux strong, with a large claw, (1)SuboscineswithMenura, lyre-bird, andAtrichia, scrub-bird, in Australia. (2)Oscines, the true singing-birds, with more than 5000 recent species, are mostly divided into some thirty “families,” few of which can be defined.