The Geographical Distribution of Animals/Chapter 8
CHAPTER VIII.
VARIOUS EXTINCT ANIMALS;—AND ON THE ANTIQUITY OF THE GENERA OF INSECTS AND LAND MOLLUSCA.
EXTINCT MAMMALIA OF AUSTRALIA.
These have all been obtained from caves and late Tertiary or Post-Tertiary deposits, and consist of a large number of extinct forms, some of gigantic size, but all marsupials and allied to the existing fauna. There are numerous forms of kangaroos, some larger than any living species; and among these are two genera, Protemnodon and Sthenurus, which Professor Garrod has lately shown to have been allied, not to any Australian forms, but to the Dendrolagi or tree-kangaroos of New Guinea. We have also remains of Thylacinus and Dasyurus, which now only exist in Tasmania; and extinct species of Hypsiprymnus and Phascolomys, the latter as large as a tapir. Among the more remarkable extinct genera are Diprotodon, a huge thick-limbed animal allied to the kangaroos, but nearly as large as an elephant; Nototherium, having characters of Macropus and Phascolarctos combined, and as large as a rhinoceros; and Thylacoleo, a phalanger-like marsupial nearly as large as a lion, and supposed by Professor Owen to have been of carnivorous habits, though this opinion is not held by other naturalists.
Here then we find the same phenomena as in the other countries we have already discussed,—the very recent disappearance of a large number of peculiar forms, many of them far surpassing in size any that continue to exist. It hardly seems probable that in this case their disappearance can have been due to the direct effects of the Glacial epoch, since no very extensive glaciation could have occurred in a country like Australia; but if the ocean sank 2,000 feet, the great eastern mountain range might have given rise to local glaciers. It is, however, almost certain that during late Tertiary times Australia must have been much more extensive than it is now. This is necessary to allow of the development of its peculiar and extensive fauna, especially as we see that that fauna comprised animals rivalling in bulk those of the great continents. It is further indicated by the relations with New Guinea, already alluded to, and by the general character of the various faunas which compose the Australian region, details of which will be found in the succeeding part of this work. The lowering of the ocean during the Glacial period would be favourable to the still further development of the fauna of such a country; and it is to the unfavourable conditions produced by its subsequent rising—equivalent to a depression of the land to the amount of two thousand feet—that we must impute the extinction of so many remarkable groups of animals. It is not improbable, that the disappearance of the ice and the consequent (apparent) subsidence of the land, might have been rapid as compared with the rate at which large animals can become modified to meet new conditions. Extensive tracts of fertile land might have been submerged, and the consequent crowding of large numbers of species and individuals on limited areas would have led to a struggle for existence in which the less adapted and less easily modifiable, not the physically weaker, would succumb.
There is, however, another cause for the extinction of large rather than small animals whenever an important change of conditions occurs, which has been suggested to me by a correspondent,[1] but which has not, I believe, been adduced by Mr. Darwin or by any other writer on the subject. It is dependent on the fact, that large animals as compared with small ones are almost invariably slow breeders, and as they also necessarily exist in much smaller numbers in a given area, they offer far less materials for favourable variations than do smaller animals. In such an extreme case as that of the rabbit and elephant, the young born each year in the world are probably as some millions to one; and it is very easily conceivable that in a thousand years the former might, under pressure of rapidly changing conditions, become modified into a distinct species, while the latter, not offering enough favourable variations to effect a suitable adaptation, would become extinct. We must also remember the extreme specialization of many of the large animals that have become extinct—a specialization which would necessarily render modification in any new direction difficult, since the inherited tendency of variation would probably be to increase the specialization in the same directions which had heretofore been beneficial. If to these two causes we add the difficulty of obtaining sufficient food for such large animals, and perhaps the injurious effects of changes of climate, we shall not find it difficult to understand how such a vast physical revolution as the Glacial epoch, with its attendant phenomena of elevations and subsidences, icy winds, and sudden floods by the bursting of lake barriers, might have led to the total extinction of a vast number of the most bulky forms of mammalia, while the less bulky were able to survive, either by greater hardiness of constitution or by becoming more or less modified. The result is apparent in the comparatively small or moderate size of the species constituting the temperate fauna, in all parts of the globe.
It is much to be regretted that no mammalian remains of earlier date have been found in Australia, as we should then see if it is really the case that marsupials have always formed its highest type of mammalian life. At present its fossil fauna is chiefly interesting to the zoologist, but throws little light on the past relations of this isolated country with other parts of the globe.
Mammalian remains in the Secondary Formations.
In the oldest Tertiary beds of Europe and North America, we have (even with our present imperfect record) a rich and varied mammalian fauna. As compared with our living or recent highly specialized forms, it may be said to consist of generalised types; but as compared with any primeval mammalian type, it must be pronounced highly specialised. Not only are such diversified groups as Carnivora, Perrissodactyle and Artiodactyle Ungulates, Primates, Chiroptera, Rodents, and Marsupials already well marked, but in many of these there is a differentiation into numerous families and genera of diverse character. It is impossible therefore to doubt, that many peculiar forms of mammalia must have lived long anterior to the Eocene period; but there is unfortunately a great gap in the record between the Eocene and Cretaceous beds, and these latter being for the most part marine continue the gap as regards mammals over an enormous lapse of time. Yet far beyond both these chasms in the Upper Oolitic strata, remains of small mammalia have been found; again, in the Stonesfield slate, a member of the Lower Oolite, other forms appear. Then comes the marine Lias formation with another huge gap; but beyond this again in the Upper Trias, the oldest of the secondary formations, mammalian teeth have been discovered in both England and Germany, and these are, as nearly as can be ascertained, of the same age as the Dromatherium already noticed, from North America. They have been named Microlestes, and show some resemblance to those of the West Australian Myrmecobius. In the Oolitic strata numerous small jawbones have been found, which have served to characterise eight genera, all of which are believed to have been Marsupials, and in some of them a resemblance can be traced to some of the smaller living Australian species. These, however, are mere indications of the number of mammalia that must have lived in the secondary period, so long thought to be exclusively "the age of reptiles;" and the fact that the few yet found are at all comparable with such specialised forms as still exist, must convince us, that we shall have to seek far beyond even the earliest of these remains, for the first appearance of the mammalian type of vertebrata.
Extinct Birds.
Compared with those of mammalia, the remains of birds are exceedingly scarce in Europe and America; and from the wandering habits of so many of this class, they are of much less value as indications of past changes in physical geography. A large proportion of the remains belong to aquatic or wading types, and as these have now often a world-wide range, the occurrence of extinct forms can have little bearing on our present inquiry. There are, however, a few interesting cases of extinct land-birds belonging to groups now quite strangers to the country in which they are found; and others scarcely less interesting, in which groups now peculiar to certain areas are shown to have been preceded by allied species or genera of gigantic size.
Palæarctic Region and N. India.—In the caves and other Post-Pliocene deposits of these countries, the remains of birds almost all belong to genera now inhabiting the same districts. Almost the only exceptions are, the great auk and the capercailzie, already mentioned as being found in the Danish mounds; the latter bird, with Tetrao albus, in Italian caverns; and a species of pheasant (Phasianus) said to have occurred in the Post-Pliocene of France, considerably west of the existing range of the genus in a wild state.
In the preceding Pliocene deposits, but few remains have been found, and all of existing genera but one, a gallinaceous bird (Gallus bravardi) allied to the domestic fowl and peacock.
The Miocene beds of France and Central Europe have produced many more remains of birds, but these, too, are mostly of existing European genera, though there are some notable exceptions. Along with forms undistinguishable from crows (Corvus), shrikes (Lanius), wagtails (Motacilla), and woodpeckers (Picus), are found remains allied to the Oriental edible-nest swift (Collocalia) and Trogon; a parrot resembling the African genus Psittacus; an extinct form Necrornis, perhaps allied to the plantain-eaters (Musophaga); Homalophus, doubtfully allied to woodpeckers, and Limnatornis to the hoopoes. The gallinaceous birds are represented by three species of pheasants, some very close to the domesticated species; Palæoperdix allied to the partridges; and Palæortyx, small birds allied to the American genus Ortyx, but with larger wings. There are also species of Pterocles allied to living birds, and a small pigeon. There are numerous living genera of Accipitres; such as eagle (Aquila), kite (Milvus), eagle-owl (Bubo), and screech-owl (Strix); with the African secretary-bird (Serpentarius), and some extinct forms, as Palæocercus, Palæohierax and Palæetus.
Aquatic and wading birds were abundant, including numerous rails, bustards, herons, sandpipers, gulls, divers, and pelicans. There were also many ducks, some allied to the genus Dendrocygna; the Oriental genus of storks, Leptoptilus; Ibidipodia, a remarkable form allied to Ibis and Ciconia; Elornis, near Limosa; Pelagornis, a large bird allied to gannets and pelicans; Hydrornis, allied to the ducks and petrels; Dolichopterus, allied to plovers. Perhaps the most interesting of these extinct birds are, however, the flamingoes, represented by forms hardly distinguishable from living species, and by one extinct genus Palælodus, which had very long toes, and probably walked on aquatic plants like the tropical jacanas.
The Miocene beds of North India have furnished few birds; the only one of geographical interest being an extinct species of ostrich, not very different from that now inhabiting Arabia.
On the whole, the birds of Europe at this period were very like those now living, with the addition of a few tropical forms. These latter were, however, perhaps more numerous and important than they appear to be, as they belong to inland and forest-haunting types, which would not be so frequently preserved as the marsh and lake-dwelling species. Taking this into consideration, the assemblage of Miocene birds accords well with what we know of the mammalian fauna. We have the same indications of a luxuriant vegetation and subtropical climate, and the same appearance of Oriental and especially of African types. Trogon is perhaps the most interesting of all the forms yet discovered, since it furnishes us with a central point whence the living trogons of Asia, Africa, and South America might have diverged.
In the Eocene we find ourselves almost wholly among extinct forms of birds. The earliest known Passerine bird is here met with, in Protornis, somewhat similar to a lark, found in the Lower Eocene of Switzerland; while another Passerine form, Palægithalus, and one allied to the nuthatch (Sitta), have been discovered in the Upper Eocene of Paris. Picariæ of equal antiquity are found. Cryptornis, from the Paris Eocene, and Halcyornis from the Lower Eocene of the Isle of Sheppey, were both allied to kingfishers; while a form allied to Centropus a genus of cuckoos, or, as Milne-Edwards thinks, to the Madagascar Leptosomus, has been found in the Upper Eocene of France. Several Accipitres of somewhat doubtful affinities have been found in the same country; while Lithornis, from the Lower Eocene of the Isle of Sheppey, was a small vulturine bird supposed to be allied to the American group, Cathartes. Among the waders, some extinct forms of plovers have been found, and a genus (Agnopterus), allied to the flamingoes; while there are many swimming birds, such as pelicans, divers, and several extinct types of doubtful affinities. Most interesting of all is a portion of a cranium discovered in the Lower Eocene of Sheppey, and lately pronounced by Professor Owen to belong to a large Struthious bird, allied to the New Zealand Dinornis and also perhaps to the ostrich. Another gigantic bird is the Gastornis, from the Lower Eocene of Paris, which was as large as an ostrich, but which is believed to have been a generalised type, allied to wading and swimming birds as well as to the Struthiones.
Beyond this epoch we have no remains of birds in European strata till we come to the wonderful Archæopteryx from the Upper Oolite of Bavaria; a bird of a totally new type, with a bony tail, longer than the body, each vertebra of which carried a pair of diverging feathers.
North America.—A number of bird-remains have lately been found in the rich Tertiary and Cretaceous deposits of the United States; but here, too, comparatively few are terrestrial forms. No Passerine bird has yet been found. The Picariæ are represented by Uintornis, an extinct form allied to woodpeckers, from the Eocene of Wyoming. Species of turkey (Meleagris) occur in the Post-Pliocene and as far back as the Miocene strata, showing that this interesting type is a true denizen of temperate North America. The other birds are, Accipitres; waders and aquatics of existing genera; and a number of extinct forms of the two latter orders—such as, Aletornis an Eocene wader; Palæotringa, allied to the sandpipers, and Telmatobius to the rails, both Cretaceous; with Graculavus, allied to Graculus; Laornis allied to the swans; Hesperornis a gigantic diver; and Icthyornis a very low form, with biconcave vertebra, such as are only found in fishes and some reptiles—also from Cretaceous deposits.
South America.—The caverns of Brazil produced thirty-four species of birds, most of them referable to Brazilian genera, and many to still existing species. The most interesting were two species of American ostrich (Rhea), one larger than either of the living species; a large turkey-buzzard (Cathartes); a new species of the very isolated South American genus Opisthocomus; and a Cariama, or allied new genus.
Madagascar and the Mascarene Islands.—We have here only evidence of birds that have become extinct in the historical period or very little earlier. First we have a group of birds incapable of flight, allied to pigeons, but forming a separate family, Dididæ; and which, so far as we yet know, inhabited Mauritius, Rodriguez, and probably Bourbon. Aphanapteryx, an extinct genus of rails, inhabited Mauritius; and another genus, (Erythromachus), Rodriguez. A large parrot, said by Prof. Milne Edwards to be allied to Ara and Microglossus, also inhabited Mauritius; and another allied to Eclectus, the island of Rodriguez. None of these have been found in Madagascar; but a gigantic Struthious bird, Æpyornis, forming a peculiar family distinct both from the ostriches of Africa and the Dinornis of New Zealand inhabited that island; and there is reason to believe that this may have lived less than 200 years ago.
New Zealand.—A number of extinct Struthious birds, forming two families, Dinornithidæ and Palapterygidæ, have been found in New Zealand. Some were of gigantic size. They seem allied both to the living Apteryx of New Zealand and the emu of Australia. They are quite recent, and some of them have probably lived within the last few centuries. Remains of Dinornis have also been found in a Post-Pliocene deposit in Queensland, N. E. Australia[2]—a very important discovery, as it gives support to the theory of a great eastward extension of Australia in Tertiary times.
Extinct Tertiary Reptiles.
These will not occupy us long, as no very great number are known, and most of them belong to a few principal forms of comparatively little geographical interest.
Tortoises are perhaps the most abundant of the Tertiary reptiles. They are numerous in the Eocene and Miocene formations both in Europe and North America. The genera Emys and Trionyx abound in both countries, as well as in the Miocene of India. Land tortoises occur in the Eocene of North America and in the Miocene of Europe and India, where the huge Colossochelys, twelve feet long, has been found. In the Pliocene deposits of Switzerland the living American genus Chelydra has been met with. These facts, together with the occurrence of a living species in the Miocene of India, show that this order of reptiles is of great antiquity, and that most of the genera once had a wider range than now.
Crocodiles, allied to the three forms now characteristic of India, Africa, and America, have been found in the Eocene of our own country, and several species of Crocodilus have occurred in beds of the same age in North America.
Lizards are very ancient, many small terrestrial forms occurring in all the Tertiary deposits. A species of the genus Chamæleo is recorded from the Eocene of North America, together with several extinct genera.
Snakes were well developed in the Eocene period, where remains of several have been found which must have been from twelve to twenty feet long. An extinct species of true viper has occurred in the Miocene of France, and one of the Pythonidæ in the Miocene brown coal of Germany.
Batrachia occur but sparingly in a fossil state in the Tertiary deposits. The most remarkable is the large Salamander (Andreas) from the Upper Miocene of Switzerland, which is allied to the Menopoma living in North America. Species of frog (Rana), and Palæophrynus an extinct genus of toads, have been found in the Miocene deposits of Germany and Switzerland.
Fresh water fish are almost unknown in the Tertiary deposits of Europe, although most of the families and some genera of living marine fish are represented from the Eocene downwards.
Antiquity of the Genera of Insects.
Fossil insects are far too rarely found, to aid us in our determination of difficult questions of geographical distribution; but in discussing these questions it will be important to know, whether we are to look upon the existing generic forms of insects as of great or small antiquity, compared with the higher vertebrates; and to decide this question the materials at our command are ample.
The conditions requisite for the preservation of insects in a fossil state are no doubt very local and peculiar; the result being, that it is only at long intervals in the geological record that we meet with remains of insects in a recognisable condition. None appear to have been found in the Pliocene formation; but in the Upper Miocene of Œninghen in Switzerland, associated with the wonderfully rich fossil flora, are found immense quantities of insects. Prof. Heer examined more than 5,000 specimens belonging to over 800 species, and many have been found in other localities in Switzerland; so that more than 1,300 species of Miocene insects have now been determined. Most of the orders are represented, but the beetles (Coleoptera) are far the most abundant. Almost all belong to existing genera, and the majority of these genera now inhabit Europe, only three or four being exclusively Indian, African, or American.
In the Lower Miocene of Croatia there is another rich deposit of insects, somewhat more tropical in character, comprising large white-ants and dragon-flies differently marked from any now inhabiting Europe. A butterfly is also well preserved, with all the markings of the wings; and it seems to be a Junonia, a tropical genus, though it may be a Vanessa, which is European, but the fossil most resembles Indian species of Junonia.
The Eocene formations seem to have produced no insect remains; but they occur again in the Upper Cretaceous at Aix-la-Chapelle, where two butterflies have been found, Cyllo sepulta and Satyrites Reynesii, both belonging to the Satyridæ, and the former to a genus now spread over Africa, India, and Australia.
A little earlier, in the Wealden formation of our own country, numerous insects have been found, principally dragon flies (Libellula, Æshna); aquatic Hemiptera (Velia Hydrometra); crickets, cockroaches, and cicadas, of familiar types.
Further back in the Upper Oolite of Bavaria—which produced the wonderful long-tailed bird, Archæopteryx—insects of all orders have been found, including a moth referred to the existing genus Sphinx.
In the Lower Oolite of Oxfordshire many fossil beetles have been found whose affinities are shown by their names:—Buprestidium, Curculionidium, Blapsidium, Melolonthidium, and Prionidium; a wing of a butterfly has also been found, allied to the Brassolidæ now confined to tropical America, and named Palæontina oolitica.
Still more remote are the insects of the Lias of Gloucestershire, yet they too can be referred to well-known family types—Carabidæ, Melolonthidæ, Telephoridæ, Elateridæ, and Curculionidæ, among beetles; Gryllidæ and Blattidæ among Orthoptera; with Libellula, Agrion, Æshna, Ephemera, and some extinct genera. When we consider that almost the only vertebrata of this period were huge Saurian reptiles like the Icthyosaurus, Plesiosaurus, and Dinosaurus, with the flying Pterodactyles; and that the great mass of our existing genera, and even families, of fish and reptiles had almost certainly not come into existence, we see at once that types of insect-form are, proportionately, far more ancient. At this remote epoch we find the chief family types (the genera of the time of Linnæus) perfectly differentiated and recognisable. It is only when we go further back still, into the Palæozoic formations, that the insect forms begin to show that generalization of type which renders it impossible to classify them in any existing groups. Yet even in the coal formation of Nova Scotia and Durham, the fossil insects are said by competent entomologists to be "allied to Ephemera," "near Blatta," "near Phasmidæ;" and in deposits of the same age at Saarbrück near Trèves, a well-preserved wing of a grasshopper or locust has been found, as well as a beetle referred to the Scarabeidæ. More remarkable, however, is the recent discovery in the carboniferous shales of Belgium, of the clearly-defined wing of a large moth (Breyeria borinensis), closely resembling some of the Saturniidæ; so that we have now all the chief orders of Insects—including those supposed to be the most highly developed and the most recent—well represented at this very remote epoch. Even the oldest insects, from the Devonian rocks of North America, can mostly be classed as Neuroptera or Myriapoda, but appear to form new families.
We may consider it, therefore, as proved, that many of the larger and more important genera of insects date back to the beginning of the Tertiary period, or perhaps beyond it; but the family types are far older, and must have been differentiated very early in the Secondary period, while some of them perhaps go back to Palæozoic times. The great comparative antiquity of the genera is however the important fact for us, and we shall have occasion often to refer to it, in endeavouring to ascertain the true bearing of the facts of insect distribution, as elucidating or invalidating the conclusions arrived at from a study of the distribution of the higher animals.
Antiquity of the Genera of Land and Fresh-Water Shells.
The remains of land and fresh-water shells are not much more frequent than those of insects. Like them, too, their forms are very stable, continuing unchanged through several geological periods. In the Pliocene and Miocene formations, most of the shells are very similar to living species, and some are quite identical. In the Eocene we meet with ordinary forms of the genera Helix, Clausilia, Pupa, Bulimus, Glandina, Cyclostoma, Megalostoma, Planorbis, Paludina and Limnæa, some resembling European species, others more like tropical forms. A British Eocene species of Helix is still living in Texas; and in the South of France are found species of the Brazilian sub-genera Megaspira and Anastoma. In the secondary formation no true land shells have been found, but fresh water shells are tolerably abundant, and almost all are still of living forms. In the Wealden (Lower Cretaceous) and Purbeck (Upper Oolite) are found Unio, Melania, Paludina, Planorbis, and Limnæa; while the last named genus occurs even in the Lias.
The notion that land shells were really not in existence during the secondary period is, however, proved to be erroneous by the startling discovery, in the Palæozoic coal measures of Nova Scotia, of two species of Helicidæ, both of living genera—Pupa vetusta, and Zonites priscus. They have been found in the hollow trunk of a Sigillaria, and in great quantities in a bed full of Stigmarian rootlets. The most minute examination detects no important differences of form or of microscopic structure, between these shells and living species of the same genera! These mollusca were the contemporaries of Labyrinthodonts and strange Ganoid fishes, which formed almost the whole vertebrate fauna. This unexpected discovery renders it almost certain, that numbers of other existing genera, of which we have found no traces, lived with these two through the whole secondary period; and we are thus obliged to assume as a probability, that any particular genus has lived through a long succession of geological ages. In estimating the importance of any peculiarities or anomalies in the geographical distribution of land shells as compared with the higher vertebrates, we shall, therefore, have to keep this possible, and even probable high antiquity, constantly in mind.
We have now concluded our sketch of Tertiary Palæontology as a preparation for the intelligent study of the Geographical Distribution of Land Animals; and however imperfectly the task has been performed, the reader will at all events have been convinced that some such preliminary investigation is an essential and most important part of our work. So much of palæontology is at present tentative and conjectural, that in combining the information derived from numerous writers, many errors of detail must have been made. The main conclusions have, however, been drawn from as large a basis of facts as possible; and although fresh discoveries may show that our views as to the past history of some of the less important genera or families are erroneous, they can hardly invalidate our results to any important degree, either as regards the intercommunications between separate regions in the various geological epochs, or as to the centres from which some of the more important groups have been dispersed.