The Geographical Distribution of Animals/Chapter 5
CLASSIFICATION AS AFFECTING THE STUDY OF GEOGRAPHICAL DISTRIBUTION.
A little consideration will convince us, that no inquiry into the causes and laws which determine the geographical distribution of animals or plants can lead to satisfactory results, unless we have a tolerably accurate knowledge of the affinities of the several species, genera, and families to each other; in other words, we require a natural classification to work upon. Let us, for example, take three animals—a, b, and c—which have a general external resemblance to each other, and are usually considered to be really allied; and let us suppose that a and b inhabit the same or adjacent districts, while c is found far away on the other side of the globe, with no animals at all resembling it in any of the intervening countries. We should here have a difficult problem to solve; for we should have to show that the general laws by which we account for the main features of distribution, will explain this exceptional case. But now, suppose some comparative anatomist takes these animals in hand, and finds that the resemblance of c to a and b is only superficial, while their internal structure exhibits marked and important differences; and that c really belongs to another group of animals, d, which inhabits the very region in which c was found—and we should no longer have anything to explain. This is no imaginary case. Up to a very few years ago a curious Mexican animal, Bassaris astuta, was almost always classed in the civet family (Viverridæ), a group entirely confined to Africa and Asia; but it has now been conclusively shown by Professor Flower that its real affinities are with the racoons (Procyonidæ), a group confined to North and South America. In another case, however, an equally careful examination shows, that an animal peculiar to the Himalayas (Ælurus fulgens) has its nearest ally in the Cercoleptes of South America. Here, therefore, the geographical difficulty really exists, and any satisfactory theory of the causes that have led to the existing distribution of living things, must be able to account, more or less definitely, for this and other anomalies. From these cases it will be evident, that if any class or order of animals is very imperfectly known and its classification altogether artificial, it is useless to attempt to account for the anomalies its distribution may present; since those anomalies may be, to a great extent, due to false notions as to the affinities of its component species.
According to the laws and causes of distribution discussed in the preceding chapters, we should find limited and defined distribution to be the rule, universal or indefinite distribution to be the exception, in every natural group corresponding to what are usually regarded as families and genera; and so much is this the case in nature, that when we find a group of this nominal rank scattered as it were at random over the earth, we have a strong presumption that it is not natural; but is, to a considerable extent, a haphazard collection of species. Of course this reasoning will only apply, in cases where there are no unusual means of dispersal, nor any exceptional causes which might determine a scattered distribution.
From the considerations now adduced it becomes evident, that it is of the first importance for the success of our inquiry to secure a natural classification of animals, especially as regards the families and genera. The higher groups, such as classes and orders, are of less importance for our purpose; because they are almost always widely and often universally distributed, except those which are so small as to be evidently the nearly extinct representatives of a once more extensive series of forms. We now proceed to explain the classification to be adopted, as low down as the series of families. To these, equivalent English names are given wherever they exist, in order that readers possessing no technical knowledge, may form some conception of the meaning of the term "family" in zoology.
The primary divisions of the animal kingdom according to two eminent modern authorities are as follows:
Classification of Animals (1869).
|Carus and Gerstaeker.|
Handbuch der Zoologie (1868).
|3. Cœlenterata||2. Cœlenterata.|
|4. Annuloida||3. Echinodermata.|
|5. Annulosa||4. Vermes.|
|6. Molluscoida||6. Molluscoida.|
|7. Mollusca||7. Mollusca.|
|8. Vertebrata||8. Vertebrata.|
For reasons already stated it is only with the fifth, seventh, and eighth of these groups that the present work proposes to deal; and even with the fifth and seventh only partially and in the most general way.
The classes of the vertebrata, according to both the authors above quoted, are: 1. Mammalia. 2. Aves. 3. Reptilia. 4. Amphibia. 5. Pisces, in which order they will be taken here.
The sub-classes and orders of mammalia are as follows:
|Huxley (1869), Flower (1870).||Carus (1868).|
|Monodelphia||1. Primates||1. Primates.|
|2. Chiroptera||2. Chiroptera.|
|3. Insectivora||3. Insectivora.|
|4. Carnivora||6. Carnivora|
|7. Ungulata||10. Artiodactyla.|
|8. Proboscidea||9. Proboscidea.|
|9. Hyracoidea||8. Lamnungia.|
|10. Rodentia||4. Rodentia.|
|11. Edentata||13. Bruta.|
|Didelphia||12. Marsupialia||14. Marsupialia.|
|Ornithodelphia||13. Monotremata||15. Monotremata.|
The above series of orders is arranged according to Professor Flower's Osteology of Mammalia, and they will follow in this succession throughout my work. Professor Huxley arranges the same orders in a different series.
In determining the manner in which the several orders shall be subdivided into families, I have been guided in my choice of classifications mainly by the degree of attention the author appears to have paid to the group, and his known ability as a systematic zoologist; and in a less degree by considerations of convenience as regards the special purposes of geographical distribution. In many cases it is a matter of great doubt whether a certain group should form several distinct families or be united into one or two; but one method may bring out the peculiarities of distribution much better than the other, and this is, in our case, a sufficient reason for adopting it.
For the Primates I follow, with some modifications, the classification of Mr. St. George Mivart given in his article "Apes" in the new edition of the Encyclopædia Britannica, and in his paper in the Proceedings of the Zoological Society of London, 1865, p. 547. It is as follows:
|Order—PRIMATES, divided into two Sub-orders:|
|3.||Cynopithecidæ||Baboons and Macaques.|
Omitting man (for reasons stated in the preface) the three first families are considered by Professor Mivart to be sub-families of Simiidæ; but as the geographical distribution of the Old World apes is especially interesting, it is thought better to treat them as families, a rank which is claimed for the anthropoid apes by many naturalists.
As no good systematic work on the genera and species of bats has been yet published, I adopt the five families as generally used in this country, with the genera as given in the papers of Dr. J. E. Gray and Mr. Tomes. A monograph by Dr. Peters has long been promised, and his outline arrangement was published in 1865, but this will perhaps be materially altered when the work appears.
The genera of Chiroptera are in a state of great confusion, the names used by different authors being often not at all comparable, so that the few details given of the distribution of the bats are not trustworthy. We have therefore made little use of this order in the theoretical part of the work.
The osteology of the Insectivora has been very carefully worked out by Professor Mivart in the Journal of Anatomy and Physiology (Vol. ii., p. 380), and I follow his classification as given there, and in the Proceedings of the Zoological Society (1871).
|14. Galeopithecidæ||Flying Lemurs.|
|15. Macroscelididæ||Elephant Shrews.|
|16. Tupaiidæ||Squirrel Shrews.|
|19. Potamogalidæ||Otter Shrew.|
|20. Chrysochloridæ||Golden Moles.|
The next order, Carnivora, has been studied in detail by Professor Flower; and I adopt the classification given by him in the Proceedings of the Zoological Society, 1869, p. 4.
|Æluroidea||23. Felidæ||Cats, Lion, &c.|
|Cynoidea||28. Canidæ||Dogs, Foxes, &c.|
|Pinnipedia||33. Otariidæ||Eared Seals.|
The Cetacea is one of those orders the classification of which is very unsettled. The animals comprising it are so huge, and there is so much difficulty in preserving them, that only a very few species are known with anything like completeness. A considerable number of genera and species have been described or indicated; but as many of these are founded on imperfect specimens of perhaps a single individual, it is not to be wondered at that those few naturalists who occupy themselves with the study of these large animals, cannot agree as to the proper mode of grouping them into natural families. They are, however, of but little importance to us, as almost all the species inhabit the ocean, and of only a few of them can it be said that anything is accurately known of their distribution. I therefore consider it best to follow Professor Carus, who makes a smaller number of families; but I give also the arrangement of Dr. Gray in his British Museum catalogue of whales and seals, as modified subsequently in the Proceedings of Zoological Society, 1870, p. 772. The Zeuglodontidæ, a family of extinct tertiary whales, are classed by Professors Owen and Carus between Cetacea and Sirenia, while Professor Huxley considers them to have been carnivorous and allied to the seals.
|Fam. (Carus).||Fam. (Gray).|
|Monodontidæ||40.||(Part of Delphinidæ.)|
The order Sirenia, comprising the sea-cows, consists of a single family:
Family 42. Manatidæ.
The extensive order Ungulata comprises the three orders Pachydermata, Solidungula, and Ruminantia of the older naturalists. The following classification is that now generally adopted, the only difference of opinion being as to whether some of the groups should be classed as families or sub-families, a matter of little importance for our purpose:
|52. Bovidæ||Cattle, Sheep, Antelopes, &c.|
The two next orders consist of but a single family each, viz.:
We now come to the Rodentia, a very extensive and difficult order, in which there is still much difference of opinion as to the details of classification, although the main outlines are pretty well settled. The foundations of a true classification of this order were laid by Mr. G. R. Waterhouse more than thirty years ago, and succeeding authors have done little more than follow his arrangement with unimportant modifications. Professor Lilljeborg, of Upsala, has however made a special study of this group of animals, and has given an original and detailed classification of all the genera. (Systematisk Öfversigt af de Gnagande Däggdjuren, Glires. Upsala, 1866.) I follow this arrangement with a few slight modifications suggested by other naturalists, and which make it better adapted for the purposes of this work.
|59. Saccomyidæ||Pouched Rats.|
|65. Echimyidæ||Spiny Rats.|
|66. Cercolabidæ||Tree Porcupines|
The Edentata have been classified by Mr. Turner, in the Proceedings of the Zoological Society (1851, p. 205), by Dr. Gray in the British Museum Catalogue, and by Professor Carus in his Handbuch. The former takes a middle course between the numerous families of Dr. Gray, seven in number, and the two families to which Professor Carus restricts the existing species. I therefore follow Mr. Turner.
|Entomophaga||72. Manididæ||Scaly Ant-eaters.|
The Marsupials have been well classified and described by Mr. Waterhouse in the first volume of his Natural History of Mammalia, and his arrangement is here followed. The suborders adopted by Professor Carus are also given.
|Rapacia (Wagner)||76. Didelphidæ||Opossums.|
|77. Dasyuridæ||Native Cats.|
|78. Myrmecobiidæ||Native Ant-eater.|
|Poephaga (Owen)||80. Macropodidæ||Kangaroos.|
|Carpophaga (Owen)||81. Phalangistidæ||Phalangers.|
|Rhizophaga (Owen)||82. Phascolomyidæ||Wombats.|
The last order, the Monotremata, consist of two families, which Professor Carus combines into one, but which it seems more natural to keep separate.
Birds are perhaps the most difficult to classify of all the divisions of the vertebrata. The species and genera are exceedingly numerous, and there is such a great uniformity in general structure and even in the details of external form, that it is exceedingly difficult to find characters by which orders and families can be characterised. For a long time the system of Vigors and Swainson was followed; but this wholly ignored anatomical characters and in many cases plainly violated well-marked affinities. Characters derived from the form of the sternum, the scutellation of the tarsi, and the arrangement of the feathers, have all assisted in determining natural groups. More recently Professor Huxley has applied the variations of the bony palate to the general arrangement of birds; and still more recently Professor Garrod has studied certain leg-muscles for the same purpose. The condition of the young as regards plumage, and even the form, texture, and coloration of the egg, have also been applied to solve doubtful cases of affinity; yet the problem is not settled, and it will probably remain for another generation of ornithologists to determine with any approach to accuracy what are the most natural divisions of the class into orders and families. In a work like the present it is evidently not advisable to adopt all the recent classifications; since experience has shown that no arrangement in which one set of characters is mainly relied on, long holds its ground. Such modifications of the old system as seem to be well established will be adopted; but the older groups will be adhered to in cases where the most recent classifications are open to doubt, or seem inconvenient as separating families, which, owing to their similarity in general structure, form and habits are best kept together for the purposes of geographical distribution.
The old plan of putting the birds of prey at the head of the class, is now almost wholly given up; both because they are not the most highly organised, but only one of the most specialised forms of birds, and because their affinities are not with the Passeres, but rather with the cormorants and some other of the aquatic groups. The Passeres therefore are placed first; and the series of families is begun by the thrushes, which are certainly the most typical and generally well-organised form of birds. Instead of the Scansores and Fissirostres of the older authors, the order Picariæ, which includes them both, is adopted, but with some reluctance; as the former are, generally speaking, well marked and strongly contrasted groups, although certain families have been shown to be intermediate. In the Picariæ are included the goat-suckers, swifts, and humming-birds, sometimes separated as a distinct order, Macrochires. The parrots and the pigeons form each a separate order. The old groups of Grallæ and Anseres are preserved, as more convenient than breaking them up into widely separated parts; for though the latter plan may in some cases more strictly represent their affinities, its details are not yet established, nor is it much used by ornithologists. In accordance with these views the following is the series of orders and families of birds adopted in this work:
|1. Passeres||Including the great mass of the smaller birds—Crows,|
Finches, Flycatchers, Creepers, Honeysuckers, &c., &c.
|2. Picariæ||Including Woodpeckers, Cuckoos, Toucans, Kingfishers,|
Swifts, &c., &c.
|3. Psittaci||Parrots only.|
|4. Columbæ||Pigeons and the Dodo.|
|5. Gallinæ||Grouse, Pheasants, Curassows, Mound-builders, &c.|
|6. Opisthocomi||The Hoazin only.|
|7. Accipitres||Eagles, Owls, and Vultures.|
|8. Grallæ||Herons, Plovers, Rails, &c.|
|9. Anseres||Gulls, Ducks, Divers, &c.|
|10. Struthiones||Ostrich, Cassowary, Apteryx, &c.|
The Passeres consist of fifty families, which may be arranged and grouped in series as follows. It must however be remembered that the first family in each series is not always that which is most allied to the last family of the preceding series. All extensive natural groups consist of divergent or branching alliances, which renders it impossible to arrange the whole in one continuous series.
|A.—Typical or Turdoid Passeres.|
|12. Phyllornithidæ||Green Bulbuls.|
|44. Dendrocolaptidæ||American Creepers.|
The preceding arrangement is a modification of that proposed by myself in the Ibis (1874, p. 406). The principal alterations are adding the families Panuridæ and Sittidæ in series A, commencing series B with Dicæidæ; bringing Vireonidæ next to the allied American family Mniotiltidæ; and placing Motacillidæ in series C next to Alaudidæ. At the suggestion of Professor Newton I place Menuridæ and Atrichidæ apart from the other Passeres, as they both possess striking peculiarities of anatomical structure.
The heterogeneous families constituting the order Picariæ may be conveniently arranged as follows:
|Intermediate||59. Leptosomidæ||The Leptosoma.|
|72. Steatornithidæ||The Guacharo.|
The Psittaci or parrot tribe are still in a very unsettled state of classification; that recently proposed by Professor Garrod differing widely from the arrangement adopted in Dr. Finsch's monograph of the order. Taking advantage of the researches of these and other authors, the following families are adopted as the most convenient in the present state of our knowledge:
|76. Cacatuidæ||The Cockatoos.|
|77. Platycercidæ||The Broad-tailed Paroquets of Australia.|
|78. Palæornithidæ||The Oriental Parrots and Paroquets.|
|79. Trichoglossidæ||The Brush-tongued Paroquets and Lories.|
|80. Conuridæ||The Macaws and their allies.|
|81. Psittacidæ||The African and South American Parrots.|
|82. Nestoridæ||The Nestors of New Zealand.|
|83. Stringopidæ||The Owl-parrots of New Zealand.|
The Columbæ, or pigeons, are also in a very unsatisfactory state as regards a natural classification. The families, sub-families, and genera proposed by various authors are very numerous, and often quite irreconcilable. I therefore adopt only two families; and generally follow Mr. G. R. Gray's hand-list for the genera, except where trustworthy authorities exist for a different arrangement. The families are:
|84. Columbidæ||Pigeons and Doves.|
|85. Dididæ||The extinct Dodo and allies.|
The Gallinæ, or game-birds, may be divided into seven families:
|87. Tetraonidæ||Partridges and Grouse.|
|Euplocaminæ||Fire-backed Pheasants, &c.|
The Opisthocomi consist of one family containing a single species, the "Hoatzin" of Guiana.
Family 93. Opisthocomidæ.
The Accipitres, or birds of prey, which were long considered to be the highest and most perfect order of birds, are now properly placed lower down in the series, their affinities being more with the aquatic than with the perching birds. The following is the arrangement adopted by Mr. Sharpe in his recently published British Museum catalogue of diurnal birds of prey:—
The Grallæ or Grallatores are in a very unsettled state. The following series of families is in accordance with the views of some of the best modern ornithologists:
|99. Rallidæ||Rails, &c.|
|100. Scolopacidæ||Sandpipers and Snipes|
|114. Plataleidæ||Spoonbills and Ibis.|
The Anseres or Natatores are almost equally unsettled. The flamingoes are usually placed in this order, but their habits best assort with those of the waders.
|118. Anatidæ||Duck and Geese.|
The last order of birds is the Struthiones or Ratitæ, considered by many naturalists to form a distinct sub-class. It consists of comparatively few species, either living or recently extinct.
In reptiles I follow the classification of Dr. Günther as given in the Philosophical Transactions, vol. clvii., p. 625. He divides the class into five orders as follows:—
|3. Rhynchocephalina||The Hatteria.|
In the arrangement of the families comprised in each of these orders I also follow the arrangement of Dr. Günther and Dr. J. E. Gray, as given in the British Museum Catalogue, or as modified by the former gentleman who has kindly given me much personal information.
The Ophidia, or Snakes, form the first order and are classified as follows:—
|Innocuous Snakes||1. Typhlopidæ||Burrowing Snakes.|
|5. Calamaridæ||Dwarf ground-snakes.|
|7. Colubridæ||Colubrine Snakes.|
|8. Homalopsidæ||Fresh-water Snakes.|
|13. Dipsasidæ||Nocturnal tree-snakes.|
|15. Lycodontidæ||Fanged ground-snakes.|
|20. Elapidæ||Cobras, &c.|
|Viperine Snakes||24. Crotalidæ||Pit-vipers.|
|25. Viperidæ||True vipers.|
The second order, Lacertilia, are arranged as follows:—
|30. Varanidæ||Water Lizards.|
|33. Lacertidæ||Land Lizards.|
|41. Gymnophthalmidæ||Gape-eyed Scinks.|
|42. Pygopodidæ||Two-legged Lizards.|
|51. Agamidæ||Fringed Lizards.|
The third order, Rhynchocephalina consists of a single family:—
53. RhynchocephalidæThe Hatteria of New Zealand.
The fourth order, Crocodilia or Loricata, consists of three families:—
The fifth order, Chelonia, consists of four families:—
|57. Testudinidæ||Land and fresh-water Tortoises.|
|58. Chelydidæ||Fresh-water Turtles.|
|59. Trionychidæ||Soft Turtles.|
|60. Cheloniidæ||Sea Turtles.|
In the Amphibia I follow the classification of Professor Mivart, as given for a large part of the order in the Proceedings of the Zoological Society for 1869. For the remainder I follow Dr. Strauch, Dr. Günther, and a MSS. arrangement kindly furnished me by Professor Mivart.
The class is first divided into three groups or orders, and then into families as follows:—
Order II.—BATRACHIA URODELA.
|5. Menopomidæ||Giant Salamanders.|
|6. Salamandridæ||Salamanders and Newts.|
Order III.—BATRACHIA ANOURA.
|7. Rhinophrynidæ||Toads.||16. Pelodryadæ||Tree Frogs.|
|8. Phryniscidæ||17. Hylidæ|
|9. Hylaplesidæ||18. Polypedatidæ|
|10. Bufonidæ||19. Ranidæ||Frogs.|
|11. Xenorhinidæ||20. Discoglossidæ|
|12. Engystomidæ||21. Pipidæ||Tongueless|
|13. Bombinatoridæ||Frogs.||22. Dactylethridæ|
These are arranged according to the classification of Dr. Günther, whose great work "The British Museum Catalogue of Fishes," has furnished almost all the material for our account of the distribution of the class.
In that work all existing fishes are arranged in six sub-classes and thirteen orders. A study of the extraordinary Ceratodus from Australia has induced Dr. Günther to unite three of his sub-classes; but as his catalogue will long remain a handbook for every student of fishes, it seems better to follow the arrangement there given, indicating his later views by bracketing together the groups he now thinks should be united.
|Teleostei||1. Acanthopterygii||47||Gasterosteidæ to Notacanthi.|
|Ganoidei||2. Do. Pharyncognathi||5||Pomacentridæ to Chromidæ.|
|3. Acanthini||6||Gadopsidæ to Pleuronectidæ.|
|4. Physostomi||29||Siluridæ to Pegasidæ.|
|5. Lophobranchii||2||Solenostomidæ and Syngnathidæ.|
|6. Plectognathi||2||Sclerodermi and Gymnodontes.|
|Ganoidei||8. Holostei||3||Amiidæ to Lepidosteidæ.|
|9. Chondrostei||2||Accipenseridæ and Polydontidæ.|
|11. Plagiostomata||15||Carchariidæ to Myliobatidæ.|
|Cyclostomata||12. Marsipobranchii||2||Petromyzontidæ and Myxinidæ.|
The families and genera of insects are so immensely numerous, probably exceeding fifty-fold those of all other land animals, that for this cause alone it would be impossible to enter fully into their distribution. It is also quite unnecessary, because many of the groups are so liable to be transported by accidental causes, that they afford no useful information for our subject; while others are so obscure and uninteresting that they have been very partially collected and studied, and are for this reason equally ineligible. I have therefore selected a few of the largest and most conspicuous families, which have been so assiduously collected in every part of the globe, and so carefully studied at home, as to afford valuable materials for comparison with the vertebrate groups, when we have made due allowance for the dependence of many insects on peculiar forms of vegetation, and the facility with which many of them are transported either in the egg, larva, or perfect state, by winds, currents, and other less known means.
I confine myself then, almost exclusively, to the sixteen families of Diurnal Lepidoptera or butterflies, and to six of the most extensive, conspicuous, and popular families of Coleoptera. The number of species of Butterflies is about the same as that of Birds, while the six families of Coleoptera selected, comprise more than twenty thousand species, far exceeding the number of all other vertebrates. These families have all been recently catalogued, so that we have very complete information as to their arrangement and distribution.
LEPIDOPTERA DIURNA, OR BUTTERFLIES.
|1. Danaidæ.||9. Libytheidæ.|
|2. Satyridæ.||10. Nemeobiidæ.|
|3. Elymniidæ.||11. Eurygonidæ.|
|4. Morphidæ.||12. Erycinidæ.|
|5. Brassolidæ.||13. Lycænidæ.|
|6. Acræidæ.||14. Pieridæ.|
|7. Heliconidæ.||15. Papilionidæ.|
|8. Nymphalidæ.||16. Hesperidæ.|
COLEOPTERA, OR BEETLES.
|1. Cicindelidæ||Tiger-beetles.||4. Cetoniidæ||Rose-chafers.|
|2. Carabidæ||Ground-beetles.||5. Buprestidæ||Metallic Beetles.|
|3. Lucanidæ||Stag-beetles.||6. Longicornia||Long-horned Beetles.|
The above families comprise the extensive series of ground beetles (Carabidæ) containing about 9,000 species, and the Longicorns, which are nearly as numerous and surpass them in variety of form and colour as well as in beauty. The Cetoniidæ and Buprestidæ are among the largest and most brilliant of beetles; the Lucanidæ are pre-eminent for remarkable form, and the Cicindelidæ for elegance; and all the families are especial favourites with entomologists, so that the whole earth has been ransacked to procure fresh species.
Results deduced from a study of these will, therefore, fairly represent the phenomena of distribution of Coleoptera, and, as they are very varied in their habits, perhaps of insects in general.
The Mollusca are usually divided into five classes as follows:—
|II.||Gasteropoda||Snails and aquatic Univalves.|
The Gasteropoda and Conchifera alone contain land and freshwater forms, and to these we shall chiefly confine our illustrations of the geographical distribution of the Mollusca. The classification followed is that of Dr. Pfeiffer for the Operculata and Dr. Von Martens for the Helicidæ. The families chiefly referred to are:—