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ilontion: C. J. CLAY AND SONS, 

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SINCE the publication of Dr Wallace's book on the geo- 
graphical distribution of animals in general, the only works which 
have appeared relating to Mammals from the same point of view 
are the small volume by Mr F. E. Beddard and the series of 
papers by Mr W. L. Sclater, referred to in the Appendix. 

Both the latter admittedly take but little account of fossil 
forms ; and there is accordingly ample room for a work which 
should collect and arrange the information on this subject, and 
indicate the deductions which may be drawn therefrom. This 
task has been attempted in the present volume. The subject 
is, however, so vast, and the information relating to it scattered 
through so many publications, that it is probable many points of 
interest or importance have not been noticed. From the mode 
of arrangement of the work, a considerable amount of repetition 
was inevitable. 

The long time that the volume was in the press will account 
for its containing no allusion to certain papers which have ap- 
peared recently. It may therefore be mentioned here that 
recent discoveries have shown a slight intermixture of northern 
types in the Palaeozoic flora of Argentina, so that the isolation 
of the northern and southern floras is .not so nearly complete 
as was supposed. A paper just published by M. Boule 1 indi- 

1 Comptes RenduS) vol. cxxii (1896). 


cates that the relationship between Cadurcotherium and the 
Astrapotheria is closer than suggested on page 82. Hence there 
is further evidence that South America received its Tertiary 
ungulates by way of Africa. The extinct Patagonian birds 
Phororhachis and Brontornis appear, according to Mr C. W. 
Andrews, to be nearly allied to the existing South American 
seriema (Cariama), and since Filholornis, of the French Phos- 
phorites, has been shown by Prof. A. Milne-Edwards to be 
related to the hoatzin (Opisthocomus), it would seem that all 
these peculiar South American types of birds have a history 
somewhat similar to that of the extinct ungulates of the same 



June ist, 1896. 




Distributional Area and Station Influence of Temperature Humidity 
Other Factors in Distribution Importance of Palaeontology Inequality 
in the Ages of different Groups of Animals Different Groups have different 
Geographical Distribution Importance of Mammals in Geographical 
Distribution Classification of Mammals Barriers to Dispersal of Mam- 
mals Influence of Man on Distribution Extinction of the larger 
Plistocene Mammals Distributional Areas of Genera and Species 
Centres of Evolution Permanency of Continents and Ocean-Basins 
Zoological Realms and Regions ....... i 


Definition and Characters of the Realm Australian Region Monotremes 
Marsupials Rodents Carnivores Ungulates Bats List of Australian 
and Papuan Genera Polynesian Region Hawaiian Region Austro- 
Malayan Region Palseontological History of Marsupials How Australia 
received its Fauna . . . . . . . . .28 



Extent and Characters Mammaliferous Deposits Monkeys Bats Insecti- 
vora Carnivores Ungulates Horses Litopterna Astrapotheria 
Toxodonts Pyrotheria Proboscideans Rodents Edentates Arma- 
dillos and Glyptodonts Sloths Anteaters Ground-sloths Marsupials 
Cetaceans Early Distinction of the Neogseic Fauna Early Separation 
of N. and S. America Incursion of Northern Mammals Distinctness of 
the existing Fauna Origin of the Santa Cruz Fauna Antarctica and the 
South American element in the Ethiopian Fauna Conclusion Sub- 
regions 64 



Features of the Arctogaeic Fauna Community of earliest Fauna Evidence of 
Secondary Reptiles Puerco Fauna Lemuroids Insectivora Carnivores 
Rodents Ungulates Summary of the Characteristics of the Mamma- 
lian Fauna of Arctogaea . . . . . . . . 1 44 



Mammalian Groups peculiar to Eastern Arctogsea Tertiary Mammalian 
Fauna of Eastern Arctogsea Oligocene Fauna Miocene Fauna Older 
Pliocene Fauna Pikermi and allied Faunas Sivvalik Fauna Higher 
Pliocene Faunas . . . . . . . . . 1/9 



Mammalian Fauna Relations of Madagascar to the Mainland . . 213 



Characteristics of the Mammalian Fauna Birds Past History of Ethiopia 
Sub-regions . . . . . . . . . . .227 



Sub-regions Characteristics of the Mammalian Fauna Past History of the 
Region Malayan Sub-region Nicobars, Mentawi, and Christmas Islands 
Philippine Sub-region ......... 264 




Characteristics of the Mammalian Fauna Mammals of the Eastern Holarctic 
Region Plistocene Fauna of the Holarctic Region Geographical Changes 
since the Plistocene Western Division of the Region Arctic Sub-region 
European Sub-region Central Asian Sub-region Tibetan Sub-region 
Mantchurian Sub-region Mediterranean Sub-region Kashmir Ca- 
nadian Sub-region Transition Zone . . . . . . 308 



Limits Characteristics of Mammalian Fauna Extinct Groups of Mammals 
characteristic of Western Arctogrea Distinctness of the Region Dual 
origin of Groups . . .. . . .-..'.'' V '' '. 363 


LIST OF WORKS AND PAPERS .- &*&, . . ..:..-, fj . 383 
INDEX . . . . V . 386 




Fig. i. The Duckbill {Ornithorhyncus anatinus] . . . .32 

2. Skull of Rat-Kangaroo 35 

,, 3. Skull of Extinct Phalanger ( Thylacoleo carnifex) 37 

4. Banded Anteater (Myrmecobius fasciatus] .... 39 

,, 5. Fore Part of Skull of Babirusa (Babirusa alfums} . . -47 

,, 6. Lower Jaw of Triconodon, or Allied Form . . . .52 

,, 7. Lower Jaw of Phascolotheriiim . . . . . -53 

,, 8. Lower Jaw of Amphilestes . . . . . . -53 

,, 9. Lower Jaw of Homunculus ....... 69 

,, 10. Skeleton of Macrauchenia patachonica ..... 76 

,, n. Skeleton of Hind Foot of Rhinoceros 78 

,, 12. Carpus and Metacarpus of Hyrax and Elephant . . -78 

,, 13. Upper Molar of Macraiichenia .80 

,, 14. Upper Molar of Oxyodontotherium ...... 80 

,, 15. Upper Molar of Astrapotherium . . . . . . 81 

16. Skull of Nesodon 83 

,, 17. Palate of Typotherium ........ 84 

1 8. External Skeleton of Armadillo ( Tatusia giganted] . . . 93 

19. Internal Skeleton of a Glyptodont ...... 95 

,, 20. External Skeleton of Glyptodon clavipes . . . . -97 

,, 21. External Skeleton of Panochthus tuberculatus . . . -99 

,, 22. Tamandua Anteater 102 

,, 23. Skull of Mylodon ......... 104 

,, 24. Skeleton of Scelidothcrium leptocephalum . . . .106 

25. Lower Jaw of Prothylacinus ....... 108 

,, 26. Lower Jaw of Acdestis oweni . . . . . . .no 

,, 27. Lower Jaw of Abderites . . . . . . rii 

,, 28. Lower Jaw of Plagiaulax 149 

,, 29. Skull of Tritylodon 150 

,, 30. Molar of Tritylodon -150 

31. Upper Cheek-Teeth of Plesiadapis 155 

,, 32. Upper Cheek -Teeth of Microcharus 156 



Fig- 33- Upper Molars of A rctocyon 159 

,, 34. Upper Molar of Ancodus . . . . . . . 161 

,, 35. Skeleton of Elotherinm 162 

36. Upper Cheek-Teeth of Palaotheriiim 166 

37. Upper Cheek-Teeth of A nchitherium , . . . . 167 

,, 38. Upper Molar of Horse 167 

39. Skull of Hipparion . . . >. , . . ... .168 

,, 40. Upper Molar of Rhinoceros . . ..' , i, . ;- . ,*.,.. 169 

,, 41. Skeleton of Titanotherium . . . .- k , . 171 

,, 42. Upper Molar of Mastodon . . .. : .;...: >.". - . 173 

,, 43. Upper Cheek-Teeth of Coryphodon . . < .,...?!.;* . 174 

,, 44. Upper Cheek-Teeth of Anoplotherium .. \ ,'. , . , ,. ... ; . 185 

,, 45. Skull of PalcBorias . +< -; *,< '.;/*- .1. . . .. '. .. . 199 

, , 46. Upper Molar of Merycopotanius . . : . 1 < c, . . 203 

,, 47. Restoration of Sivatheriutn . . . . -. . v /... . 205 

,, 48. Skull of Lemur . , ' . . . . .; ' ; . 217 

,, 49. Ring-tailed Lemur (Lemur catta) * . ; '\ '. '{. rt&l . . 218 

,, 50. Skull of Aye-aye (Chiromys madagascariensis} .,,.. . 219 

,, 51. The Fossa (Cryptoproctaferooc] . ... *,;,., 221 

,, 52. African Jumping-Shrew (Macroscelides tetradactylus} . . 232 

> 53- West African Potamogale (Potamogale velox] ;.-. ... . 233 

54. Cape Hunting-Dog (Lycaon pictus] . .i ....... . 236 

,, 55. Fulgent African Flying Squirrel {Anomahirus fiilgens) . . 237 

,, 56. Head of African Wart- Hog (Phacochcerus cethiopicus) . . 241 

,, 57. Water-Chevrotain (Dorcatherium aquaticuni) .... 243 

,, 58. Head of Gemsbok (Oryx gazelld) . . . . . . 246 

,, 59. Cape Hy rax (Procavia capensis) .... . . 248 

60. Slow Loris (Nycticebus tardigradus} ..... 269 

,, 6r. Tree-Shrew (Tupaia tana) 271 

,, 62. Indian Sloth-Bear (Melursus ursinus) 275 

,, 63. Indian Ratel (Mellivora ratel) . 276 

,, 64. Japanese Serow (Nernorh&dus crispus] 279 

,, 65. White-bellied Pangolin (Manis trictispis) .... 283 

,, 66. Russian Desman (Myogale moschatd] . . . . 320 

,, 67. Head of Spanish Ibex (Capra pyrenaica} 323 

,, 68. Musk-Deer (Moschus moschiferus] . . . . . 325 

,, 69. Rocky Mountain Goat (Haploceros montanus] .... 343 

,, 70. Musk-Ox (Ovibos moschatus) . . . . . i . 346 

, , 71. Musquash (Fiber zibethicus) . 362 

,, 72. Face of Geomys bursarius . . . . * . . . . 366 

,, 73. Face of Thomomys talpoides . . . . ... . 366 

74. Foot of Geomys 377 

,, 75. Head of Mule-Deer (Cariacus macrotis) 368 

,, 76. Head of Prong-buck (Antilocapra americana) .... 369 



Fig. 77. 

Skeleton of Patriofelis ferox ..... 

37 2 

,. 78. 

Skeleton of Agriochcerus latifrons .... 



Hind-foot of Agriochcerus ..... 


,. 80. 

Skull of Protoceras ...... 


,, 81. 

Molar of Palceosyops 



Extremity of Skull of Uintatherium 


Figs, i, 4, 22, 28, 32, 48, 49, 50, 52, 53, 54, 55, 57, 59, 60, 63, 65, 66, 68, 
70, 71 are from The Study of Mammals > Living and Extinct, by Flower and 
Lydekker. Figs. 13, 14, 15, 16, 17, 18, 19, 20, 21, 23, 24 are taken from the 
author's work on Argentine Fossil Mammals, published in the An. Mus. La 
Plata. Figs. 9, 25, 26, 27 are from Senor Ameghino. Fig. 67 is from a photo- 
graph by Mr Abel Chapman. Fig. 82 is from Prof. Cope, fig. 81 from Prof. 
Earle, fig. 40 from Prof. Boyd Dawkins, fig. 30 from Prof. Fraas, and figs. 7, 8 
from Prof. Goodyear. Fig. 46 is taken from a plate in Hutchinson's Extinct 
Monsters ; 58 from a photograph by Mr Eccles, of Reading, and 75 from one 
in the possession of Mr J. E. Harting. Figs. 6, 35, 41, 43 are from Prof. 
Marsh; and n, 12, 31, 33, 37, 78, 79, 80 from Prof. H. F. Osborn. For the 
blocks of figs. 72, 73, 74, the author is indebted to Dr C. H. Merriam. 
Fig. 45 is from Nicholson and Lydekker's Manual of Paleontology ; fig. 29 
from Owen; fig. 64 from Dr Sclater in the Proc. Zool. Soc.; fig. 77 from Dr 
Wortman, and 39 from Prof, von Zittel. 




Distributional Area and Station Influence of Temperature Humidity 
Other Factors in Distribution Importance of Palaeontology Inequality 
in the Ages of different Groups of Animals Different Groups have different 
Geographical Distribution Importance of Mammals in Geographical 
Distribution Classification of Mammals Barriers to Dispersal of Mam- 
mals Influence of Man on Distribution Extinction of the larger 
Plistocene Mammals Distributional Areas of Genera and Species 
Centres of Evolution Permanency of Continents and Ocean-Basins 
Zoological Realms and Regions. 

THAT there are differences in the animals and plants of different 
districts and different countries is a fact apparent to every person 
who has travelled at all; while to those who have travelled ex- 
tensively it will further be evident that the amount of this difference 
is by no means correlated with the distance of one country from 
another, the fauna of Japan, for instance, being much more like 
that of England and France than is the fauna of Eastern Africa to 
that of the adjacent island of Madagascar. Unfortunately, among 
persons who are not conversant with the principles of zoological 
science, the distinction between the faunas of different countries 
has been much obscured by the practice common to almost all 
the old voyagers and colonists of bestowing upon the animals of 
new countries the names of such Old World creatures as they 
appeared most nearly to resemble. The puma of America was, 
for instance, called the lion, and the jaguar of the same country, 
the tiger; while the koala of Australia was christened the native 
bear, and its marsupial allies the dasyures are still commonly 
spoken of as native cats. To students of the science of 
L. I 


Geographical Distribution it is, therefore, essential to discard such 
misleading popular titles, and to speak of animals by their correct 

Apart from the specific or generic distinctions between the 
animals of one country and another, the observer 


Area and sta- will not fail to notice more or less well-marked 
differences between those inhabiting different dis- 
tricts of a single country ; such differences being most intensified 
when a country presents great variation in its physical features. 
An excellent instance of this is afforded by South America, where 
there are the open grassy plains of the Argentine, the dense tropi- 
cal forests of Paraguay and Brazil, and the snow-clad heights of 
the Andes. In the former tract the traveller will meet with the 
peculiar rodents known as viscachas, the Patagonian cavy, a species 
of deer, numerous armadillos, and the rhea (miscalled the American 
ostrich). In the Brazilian forests, on the other hand, he will find 
monkeys, marmosets, tapirs, tree-porcupines, sloths, and anteaters, 
together with certain armadillos which are for the most part speci- 
fically or generically distinct from those of the pampas. If, on 
the other hand, he ascend high on the Andes, he will leave 
behind the animals of the forest, to be confronted with chinchillas, 
guanacos and vicunas. Different, however, as are the animals of 
these various districts, yet an acquaintance with their zoological 
affinities will prove that many of them belong to closely allied 
groups, some of which are met with in no other parts of the world. 
This will serve to show that they belong to what is known as one 
zoological province or region, and that the differences between the 
faunas of different districts of that province are due to the physical 
variations between its component districts. 

Perhaps this point may be still better illustrated by the cases 
where the same species of animal is restricted to different districts 
of one country or continent. For instance, the common squirrel 
is only found in the wooded districts of Europe, and is entirely 
absent from open plains. The chamois, again, is only met with 
in the isolated mountain ranges of the Pyrenees, the Alps, and the 
Caucasus ; while the Siberian ibex of the Altai reappears in Tibet 
and the Himalaya, but is wanting in the intervening tracts. In 
these instances Europe would be spoken of as the distributional 


area of the squirrel and the chamois, and Central Asia as that of 
the Siberian ibex ; but the particular districts suited to the exist- 
ence of each would be termed its station. And here it may be 
mentioned that whereas the distributional area of a species is 
generally continuous, its various stations may be partially or com- 
pletely isolated, as in the instances of the aforesaid mountain 
animals, which cannot live on the plains below. 

Station is thus seen to be very intimately connected with 
temperature; and of this a very striking example may 
be found among the mammals of South America. 
As already mentioned, the llama-like animals respec- 
tively known as vicunas and guanacos are met with in company 
on the highlands of the Cordillera in Peru and Ecuador, but as 
we go further south the latter are found on the plains of southern 
Argentina and Patagonia, as well as in the island of Tierra-del- 
Fuego, at the sea-level. Here, then, there is a clear proof of the 
intimate connection existing between temperature and station; 
the guanaco, being an animal which can live only in cold or 
temperate climates, finds suitable conditions for its existence 
in tropical latitudes solely at a height of many thousands of feet, 
although further south it is able to thrive at the sea-level. 

This being so, it is obvious that temperature must likewise 
exert a very considerable influence on the whole distributional 
area of many animals. Of this, the most marked instance is found 
in the fauna of the Arctic regions, which forms a circumpolar zone 
of animals more or less markedly distinct from those dwelling 
further south. And if the whole land-area of the world were con- 
nected, and not broken up by mountain-chains, its faunas might 
probably be divided into zones or belts, whose limits would mainly 
depend upon temperature. 

In this connection it may be mentioned that the instance of the 
range of the guanaco is of considerable importance in regard to 
a decided difference between the Old and New Worlds in respect 
to the influence of mountains on the present distribution of the 
animals of the two areas. In the Old World the chief mountain- 
ranges, such as the Pyrenees, Alps, Carpathians, Caucasus, Hindu- 
Koh, Himalaya, Thian Shan, and Altai, run in a more or less 
decided east-and-west direction; whereas in America, and more 

I 2 


especially in the southern half of that continent, they have a north- 
and- south trend. Consequently, whereas in the former area the 
mountain-ranges have acted as barriers to the dispersal of animal 
life, there has been no such obstacle to diffusion in America, and 
animals have been able to distribute themselves according to tem- 
perature-conditions. It is in consequence of this physical feature 
that a single species of cold-loving animal like the guanaco can 
range at the present day from the equator to latitude 55 south, 
whereas in the Old World the common ibex is now restricted to 
the isolated mountain-ranges of Europe, and the Siberian ibex is 
confined to the systems of the Himalaya and the Altai. Such 
isolated stations could, of course, only have been reached during 
a period when the general temperature of the northern hemisphere 
was much colder than it is at present, and the animals were thus 
enabled to cross the lowlands from one chain to the other ; and 
that such a cold period formerly existed, there is abundant evidence 
in the traces of an extensive glaciation found over a large portion 
of Europe and Asia. Although, therefore, strictly speaking, tem- 
perature has really had as much effect in the distribution of 
animals in the Eastern Hemisphere as in the Western, yet, since 
the glacial epoch, its influences have been considerably masked 
by the trend of the chief mountain-ranges, and likewise by the 
greater isolation of the different countries of the former area as 
compared with the latter. 

These essential differences thus render it impossible to mark 
out the Old World in the zones of animal distribution which have 
been attempted for North America ; and they will likewise serve 
largely to explain the divergence of views on this point which may 
be noticed between the writings of Drs Wallace and Merriam. 

The latter writer 1 , whose conclusions are mainly based on the 
evidence of North American animals and plants, is of opinion 
that in the northern hemisphere " animals and plants are restricted 
in southward distribution by the mean temperature of a brief 
period covering the hottest part of the year"; and it is added that 
in certain districts the mingling of essentially northern types with 
those characteristic of a more southerly zone is due to the mean 
temperature of the hottest part of the year being sufficiently low 
1 Appendix, No. 20. 


for the existence of the former, while the total quantity of heat 
suffices for the latter. In other words, there is a low summer 
temperature combined with a high total sum of heat. 

Among the secondary causes affecting distribution, humidity, 
according to the same observer, may occupy the first 
place. "Humidity," he writes, " governs details of 
distribution of numerous species of plants, reptiles and birds, and 
of a few species of mammals, within the several temperature-zones. 
. . . Humidity and other secondary causes determine the presence 
or absence of particular species in particular localities within their 
appropriate zones, but temperature predetermines the possibili- 
ties of distribution ; it fixes the limits beyond which species 
cannot pass ; it defines broad trans-continental belts within 
which certain forms may thrive if other conditions permit, but 
outside of which they cannot exist, be the other conditions never 
so favourable." 

Important as the influence of temperature and, in a smaller 
degree, that of humidity, has undoubtedly been in other Fac 
determining the distributional limits of the species tors in Distri- 

f. , , , . , , . . bution. 

or genera ot animals and plants now inhabiting 
particular countries, and large as has been the part played by the 
glacial epoch in producing the present condition of things, it is 
evident that temperature has been by no means the only, even if 
it be the chief factor in distribution. In the first place there are 
several species, more especially among the carnivorous mammals, 
which seem quite independent of both station and temperature, 
the New World puma ranging from Patagonia to Canada, while 
the tiger inhabits alike the burning jungles of India and Burma, 
and the Arctic tundras of Siberia. Striking as such cases are, they 
are, however, to be regarded merely as examples of the individual 
adaptability of certain species, which, like the carnivores named, 
are able to obtain suitable food in any part of the world, and 
they do not throw any discredit on the power of temperature as 
a controlling factor in animal distribution generally. 

Of the utmost importance in this respect are the changes 
which the surface of the globe itself has undergone in past 
epochs, whereby continents that are now more or less completely 
sundered from one another were formerly connected, while what 


are now islands were once parts of continents, and vice versa. 
Such connections and disconnections, by allowing migrations at 
one time, and preventing them from taking place in the reverse 
direction at a subsequent epoch, have been the chief factors which 
have resulted in the present very remarkable difference in the 
faunas of different parts of the globe. And it is solely due to such 
changes that many of the lower types of mammalian life, like the 
marsupials of Australia, and the lemurs and insectivores of Mada- 
gascar, have been preserved at the present day ; their insulation 
having afforded them protection from the invasion of the larger 
and more specialised mammals of other parts of the world, by 
which they would inevitably have been swept away, had the two 
groups ever come in contact. It is in regard to these migratory 
movements of animals and changes in the land-surface of the 
globe that zoology and geography are brought into such close 
relationship ; the former science sometimes helping to explain the 
alterations that have taken place in the contours of the land, while 
in other cases the present distribution of the land explains the 
past history of the animals by which it is inhabited. 

To understand rightly the present distribution of animals, it is, 
im ortance however, essential to study their past history as 
of Paiaeonto- recorded by the preservation of their fossilised 
remains in the strata of the earth's crust ; as with- 
out such history it would be quite impossible to grasp the reason 
of many apparent anomalies in their present distribution. How, 
for instance, without the aid of palaeontology would it be possible 
to understand how it came about that tapirs are now found only 
in tropical America and the Malayan countries, or that marsupials 
occur solely in America and Australasia at the present day ? And 
here it may be well to mention that the science of geographical 
distribution depends essentially upon a belief on the part of the 
student that all animals are genetically connected one with 
another, and that the existing forms have originated from earlier 
kinds by some mode of evolution. Were this belief not accepted, 
the whole science of distribution would fall to pieces ; as if animals 
were separately created, there would be nothing calling for special 
explanation in the fact of tapirs being restricted to the two areas 



To those readers who may not be geologists, the following 
table of the leading divisions into which the strata of the earth's 
crust have been divided will probably be advantageous. Com- 
mencing with the highest beds, the series will run in descending 
order as follows, viz. : 

Tertiary. PLISTOCENE. Cavern and River Deposits. 

PLIOCENE. The "Crags" of the East Coast. 
MIOCENE. (Eningen beds of Baden. 
OLIGOCENE. Gypsum of Paris Basin; Phos- 
phorites of Central France. 
EOCENE. London Clay. 
Secondary. CRETACEOUS. Chalk, Upper Greensand, Gault, 

Lower Greensand, and Wealden. 
JURASSIC. Purbeck beds, Portland series, Kim- 
meridge Clay, Coral Rag, Oxford 
Clay, Great Oolite, Stonesfield 
Slate, Inferior Oolite, Lias. 
TRIASSIC. New Red Sandstone of Cheshire. 
Paleozoic. PERMIAN. Red Marls. 

CARBONIFEROUS. Coal-Measures and Moun- 

Older rocks of Wales and the 

Before attempting to draw any conclusions as to the former 
configuration of the surface of the earth from the 

,. . .,,...,... Inequality in 

distribution of the animals now inhabiting its dif- the Ages of dif- 

ferent countries, it is essential to understand that 
the different classes into which vertebrate animals 
are divided (and these only will be taken into consideration in the 
present volume) have a very different past history; the lower 
groups, such as fishes, reptiles, and amphibians being much older 
types than mammals and birds, and having attained their maximum 
development at a time when the two latter formed but a small 
minority of the earth's population. 

There is a considerable probability that at least a very 
large proportion of the animals that have populated the globe in 


the later geological epochs originated high up in the northern 
hemisphere, if not, indeed, in the neighbourhood of the pole itself 
(which is known to have enjoyed a genial climate during the 
Tertiary period), and that they gradually migrated southwards in 
a series of waves, probably under pressure of the development of 
new and higher types in high latitudes; and it is to such southerly 
migrations that the present marked differentiation of the fauna of 
different parts of the earth's surface is chiefly due. Whether such 
a northerly origin held good for the terrestrial life of the Secondary 
epoch, there are no means of determining ; but it would appear 
that the higher animals (which were chiefly reptiles) of that epoch 
were very similar throughout the world, and that the differentiation 
of faunas had scarcely, if at all, commenced. Instances of this 
are afforded, as noticed in the sequel, by the occurrence of an 
identical genus of mammal (Tritylodori) in the lower Secondary 
rocks of Europe and South Africa ; as well as by the close alliance 
between the dinosaurian reptiles from the Jurassic rocks of 
Europe, North America, Argentina, India, and Madagascar (the 
genera being in some cases identical), and likewise between 
the anomodont reptiles of the Trias of Europe and the early 
Secondary rocks of South Africa and India. 

Reptiles belonging to orders still existing, such as crocodiles 
and chelonians (tortoises and turtles), had already attained a high 
degree of development in the Eocene division of the Tertiary 
period, when many genera now living had already made their 
appearance, whereas at that time the mammals were quite differ- 
ent from the modern forms. At the same time the side-necked 
tortoises (Pleurodira) were the dominant forms in the northern 
hemisphere, whereas they have now all migrated to southern lands, 
their place in the north being taken by the more specialised 
S-necked group (Cryptodira). This, however, is not all, for the 
rhynchocephalians, of which the sole existing representative is 
the New Zealand tuatera (Sphenodori), attained their maximum 
development in the northern hemisphere during the early part of 
the Secondary epoch, and their southern migration must have 
taken place during some portion of the same period. The 
palaeontological history of amphibians is still very imperfectly 
known, but since the group as a whole is an ancient one, the 


migrations of the earlier forms must likewise probably have taken 
place at an early epoch. 

With mammals the case is very different. The earliest known 
forms, which date from the Triassic and Jurassic rocks, are chiefly 
marsupials and forms apparently allied to the monotremes, and it 
is probable that most of the descendants of these, as is more fully 
indicated in the sequel, migrated southwards during the early part 
of the Tertiary epoch, to find in Australasia a refuge from the 
competition of higher forms. Of the higher placental mammals, 
none of the modern types make their appearance before the 
Oligocene and Miocene periods, while many do not antedate the 
Pliocene. Their southern migrations accordingly took place later 
on in the Tertiary period, one of the earliest movements being the 
wandering of lemuroids, insectivores, and civet-like carnivores 
into South Africa and Madagascar. On the other hand, many 
other higher types, such as the hippopotami, giraffes, and antelopes, 
which were abundant in Europe and southern Asia during the 
Pliocene, only left their more northern homes to find a permanent 
abiding place in Africa at a very late epoch in the earth's history. 

Although the glacial epoch probably had a large share in the 
southern movements of the later Tertiary mammals, some cause 
with which we are unacquainted would appear to have been the 
impelling power at earlier epochs. But be this as it may, it is 
quite evident that a continuous series of waves of migrations of 
animal life has taken place throughout a very long portion of the 
earth's history. Similar migrations are also evident in the case of 
birds (which are likewise a modern group), many forms, such as 
secretary-birds and trogons, now exclusively southern in their 
distribution, being represented in Europe during the middle part of 
the Tertiary period. 

From this inequality in the ages, and consequently in the date 
of migration, of different groups of animals, it is 
manifest that there will be great differences in the 

present distribution of such groups; and hence it different 

. . .. , Geographical 

will be evident that zoological provinces indicated Distribution. 
by one group will not hold good for others. Notable 
instances of this are afforded by the very different divisions into 
which the globe is divided by those who take mammals, reptiles, 


or amphibians as their standards. Between birds and mammals, 
as might have been expected from the comparatively recent high 
development of these groups, there is much greater accord. 
Birds, however, differ from mammals (with the exception of bats) 
in their power of flight, which enables many of them to cross 
wide ocean-tracts, and therefore renders them less valuable as 
indicative of the changes that have taken place in the distribution 
of land and water than the latter, which, as a rule, require 
direct means of land transit for their wanderings. 

Excluding man (and for the most part bats) and likewise the 

aquatic forms, such as seals, whales, and porpoises, 

o/ Mamma" niammals are the animals best adapted for parcelling 

in Geographi- out the globe into zoological provinces for two chief 

cal Distribu- 3 _. . , r , . , , 

tion. reasons. Firstly, they form a group which only at- 

tained its maximum development at a comparatively 
late epoch of the earth's history; and, secondly, their movements 
are mainly limited by the extent of the land-surfaces of the globe 
which were in actual communication at the time of such migra- 
tions. Consequently they afford the safest and truest indications 
of the latest changes which have taken place in the distribution 
of land and water. 

As reference in the following pages will constantly have to be 

. fi made to the various groups of mammals, it will be 

tion of Mam- well to give a list in this place of the chief ordinal 

and subordinal groups into which the class is 

divided ; such as are now extinct being indicated by an asterisk. 

The list stands as follows, viz. : 

i. Order PRIMATES. Apes, Monkeys, and Lemurs. 

1 Suborder ANTHROPOIDEA. Apes and Monkeys. 

2 LEMUROIDEA. Lemurs. 

ii. Order CHIROPTERA. Bats. 

1 Suborder MEGACHIROPTERA. Fruit-bats. 

2 ,, MICROCHIROPTERA. Insectivorous Bats. 

iii. Order INSECTIVORA. Insectivores. 

1 Suborder DERMOPTERA. Flying-Lemurs. 

2 ,, INSECTIVORA VERA. Shrews, Moles, 

Hedgehogs, etc. 


iv. Order CARNIVORA. Carnivores. 

1 Suborder CARNIVORA VERA. Cats, Dogs, Bears, 

Weasels, etc. 

2 PINNIPEDIA. Seals and Walruses. 

3 *CREODONTA. Hyanodon, etc. 

v. Order RODENTIA. Rodents. 

1 Suborder SCIUROMORPHA. Squirrels, Marmots, and 


2 MYOMORPHA. Dormice, Mice, and Jer- 


3 HYSTRICOMORPHA. Porcupines, Agutis, 

Cavies, etc. 

4 ,, LAGOMORPHA. Picas and Hares. 

vi. Order UNGULATA. Hoofed Mammals. 

1 Suborder ARTIODACTYLA. Antelopes, Camels, Pigs, 


2 ,, PERISSODACTYLA. Horses, Tapirs, and 


3 ,, *LITOPTERNA. Macrauchenia, etc. 

4 *ASTRAPOTHERIA. Astrapotherium, etc. 

5 *PYROTHERIA. Pyrotherium. 

6 *TOXODONTIA. Toxodon, etc. 

7 ,, HYRACOIDEA. Hyraces. 

8 ,, PROBOSCIDEA. Elephants and Mastodons. 

9 ,, *AMBLYPODA. Coryphodon, etc. 

vii. Order SIRENIA. Manatis and Dugongs. 
viii. CETACEA. Whales and Porpoises, 
ix. EDENTATA. Edentates. 
x. ,, EFFODIENTIA. Aard-varks and Pangolins. 
xi. ,, MARSUPIALIA. Pouched Mammals. 

1 Suborder DIPROTODONTIA. Kangaroos, Phalangers, 

and Wombats. 

2 ,, POLYPROTODONTIA. Dasyures, Bandicoots, 


xii. Order MONOTREMATA. Egg-laying Mammals, 
xiii. *MULTITUBERCULATA. Pldgiciulax, Tritylodon, etc. 


These orders are further subdivided into families and genera. 
In regard to the number of the latter of these, there is still a con- 
siderable difference of opinion among naturalists ; but in the 
present work those adopted in Flower and Lydekker's Study of 
Mammals 1 will be in the main adhered to, with such corrections 
and additions as are necessary owing to recent emendations in 
nomenclature or to the discovery of new forms. 

Omitting from consideration the purely aquatic and volant 

members of the class, the most effectual barriers to 

Dispersal of the dispersal of mammals are formed by channels 


their being crossed by swimming. And it is this inability to 
traverse any extent of water that renders what are known as 
oceanic islands practically devoid of all mammalian life, with 
the exception of a few bats and small rodents ; the latter animals 
having apparently some means of dispersal not common to other 
members of this class. Oceanic islands, it may be explained, are 
such as rise from great depths in the ocean, and are composed, 
almost invariably, either of volcanic rocks or of coral. They show, 
for the most part, no decisive evidence of having been connected 
with any continental land, and thus have never been enabled to 
receive a mammalian fauna 2 . In marked distinction to these are 
the so-called continental islands, such as Madagascar and Great 
Britain, which, both from the evidence of their mammalian fauna 
and their geological conformation, have indubitably been in direct 
communication with the adjacent continent at no very distant 
epoch. As a rule, the channels between such islands and the 
mainland are comparatively shallow, so that a moderate degree of 
upheaval would place the two in direct connection. 

The relative depth of the channel between two islands, or 
between an island and a continent is indeed of much more im- 
portance in regard to the dispersal of mammals than is its width. 
This is best exemplified by the well-known case of "Wallace's 
line " in the Malayan archipelago ; that name being applied to 

1 London, 1891. 

2 There is a possibility that some oceanic islands may have been connected 
with continents, and that their original mammalian fauna has been destroyed 
by submergence. 


the narrow strait separating the islands of Bali and Lombok, and 
its northward continuation, the Makassar Strait, dividing Borneo 
from Celebes. Although the two former islands are extremely 
close together, while Celebes is much less widely separated from 
Borneo than is the latter from Sumatra, yet the faunas of Lombok 
and Celebes are markedly distinct from those of the islands lying 
to the north and west of Wallace's line. Soundings show that the 
Makassar Strait, and likewise the Bali-Lombok Strait are of greater 
depth than the channels separating the other islands of the archi- 
pelago; and consequently that Wallace's line indicates a very 
old barrier which has long been impassable to the majority of 

That continental islands have received the great bulk of their 
mammalian fauna by means of a more or less complete land- 
connection with the mainland, is perfectly evident. Nevertheless, 
there are cases where certain mammals have crossed the inter- 
vening channel, either by swimming, or by having been carried 
across on natural rafts of some kind ; an instance of this nature 
being exemplified by the occurrence of an African type of pig in 
Madagascar. It thus becomes a question of considerable interest 
to ascertain what stretches of sea large mammals are capable of 
crossing. It is stated that the jaguar has been known to swim 
across the Rio de la Plata, which at its mouth is something like 
eighty miles across ; and a polar bear has been observed swim- 
ming at a distance of twenty miles from land in Bering Strait. 
The tiger frequently crosses the narrower channels in the Sandar- 
bans of Lower Bengal; and both deer, pigs, and elephants are 
good swimmers. The latter animals have, indeed, been known to 
swim for six hours at a stretch, and, with a rest, for upwards of 
nine ; but their rate of progress is extremely slow, and probably 
exceeds but little, if at all, a mile an hour. The Palk Strait, 
which is considerably less than forty miles in width at its narrowest 
part, has formed an effectual barrier to the passage of the tiger 
from India into Ceylon; and it may accordingly be assumed 
that about twenty miles is the utmost limit which mammals are 
likely to cross by swimming, even when favoured by currents. 

Such passages as these must, however, be of very rare occur- 
rence, for a terrestrial mammal is not likely to take it into its head 


to swim straight out to sea in an unknown direction. Moreover, 
supposing a mammal new to a particular island to have arrived 
there by swimming, unless it happen to be a pregnant female, or 
unless another individual of the same species but of the opposite 
sex should arrive soon after (a most unlikely event), it would in 
due course die without being able to propagate its kind. And 
even if it should happen to be a pregnant female, there would be 
no certainty that its offspring, if but one in number, should be of 
the opposite sex to its parent. Accordingly, it would seem that 
the population of islands by mammals that have arrived by swim- 
ming must be a very rare event indeed. Rafts may be of more 
importance. Mr Aplin, in the Proc. Zool. Soc. for 1894, mentions 
that jaguars and pumas are frequently transported by them from 
one side of the Rio de la Plata to the other; and in the rainy 
season many are to be seen off the northern coast of Borneo, 
some of which may be as much as thirty yards in length. Still 
such rafts are not likely to cross straits, except when there is a 
current setting from one bank to the other. 

Before leaving this part of the subject, it must be mentioned 
that the degree of difference between the fauna of an island and 
that of the adjacent continent, or between the faunas of two 
islands, affords a most important clue as to the relative date of the 
land-connection between them. Madagascar, for instance, has a 
mammalian fauna which although clearly derived from Africa, is 
yet so different from that now inhabiting the mainland, that the 
land-connection between the two must have been broken up at an 
epoch comparatively remote. Ceylon and India present a condition 
in respect of their faunas intermediate between the last and that of 
Great Britain as compared to the Continent. In the latter instance, 
with the exception of a single Irish weasel, all the mammals are 
identical with species now inhabiting the Continent, thus proving 
that the connection has been a comparatively recent one ; although 
the peculiar weasel indicates that the separation between Ireland 
and Britain has been of sufficient duration to admit of the develop- 
ment of a distinct specific type in the former country. 

Although large rivers like the Amazon and La Plata un- 
doubtedly form serious barriers to the migration of mammals, yet 
these are not so insuperable as might at first sight be supposed, 


owing to the fact that in districts where vegetation is luxuriant, 
huge natural rafts are formed by the trunks of trees intermingled 
with vegetable matter, upon which numbers of animals may be 
borne down stream, and thus transferred from one bank to the 
other. Nevertheless, in treeless districts, or near their mouths, 
large rivers afford absolutely impassable barriers to the movements 
of mammals. In South America, for instance, even such an 
aquatic creature as the carpincho, or capibara (ffydrochcerus) has 
been unable to cross the Plata river from Uruguay into the 
Argentine, while, conversely the viscacha (Lagostomus) of Argentina 
is prevented by the same river from reaching Uruguay. 

Deserts are, perhaps, even more impracticable than rivers ; the 
Sahara which was long supposed to have been the site of an 
ancient sea, although it has really been a desert since very remote 
ages having apparently formed a barrier preventing the fusion of 
the mammals of North Africa with those to the south of that tract 
since at least the Pliocene epoch. It must not, however, be sup- 
posed that what are desert-tracts at the present day have always 
been such, the existence of a fossil chimpanzee in North-Western 
India during the Pliocene period indicating that the open sandy 
plains of the Punjab were at that time covered with dense tropical 
forests, and probably that the same was the case with parts of 
Syria and Arabia. 

Before taking leave of seas and deserts, it should be mentioned 
that in the polar regions ice may act in lieu of a land-connection 
to enable mammals to pass from one country to another. On this 
subject Dr Heilprin writes that " the reindeer is stated to cross 
the Bering Strait by way of the Aleutian Islands and the Frozen 
Sea, and in a somewhat similar manner the musk-ox finds its way 
to Melville Island; it is, however, somewhat singular that the last- 
named animal, despite its long ice-journeys, never manages to 
reach either the continent of Asia or Greenland." 

High mountain ranges form an effectual barrier to the migration 
of mammals, not only on account of the physical difficulties of 
crossing them, but likewise by the lowness of the temperature at 
great altitudes, coupled with the absence of proper food, being 
fatal to the existence of many. As already stated, however, 
mountain-ranges are much more far-reaching in their effects on such 


migrations when, as in the Old World, their trend is from west to 
east, than when, as in America, they run from north to south, and 
thus do not interfere with the free movements of plain-dwelling 
animals on either side. In many instances the mammals inhabit- 
ing each of the isolated mountain-chains, as those of Europe, are 
to a great extent specifically identical one with another, not having 
had time to become modified into distinct species since they 
reached their present haunts at the close of the glacial period. 
Even among these, however, there are indications of the com- 
mencement of specific differences, the chamois of the Caucasus 
forming a variety differing somewhat from the typical Alpine race. 
Where the isolation has been longer, as in the case of the fauna 
of the highlands of Tibet, the difference is much more strongly 
marked ; the mammalian fauna in this instance being as peculiar 
and distinct as that of many ancient continental islands. 

Probably ever since man has existed in any numbers on the 
influence of glbe he has been exerting a more or less strongly- 
ManonDistri- marked influence on the distribution of animals, 
either by destroying them, or by conveying them to 
countries or districts which are not their natural home. By the 
involuntary aid of man the common rat and mouse, which belong 
to a genus unknown in the New World, have been conveyed to 
every country in the globe ; while the rabbit has been carried to 
the Antipodes, where it has flourished and increased in an 
unprecedented manner. Cattle and horses have been introduced 
into South America, Australia, and other countries where they 
were naturally unknown, and by their rapid increase have shown 
that the absence of particular animals from particular districts is 
not necessarily due to their being unsuited to live there, but rather 
to the fact that they have been unable to find their way thither. 
The fallow-deer, again, has been imported from its Mediterranean 
home into England and other countries of northern Europe; while 
goats and pigs have been carried to a number of oceanic islands, 
where they have done irreparable harm in exterminating the native 
fauna and flora. 

In all these instances the fact of the introduction has always 
been more or less clearly known, and therefore no difficulty arises 
as to what are native and what are introduced forms. Very 


different, however, is the case with the islands of the Malay Archi- 
pelago, where the natives, who have a wonderful facility for 
taming animals, have carried a species peculiar to one district or 
island to localities where it is quite unknown as a native ; and in 
consequence of this transportation and acclimatisation it is pro- 
bable that several mammals have been given a habitat to which 
they have not the most remote right. To the Malays is due the 
introduction of the small civet known as the rasse into Mada- 
gascar. Whether the dingo, or native dog of Australia, was intro- 
duced at an exceedingly remote era by the original colonisers of 
that island, or whether it is truly indigenous, is a question that 
will probably never be decisively answered. It is likewise quite 
impossible to say what part man may have played in the extermi- 
nation of the large mammals that inhabited Europe about the 
close of the glacial period, but it seems quite probable that he 
may have had a considerable share in their destruction. Be this 
as it may, the domestication of certain mammals has undoubtedly 
had the effect of destroying the wild race, as is remarkably ex- 
emplified by the two existing species of camel, of neither of which 
do we know the original habitat. The original European wild ox 
unless, indeed, the half-wild cattle of the British parks be its 
direct descendants has likewise disappeared at some unknown 
epoch owing to the hand of man. Although other mammals, such 
as the quagga (Equus quagga}, Burchell's rhinoceros (Rhinoceros 
simus), and the blaubok (Hippotragus leucophaus) have been 
almost or completely exterminated by human agency in South 
Africa, while the American bison has been practically swept away 
from its native prairies, yet in all these instances there is a more 
or less full record of the original range of the creatures. In 
other cases also mammals have been utterly exterminated by 
human agency from countries of which they were originally in- 
habitants, as is exemplified by the disappearance from the British 
Islands of the bear, the wolf, the beaver, and the wild boar within 
the historic period, although they still survive in other parts of 
their habitat. In these particular instances there is fortunately 
full evidence as to the former existence of these animals in 
Britain ; but it is highly probable that in more remote countries 
mammals have been exterminated without any record being left 
L. 2 


of their existence, so that the full extent of their range, if they be 
surviving forms, can now never be ascertained. 

To quote the words of Dr Wallace, it is evident that in the 
present day we live in an impoverished epoch, so 


of the larger iar as the larger mammals are concerned, as com- 

pared with the Plistocene era ; this being true not 
only as regards the northern half of the Old World, 
but likewise North and South America, as well as Australia. 
From the northern half of the Old World have disappeared the 
mammoth, the elasmothere, the woolly and other species of 
rhinoceros, the sabre-toothed tigers, etc.; North America has lost 
the megalonyx and the Ohio mastodon ; from South America the 
glyptodonts, mylodons, the megalothere, and the macrauchenia have 
been swept away ; while Australia no longer possesses the dipro- 
todon and various gigantic species of kangaroos and wombats. 
In the northern hemisphere this impoverishment of the fauna has 
been very generally attributed to the effects of the glacial period, 
but although this may have been a partial cause, it can hardly be 
the only one. The mammoth, for instance, certainly lived during 
a considerable portion of the glacial epoch, and if it survived thus 
far, why should it have disappeared at the close? Moreover, all 
the European mastodons and the southern elephant (Elephas 
meridionalis] died out before the incoming of glacial conditions ; 
and the same is true of all the extinct elephants and mastodons of 
southern Asia. Further, a large number of English geologists 
believe the brick-earths of the Thames valley, which contain 
remains of rhinoceroses and elephants in abundance, to be of 
post-glacial age. As regards the southern hemisphere, it can 
hardly be contended that glacial conditions prevailed there at the 
same time as in the northern half of the world. 

It is thus evident that although a very great number of large 
mammals were exterminated (perhaps partly by the aid of human 
agency) at the close of the Plistocene period, when the group 
had attained its maximum development as regards the bodily size 
of its members, yet other large forms had been steadily dying 
out in previous epochs. And it would seem that there must 
be some general deep-seated cause affecting the life of a species 
with which we are at present' unacquainted. Indeed, as there 


is a term to the life of an individual, what is more natural than 
that there should also be one to the existence of a species ? It still 
remains, indeed, to account for the fact that the larger Plistocene 
mammals had no successors in the greater part of the world, but 
perhaps this is in some way connected with the advent of man. 

Before coming to the consideration of the zoological divisions 
into which, from the present geographical distribu- 
tion of mammals, the world may be mapped out, it tionai Areas of 
is necessary to devote a brief space to the considera- Genera^ and 
tion of two other points ; the first relating to the 
relative size of the distributional area of genera and species, and 
the second to the permanency of ocean-basins and continents. 

As regards the first point, it appears to be true in the case of 
mammals (although not of all other groups) that every species has 
a continuous distributional area, except where this has been broken 
up by human destructiveness. It is not meant by this that every 
part of such area is inhabited by the particular species, as 
"station" renders this impracticable; but merely that the whole 
area is ranged over by the species in such spots as are suited to 
its particular mode of life. Great variation obtains, however, in 
regard to the size of such distributional area; and it will be 
obvious that the size of the area varies directly as the adaptability 
of the species to different climatic and other physical conditions. 
Perhaps the most important condition of all is the possibility of 
obtaining suitable food ; and in this respect carnivorous mammals 
are in a far better position than any other members of their class, 
since the kind of animal on which they prey is immaterial. This 
will readily account for the extensive geographical ranges enjoyed 
by the puma and the tiger, which, as stated on page 5, embrace 
almost every degree of latitude. Animals with such a wide dis- 
tribution are of but little use to the student of geographical 
distribution. Moreover, it will generally be found that species 
with a wide range belong to large genera having a still more 
extensive distributional area; this being markedly the case with 
the puma and the tiger ; the genus Felts being one of the largest 
in the class, and ranging over the whole world with the exception 
of Australasia. Such cosmopolitan genera are likewise almost 
valueless to the distributionist. 

2 2 


On the other hand, species with a small distributional area 
usually belong to small genera, of which they may be the only 
representatives. Instances of this nature are afforded by the 
panda (sElurus] and binturong (Arctictis) of the eastern Himalaya, 
and the parti-coloured bear (^Eluropus] and chiru antelope (Pan- 
tholops] of the Tibetan plateau. Although such single representa- 
tives of genera are highly important to the study of distributional 
zoology, of vastly greater importance are small genera having two 
or more species living in widely separated areas. Examples of 
such are to be found among the porcupines of the genus Atherura, 
of which one species is Malayan and the other two West and 
Central African ; in the mice of the genus Golunda, with one 
African and one Indian representative ; and likewise by the tapirs 
(Tapirus\ of which there is one Malayan, and several tropical 
American species. These examples of " discontinuous distribu- 
tion " among genera are of the very highest import to the science ; 
since they clearly indicate that some of the lands lying between its 
present disconnected distributional areas must have formerly been 
the habitat of the genus, and thus enable important conclusions to 
be drawn as to the former land-connections between such areas. 
Both in the case of the tapirs and of the brush-tailed porcupines, 
remains of extinct species have been discovered in the intermediate 

Equally important are families, either large or small, which 
contain two or more closely allied small genera respectively con- 
fined to distant areas. As an instance of a large family containing 
such allied genera, may be cited the Viverridce, among which the 
true linsangs (Linsangd) are represented by several species from 
the Eastern Himalaya and the Malayan countries, while the 
closely-allied Poianais confined to West Africa. The chevrotains 
(Tragulidee), on the other hand, form a small family with a dis- 
continuous distribution ; one genus (Tragulus) being now Oriental, 
while the other (Dorcatheriuni) is West African. Here it is quite 
evident, of course, that the distributional area of the family must 
once have been continuous ; and, as a matter of fact, remains of 
both genera occur in the Pliocene of India, those of the latter being 
also found in the European Miocene. 

In other families with a discontinuous distribution, as in the 


rodent family Octodontida, which is now mainly confined to Africa 
south of the Sahara, and Central and South America, the genera 
may be less closely allied, although sufficiently so to indicate 
a continuous distributional area, or rather a common centre of 
dispersal, at no very remote epoch. 

Allied families, with a small number of genera, severally con- 
fined to distant localities are likewise of the highest value in 
building up the former history of the globe. As examples of this 
nature may be cited the tree-shrews (Tupaiida) of the Oriental 
countries and the jumping shrews (Macroscelidida) of Africa on the 
one hand, and the Solenodontidce of the West India islands, and 
the Centetidce of Madagascar on the other. Such families must 
clearly have had a common centre of origin and dispersal ; the 
available evidence suggesting that in the case of the two former 
such centre was Europe. 

Although of far less common occurrence than among families 
or genera, discontinuous distribution in an order is perhaps of even 
more importance than either of the other cases, as it implies a 
greater interval of time since the original dispersal took place, and, 
therefore, carries back such conclusions as can be drawn in regard 
to former land-connections to a still earlier epoch. Among 
mammals the only instance of this nature is to be found in the 
marsupials 1 , of which two families are American (and mainly 
South and Central American), while all the others are confined to 
Australasia and some of the adjacent Malayan islands. In this 
case also there is abundant evidence of the wide distribution of 
the whole group in former epochs of the earth's history. 

This last instance leads on to the consideration of what may 
be termed "centres of evolution." In a previous 

paragraph it has been stated that, according to the " f 

available evidence, a very large proportion, if not 
the whole, of the terrestrial mammalian life of the globe has 
originated in the northern hemisphere, from which it has spread 
southwards in a continuous successive series of waves. When, 
however, certain groups of mammals had once reached the more 

1 In this work the Effodientia are separated from the Edentata; but when 
these are united, there is a second instance. 


remote parts of the southern hemisphere, where they were free 
from the competition of the higher forms, and met with favourable 
conditions, they seemed to take a new lease of life, and attained a 
fulness and variety of development which they had never reached 
before. As a rule, more or less complete isolation has been a 
dominant feature of this development; of which the best and most 
striking instance is that of the marsupials in Australasia. That 
area may accordingly be called the marsupial evolutionary centre. 
Scarcely less striking is the instance of the edentates (of which the 
original derivation is unknown) in South America, where, in 
company with certain peculiar extinct groups of ungulates, they 
attained an extraordinary development, both as regards the 
number of specific, generic, and family types, and likewise in 
respect of the bodily size of some of its members. This second 
area may be termed the evolutionary centre of the edentates. A 
third great centre is constituted by Europe, Asia, and North 
America, which appear to have been the main developmental 
centre of the higher mammals, and may accordingly be named the 
placental evolutionary centre. Two other minor centres are 
respectively indicated by Madagascar and Africa south of the 
Sahara : the former as being the headquarters of the lemurs, may 
appropriately be spoken of as the lemuroid centre, while the 
great development of the antelopes in Africa suggests the name of 
the antelopine evolutionary centre for that continent. 

The circumstance that throughout the greater part of North 

America and Europe a very large proportion of the 

of Conine nts y continents are built up of sedimentary strata of 

and Ocean- marine origin, naturally led geologists in the early . . . 1 1*1 

days of their science to the conclusion that every 
part of the land had at one time been deep ocean, and every 
stretch of ocean dry land. More careful study led, however, 
to the belief that this idea was not founded on fact, and that 
although it was perfectly true that what are now continents had 
been many times under the sea, yet that such areas had never 
formed abyssal ocean-depths ; and, conversely, that such ocean- 
depths had never been dry land. In addition to many other lines 
of evidence, this view of the permanency of continents and ocean- 
basins is strongly supported by the circumstance that nearly all 


oceanic islands are either of volcanic or coral origin, and do not 
contain sedimentary rocks; and also that deposits analogous to 
those laid down in the deepest ocean beds are generally wanting 
from among the sedimentary series of rocks of which the continents 
and islands are composed. A further argument was afforded by 
the discovery that the greater portion of peninsular India and 
South Africa has been dry land since the Palaeozoic epoch. 

As is so commonly the case in similar instances, the promulga- 
tors of the doctrine of the permanency of continents and ocean- 
basins pressed their hypothesis too far; and it is now evident 
that although the doctrine is true as a whole, and more especially 
as regards the later stages of the earth's history, yet it requires 
very considerable modification from the original form in which it 
was advanced. In the first place, it has been shown that crystal- 
line granitic and gneissic rocks occur in the Seychelles, which were 
formerly regarded as true oceanic islands ; and, secondly, deep-sea 
deposits have been discovered in the West Indies and the Solomon 
Islands. Moreover, various lines of evidence indicate that during 
the Jurassic and Cretaceous epochs there was a continuous land- 
connection between Africa (by way of Madagascar and the 
Seychelles) and India; while at some time in the Secondary era, 
in the opinion of Drs Neumayr and Blanford, South America and 
South Africa were in communication across the South Atlantic. 
The latter connection appears, indeed, to have been a survival 
from an older Palaeozoic girdle of land which, from the evidence of 
fossil floras, seems to have existed in low latitudes round nearly 
three-quarters the circumference of the globe, and which was cut 
oft" from the land to the north. There is, moreover, the possibility 
of a Tertiary connection of Australia with Patagonia by way of 
Polynesia, to which allusion is made in the third chapter. Then, 
again, the recent investigations of Dr J. W. Gregory 1 on the fossil 
corals of the West Indies have afforded strong support to the view 
that the Atlantic is of comparatively recent origin. After referring 
to the remarkable resemblance between the existing fauna of the 
West Indian seas and that of the Miocene deposits of the Mediter- 
ranean basin, Dr Gregory 2 observes that the sea-urchins, or 

1 Quart. Joitrn. Geol. Soc. vol. LI. pp. 255 312 (1895). 

2 Ibid., pp. 306, 307. 


echinoderms, yield still more conclusive evidence. "As I have 
previously pointed out," he writes, "the intimate affinity between 
those of the West Indies and the Mediterranean can only be 
explained by the assumption of the existence of a shallow-water 
connection across the Central Atlantic in at latest Miocene 
times. That the fauna did not follow along the shores of the North 
Atlantic basin, is shown by its absence from the northern Miocenes 
of Europe and North America. The evidence now adduced from 
the fossil corals of Barbados lends support to this view, as showing 
that the West Indian fauna is only a fragment of that of the 
Mediterranean Miocene, and has received nothing from the 
Pacific. This is in full agreement with Prof. Suess's theory that 
the Atlantic is of comparatively recent geological age, and arose 
by the gradual enlargement of two bays which ran north and south 
from a sea that once extended across the Mid-Atlantic from 
Europe to America, including both the Mediterranean and the 
Caribbean Sea." 

The question of the southward extension of America, Africa, 
and Australia to join the Antarctic continent during Tertiary times 
is alluded to in the sequel. 

Summing up the evidence in regard to the permanency of 
oceans and continents, Dr Blanford 1 several years ago observed 
" that whilst the general permanence of ocean-basins and conti- 
nental areas cannot be said to be established on anything like 
firm proof, the general evidence in favour of this view is very 
strong. But there is no evidence whatever in favour of the 
extreme view accepted by some physicists and geologists that 
every ocean-bed now more than 1000 fathoms deep has always 
been ocean, and that no part of the continental area has ever been 
beneath the deep sea. Not only is there clear proof that some 
land-areas lying within continental limits have at a comparatively 
recent date been submerged over 1000 fathoms, whilst sea-bottoms 
now over 1000 fathoms deep must have been land in part of the 
Tertiary era, but there are a mass of facts both geological and 
biological in favour of land-connection having formerly existed in 
certain cases across what are now broad and deep oceans." 

1 Appendix, No. 8, p. 107. 


Although much previous work had been done on the subject, 
the first real attempt to divide the land-areas of the 

, , . i-i Zoological 

globe into zoological provinces, or regions, was made Realms and 
by Dr P. L. Sclater 1 in 1858. According to this Regions ' 
scheme, which was mainly based on the study of Passerine birds, 
the world was parcelled out into the following six zoological 
regions, viz.: 

1. Palczarctic ; Europe, Northern Africa, Northern and Cen- 
tral Asia. 

2. Ethiopian ; Africa south of the Atlas, and Madagascar. 

3. Indian, renamed Oriental by Dr Wallace; India, South- 
eastern Asia, and part of the Malay Archipelago. 

4. Australian ; Australia, with New Guinea and the adjacent 
islands, New Zealand, and Polynesia. 

5. Nearctic ; America as far south as Mexico. 

6. Neotropical ; Central and South America, with the West 

This scheme, which has been adopted and developed in the 
brilliant writings of Dr Wallace, has the important merit that it 
coincides to a great extent with the leading geographical divisions 
of the globe. It has, however, the serious drawback that it gives 
no greater rank to Australasia and South America than to the other 
divisions ; whilst the remarkable difference between the fauna of 
Africa and Madagascar is overlooked. Further, the northern parts 
of America are widely separated from those of Europe and Asia 
to which they are faunistically extremely close. 

It should be added that in Dr Sclater's scheme the first four 
regions, or those belonging to the Old World, were brigaded 
together under the title of PAL^OG^EA, while the two last, or New 
World regions, were bracketed as NEOG^EA. 

The next important classification was one propounded in 1868 
by Professor Huxley 2 , who, basing his conclusions on the distri- 
bution of the game-birds, divided the world into a northern and a 
southern division, taking the name of ARCTOG^EA for the former, 
and NOTOG^EA for the latter ; Notogaea being further sub-divided 
into a Novo-Zelanian (New Zealand), Australian, and Austro- 

1 Appendix, No. 26. ' 2 Ibid., No. 18. 


Columbian region, the latter being equivalent to the Neotropical 
of Sclater. 

Six years later, Dr Sclater 1 , who had by this time turned his 
attention to the distribution of mammals, proposed to group the 
regions he had previously named under three larger divisions, 
making a fourth division for New Zealand and Polynesia. This 
scheme is as follows, viz. : 

I. ARCTOG^EA. Palsearctic, Nearctic, Oriental, and Ethiopian 

II. DENDROG^A. Neotropical region. 

III. ANTARCTOG^EA. Australian region (exclusive of New 
Zealand and Polynesia). 

IV. ORNITHOG^EA. New Zealand and Polynesian region. 

So far as mammals are concerned, this scheme was a great 
advance on the first one, although the distinctness of Madagascar 
was not recognised, while the Palasarctic and Nearctic regions 
were still maintained. Most of the names for the major divisions 
are, however, open to objection. 

In 1878 Dr Heilprin 2 , who does not employ these larger 
groups, proposed, after a suggestion of Professor A. Newton, to 
unite Dr Sclater's Palsearctic and Nearctic regions under the 
common title of the Holarctic region ; separating, however, from 
the former a "transitional" Mediterranean region, and from the 
latter a similar Sonoran region. 

A further step was made in 1890 by Dr Blanford 8 , who pro- 
posed the following scheme, viz. : 

I. Australian region. 

II. South American region. 

III. Arctogaan region ; this being divided into Malagasy, 
Ethiopian, Oriental, Aquilonian ( Palaearctic and northern part 
of Nearctic), and Medio-Columbian ( = Sonoran). 

Several other minor modifications have been suggested from 
mammalian evidence, Dr Allen 4 in 1892 reviving the view that the 
Oriental and Ethiopian regions should be united, under the name 
of the Indo-African ; but the next most important memoir is that 

1 Appendix, No. 27. 2 Ibid., No. 17. 

3 Ibid., No. 8, p. 76. 4 Ibid., No. 2. 


of Dr Hart Merriam 1 in 1892, whose views are fully discussed in 
the sequel. 

It will accordingly suffice for our present purpose to say that in 
1893 an anonymous writer 2 proposed to take the terms NOTOG^EA, 
NEOG^A, and ARCTOG^A to indicate the three major divisions of 
Dr Blanford's classification ; the same terms being used by Mr 
W. L. Sclater 3 in a nearly similar sense. 

The following scheme is the one adopted in the present 
volume, viz. : 

I. NOTOG^EIC REALM. i. Australian Region. 

2. Polynesian Region. 

3. Hawaiian Region. 

4. Austro-Malayan Region. 

II. NEOG^EIC REALM. Neotropical Region. 

III. ARCTOG^EIC REALM. i. Malagasy Region. 

2. Ethiopian Region. 

3. Oriental Region. 

4. Holarctic Region. 

5. Sonoran Region. 

It will be noticed that the three realms correspond to the three 
great evolutionary centres of mammals alluded to in an earlier 

It may be added that, in a work expressly devoted to the 
geographical distribution of mammals, it will be unnecessary to 
allude to the schemes proposed on the evidence of other groups of 
animals, and we may accordingly proceed forthwith to the con- 
sideration of the distinctive features of the various realms and 
regions here adopted. 

1 Appendix, No. 19. 2 Ibid., No. 4, p. 289. 3 Ibid., No. 28. 



Definition and Characters of the Realm Australian Region Monotremes 
Marsupials Rodents Carnivores Ungulates Bats List of Australian 
and Papuan Genera Polynesian Region Hawaiian Region Austro- 
Malayan Region Palaeontological History of Marsupials How Australia 
received its Fauna. 

THE term Notogaea was first proposed, as stated in the preced- 
ing chapter, by Professor Huxley 1 , to include not only the 
Australian region of Dr Sclater, but likewise the Neotropical region 
(Austro-Columbia) ; but an anonymous writer 2 appears to have 
been the first to restrict it to the former of these areas 3 . This 
view, as being, on the whole, the most convenient, is adopted here; 
and the Notogasic realm may accordingly be taken as the first of 
the three primary zoological divisions of the globe, and as equiva- 
lent to the Australian region of Drs Sclater and Wallace. Accord- 
ing to the latter writer 4 , " its central and most important masses 
consist of Australia and New Guinea, in which the main features 
of the region are fully developed. To the north-west it extends to 
Celebes, in which a large proportion of the Australian characters 
have disappeared, while Oriental types are mingled with them to 
such an extent that it is rather difficult to determine where to 
locate it. To the south-east it includes New Zealand, which is in 
some respects so peculiar that it has even been proposed to con- 
stitute it a distinct region. On the east it embraces the whole of 
Oceania [Polynesia] to the Marquesas and Sandwich Islands, 

1 Appendix, No. 18. ' 2 Ibid., No. 4. 

3 The term Antarctogsea has been proposed by Dr Sclater (Appendix, 
No. 27, p. 214), for this area, but it is not a happy one. 

4 Appendix, No. 32, vol. i., p. 387. 


where a very scanty and often peculiar fauna must be affiliated to 
the general Australian type." To the north-east the line of de- 
marcation of the realm from the Oriental region of Arctogsea has 
been finally fixed at the deep channel separating the islands of 
Celebes and Lombok on the one side from those of Borneo and 
Bali (at the extremity of Java) on the other ; this division being 
now well known under the name of " Wallace's line." 

All writers are, however, by no means agreed as to the right of 
the whole of the area thus indicated to form a single zoological 
division. Before the publication of Dr Wallace's great work, 
Professor Huxley had proposed to separate New Zealand as a 
region of equal rank with his Australasian region. At a later date 
Professor Heilprin 1 suggested that the Australian realm should 
include only Australia, Tasmania, New Guinea, with the smaller 
Papuan islands, and New Zealand ; Polynesia, including all the 
islands lying to the east of the Coral Sea, being raised to the rank 
of a distinct realm (the Polynesian), while the Austro-Malayan 
islands were regarded as forming a transitional tract between the 
Australian realm and what is here termed the Oriental region. In 
this connection it may be well to notice that the Austro-Malayan 
sub-region of Dr Wallace is by no means coterminous with the 
Austro-Malayan transition-tract of Heilprin, the former including, 
and the latter excluding New Guinea. 

So far as mammals alone are concerned, Notogaea is widely 
separated from the whole of the rest of the world by being the sole 
habitat (both now and in the past) of the typical diprotodont 
marsupials and the monotremes; although it must not be sup- 
posed that either of these groups is distributed over the entire 
area. As a matter of fact, apart from introduced rodents, 
Polynesia is devoid of mammalian life with the exception of bats 
and a rat 2 , while New Zealand has but two representatives of the 
former group, and a rat which may or may not be indigenous. 
But wherever we meet with a fully developed mammalian fauna, 
as in the transitional Austro-Malayan islands, there a certain 
number of marsupials are met with, although the monotremes 

1 Appendix, No. 17. 

2 Mus exulans, see Proc. Zool. Soc. 1895, p. 338. 


are restricted to Australia, with Tasmania ; and New Guinea, with 
the adjacent islands, such as the Aru group. 

The Notogaeic realm, as denned above, may be conveniently 
divided into four distinct regions, as follows. Firstly the Australian 
region, comprising Australia, Tasmania, New Guinea and the adja- 
cent Papuan islands; characterised by marsupials and monotremes 
forming by far the predominant element in the mammalian fauna. 
Secondly the Austro-Malayan region, embracing Lombok, Celebes 
and the other islands lying between them and the Australian region; 
this area being characterised by the absence of monotremes and by 
the marsupials (all of which belong to the diprotodont division of 
the order) forming only a small minority of the mammalian fauna. 
Thirdly, there is the Hawaiian region, including only the Sandwich 
Islands; and, lastly, the Polynesian region, which maybe taken to 
include all the islands, save those last named, lying to the eastward 
of the Coral Sea, together with New Zealand, and is characterised 
by the general absence of terrestrial mammals. There is some 
difficulty in deciding whether the islands of the Solomon group 
should be included in this region, or classed with the Papuan 
division of the Australian region, seeing that, in addition to a 
considerable number of bats, they have four species of mice, and 
one diprotodont marsupial (Phalanger) 1 . When, however, the 
poverty of this fauna as compared with that of Papua is taken 
into consideration, and it is also borne in mind that Mr C. Hedley 2 
has come to the conclusion that the Solomon Islands, together 
with the New Hebrides, New Caledonia, Fiji, and Norfolk Island, 
are closely connected by means of their flora with New Zealand, 
and have but little in common with Australia and New Guinea, it 
seems preferable to include the former group in the Polynesian 
region. On the same grounds, New Zealand is regarded, in 
accordance with the views of Heilprin, as also forming a portion 
of the same zoological region, and not as the representative of a 
separate region by itself. The fauna of the Solomon Islands has 
doubtless been derived directly from that of the Duke of York 
group, which clearly belongs to the same region as New Guinea, 
and shows a much more strongly marked Papuan facies, having 
three species of mice, and four marsupials. 

1 See Thomas, Appendix, No. 30. 2 Appendix, No. 16. 


Although the northern half of Australia lies within the tropics, 
yet few portions of this great island present that 
luxuriance of vegetation which we are accustomed 
to associate with tropical scenery ; and large tracts 
of the interior, owing doubtless to the absence of elevated moun- 
tain ranges in the central districts, form arid sandy deserts more or 
less unsuited to the maintenance of animal life. The coast regions 
and the borders of the larger rivers are accordingly those where 
vegetation flourishes best ; the finest tracts of pasture-country, well 
supplied with water, lying to the east and south-east, and Victoria 
possessing a mountain range whose summits are perpetually 
clothed with snow. Mountains also occur in the dry and hot 
western districts. Although Tasmania enjoys moister conditions, 
Australia as a whole is characterised by the lack of water and the 
general dryness of its climate ; and it is probable that to this 
aridity the number of jumping animals, such as kangaroos, rat- 
kangaroos, and jerboa-rats, now characteristic of this part of the 
region is due, since such creatures are admirably adapted for 
traversing long distances in search of food and water. On the 
other hand, New Guinea, together with the Papuan islands, has a 
moist tropical climate, essentially different from that of Australia, 
but similar to the conditions obtaining in a large portion of 
the Austro- Malayan islands. Hence it is not to be wondered 
at that the mammals of New Guinea differ very markedly 
from those of Australia ; this being especially noticeable in the 
paucity of typical jumping kangaroos, and the proportionately 
large number of arboreal members of this group. Nevertheless the 
mammalian fauna of Queensland and North Australia exhibits a 
marked approximation to that of New Guinea, one species of 
kangaroo, as well as a cuscus, a striped phalanger (DaJylopsila), a 
flying phalanger (Petaurus), a pouched-mouse (Phascologale), and 
an echidna, being common to the two areas, and it is in these 
countries alone that tree-kangaroos are met with. From these 
resemblances in their faunas and especially from the restriction 
of the monotremes to these two areas, there can be no question 
as to the propriety of including Australia and New Guinea in the 
same zoological region, and thus separating the latter country from 
the Austro-Malayan region. 



The egg-laying mammals, or monotremes, constitute not only 
a distinct order (Monotremata), but likewise a 

Monotremes. . N 

separate sub-class (Prototheria); and are broadly 
distinguished from all other members of their class by laying eggs, 
from which the young are in due course hatched; as they are likewise 
by the milk-glands of the female opening on the surface of the 
skin by means of a number of minute perforations, without being 
furnished with nipples. The group is represented by three genera, 

FIG. i. THE DUCKBILL. (Ornithorhynchus anatinus.} 

one of which is widely different from the other two and forms a 
family by itself, while the latter constitute a second family. The 
duckbill {Ornithorhynchus anatinus}, as the single representative 
of the first family (OrnithorhynchidcB) is commonly termed, is an 
aquatic, somewhat mole-like, burrowing animal, easily recognised 
by the expansion of the muzzle into a broad duck-like beak 
covered during life with a sensitive skin, and also by the broadly 
webbed feet, of which the soles are naked and devoid of pads. 


Although in the adult the mouth is furnished only with horny 
plates, in young individuals the sides of the jaws are provided 
with three pairs of molar teeth, quite unlike those of any other 
living mammals. At the present day the duckbill is confined to 
Queensland south of latitude 18, New South Wales, Victoria, 
South Australia, and Tasmania; it is represented by an extinct 
species from the Plistocene of Queensland, but otherwise the 
pakeontological record of the group is a complete blank. 

Just the same is the case with the echidnas, or spiny anteaters 
(Echidnidce], of which the only fossil remains known have been 
obtained from the superficial deposits of New South Wales. 
Terrestrial and fossorial in their habits, the echidnas differ from 
the duckbill in having the muzzle in the form of an exceedingly 
slender cylindrical toothless beak, furnished with an extensile 
worm-like tongue ; while the fur is thickly mingled with short 
spines, the tail being rudimental, and the unwebbed toes provided 
with extremely powerful claws. Of the two species, the common 
five-clawed echidna (Echidna aculeata] extends from south-eastern 
New Guinea throughout Australia to Tasmania; whereas the 
three-clawed echidna (Proechtdna 1 bruijni} is restricted to New 

With the exception of the Plistocene forms already alluded to, 
no fossil monotremes whatever are known to science. It is, how- 
ever, not improbable that certain extinct mammals from the 
Secondary and lower Tertiary rocks of Arctogaea, commonly termed 
Multituberculata, which will be more fully alluded to in the sequel, 
may indicate a second order of the sub-class Prototheria. Both 
the extinct and the living groups are, however, of a highly special- 
ised type, so that the one cannot apparently be regarded as 
ancestral to the other ; but if the presumed distant relationship 
between the two be substantiated, it will indicate that we are to 
look to a northern origin for the existing monotremes. 

The marsupials, which likewise represent both a separate 
order (Marsupialia) and a sub-class (Metatheria) 


oy themselves, differ from the monotremes by 

producing living young, and by the milk-glands of the female 

1 It has recently been proposed to substitute the term Zaglossus, which is 
stated to be earlier, for this genus. 

L. 3 


discharging their secretion by means of nipples. From the higher 
mammals (Eutheria) they are distinguished by the imperfectly 
developed condition of the newly-born young, and the absence 
of any prenatal connection between the vascular system of the 
foetus and the maternal parent by means of the organ known 
as the placenta. Very generally the young are carried about for 
some time after birth in a pouch situated on the abdomen of the 
parent, where they at first remain immovably fixed to the nipples, 
the milk being injected into their throats by the action of a special 
muscle. In the carnivorous and insectivorous forms the number 
of incisor teeth in the upper jaw usually exceeds the three pairs 
which form the general maximum limit in the higher mammals. A 
further peculiarity of the order is to be found in the replacement 
of the teeth. Instead of the whole or nearly the whole of the first, 
or milk-set of teeth in advance of the true molars or hinder cheek- 
teeth being replaced by a second set of permanent teeth, only one 
tooth is thus (and that by no means invariably) replaced. The 
tooth thus replaced was long regarded as corresponding to the last 
or fourth milk-molar of the higher mammals, while the apparently 
replacing tooth was identified with the last or fourth premolar of 
the same. From recent researches, however, it would seem that 
in reality this is not a case of true replacement, and that the tooth 
which makes its appearance late in life is a retarded premolar, 
representing the fourth in that series, while the replacing tooth is 
the fifth. 

Marsupials may be divided into two main sections or sub- 
orders, readily distinguished from one another by their dentition, 
both of which are represented in Notogaea. In the first of these, 
or Diprotodont sub-order, which is the more specialised of the two, 
the incisor teeth are separated by a gap from those of the cheek- 
series, and do not usually exceed three in number on each side of 
the upper jaw 1 , and in the lower jaw are generally reduced to a single 
pair, while the tusks, or canines (<:), are either small or wanting. In 
their habits the members of this section are more or less exclusively 
herbivorous. On the other hand, in the Polyprotodont mar- 
supials, all of which are mainly carnivorous or insectivorous in 

1 The only exception to this occurs in the South-American forms. 


their diet, the incisor teeth are numerous and pointed, the canines 
are large and well developed, and the whole of the anterior teeth 
form a series more or less nearly continuous with those on the 
sides of the jaws. The typical diprotodonts, or those in which 
two of the toes of the hind foot are enclosed in a common integu- 
ment 1 , are exclusively confined to the Notogaeic realm, where they 
attain their maximum development in the Australian region ; but 
the polyprotodonts and an aberrant group of diprotodonts are 
still represented in the Neogaeic realm, while during the Secondary 
and earlier part of the Tertiary period the former were widely 

(To exhibit Diprotodont type of dentition.} 

spread over Arctogaea. In the Australian region marsupials play 
the part of the eutherians of other regions, and show a remark- 
able diversity of external form and structure, adapting them to all 
modes of life with the exception of the aquatic. And it is fairly 
evident that within the limits of this region the diprotodonts were 
originally evolved from the more generalised polyprotodonts. 

Of the three existing family groups into which the Notogaeic di- 
protodonts are divided the first is the Macropodidce, or the kangaroos 
and their allies ; this being in some respects the most specialised 
group of all, and characterised by certain peculiar features in the 
skull and dentition. Among these the typical genus Macropus, in- 
cluding the true kangaroos, comprises a total of twenty-three 
species, out of which twenty are confined to Australia and 

1 The term syndactylous is applied to this type of foot. 



Tasmania, one (M. agilis) is common to Australia and Queens- 
land, and two others (M. bruijni and M. browni} are confined to 
New Guinea or the adjacent islands. Of the six species of rock- 
kangaroos (Petrogale), none are found out of continental Australia, 
and the same is true with regard to the three representatives of 
the nail-tailed wallabies (Onychogale), and likewise with the three 
hare-wallabies (Lagorchestes}. On the other hand, the three kinds of 
dorca kangaroo (Dorcopsis) are exclusively Papuan ; while of the 
climbing tree-kangaroos (Dendrolagus), three are from Papua 
and two from Queensland. The single species of banded wallaby 
(Lagostrophus) is Australian ; as are also the whole of the rat- 
kangaroos, forming the genera Potorous, Caloprymnus, Bettongia, 
and sEpyprymnus, and likewise the peculiar musk-kangaroo 
(Hypsiprymnodon), which serves to connect the other members of 
the family with the phalangers. 

Several of the existing representatives of the above-mentioned 
genera are found in a fossil state in the cavern-deposits of New 
South Wales and the Plistocene formations of Queensland, in 
addition to which there are likewise several extinct representatives 
of the genus Macropus, some of which considerably exceed the 
largest living forms in point of size. The same formations have 
also yielded the remains of three extinct genera, namely Palor- 
chestes, Procoptodon, and Sthenurus, all of which appear to have 
been allied to the wallabies, although some of the species were 
vastly larger than any existing kangaroo. Another, but very im- 
perfectly known genus Triclis, seems to have connected the 
musk-kangaroo so closely with the phalangers, that it is scarcely 
possible to draw any distinction between these two families. 

In the family of the phalangers (Phalanger idcz], which 
differs from the more typical representatives of the preceding 
by the more generalised characters of the skull, teeth, and 
limbs, there is an exclusively Australian form in the koala, 
forming the sole representative of the genus of the same name. 
Of the five species of cuscuses (Phalanger}, one is, however, 
common to northern Australia, New Guinea, and the Austro- 
Malayan Islands, while the other four are restricted to the two 
latter areas. The two species of true phalanger (Trickosurus) are, 
on the other hand, exclusively Australian ; while the ring-tailed 


phalangers, constituting the genus Pseudochirus, are common to 
Australia and New Guinea. Another exclusively Australian type 
is to be found in the taguan flying phalanger (Petauroides) ; but of 
the non-volant striped phalangers (Dactylopsila) one species is 
common to Queensland, the Aru Islands, and New Guinea, while 
the second is exclusively Papuan. The true flying phalangers of 
the genus Petaurus include two Australian species, and a third, 
common to northern and eastern Australia and New Guinea and 
the adjacent islands. Leadbeater's phalanger (Gymnobelideus), 
which appears to be closely related to the ancestral stock 
from which were evolved the members of the last genus, is 
restricted to Victoria ; but the dormouse-phalangers of the genus 

FIG. 3. SKULL OF EXTINCT PHALANGER (Thylacoleo carnifex). 

Dromicia have both Australian and Papuan representatives, while 
the pen-tailed phalanger (Distachurus) is exclusively from New 
Guinea, and of the two pigmy flying phalangers (Acrobates), one is 
Australian and the other Papuan. Lastly, the aberrant long- 
snouted phalanger ( Tarsipes), representing a sub-family by itself, 
is confined to Western Australia. Remains of species belonging 
to some of the existing genera have been disinterred from the 
caves of New South Wales and the Plistocene deposits of Queens- 
land; while several more or less imperfectly known extinct 
generic types have been described. Among the latter, by far the 
most remarkable is Thylacoleo, which was a gigantic phalanger 
comparable in size to a large leopard, and distinguished by the 
great development of the last premolar tooth in each jaw. The 


tooth in question has an elongated cutting-blade, adapted to work 
against its fellow in the opposite jaw with a scissor-like action, 
somewhat after the fashion of the carnassial teeth of a tiger ; but 
the other cheek-teeth were all relatively small, although the tusks 
were large. The giant among the marsupials was the extinct 
Diprotodon of the Australian Plistocene, a creature rivalling in size 
the extinct South American Megalotherium, and allied on the one 
hand to the kangaroos, and on the other to the phalangers. It 
was not, however, endowed with the leaping powers of the former, 
and doubtless walked on the ground in the ordinary manner, its 
toes having apparently been covered with structures intermediate 
between hoofs and nails. Nearly related, but likewise repre- 
senting a family by itself, is the somewhat smaller, but still 
gigantic Nototherium, which in the conformation of its limb-bones 
appears to approximate to the wombats, and may consequently 
have been, like those animals, of fossorial habits. 

The last Notogaeic family of the Diprotodont section is that of 
the wombats (Phascolomyidcz\ distinguished from all the preceding 
forms by the presence of only a single pair of incisor teeth in both 
the upper and lower jaws ; canines being absent, and the whole 
dentition thus curiously simulating that of the rodents among 
the higher mammals. All the three existing species, which 
are included in the single genus Phascolomys, are confined to 
Australia and Tasmania; and, except certain extinct species 
belonging to the same genus, the only other member of the 
family is the extinct Phascolonus (Sceparnodoti) from the Australian 
Plistocene, distinguished by the peculiarly flattened and chisel- 
like form of the upper incisor teeth. This, the only known 
species, attained much larger dimensions than either of the 
existing wombats. 

The Polyprotodonts likewise include three existing families 
found within the limits of the Australian region, none of the mem- 
bers of which stray either into the Austro- Malay an or Polynesian 
regions, although the separate family of the opossums (Didelphyidcz} 
inhabits the New World. In the family of the bandicoots (Pera- 
melidtz}, the two species of rabbit-bandicoot (Peragale) are ex- 
clusively Australian, whereas the true bandicoots (Perameles] have 
both Papuan and Australian representatives ; the third genus 




( Chceropus\ which includes only the pig-footed bandicoot, being 
confined to Australia. 

In the second family, or Dasyurida, the genus Thyladnus is 
now confined to Tasmania, but it was represented during the 
Plistocene period on the Australian mainland, where one species 
is stated to have been obtained from beds of Pliocene age. A 
similar distribution also obtains in the case of the genus Sarcophifas, 
now represented only by the well-known Tasmanian devil. 

FIG. 4. BANDED ANTEATER. (Myrmecobius fasciatus.) 

Although mainly Australian, the smaller animals known as dasyures 
(Dasyurus] have, however, a single Papuan representative ; while 
the pouched mice (Phascologale) are likewise common to the two 
areas, one of the species ranging from New Guinea to eastern and 
southern Australia. On the other hand, both the narrow-footed 
pouched mice (Sminthopsis) and the jumping pouched mouse 
(Antechinomys] are exclusively Australian. The same is the case 


with the aberrant banded anteater (Myrmecobius), which although 
generally included in the Dasyuridce, should perhaps form the 
type of a family by itself; this animal differing from all the fore- 
going in the number, and also in the structure of the cheek-teeth, 
and thereby making a marked approximation to certain Marsupials 
of the Jurassic epoch noticed in the sequel. 

Before leaving this family, it should be mentioned that certain 
extinct Marsupials from the Tertiaries of Patagonia, referred to in 
the next chapter, seem to be inseparable from it, while there are 
strong reasons for regarding one of them (Prothylacinus) as very 
nearly allied to the existing genus Thylacinus. 

The last of the Australian families {Notary ctidce] of the sub- 
order is represented solely by the marsupial mole (Notoryctes), 
from the sandy deserts of central South Australia ; this being the 
only member of the order which has taken to a subterranean mode 
of life. There are no extinct Australian genera of the sub-order. 

Exclusive of the bats, the only other order of mammals well 
represented in the Australian region is that of the 
Rodentia, or Gnawing Mammals, which bear, how- 
ever, a small proportion to the marsupials, and all of which belong 
to the mouse-family (Murtdce). And it is noteworthy that although 
several of these belong to generic types unknown elsewhere, the 
whole of them are animals of comparatively small size, so that it 
is possible that their ancestors may have been introduced without 
a direct land-connection with any other part of the world. A 
curious feature in connection with this group is that two of the 
Australian species, namely Hydromys chrysogaster and Musfuscipes, 
are aquatic in their habits ; whereas, as we have seen, none of the 
Australian marsupials are natatorial, although the duckbill is 
eminently so. The Australian water-rat (Hydromys), which is 
common to Australia and New Guinea, belongs to a sub-family 
typically distinguished from all other Murtdce by the reduction of 
the molar teeth to two pairs in each jaw. While this animal has 
partially webbed toes, and is strictly aquatic in its habits, the 
allied Xeromys from Queensland is terrestrial, and approximates 
to the more typical members of the family, although to which 
group is still uncertain. The only other representatives of the 
sub-family Hydromyina are met with in the mountains of Luzon, 


in the Philippine group, where there is one genus allied to 
Hydromys, while other species have been assigned to the genus 
Xeromys. Whereas the typical Australian representative of the 
latter has but two pairs of molar teeth, one of the Philippine 
forms has three, thus approximating to more ordinary murines. 
The occurrence of these Australian types of rats in the Philippines 
is of the utmost importance in respect to Australia having received 
its mammalian fauna from south-eastern Asia. 

Of the typical genus Afus, whose geographical distributional 
area includes the whole of the eastern hemisphere with the ex- 
ception of Madagascar and many of the Polynesian islands 1 , 
Australia has upwards of twenty-six representatives, while two 
species occur in the Duke of York group, and others probably on 
the Papuan mainland. One of the Duke of York species (M. 
prcetor) ranges eastwards into the Solomons, where three other 
kinds are also found. The jerboa-rats (Conilurus*) form a pe- 
culiar saltatorial group restricted to the sandy deserts of the 
mainland of Australia, where they are represented by about a 
dozen species ; while the broad-toothed rat (Mastacomys) is con- 
fined to Tasmania, although its fossilised remains, like those of 
the other genus, are met with in the caverns of New South Wales. 
More nearly allied to the true rats and mice, the mosaic-tailed rats 
(Uromys) inhabit Queensland and the Aru Islands, one of the 
species from the former area also occurring in the Solomons. 
Lastly, the prehensile-tailed rat (Chiruromys], from the mountains 
of south-eastern New Guinea, represents a genus distinguished 
from all other placental mammals of the eastern hemisphere, 
with the exception of the British harvest-mouse and the Oriental 
binturong, by the prehensile nature of its tail. 

In connection with these rodents it is important to observe 
that fossil Murida are unknown from any part of the world earlier 
than the Miocene epoch, so that it is evident the living Australian 
representatives of the family are comparatively recent immigrants 
into the region they inhabit. 

1 The Pacific rat (Mus exulans) appears to be widely distributed in these 
islands, see note on p. 29. 

2 Commonly known by the preoccupied name Hapalotis. 


Much discussion has taken place with regard to the date of 
introduction of the native dog, or dingo ( Canis dingo] 

Carnivores. . ,. . . -jj^i^-j. 

into Australia, and it was long considered that it 
was imported by human agency. Seeing, however, that its remains 
have been found in association with those of extinct kangaroos 
and Diprotodon, there seems considerable probability of its being 
an indigenous inhabitant of the country l . 

The only other non-volant mammal found in the Australian 

region is a species of pig (Sus papuensis). This 

animal is, however, so closely allied to certain 

Malayan species that it seems quite possible that its introduction 

may be due to human agency. 

The Australian region contains representatives of all the families 

of Bats with the exception of the Neogaeic Phyllo- 

stomatid(z\ some of the genera, such as the tube- 

nosed bats (Uronycteris*}, among the Pteropodida, being peculiar 

to this and the Austro- Malayan region, while others are more or 

less widely spread, or even cosmopolitan. It will be unnecessary 

to mention the various genera by name ; but the affinity of the 

Notogaeic Chiroptera to those of Eastern Arctogaea, as exemplified 

by the abundance of fruit-bats (Pteropodidce) and the absence of 

the PhyllostomatidcR) is noteworthy. 

In the following synoptical list the higher groups and genera 
(exclusive of Bats) peculiar to the Notogaeic realm 
are printed in italic type ; the letters A, P, and M 

Papuan following the names respectively indicate that the 


groups in question occur in Australia (inclusive of 
Tasmania), New Guinea (with the adjacent Papuan islands), or the 
Austro-Malayan region ; extinct groups have an asterisk prefixed. 
Ornithorhynchidce, A. 
OrnithorhynchuS) A. 
EchidnidcB, A. P. 

Echidna, A. P. (The living species common to 

both areas.) 
Proechidna, P. 

1 See Ogilby, "Catalogue of Australian Mammals," Sydney, 1891 92. 

2 This name replaces the preoccupied Harpvia. 




II. Marsupialia. 


Macropodidce, A. P. 

Mac r opus, A. P. 

Petrogale, A. 

Onychogale, A. 

Lagorchestes, A. 

Dorcopsis, P. 

Dendrolagus, A. P. 

Lagostrophus, A. 

Potorous, A. 

Caloprymnus, A. 

Bettongia, A. 

sEpyprymnus, A. 

Hypsiprymnodon, A. 
*Palorchestes, A. 
* Procoptodon, A. 
*Sthenurus, A. 
*Triclis, A. 

Phalangerida, A. P. M. 
Koala, A. 

Phalanger, A. P. M. 
Trichosurus, A. 
Pseudochirus, A. P. 
Petauroides, A. 
Dactylopsila, A. P. 
Petaurus, A. P. M. 
Gymnobelideus, A. 
Dromicia, A. P. 
Distcechurus, P. 
Acrobates, A. P. 
Tar sip es, A. 
*Thylacoleo, A. 

*Diprotodontidcz, A. 
* Diprotodon, A. 

(Elsewhere represented only 
by an aberrant group in 
South America.) 


4. *Nototheriid(K) A. 

* Nototherium^ A. 

5. Phascolomyid&i A. 

PhascolomyS) A. 
*Phascolonus, A. 


1. Peramelid&i A. P. 

Peragale, A. 
Perameles, A. P. 
Chc&ropuS) A. 


ThyladnuS) A. 
SarcophiluS) A. 
Dasyurus, A. P. 
Phascologale, A. P. 
Smmthopsis, A. 
AntechinomyS) A. 
MyrmecobiuSj A. 

3. Notoryctidce, A. 

Notoryctes, A. 

III. Rodentia. 

i. MuRiDjE. Cosmopolitan. 
Hydromys, A. P. 
Xeromys, A. and Philippines. 
Mus, A. P. M. 
Conilurus, A. 
Mastacomys, A. 
Uromys, A. P. 
Chiruromys, P. 

IV. Carnivora. 

CANID^:, Cosmopolitan. 
Canis, A. Cosmopolitan. 

V. Ungulata. 

SUID.E. Throughout Eastern Hemisphere, except 


Sus, P. Elsewhere throughout greater part of 
Eastern Hemisphere. 


VI. Chiroptera. All the families, with the exception of 

the Neogaeic Phyllostomatidae, well re- 

The Polynesian region, as already said, is characterised by the 
general absence of non-flying mammals, and there- 
fore claims but little notice here. The only mar- 
supial occurring within the region is a variety of the 
widely-spread grey cuscus (Phalanger orientalis], which occurs in 
the Solomon Islands, where four species of Mus are likewise met 
with. As the cuscus, together with one of the rats, is also found 
in the Duke of York group, it may be inferred that the non-volant 
mammals of the Solomons have been derived from the latter area. 
In addition to members of widely-spread types, the Solomons 
possess two peculiar genera of bats. 

New Zealand appears to be inhabited only by two peculiar 
generic types of bats, each represented by a single species, and a 
rat (Mus maorimri}, but whether the latter is indigenous or intro- 
duced appears doubtful. 

Although a work devoted to mammals has little to do with an 
area where the sole member of the class is a bat of 
the genus Atalapha, brief mention must be made of R^on*"' 
the Sandwich Islands, which from their bird-fauna 
are regarded as entitled to distinction from the Polynesian region. 
Of the birds of this area, Mr W. L. Sclater ' writes that the greater 
number not only of the species, but even of the genera "are 
peculiar and wholly restricted to these islands. It is, of course, 
among the smaller land-birds (Passeres) that this individuality is 
most marked; but even in the other groups, where the distribu- 
tion is generally wider, the Hawaiian birds are, in many cases, 

Poverty, and an admixture of Australian and Malayan types, 
with a very marked preponderance of the latter, are 
the leading features in the mammalian fauna of the lay^n Region. 

Austro-Malayan region. This area includes the 

islands lying between Makassar Strait and the narrow channel 

separating Lombok from Bali on the west and the Australian region 

1 Appendix, No. 28. 


on the east. The largest of these is Celebes, while those of the 
Moluccan group, such as Gilolo, Buru, Ceram, and Timor-Laut, 
together with Timor and Sumbawa, are of smaller size. Unfor- 
tunately no complete lists of the fauna of these islands, so far as I 
am aware, have yet been published. 

Commencing with Timor and the Moluccas, we find several 
of the latter group of islands inhabited by four species of 
cuscus (Phalanger\ two of which are common to the Australian 
region, while the third (P. ornatus] is peculiar, and the fourth 
(P. celebensis], which in this group is found only in Sanghir Island, 
is an inhabitant of Celebes, where the other three are unknown. 
The only other Austro-Malayan marsupial 1 is a variety of the 
Australian lesser flying-phalanger (Petaurus breviceps\ this variety 
ranging eastwards from Gilolo to the New Britain group. With 
the possible exception of certain shrews, most of the few Moluccan 
species of eutherian mammals appear to be identical with those 
of Celebes, whence they were probably introduced. A deer from 
Timor has received a distinct name (Cervus timoriensis\ and the 
same island is also inhabited by a common Malayan monkey 
(Macacus cynomolgus), a palm-civet (Paradoxurus hermaphroditus], 
and a true civet ( Viverra tangalungci], the latter being common to 
the Moluccas. The common Javan porcupine (Hystrix javanica), 
which is widely spread in the Malayan islands, is also found in 
Timor. There is likewise a cat in the same island, which although 
described under the name of Felis megalotis as a distinct species, 
and regarded by Mr Jentink as such, has been identified by 
Mr W. L. Sclater in his " Catalogue of the Mammalia in the 
Indian Museum " as a mere variety of the domestic species. In 
regard to all the Timorese forms which are closely allied to, or 
identical with well-known Malayan species, it is necessary to take 
into consideration the well-known partiality of the Malays for 
taming wild animals and carrying them about during their voyages; 
and it is highly probable that all or most of such animals found 
in Timor have been thus introduced. From the small island of 
Flores Mr Jentink has described a rat (Mus armandvillei), which is 
the largest member of its genus. 

1 The Kei Islands, like the Aru group, may be best affiliated to Papua. 




In addition to the above-mentioned cuscus, which appears to 
be its only marsupial, Celebes possesses several peculiar types of 
eutherian mammals. Among these is a black and nearly tailless 
ape (Cynopithecus niger) representing a genus by itself; while there 
is also a species of macaque (Macacus maurus) peculiar to the 
southern portion of the island. A species of the lernuroid tarsiers 
(Tarsius fuscomanus) is found both in Celebes and the neighbour- 
ing islands of Salayer and Sanghir, although represented by an 

FIG. 5. FORE PART OF SKULL OF BABIRUSA (Bcibirusa alfurus], 

allied form in the Philippine group. In the Carnivora there is a 
Malayan species of civet ( Viverra tangalunga), and also a peculiar 
species of palm-civet (Paradoxurus musschenbroecki). In the pig- 
tribe the babirusa (Babirusa alfurus), characterised by the extra- 
ordinary development of its tusks, is the sole representative of a 
genus confined to this island and Bum; while scarcely less peculiar 
is the small and somewhat antelope-like buffalo known as the anoa 
(Bos depressicornis\ which although allied to the tamarao (B. mindo- 


rensts) 1 of the Philippines, has its nearest relatives in certain 
extinct species from the Pliocene of Northern India. There is 
also a true pig (Sus celebensis], nearly allied to Malayan forms ; as 
well as a deer forming a variety of the widely spread sambar 
(Cervus unicolor}. Among the rodents, a rat with an extremely 
long muzzle constitutes a peculiar genus (Echinothrix)^ and there 
are also other Muridce, as well as squirrels (Sciurid<K), in addition 
to numerous bats, mostly belonging to Oriental types ; certain of 
the squirrels, such as Sciurus prevosti, being common to the 
Malayan countries. 

Unfortunately there is absolutely no palseontological evidence 
to help us in regard to the past history of these islands ; but from 
the living mammalian fauna we should be inclined to place the 
whole area within the limits of the Oriental region. On the other 
hand, a large number of the birds both of the Moluccas and 
Celebes are peculiar; and Australian affinities are displayed by the 
presence of a bird of paradise (Semioptera) in Gilolo and Batjan, 
and of a cassowary (Casuarius) in Ceram. Accordingly, it may be 
well to include not only the Moluccas, but likewise Celebes within 
the limits of the Notogseic realm, which will then embrace the 
whole of the countries where monotremes, typical diprotodont 
marsupials, birds-of-paradise, and cassowaries occur. Still it 
must be confessed that this is, after all, mainly a matter of con- 
venience, seeing that since, as will be shown below, the diprotodont 
marsupials have in all probability originated in the Australian 
region, those inhabiting the Austro-Malayan region must ap- 
parently be regarded as comparatively late immigrants from the 
south-east 2 ; the same being also true with regard to the single 
bird-of-paradise and the cassowary inhabiting this area. And it 
is noteworthy that both genera of Austro-Malayan marsupials are 
precisely such as, from their arboreal habits, would be likely to be 
transported on floating timber. Dr Wallace has suggested that 
Celebes has been separated from the Oriental region since the 

1 It has been suggested that this animal is a hybrid between the anoa and 
the Indian buffalo. 

2 In this view I am in accord with Dr Blanford, who (GeoL Mag. decade 
3, vol. IX. p. 165, 1892) writes that the marsupials of Celebes "are probably 
of later introduction than the mammals with Oriental affinities." 


Miocene ; this, however, is obviously too early a date, since the 
only known allies of the anoa are met with (in common with the 
earliest of all the oxen) in the Siwalik Pliocene of northern 

In regard to the Moluccas, Dr Wallace 1 observes that the 
absence of many characteristic groups of Papuan birds, and 
likewise of kangaroos and the smaller marsupials of New Guinea, 
leads to the conclusion that these islands " cannot be mere frag- 
ments of the old Papuan land, or they would certainly, in some 
one or other of their large and fertile islands, have preserved a 
more complete representation of the parent fauna. Most of the 
Moluccan birds are very distinct from the allied species of New 
Guinea; and this would imply that the entrance of the original 
forms took place at a remote period. The two peculiar genera 
with clearly Papuan affinities, show the same thing. The casso- 
wary, found only in the large island of Ceram and distinct from 
any Papuan species, would however seem to have required a land 
connection for its introduction, almost as much as any of the 
larger mammalia." 

In another work 2 , summing up the general conclusions with 
regard to the fauna of Celebes, the same writer observes that " we 
are fully justified in classing it as an ' anomalous island,' since it 
possesses a small but very remarkable mammalian fauna, without 
ever having been directly united [during Tertiary times] with any 
continent or extensive land ; and, both by what it has, and what 
it wants, occupies such an exactly intermediate position between 
the Oriental and Australian regions that it will perhaps ever 
remain a mere matter of opinion with which it should properly 
be associated. Forming, as it does, the western limit of such 
typical Australian groups as the marsupials among mammalia, 
and the Trichogfassida* and Mdiphngidct* among birds, and being 
so strongly deficient in all the more characteristic Oriental families 
and genera of both classes, I have always placed it in the Austra- 

1 Geographical Distribution of Animals, Vol. I. p. 419. 

2 Island Life, p. 432. 

3 Equal LoriidfB ; includes the brush-tongued lories and loriquets. 

4 Honey-suckers. 

L. 4 


lian region 1 ; but it may perhaps with equal propriety be left 
out of both till a further knowledge of its geology enables us to 
determine its early history with more precision." 

Having now briefly surveyed the leading features of the terres- 
trial mammalian fauna of the whole Notogaeic realm, 


logicai History and discussed the relationship of the mammals of 
the Austro-Malayan to those of the Australian region 
(in the restricted sense of the term), we are in a position to enter 
upon the consideration of the probable past history of Australia 
and New Guinea, so far as the same group of animals is con- 
cerned. Before doing so, it is, however, essential to state what 
is known concerning marsupials from other regions of the world. 
Here it may be premised that in regard to Australia mammalian 
palseontological history is a total blank previously to the Pliocene 
epoch ; while apparently but little is known even of that period, 
the great majority of the fossil mammals of that country belonging 
to the Plistocene epoch of the earth's history. As to the past 
history of the mammals of New Guinea, we know absolutely 
nothing ; and, as already mentioned, the same is the case with 
regard to those of the Austro-Malayan islands. This, however, by 
no means exhibits the whole depth of our deficiency of informa- 
tion. Throughout the whole of eastern Asia, to say nothing of 
Alaska and western Canada, we have no information whatever as 
to mammalian life previous to the Pliocene era; while even in 
that period our sole knowledge relates to a portion of northern 
India and China. If, therefore, some of the modern types of 
marsupials originated in eastern Asia from the older forms, the 
blank in the palaeontological history of the group relates to just 
the very countries where these animals might naturally be expected 
to occur during the Tertiary period. While there is no record of 
their existence in Asia, in Europe fossil Tertiary marsupials are 
unknown at a later date than the upper Oligocene, and in North 
America than the middle Oligocene, and the whole of those 
hitherto discovered belong to the existing American group of 
opossums. If, however, we were to infer from this that the whole 
order (with the exception of that group) never existed in conti- 

1 Equivalent to the Notogaeic realm of this work. 


nental Arctogaea after the Secondary epoch, a very serious error 
might be committed. And although it is improbable that any 
marsupials of an Australian type ever existed in Europe or North 
America, there is no reason why they should not have occurred 
in south-eastern Asia. 

The extinct dasyurids of the Patagonian Miocene have been 
already mentioned, and these, together with another S. American 
group, are more fully noticed in the next chapter. With regard 
to the opossums, it will suffice to state that while they are 
unknown in the aforesaid Patagonian deposits, certain species 
occur in the middle Oligocene White River beds of the United 
States, and others in the lower, middle, and upper Oligocene 1 beds 
of Europe. Although the number of their incisor teeth is unknown, 
there is little doubt that the European Oligocene opossums 2 (which 
have been very generally separated as Pcrathtrium\ should 
be included in the existing genus Didelphys. Remains of 
these animals have been obtained from the upper Oligocene of 
Cournon in France, from the middle Oligocene beds of Hordwell 
in Hampshire, from the equivalent deposits of Debruge in 
Vaucluse, and of Montmartre near Paris, and likewise from 
the Quercy Phosphorites in the south of France. With the 
exception of a peculiar South American group of diprotodonts, 
the remaining fossil mammals which can be referred to the 
Marsupialia are mainly if not exclusively confined to the Second- 
ary period ; all being of small dimensions, and many of them 
exceedingly minute. While many of them evidently died out 
without leaving any existing descendants, one group seems to 
have been the ancestral type from which the existing Dasyurida 
have originated. Among the former, we have the family Tricono- 
dontidce, as represented by the genus Triconodon of the upper 
Jurassic of England and also by nearly allied forms from the 
corresponding rocks of the United States. In this family the 

1 It may be well to mention that the beds of St Gerand-le-Puy, in France, 
which many writers reckon as lower Miocene, are here classed as upper Oligo- 
cene. See Lydekker, Cat. Foss. Mamm. Brit. Mus. Pt. ,iv. p. xvii. 

2 The existing Didelphyida differ from the Dasyurida in the presence of four, 
in place of three, pairs of incisor teeth in the lower jaw, and of five pairs in the 
upper jaw instead of four. 



molar teeth consist of three simple compressed trenchant cusps 
arranged in a longitudinal line ; the upper ones biting on the 
outer side of those of the lower jaw. In the upper jaw the 
number of teeth is still unknown, but the lower jaw carries three 
pairs of incisors, four of premolars, and either three or four of molars, 
in addition to the tusks or canines, which are implanted by two 
distinct roots. In this respect the latter teeth present an approxi- 


mation to those of the bandicoots, where the root of the canine is 
partially divided by a longitudinal groove. A second family 
(Spalacotheriidce), likewise represented in the upper Jurassic rocks 
both of Europe and the United States, is distinguished by the 
cusps of the molars being arranged in a triangle, with the apex 
pointing inwards in the upper, and outwards in the lower jaw ; 
these teeth being similar in structure to those of the marsupial 

Of more interest is the large family of the Amphitheriida, 
which may be taken to include a great number of forms apparently 
agreeing with the opossums in having four pairs of lower incisor 
teeth. The lower molars never consist solely of three simple 
cusps arranged in a straight line like those of the Triconodontida, 
or in a triangle like those of the Spalacotheriidce. Among these 
forms the genus Phascolotherium, from the lower Jurassic Stones- 
field Slate of Oxfordshire, appears to have had only seven pairs of 
cheek-teeth ; the lower molars having three cusps arranged in a 
longitudinal line, of which the middle one is considerably larger 




than the other two, while there are minute additional cusps at the 
two extremities, and a distinct ledge at the base of the inner side 

Nat. size. 


of each tooth. In the allied Amphilestes, from the same forma- 
tion, of which the imperfect lower jaw is shown in the annexed 
figure, the molars are of the same general type as in the preceding, 
but much more numerous, their total number being probably 
nearly the same as in the next genus. 

Nat. size. 


Another type is represented by the genera Amphitherium of 
the Stonesfield Slate and Amblotherium of the upper Jurassic of 
Dorsetshire, in which, in addition to the canines, there are from 
six to eight molar teeth, four premolars, and four incisors in each 
half of the lower jaw. The lower molars differ from those of the 
preceding genera, and thereby resemble the corresponding teeth 


of the opossums and bandicoots, in that they consist of an anterior 
portion carrying three cusps in a triangle, and of a posterior 
moiety or heel. Several more or less nearly related genera have 
left their remains in the upper Jurassic rocks of the United States, 
among which Dryolestes may be specially mentioned ; and it 
appears that in North America the group survived till the succeed- 
ing Cretaceous epoch. The especial interest attaching to these 
marsupials is that they, and they alone, have molar teeth com- 
parable in number, and to a certain extent in structure, with those 
of the living Australian Myrmecobius ; and there can be but little 
hesitation in regarding the latter as the specially modified descen- 
dant of these ancient forms of mammalian life. It is, however, 
important to notice that all the Jurassic types have four pairs of 
lower incisor teeth, which are now retained by the opossums alone, 
having been reduced in all the Australian polyprotodonts to three. 
Although a very low type of mammal (Dromatherium) occurs 
HOWAUS- i n t ^ ie Triassic rocks of North America, the fore- 
traiia received going include all the leading extinct forms that can 
be included among the marsupials. During the 
Jurassic epoch the group seems to have been widely distributed 
over Europe and North America ; it is known to have existed in 
the latter area during the Cretaceous epoch, and it probably also 
survived to that date in some part of the northern half of- the 
Old World. After that, our knowledge is a blank till we meet 
with the Oligocene opossums of Europe and North America, and 
the Miocene Patagonian marsupials ; so that as regards the 
Eocene epoch there is absolutely no record whatever of the group. 
That Australia received its original fauna of polyprotodont 
marsupials from the northward may be regarded as practically 
certain ; and the question as regards the Notogseic realm accord- 
ingly narrows itself to the approximate date of the immigration. 
On this point Dr Wallace 1 writes that "it was probably far back 
in the Secondary period that some portion of the Australian 
region was in actual connection with the northern continent, and 
became stocked with ancestral forms of marsupials ; but from that 
time till now there seems to have been no further land-connection, 

1 Geographical Distribution of Animals, Vol. I. p. 465. 


and the Australian lands have thenceforward gone on developing 
the marsupial and monotremate types into the various living and 
extinct races we now find there." 

Since this passage was written, the case has been somewhat 
materially altered by the discovery of the dasyuroid marsupials of 
the Patagonian Tertiaries ; while recent researches have tended to 
show that the alliance between the Dasyuridce and the Didelphyida 
is much more intimate than was formerly supposed to be the 
case 1 . This being so, it is a fairly safe assumption that both 
families are descended from a single common ancestral stock 
which, apart from any question of a connection between Australia 
and South America, can hardly have originated anywhere than in 
the northern hemisphere, seeing that the Didelphyidce are totally 
unknown in Notogaea. There is, however, as already stated, no 
evidence of the existence of opossums before the Oligocene ; and 
it is in the highest degree improbable that the two families were 
differentiated as far back as the Jurassic, or even the Cretaceous 
epoch. Not improbably polyprotodont marsupials survived in 
south-eastern Asia till the early portion of the Eocene division of 
the Tertiary epoch, and in this region both Dasyuridce and 
Didelphyidce. were differentiated. Representatives of the former 
family soon afterwards found their way into Australia and New 
Guinea, while the opossums would appear to have dispersed 
in one direction into Europe and in the other into North America, 
eventually making their way from the latter country at a late 
epoch in the Tertiary period into South America. 

Assuming that the Patagonian dasyurids are more or less 
closely allied to the Australian forms (and this certainly appears 
to be the case), it may be taken for granted that they have not 
originated independently. From considerations advanced in the 
next chapter, it is almost impossible to believe that they travelled 
by way of North America ; and if this be so, their only mode of 
migration would be by means of a land-bridge between South 
America and Australia by way of the Antarctic continent, or 

1 In the British Museum "Catalogue of Marsupials and Monotremes," 
p. 315 (1888), Mr O. Thomas writes that the family Didelphyidce "is, on the 
whole, very closely allied to the Dasyurida, from which, were it not for its 
isolated geographical position, it would be very doubtfully separable." 


possibly in a zone nearer the equator 1 . Assuming such a connec- 
tion to have existed in Tertiary times (and there is no reason why 
it should not have existed), it must either have taken place 
before the development of the diprotodonts in Australia, or must 
have been in such high latitudes, or so transitory, as to permit of 
the passage of only a few forms. It is true that there is no 
definite evidence that land mammals ever existed on the Antarctic 
continent, but during a recent expedition certain seals were killed 
bearing on their hides marks which appeared to have been inflicted 
by the claws of a land carnivore. If this be substantiated by 
future discoveries, it would be not only probable, but essential 
that there should have been a Tertiary connection between 
'Antarctica' and other lands. With regard to the probability that 
'Antarctica' is of continental origin, in summarising what is known 
with regard to the geology of 'Antarctica,' Messrs David and 
Smeeth observe that whether a continent, or an archipelago the 
islands of which are united by thick sheets of ice, the southern land 
is considered to have a superficial area of 4,000,000 square miles, 
being, therefore, larger than Australia. A great chain of volcanoes 
has been described, which in Victorialand rise over 15,000 ft. 
above the sea. On the South American side of Antarctica may 
be specially noticed the active volcano of Bridgman, and the 
large and partially-submerged volcano of Deception Island, with 
its crater over five miles in diameter, the wall of which, built up of 
alternating layers of ice and volcanic scoriae, rises to 1,800 ft. 
above the sea. Sedimentary rocks of Eocene age, with fossil trees, 
were discovered in 1893 at Seymour Island; and the French ship 
Talisman many years previously dredged off the Antarctic conti- 
nent fragments of rock containing Gyroporella, a fossil plant very 
characteristic of the Triassic rocks of Europe. Near Laurie Island, 
in the South Orkneys, limestone occurs. The rocks collected by 
Mr Borchgrevink are of especial interest as confirming the theory 
that Antarctica is a continent rather than an archipelago, for the 
microline-granite with garnet and tourmaline, and the mica-schists 
must have had a continental origin, such rocks being almost 
unknown in oceanic islands, but being of frequent occurrence in 
continental areas. 

1 See Chapter III. 


With regard to the presumed survival of marsupials in south- 
eastern Asia till the Tertiary epoch, it may be mentioned that 
although there is a total lack of knowledge of the early Tertiary 
mammals of Asia, yet there are not wanting indications of an 
affinity between the fauna of the eastern portion of the latter 
continent and that of North America which points to a migration 
from a common centre along the two sides of the Pacific; a 
migration which in the early Tertiary period received on the 
American side an abrupt check by the sea then dividing North 
and South America. There is, for instance, living in Central Asia 
a species of deer so closely allied to the North American wapiti, 
that it is a question whether the two are really entitled to specific 
distinction ; while the Chinese alligator has its nearest living ally 
in the species inhabiting the Mississippi. Another piece of 
evidence is furnished by the occurrence in the Tertiaries of the 
Balkan Peninsula of remains of the perissodactyle genus Titano- 
therium, belonging to a family only known elsewhere in North 
America. Quite recently remains of the North American masto- 
don {Mastodon americanus] have been discovered in eastern 
Russia 1 . 

The existence of such essentially American types in Eastern 
Europe and Central Asia clearly seems to point to a more intimate 
connection between the faunas of those regions than exists 
between those of Western Europe and North America; and thus 
lends countenance to the idea that marsupials may have lingered 
on in Eastern Asia till long after the earlier forms had disappeared 
from Western Europe. On this view, it is quite probable that the 
opossums of the Oligocene of Europe may have been immigrants 
into that area from the south-east ; this being confirmed by the 
absence of the group from the Ethiopian and Malagasy regions. 
As already said, the existence of Australian types of Muridce 
in the Philippines affords a pretty clear proof that Notogaea 
received its fauna from south-eastern Asia, where types that had 
died out elsewhere at an earlier epoch appear to have survived. 
Doubtless, however, the Muridce effected their entrance into 
Australia at a later epoch than the marsupials. 

1 See Pavlow, Mem. Ac. St Petersbourg, ser. 8, vol. i. pt. 3 (1894). 


The case of the ratite, or flightless struthious birds of Notogaea, 
as represented by the extinct moas (Dinornithidcz) and the living 
kiwis (Apterygidce) of New Zealand, and the cassowaries and 
emeus (Casuariidcz) of Papua and Australia, seems to confirm the 
conclusions drawn from the evidence of the marsupials as to the 
isolation of the Notogseic realm not being so ancient as supposed 
by Dr Wallace. It is to be presumed that all will agree that 
more or less complete land-connections must have been necessary 
for the migration of these birds ; and if it can be shown that the 
group is a comparatively modern one, it cannot but support the 
marsupial evidence. Before proceeding to do so, allusion must, 
however, be made to the views of other writers as to the relation- 
ships of the different Notogeeic lands. 

In Island Life 1 , Dr Wallace considers that during the 
Cretaceous, and probably also for a considerable portion of 
Tertiary times, Western Australia was cut off by a deep sea from 
the eastern margin of the continent, which was united with Tas- 
mania, and possibly also with New Guinea. The eastern and 
western islands, he writes, " would then differ considerably in 
their vegetation and animal life. The western and more ancient 
land already possessed, in its main features, the peculiar Australian 
flora, and also the ancestral forms of its strange marsupial fauna, 
both of which it had probably received at some earlier epoch by 
a temporary union with the Asiatic continent over what is now 
the Java Sea. Eastern Australia, on the other hand, possessed 
only the rudiments of its existing mixed flora derived from three 

distinct sources The Marsupial fauna had not yet reached the 

eastern land, which was, however, occupied in the north by some 
ancestral struthious birds, which had entered it by way of New 
Guinea through some very ancient continental extension, and of 
which the emeu, the cassowaries, the extinct Dromor?iis of Queens- 
land, and the moas and kiwis of New Zealand, are the modified 
descendants." He further concludes that a large area of what is 
now the Tasman Sea was upheaved, and nearly, or quite, con- 
nected New Zealand with Australia, whereby the fauna and flora 
then existing in Eastern Australia were enabled to colonise New 

1 Pages 465 et seq. 


Zealand. Finally, this hypothetical bridge sank, isolating such 
forms as had reached New Zealand, and, soon after, Eastern and 
Western Australia became connected by land, and thus assumed 
a homogeneous fauna. From this and other passages, we are led 
to conclude that the author believes that the Notogaeic flightless 
birds immigrated, if not in Cretaceous, at least in early Tertiary 
times 1 , from more northern regions. 

Dr Wallace's conclusions are, however, challenged by Mr C. 
Hedley 2 , who, from a study of the floras of these regions, supple- 
mented by the distribution of land molluscs, and recent geological 
observations, refuses to admit that Western Australia ever pos- 
sessed a monopoly of characteristic Australian animals or plants. 
Although he considers the separation of the western and eastern 
portions of the continent by a Cretaceous sea may be granted, 
yet the land representing Western Australia was much smaller 
than Wallace supposes. " The shallow inland Cretaceous sea was 
studded with islands, large and small, which served the fauna and 
flora as stepping-stones in their migrations from west to east and 
from east to west." During a late era in the Tertiary epoch he 
believes New Guinea to have been in connection with Australia ; 
and further urges " that an ancient continent, separated on the 
west from Australia by the abysses of the Coral and of the Tasman 
Sea, is represented by the Solomons, the Fijis, the New Hebrides, 
New Caledonia, Lord Howe Island, and New Zealand, with its 
outlying islands In conclusion, I would contend that New 
Zealand is associated with the Solomons and the New Hebrides, 
firstly, as a member of their volcanic system ; secondly, by com- 
munity of fauna and flora; whereas to Australia it is related not 
at all physically, and to a foreign and intrusive element bio- 
logically ; and that a theory which derives the fauna and flora of 
New Zealand primarily from these archipelagoes and remotely 
from New Guinea, necessitates fewer unproved assumptions than 
that which derives them from Australia." 

To return to the ratite birds, it may be observed in the first place 

1 If the immigration into Eastern Australia was Tertiary, what becomes of 
the author's statement that Australia has been isolated since the Secondary 
epoch ? 

a Appendix, No. i6> 


that the giant flightless species such as Phororhachis and Bron- 
tornis of the Patagonian Tertiaries have been recently shown to 
form a totally distinct group the Stereornithes, and it is quite 
probable that the same may prove to be the case with Gastornis, 
Dasornis, and Diatryma of the lower Eocene of the northern 
hemisphere. Apart from these, the earliest known ratites are 
Hypselornis of the Pliocene of India which appears to be allied 
to the emeus and cassowaries and the Australian Dromornis, 
one species of which is likewise of Pliocene age; all the other 
forms being Plisiocene. Moreover, it is now tolerably certain 
that the true ratites have originated from flying birds, and it is 
therefore highly probable that the group is an essentially modern 
one 1 . Accordingly, there is a strong presumption that the ances- 
tors of these birds did not enter Notogsea till comparatively late 
in the Tertiary period ; and that, in fact, their southern migration 
was not far removed in time from that of the giant land-tortoises, 
noticed in the next chapter. Possibly they may have entered 
Australia by way of New Guinea during the connection which 
Mr Hedley believes to have existed between those two countries 
late in the Tertiary epoch ; while the New Zealand forms may 
have made their way by means of the presumed land-connection 
between these islands and the Solomons, New Hebrides, etc. 

Of course there is the difficulty as to why mammals did not 
enter the Polynesian region at the same time; but it is conceivable 
that even at this date the mammalian fauna of South-eastern Asia 
may have been very poor in Eutherians, while it is quite possible 
that the ancestral forms of these birds may not have required the 
complete land-connection necessary for the passage of the higher 
mammals. Such connection as served for these birds, however, 
may have well sufficed for the transit of the ancestors of the 
Australian murine rodents, which almost certainly entered the 
country at a later date than the original marsupials and mono- 

Assuming, then, that the marsupials and monotremes of the 
Australian region did not reach their present home till the early 
part of the Tertiary epoch, we must make the further assumption 

1 Captain Hutton is of opinion that the moas originated directly from flying 
birds in New Zealand, but the evidence in favour of this view appears insufficient. 


that at this period South-eastern Asia was entirely, or to a great 
extent, devoid of higher mammals. Nor is this unlikely, seeing 
that the ungulates and carnivores of the lower Eocene of the 
northern hemisphere would clearly have required time to spread 
themselves to the southward. Hence it may be suggested that, 
towards the close of the Cretaceous epoch there was first a migra- 
tion towards the south-east of the ancestral marsupials (and 
monotremes) inhabiting the northern hemisphere during the 
Secondary epoch ; and that similar migrations of the higher mam- 
mals took place during Tertiary times. 

When once the ancestral polyprotodont marsupials obtained a 
footing in New Guinea and Australia, where they have since been 
isolated from any serious competition with the higher mammals, 
they flourished and developed to a degree which they could not 
possibly have attained in any other part of the world under 
existing conditions. And it is doubtless within this region that 
the more specialised diprotodont types were evolved. Remark- 
able as it undoubtedly is, the present state of development of the 
Australian marsupials is nothing to what it was during the 
Plistocene period, when there lived the giant kangaroos, pha- 
langers, wombats, diprotodons, and nototheres already alluded to, 
by the side of which the largest existing species would appear 
almost dwarfs. The cause of this universal extinction (for uni- 
versal it is) of all the larger types of mammalian life throughout 
the world soon after the appearance of man, is one of those 
problems which at present is not capable of being satisfactorily 
solved, as not even a glacial period could have made a clean 
sweep of the whole globe. It may be added that the evolution of 
the diprotodont marsupials within the limits of the Australian 
region, points to the conclusion that the outlying discuses of the 
Austro-Malayan regions are immigrants from the south-east. 

With regard to the monotremes, it has already been mentioned 
that there is no record of their past history beyond the limits of 
the Australian region. It can, however, scarcely be doubted that 
their ancestors came from the north with the primitive marsupials ; 
and if, as is not improbable, the Secondary and early Tertiary 
Multituberculata of the northern hemisphere are an allied type, 
there can be no doubt whatever as to this having been the case. 


That Notogaea, as typified by the Australian region, is entitled 
to form one of the three primary zoological divisions of the globe, 
the distinctness of its mammalian fauna from that of any other 
area, not only at the present day, but likewise during the Plisto- 
cene, and probably also the Pliocene epoch, amply demonstrates. 
The inclusion within the same realm of the Polynesian region, 
which evidently never had such a close connection with south- 
eastern Asia during the time that area was mainly populated with 
marsupials, is justified partly on account of its containing more 
or less similar types of birds, and partly by the practical absence 
of terrestrial mammals. On the other hand, the Austro-Malayan 
region, which is really a kind of zoological No-man's-land, is 
placed within the limits of the same great realm more as a matter 
of convenience than anything else, although it is undoubtedly 
sharply differentiated from the Oriental region by Wallace's line. 

In conclusion, a few lines may be devoted to showing that 
certain other groups indicate that the vertebrate fauna of Notogaea, 
as a whole, has had a northern origin. Among the lizards, the 
family of iguanas (IguanidcB], which in this realm occurs only in 
the Fiji and Friendly Islands, is represented in a fossil state in 
the Oligocene beds of France ; while the gigantic extinct monitor 
( Varanus priscus) of the Australian Plistocene appears to have its 
nearest ally in the smaller V. sivalensis of the Pliocene of northern 
India. The Notogseic Chelonians, which are confined to Australia 
and New Guinea, all belong to the side-necked group (Pleurodira) 
of the order, and are represented by the families Chelyidce. and 
Carettochelyidce, the latter containing only a single species from 
the Fly River. Now, although none of the Australian genera have 
been detected in the northern hemisphere, the side-necked chelo- 
nians, as shown in the next chapter, were abundantly represented 
there during the early Tertiary and Secondary epochs ; and it 
is a remarkable fact that an extinct genus believed to be allied 
to Carettochelys occurs in the Eocene of northern India. Although 
from their aquatic, and sometimes partially marine habits, the 
crocodiles are of less importance than some other groups from a 
distributional point of view, yet it is noteworthy that the single 
representative of that group ( Crocodilus porosus] inhabiting Noto- 
gaea (where it is found in North Australia, the Solomons, and 


Fiji) is spread over India, Ceylon, and the south of China ; while 
the absence of caimans and jacaras from Notogaea aifords, so far 
as it goes, an additional argument that any land connection which 
may have existed in Tertiary times between Australia and South 
America must either have been very transitory, or must have been 
situated in such latitudes that tropical forms could not have used 
it as a means of transit. 

Of more importance than all is the tuatera lizard (Sphenodon) of 
New Zealand the sole existing representative of the order 
Rhynchocephalia, since this curious creature is closely allied to 
the extinct Rhynchosanrus and Hyperodapedon of Triassic strata of 
the northern portion of the Old World. Finally, the Port Jackson 
shark (Cestracion philippi) belongs to a genus which was living in 
the seas of Europe during the Jurassic and Cretaceous periods ; 
and the sole living survivor of the. swarms of species of the 
Molluscan genus Trigonia inhabiting the same seas occurs in 
Australian waters. 



Extent and Characters Mammaliferous Deposits Monkeys Bats Insecti- 
vora Carnivores Ungulates Horses Litopterna Astrapotheria 
Toxodonts Pyrotheria Proboscideans Rodents Edentates Arma- 
dillos and Glyptodonts Sloths Anteaters- Ground-sloths Marsupials 
Cetaceans Early Distinction of the Neogaeic Fauna Early Separation 
of N. and S. America Incursion of Northern Mammals Distinctness of 
the existing Fauna Origin of the Santa Cruz Fauna Antarctica and the 
South American element in the Ethiopean Fauna Conclusion Sub- 

THE second primary zoological division of the globe may be 
known as Neogsea 1 , or the Neogseic realm. It 
characters"** includes only the Neotropical region. Comprising 
not only the whole of South and Central America, 
as well as the West Indian Islands, this area also embraces the 
lowlands lying on either side of the Mexican plateau the so- 
called tierras calientes thus running up in a fork-like manner to 
the lower extremity of North America. While, therefore, the 
greater part of this vast area is sharply delineated by its coast- 
boundary, to the north it has a kind of No-man's-land connecting 
it with the Sonoran region of Arctogaea, and, as will be shown 
later, through this transitional area there has been a certain 
amount of intermixture of the proper faunas of the Neotropical 
and Sonoran regions. Neogaea, as a whole, may be characterised 
as a country of extensive tropical forests or open grassy plains ; 
deserts occupying only a few scattered areas in the upper Argen- 
tine (Tucuman, etc.), and certain parts of the coasts of Chili 

1 This term was originally proposed by Dr Sclater to include the whole of 
the New World, but has been used by an anonymous writer (Appendix, No. 4) 
in the present sense. Dr Sclater 's term Dendroggea (Appendix, No. 27, 
p. 214) is open to considerable objection, as the greater part of Argentina is 


and Peru ; the whole of the rest of the area, with the exception of 
the higher regions of the Andes, being thus admirably adapted for 
the support of animal life. At least one half of the whole area is 
occupied by a dense tropical forest, attaining its richest develop- 
ment in the hot steamy tracts of Brazil and Paraguay, and being 
unequalled in extent in any other part of the globe. With a width 
of some three thousand miles from the Atlantic seaboard at Per- 
nambuco to the foot of the Andes, this forest extends north and 
south for nearly thirty degrees of latitude ; while not only does it 
clothe the lowlands and valleys, but extends high up the mountain- 
sides, as may be seen in the exquisitely lovely harbour of Rio de 
Janeiro, where the forest-vegetation commences immediately above 
the wash of the waves, and thence extends in one continuous leafy 
mass to the summits of mountain-ranges at an elevation of eight 
or nine thousand feet. In the northern part of the area open 
grass-lands, like the "campos" of Brazil and the savannas of 
Venezuela, alternate with the forest ; while in the neighbourhood 
of Buenos Aires the open pampas 1 forms one extensive sea of grass. 
The Andes, constituting the backbone of the country, run in one 
continuous chain from north to south on the Pacific seaboard, and 
present the usual varieties of climate and physical conditions 
common to other elevated mountain-ranges. Such climatic 
variations are, however, only an epitome of those met with in 
travelling from the northern to the southern extremity of the area ; 
the steamy valley of the Amazons having a tropical climate, whereas 
when we reach the southern point of Patagonia and Tierra del 
Fuego we are in the midst of snows and glaciers. To the hot 
forest-regions are restricted the monkeys, marmosets, sloths, ant- 
eaters, and tree-porcupines ; while the open plains of the south are 
tenanted by guanaco, deer, viscachas, and rheas. Moreover, in 
the forests, the variety of mammalian life, especially as regards the 
larger forms, is in marked contrast to its comparative paucity in 
the open plains ; not but that, till civilised man made his appear- 
ance on the scene, the number of individuals may have been 
nearly, if not quite as large in the latter area as in an equal extent 
of the former. 

1 Although in Spanish the term 'pampas' is plural, in English it seems 
preferable to use it as singular. 

L. 5 


At the present day the mammalian fauna of Neogaea is mark- 
edly distinct not only from that of Notogaea but likewise from that 
of the whole of the rest of the globe (Arctogsea), although the dis- 
tinction is now, owing to free communication with the north, 
much less marked than it was in Tertiary times, and it is accord- 
ingly essential to enter at once into the consideration of the 
extinct forms in order to show why this part of the world is 
entitled to rank as one of three primary zoological regions. 

There are several districts in South America where fossil 
Mammal- remains of mammals have been found ; most of these 
iferous being remarkable for the extraordinary profusion in 

which the bones occur. The first that may be men- 
tioned are the celebrated caves of Lagoa Santa, in the province of 
Minas Geraes, to the northward of Rio, which have yielded re- 
mains of a great variety of Plistocene genera and species, inclusive 
of those of man. Probably contemporaneous with these are the 
sand-dunes on the coast of Buenos Aires, which likewise contain 
human remains in association with those of extinct mammals ; 
while the so-called Pampean beds of the Argentine pampas are 
apparently somewhat older, although still pertaining to the Plisto- 
cene period. As these Pampean deposits are exceedingly rich in 
fossil mammals, they may be described in some detail. They 
form the great level tract of country extending southwards from 
the Rio de la Plata and the Parana to the Rio Colorado, south of 
Bahia Blanca, and westwards from the Atlantic seaboard about 
half the distance to the Andes; thus occupying some 200,000 
square miles of country. The pampas is an almost level grass- 
covered plain, intersected by water-courses, and penetrated near 
its margins by small mountain-ranges, while it is almost entirely 
barren of trees. It is composed of a- rich black alluvial mud, 
mingled with beds of sand, and underlain by, or in some places 
interstratified with layers of a hard white calcareous deposit 
known as tosca\ but in certain spots it contains beds of marine 
shells belonging to species still living in the adjacent seas. Except 
in those spots where the tosca comes to the surface, there is not a 
stone or a pebble to be seen in the whole deposit, and near 
Buenos Aires the formation has been bored to a depth of ninety 
feet without touching bottom. From its composition it is evident 


that the deposit has been carried down from the interior of the 
north by the Parana, Paraguay, and other tributaries of what is 
now the Rio de la Plata ; but since there is no splitting of the 
latter river at its estuary, it is evident that the formation cannot 
properly be called a delta. That it is mainly of freshwater origin 
seems evident not only from its intrinsic character, but likewise 
from the vast number of entire skeletons of mammals buried 
within it, since these creatures must certainly have lived very near 
to the places where their bones are now entombed. In the more 
southern part of its area the pampas is, however, probably to 
a large extent of estuarine origin ; and the presence of layers of 
marine shells in its uppermost horizon near Buenos Aires proves 
that at least a portion was submerged beneath the sea before its 
final upheaval. Whereas the Rio de la Plata now flows in a single 
channel in a south-easterly direction near the northern limit of the 
coast-portion of the pampas, it would seem probable that the 
Parana and Paraguay rivers may have originally continued their 
southerly course across the southern pampas, through which they 
may have flowed in a number of streams. Most likely the Pam- 
pean formation was laid down in a slowly subsiding area, in which 
the rate of deposition approximately counterbalanced the sinking, 
so that the greater part of it has been always land until the period 
of the great submergence. After the latter, the entire area was 
upheaved to a small degree above the sea-level, when the rivers 
assumed their present approximate courses. Whereas in certain 
localities the deposit is barren of mammalian remains, in other 
spots it appears absolutely crowded with them, and the number of 
entire skeletons that are entombed in it must doubtless be counted 
by thousands. 

Somewhat older than the Pampean is a mammaliferous deposit 
occurring on the coast near Bahia Blanca in a small hill known as 
Monte Hermoso ; and beds of approximately equivalent age occur 
in Catamarca at the foot of the Andes. Probably these beds should 
be regarded as of Pliocene age ; and it may be mentioned that 
equivalents of these deposits are met with in other parts of 
Argentina, while representatives of the Pampean occur in Pata- 
gonia, Chili, Bolivia, etc. Still older than the Monte Hermoso 
deposits are the Santa Cruz beds of Patagonia, occurring not only 



on the river of that name, but likewise further north in the Chubat 
district. These Santa Cruz beds are exceedingly rich in mamma- 
lian remains, which are stained of a deep black colour ; and while 
they contain many groups common to the Pampean formation, 
they lack those forms found in the latter which have an Arctogseic 
fades, thus indicating that we have reached an epoch when the 
fauna of South America was far more completely isolated from 
that of the rest of the world than is at present the case. By Dr 
Ameghino the Santa Cruz beds were at first correlated with the 
lower Eocene of Europe, although he subsequently admitted that 
they must occupy a somewhat higher position in that period 1 . 
From the nature of their entombed mammals 2 it is, however, 
certain that they must be still newer, and they cannot be regarded 
as older than lower Miocene. Indeed, they are underlain by the 
so-called Patagonian beds 3 , which are of marine origin, and 
contain numerous cetaceans, among which are whalebone whales 
(Mystacoceti). From equivalent beds in the Chubat district of 
Patagonia, a large number of such cetaceans have been described 
by the present writer 4 , and it is quite evident that the oldest age 
that can be assigned to these beds is upper Oligocene, seeing that 
in Europe whalebone whales are unknown till a considerably later 
epoch. Probably the freshwater beds containing the peculiar 
mammal alluded to below under the name of Pyrotherium are the 
freshwater equivalents of the Patagonian horizon 5 . 

Additional evidence in support of the comparatively late age 
of the Patagonian beds is afforded by the researches of Prof. Cope 6 
on the cetaceans of the Miocene (or Upper Oligocene?) of the 
United States. From these beds have been obtained remains of 
Hypocetus (Paracetus) a genus elsewhere known only from the 
Patagonian beds together with Cetotherium (also occurring in the 
latter deposits), Balcenoptera, and a species of Balcena identified 
with one from the European Pliocene ; the North and South 
American species of Hypocetus being closely allied. 

1 Bol. Ac, Cordoba, Vol. xin. p. 260 (1894). 

2 Remains of the existing genus Dasypus occur in these beds. 

3 Ameghino, loc. cit. 

4 Ann. Mus. La Plata, Pal. Argent. Pt. II. (1893). 

5 Ameghino, op. cit. p. 262. 

6 Prof. Amcr. Phil. Soc. Vol. XXXIV. pp. 135 155 (1895). 


With these preliminary considerations, we pass to a critical 
examination of the recent and fossil mammalian fauna of Neogaea, 
which will show how intimately investigations of this nature are 
connected with the present and past configuration of the land- 
areas of the globe. 

Like as are many of them superficially to their cousins of the 
Old World, the monkeys of the New World, which 

,, , , .... . - Monkeys. 

are now confined to the tropical forest-regions of 
the Neogaeic realm, are structurally quite different to the former ; 
and this circumstance, coupled with their isolated distribution, in- 
dicates that their relationship is, at most, but a very distant one. 
Indeed it is not improbable that the Old and New World monkeys 
may have originated independently from the group of lemurs, which 
were formerly very widely distributed over Arctogaea. Of the 
two families of Neotropical monkeys, the first is represented by 
the beautiful little marmosets, constituting the family Hapalida. 
Although having the same number (32) of teeth as the Old World 
monkeys, the marmosets differ in that the number of premolars on 
each side of both jaws is three, and that of the molars two ; these 
numbers being transposed in the other group. The marmosets 
are further distinguished by the broad septum between the nostrils, 
the absence both of pouches in the cheeks and of callosities on 
the buttocks, and the want of any prehensile power in the tail ; in 
addition to which it may be noticed that the thumb is not op- 
posable to the other fingers, while all the digits, with the 
exception of the first on the hind foot, are provided with long 
claws. None of these animals are known from the Santa Cruz 


deposits. In the second family or Cebidce, which includes much 
larger species than the diminutive marmosets, the total number of 


teeth is 36, owing to the retention of the three pairs of molars 
found in the Old World group ; these monkeys being further dis- 
tinguished from the marmosets by the presence of nails on all the 
fingers, and by the tail being frequently prehensile. The Santa 
Cruz beds have yielded remains of monkeys (JFTomunculus) refer- 
able to this family, showing that they belong to the original South 
American fauna, but beyond this nothing is known as to the 
palseontological history or origin of the group. Lemurs, both in 
the past and the present, are quite unknown in the realm under 

Although bats might be thought of comparatively little im- 
portance from a distributional point of view, yet the 

Bats. :_ 

Neogseic realm presents some very remarkable 
features in this respect. In the first place the two Old World 
families of fruit-bats (Pteropodtdce) and horse-shoe bats (Rhinolo- 
phidce] are entirely wanting, whereas the great family of vampire- 
bats (Phyllostomatida) is mainly restricted to it, although a few 
representatives straggle northwards along the Pacific coast of 
North America. The family of Emballonuridtz is also more 
strongly represented here than in any other part of the world. 
Probably on account of their small size, there is no palaeonto- 
logical record of the South American bats from the earlier 
Tertiary deposits. 

The Insectivora are almost unrepresented in the continental 
portion of the realm, although a shrew (Sorex) 

Insectivora. ' . 

reaches Guatemala and Costa Rica, and a member 
of the allied N. American genus Blarina is also found in the 
last-named country; both these being doubtless very recent 
immigrants from the north. Very remarkable is the occurrence in 
the West Indian Islands of the two species of Solenodon, consti- 
tuting a distinct family (Solenodontidce) by themselves. These 
have generally been considered as very nearly allied to the tenrecs 
(Centetidce) of Madagascar, but Mr O. Thomas 1 is of opinion 
that the relationship is not really very close, the similarity in the 
structure of their molars being merely a generalised character. 
Both, however, probably indicate an ancient group, which has 

1 Proc. Zool. Soc. 1892, p. 500. 


migrated from the higher latitudes of the northern hemisphere to 
find a refuge in these two far distant localities. It may be 
mentioned that the Solenodontidce and Centetidce, together with the 
Potamogalida of western, and the Chrysochloridcz, or golden moles, 
of southern Africa, constitute a section of the order distinguished 
from all the other forms by having the cusps on their upper molar 
teeth arranged in the form of the letter V, instead of in that of a 
W ; and it will not fail to be noticed that the whole group is now 
exclusively a southern one. Whether it is represented in the Santa 
Cruz beds is not quite certain, although one lower jaw has been 
referred to it by Dr Ameghino 1 under the name of Necrolestes. 
Be this as it may, it is clear that, with the exception of the afore- 
said shrews, insectivores with W-shaped molars are entirely want- 
ing in the realm ; and that such V-shaped types as still exist, or 
formerly occurred, evidently indicate an ancient northern group. 
Other instances of the survival in South America and Madagascar 
of allied forms will be noticed in the sequel, and admit of a 
somewhat similar explanation. 

Although fairly well represented at the present day, the carni- 
vores of this realm have but few absolutely charac- 

. Carnivora. 

tenstic forms, and as no remains of the true Carni- 
vora occur in the Santa Cruz beds, the whole of them may be 
regarded as comparatively late immigrants from the north. The 
civet family ( Vsvtrrida) and hyaenas (Hyanidcz) are totally wanting 
at all epochs, as indeed they are throughout the whole of the New 
World; but the weasel tribe (Mustelidce) have a comparatively 
small number of representatives. Cats (Felidcz) on the other 
hand are numerous, although several species, and more especially 
the puma (Felts concolor), range to a greater or less extent into 
North America. Such a cosmopolitan genus is, however, of no 
importance whatever from a distributional point of view; and 
much the same may be said of the extinct sabre-tooths (Machar- 
odus), which were distributed in Tertiary times throughout the 
entire northern hemisphere. Unknown in the deposits of Monte 
Hermoso and Santa Cruz, this genus is represented by a gigantic 
species in the Pampean, which undoubtedly reached the country 


1 Bol. Ac. Cordoba, Vol. xm. p. 364 (1894). 


from the north in company with the other late immigrants. The 
dog tribe (Canidcs) likewise comes under the category of cosmo- 
politan groups, and has numerous South American species, 
although only one from the Monte Hermoso beds dates earlier 
than the Pampean. It may be mentioned that while true wolves 1 
are absent from this realm, all the continental living members of 
the genus Cants found in it form a group by themselves, quite 
distinct from the true foxes of other regions. Very remarkable is the 
occurrence in the Pampean of a large species (C. moreni) perfectly 
distinct from all those now inhabiting the country, and presenting 
some curious approximations in the structure of the skull to 
domestic dogs. Peculiar to the realm is the bush-dog (Icticyori), 
of Guiana and Brazil, a small short-haired and short-legged 
species differing from all other members of the family by the 
small size and reduced number of the molar teeth. Its remains 
occur in the Brazilian caves, but are unknown from earlier 
deposits ; and it may thus be regarded as a comparatively late 
immigrant from the north, which perhaps developed its special 
characters after its arrival in the country. Of the Ursidce. there is 
but a single existing species in Neogaea, namely the spectacled 
bear ( Ursus ornatus) of the highlands of Chili and Peru ; but 
there occur in the Pampean formation of the Argentine remains of 
an allied extinct genus known as Arctotherium, another species of 
the same genus being recorded from the superficial deposits of 
California. Far more characteristic of the realm are the raccoons 
and coatis (Procyonid<z\ although several of these are common to 
North America. Till recently it was considered that this family 
was entirely restricted to the New World, but the Oriental cat-bear 
or panda (Ailurus) is now included ; and since fossil remains of 
the latter genus have been discovered in the Pliocene of England, 
while those of raccoons and of the extinct genus Leptarctus occur 
in that of the United States, it is practically certain that the group 
was formerly widely distributed over the northern hemisphere. 
Among this family the raccoons (Procyon) extend over most parts 
of both North and South America ; the coatis (Nasua} range from 
Mexico to Paraguay; the kinkajou (Cercoleptes) is found from 

1 The Falkland Island wolf (Cants antarcticus) forms a remarkable ex- 


Mexico to Peru and Brazil; while the two species of the genus 
Bassariscus are inhabitants of the southern United States, Mexico, 
and Central America. From the Tertiaries of Catamarca and 
Parana there has been described the extinct genus Cyonasua, 
differing from the c.oatis in the form of its teeth ; this genus 
showing that the family had obtained an entrance into South 
America as early as the Pliocene period, although it is quite 
unknown in the Santa Cruz epoch. In the Mustelidce. where all 
reference to the cosmopolitan otters, of which there is one Brazilian 
species, may be omitted the southern skunks (Conepatus], which 
have mostly but thirty-two teeth, are now practically character- 
istic of this realm although the common species ranges into 
Texas and their remains occur fossil in the caverns of Brazil. 
The true skunks (Mephitis), on the other hand, which have 
thirty-four teeth, are North American, although one species 
ranges as far south as Guatemala ; and since the whole group is 
unknown in the earlier Tertiary deposits of the Argentine, it may 
likewise be considered a recent immigrant from the north. The 
same is true of the genus (Galictis) now represented by the 
South American grison and tayra, since remains of this group of 
mustelines occur in the Plistocene of the United States, as well 
as in the Brazilian caves. 

Of far more importance than either of the preceding orders 
are the hoofed mammals or ungulates, since here we 
meet with certainly three, and probably four extinct 
subordinal groups exclusively confined to this realm, while a large 
number of the existing families are unrepresented even in the 
Pampean formation. Whereas this order is entirely absent from the 
West India Islands, South America at the present day is singularly 
poor in ungulates. The only existing forms are the peccaries 
(Dicotyles), which are peculiar to the New World; certain deer 
belonging to a genus ( Cariacus) which is likewise confined to the 
western hemisphere, and a Chilian form constituting a genus 
(Pudud] by itself; the exclusively South American guanacos and 
vicunas, which, together with their domesticated allies constitute 
the genus Lama; and tapirs (Tapirus). The first three of these 
belonging to the Artiodactyla, while the last alone represents the 
Perissodactyla, or odd-toed division of the order. At all periods 


of its history true pigs (Sus), hippopotami (Hippopotamus], camels 
(Came/us), chevrotains (Tragulida\ giraffes (Giraffida), true deer 
(Cervus), antelopes, sheep, goats, and oxen, together with a 
number of extinct forms connecting the ruminants with the pigs, 
have been conspicuous for their absence. The same is true, 
among the perissodactyles, of the rhinoceroses (Rhinocerotidce) 
and the extinct palseotheres (Pal&otheriida) and lophiodons 
(Lophiodontida) of Europe, and the uintatheres {Uintatheriida) 
and titanotheres ( Titanotheriidce] of the United States ; while the 
true elephants (Elephas] among the Proboscidea have likewise 
been always absent. 

Of the existing South American ungulates the peccaries 
belong to a family (DicotylidcB) which is abundantly represented in 
the Tertiary formations of the United States ; while in those of 
Europe and Asia there occur allied forms apparently connecting 
the peccaries with the true pigs. On the other hand, in South 
America their remains occur only in the superficial and cavern- 
deposits, so that there can be no doubt as to their late intrusion 
into the country from the north. The vicunas and guanacos are 
the western representatives of a family (Cameltda) whose other 
members are Asiatic and African, and of which the past history 
seems to have been very similar to that of the last group. Fossil 
camels occur in the Pliocene of India, while a host of extinct 
genera more or less closely allied to the living South American 
forms occur in the Tertiaries of the United States ; and since in 
Argentina and Brazil remains of Lama and the related types 
occur only in the Monte Hermoso, Pampean, and cavern de- 
posits, there can be no hesitation in regarding the group as com- 
paratively recently immigrant into the country. The deer of the 
genus Cariacus are likewise only known in South America from 
the Pampean and some other of the later Tertiaries, as well as the 
Brazilian caves, while in the Pliocene of the United States they 
appear to be represented by the ancestral Blastomeryx; and these 
accordingly come under the category of intruders from the north. 
As regards the tapirs, the genus and family now presents a 
remarkable instance of discontinuous distribution, one species 
being confined to the Malayan countries, while all the others are 
South American. Whereas in the realm under consideration re- 


mains of these animals occur only in the superficial deposits, they 
are met with abundantly in the Miocene and Pliocene deposits 
of Europe and Asia, as well as in the United States; and it is 
accordingly clear that the group was once widely distributed over 
the northern hemisphere, whence its surviving members have 
wandered southwards to Malaysia in the east and South America 
in the west. 

Till introduced by the early Spanish settlers, horses, which are 
now so abundant in the pampas, were totally un- H orses 
known in South America in a living state, although 
their fossilised remains occur commonly in the Pampean, as well 
as in the somewhat older deposits of Parana, and Monte Hermoso. 
They are, however, entirely unknown in the Santa Cruz beds. 
Some of these fossil Argentine horses belong to the typical genus 
Equus\ while others, on account of the simpler structure of their 
molar teeth and the great length of the slits in the skull beneath 
the nasal bones, are referred to a separate genus, under the name 
of Hippidium. A third genus (Onohippidimn) is distinguished 
from the latter by a large lachrymal depression in the bones of the 
sides of the face, corresponding to the so-called larmier of the deer. 
Fossil species of Equus occur in the Plistocene deposits of the 
whole northern hemisphere, while those of the United States yield 
remains of a genus (Pliohippus] nearly allied to the South American 
Hippidium; and as both these genera are the descendants of 
extinct forms whose remains occur in the older Tertiaries of the 
same hemisphere, the extinct South American horses must likewise 
be classed with the groups that have entered the country from the 
north. Why these Plistocene South American Equida became 
extinct in a country so admirably suited to their existence as 
Argentina, is a question to which it is impossible to find a 
satisfactory answer. With the horses we reach the last of the 
Neogaeic representatives of the more typical ungulates, that is to 
say the Artiodactyla and Perissodactyla ; and we pass on to the 
other extinct subordinal groups, at least three of which, as already 
said, are peculiar to it. 

From the preceding paragraphs it will be apparent that, if we 
except the deer, horses, and the guanacos and their 
allies, South America, so far as the Artiodactyla and 




Perissodactyla are concerned, was exceedingly poorly off for 
ungulates during the Plistocene epoch. Nevertheless, the 
country was very rich in hoofed mammals, not only during the 
Pampean, but likewise during the Santa Crucian epoch, and in 
this respect it was quite as peculiar as it is in its edentates. It is 
not a little remarkable that three of the extinct subordinal groups 
of the order which are confined to this realm exhibit a more or 
less decided approximation, more especially in the structure of 
their molar teeth, to the earlier northern Tertiary representatives 
of the Perissodactyla. Hence it is probable that all the four 
suborders in question have originated from a common ancestral 
stock, although apparently before the perissodactyles were differen- 
tiated from an earlier group known as the Condylarthra. The 
date when the ancestors of the South American forms reached 
their present home is, however, enveloped in mystery, and 
although it is fairly certain that such ancestors had a northern 
origin, yet it is highly improbable that they entered South America 
from that direction. 

The first of the three extinct subordinal groups in question, 
for which the name of Litopterna has been proposed, is the one 
showing the nearest parallelism with the Perissodactyla, and is 
typified by the genus Macrauchenia, of which the skeleton is 
figured in the accompanying illustration. In this group the cheek- 
teeth approximate in general structure to those of the well-known 
European Oligocene genus Palaotherium, although in the typical 
genus Macrauchenia they have been so modified as to render the 
resemblance obscure. An essential characteristic of the upper molar 
teeth (fig. 13) is to be found in the presence of two distinct lobes 
to their outer walls. The toes of both fore and hind feet are 
elongated, and constructed on the same general plan as those of 
the Perissodactyla, never exceeding three in number on each foot, 
and the middle one being symmetrical in itself. Moreover the 
astragalus of the tarsus or heel-joint resembles the corresponding 
bone of the latter group in having a deep pulley-like groove on its 
upper surface for the articulation of the tibia, although inferiorly 
it is unlike. The calcaneum, or heel-bone, on the other hand, 
resembles that of the Artiodactyla in bearing a small facet for the 
articulation of the fibula, or small bone of the leg. A more 




(To exhibit Perissodactyle type of structure.} 


(Showing linear and alternating arrangements of carpal bones.} 


important difference, from both the Perissodactyla and Artio- 
dactyla, occurs in the structure of the carpus (wrist) and tarsus 
(ankle), in both of which the two rows of small component bones 
are arranged in vertical series one above another, instead of 
overlapping and interlocking; this so-called linear arrangement 
being a more primitive type than the interlocking or alternating 
arrangement characterising the two existing suborders. The 
vertebrae of the neck are elongated, with flat terminal faces, and 
have the vertebral artery piercing the sides of the neural arch 
in a manner elsewhere found only in the camel family and the 
great anteater. In the femur, or thigh-bone, the projection known 
as the third trochanter is much less developed than in the Perisso- 
dactyla. All the members of the group were long-limbed, long- 
necked, and slenderly built animals, Macrauchenia itself being 
as large as a camel, although the genera from the Santa Cruz 
beds of Patagonia were represented by species of much smaller 

Of the typical genus Macrauchenia fossil remains occur not 
only in the Pampean formation, but likewise in the superficial 
deposits of both Patagonia and Bolivia. The most curious feature 
about this remarkable animal is the position in the skull of the 
aperture for the nostrils, which instead of being situated at the 
extremity of the muzzle, is placed in the middle of the forehead, 
between the eyes. Otherwise the skull is not unlike that of a 
horse in general contour. The teeth, which are forty-four in 
number, and form a continuous uninterrupted series, are a special- 
ised modification of the type of those of the undermentioned 
genus ; their crowns being taller, with a more complicated ar- 
rangement of folds. In nearly all points of its structure, especially 
in the number of the teeth, and the absence of large tusks, as well 
as in the structure of the wrist- and ankle-joints, Macrauchenia 
is a very primitive kind of animal. In the Santa Cruz beds of 
Patagonia the family to which this genus belongs is represented 
by several smaller animals, such as those named Oxyodonto- 
therium, in which the aperture for the nostrils occupied a more 
normal position in the skull, and the crowns of the molar teeth 
were shorter and of a more simple structure. The crown-surfaces 
of worn upper molar teeth from the right side of the jaw of both 




genera are shown in the accompanying figures (13 and 14); the 
letters indicating the corresponding elements in each. 



Oxyodontotherium . 

The second family, or Proterotheriidce, is confined to the Monte 
Hermoso, Parana, and Santa Cruz beds of Patagonia, in the last 
of which it is represented by the genera Proterotherium and Dia- 
diaphorus. In these animals, none of which much exceeded a 
sheep in size, the upper molar teeth are much more like those of 
the European Palceotherium ; and, in place of forming a continuous 
series, the teeth are interrupted; one pair, both in the upper and 
lower jaw, being developed into tusks. In structure the feet pre- 
sented a general resemblance to those of the extinct three-toed 
horses (Hipparion) of the northern hemisphere, but in some in- 
stances the toes were reduced to a single one, thus showing a 





curious parallelism in the development of this group to that of the 

A second sub-order of ungulates, entirely confined to the Santa 
Cruz and Patagonian beds of Patagonia, is repre- 
sented by the two generic types known as Astrapo- 
therium and Homalodontotherium, each of which constitutes a 
family by itself. Rivalling the rhinoceros in bulk, both of these 
extraordinary creatures differ from the last group by the structure 
of their molar teeth, which approximate to those of the Rhino- 
cerotidce, those of the upper jaw having a continuous outer wall, not 
divided into lobes. In the ankle-joint the astragalus differs from 
that of the preceding group in having its upper surface flat, thus 
resembling that of the elephants. In both the wrist and ankle 


the component bones were arranged on the linear plan, and not 
improbably each foot had five toes. The vertebrae of the neck 
were short, with flat terminal faces ; and in correlation with this 
shortness of the neck, the limbs and feet were more or less 
abbreviated. In the first-named of the two genera each jaw was 
provided with an enormous pair of tusks, wearing obliquely against 
L. 6 


one another like those of a wild boar ; and although there were 
no small teeth in the upper jaw, the lower jaw carried three pairs of 
spatulate incisors quite unlike those of any other known animal. 
On the other hand, Homalodontotherium takes its name from having 
its forty-four teeth arranged in a continuous even series, unbroken 
either by tusks or by gaps. Whereas the molar teeth of Astrapother- 
ium present a marked resemblance to those of the rhinoceroses, 
it is noteworthy that those of the allied genus make a certain 
approximation to the corresponding teeth of an extinct ungulate 
from the European Oligocene known as Cadurcotherium, ap- 
parently more or less closely allied to the rhinoceroses. It is, 
however, quite improbable that it is to these allied forms we 
have to look for the origin of the peculiar South American 
ungulates, seeing that they must have branched off from the primi- 
tive stock at an earlier stage. 

The third extinct subordinal group, or Toxodontia, takes its 
name from a gigantic species from the Pampean 


formation to which Owen, on account of the pecu- 
liarly curved form of the molar teeth, gave the name of Toxodon. 
Rivalling the largest existing rhinoceros in point of size, the 
Toxodon has at first sight very much the general appearance of 
one of those animals, having a massive skull, short limbs and 
neck, and three-toed feet. The middle toe is, however, scarcely 
larger than the lateral ones ; and while the bones of the wrist are 
arranged in the alternating manner, those of the tarsus are placed 
in a linear series, and the astragalus has a nearly flat, instead of a 
grooved, superior surface ; the terminal faces of the vertebrae of 
the neck being also flat, instead of articulating together by ball- 
and-socket joints. Omitting mention of other more or less strongly 
pronounced peculiarities in the structure of the limbs, attention 
may be specially directed to the teeth, which differ from those of 
all existing ungulates in that the whole of them grow continuously 
throughout the life of their owner, without ever forming roots. 
The front or incisor teeth are chisel-like, and thus resemble in 
form, although not in number, those of a beaver or rabbit; and, 
indeed, in the general conformation of the teeth, as well as in the 
non-development of roots, the whole dentition of this animal 
presents a most curious similarity to that of the rodents. Some 



idea of the huge size of these animals may be gathered from the 
fact that the skull may measure as much as a yard in length. 
Whereas Toxodon itself is confined to the Pampean beds, it is 
represented in the somewhat older deposits of Monte Hermoso 
and Catamarca by allied forms known as Toxodontotherium and 

FIG. 1 6. SKULL OF Nesodon. Much reduced. 

In the Santa Cruz beds of Patagonia the place of the fore- 
going is taken by the allied genus Nesodon, likewise belonging to 
the same family Toxodontidce. In these animals, of which the 
skull is shown in the figure, the molars retain marked resem- 
blances to those of the rhinoceroses, which have been lost in the 
more specialised Toxodon ; these teeth growing for a considerable 
period, yet late in life developing roots in the ordinary manner. 
The front teeth are very peculiar, the canines remaining small 
throughout life, but the second pair of incisors in the upper, and 
the third in the lower jaw growing beyond the others to form large 
tusks in the adult, and never developing roots. In their rooted 
molars the Nesodons depart less widely from the primitive Peris- 


8 4 



sodactyle type than do their more specialised descendants the 
Toxodons, although there are no forms known from other parts of 
the world to which they present any direct relationship or show any 
marked resemblance. 

A second family ( Typotheriidce) of the sub-order is represented 
in the Pampean by the typical genus Typotherium and in the 
Santa Cruz, or Patagonian, beds by the allied Trachytherus. Both 
these have a dentition still more like that of the rodents ; the 
front, or incisor teeth being reduced to a single chisel -like pair in 



each jaw, while the molars are narrow, rootless, and of compara- 
tively simple structure and relatively small size. Moreover, in- 
stead of hoofs, these highly modified ungulates appear, like many 


members of the rodent order, to have had their toes protected 
by nails. Like as are these animals to rodents, the resemblance 
to that group is carried to a still greater extent in certain small 
forms from the Santa Cruz beds which have been named Pachy- 
ruchus and Hegetot her turn ; these constituting a third family, the 
Pachyruchida. From the features referred to, the reader might 
be disposed to consider that there is some direct genetic affinity 
between rodents and the group under consideration, but this is 
clearly not the case, such resemblances as exist being solely the 
result of that parallelism in development which appears to have 
been such an important factor in the evolution of mammals. 

It may be well to mention here that in a recent paper 1 Dr 
Noack suggests an intimate affinity between the toxodonts and 
the existing hyraces (Hyracoidea) of Africa and Syria; adding 
that the presumed affinity affords evidence of a land-connection 
between Africa and South America. The two groups are, however, 
essentially different in regard to the structure of the fore-foot, in 
which the bones of the carpus or wrist are arranged, as already 
shown, in the alternating manner among the toxodonts, whereas 
in the Hyracoidea they form a linear series. Moreover, the molar 
teeth of the two groups are markedly distinct, although in both 
they approximate to the perissodactyle type. Still there is a pos- 
sibility that the Hyracoidea may represent a less specialised branch 
which has originated from the primitive toxodont stock, but has 
retained the linear type of carpus ; and if this really be the case, 
it would afford very strong confirmation of the view that South 
America received its earliest ungulates by way of Africa and the 
Antarctic Continent. 

A remarkable ungulate from the Patagonian beds, with molar 
teeth very like those of the extinct European genus 
Dinotherium, and bearing a pair of large tusks in 
the lower jaw at least, has been tentatively assigned by the present 
writer to the Proboscidea ; but judging from the distinction of the 
other ungulates of the older Argentine Tertiaries from those of 
the rest of the world, it is more probable that it represents a 
sub-order by itself. Pyrotherium, as the animal is called, was 

1 ZooLJahrb.-AbtheilfurSystemat. vol. vil. pp. 540 542. 


associated with nesodons, astrapotheres, and homalodontotheres, 
which have been assigned by Dr Ameghino to genera distinct 
from those of the Santa Cruz beds. But from my own observations 
on a series of remains of these animals in the La Plata Museum 
obtained in association with those of Pyrotherium, they appear to 
be genetically inseparable from their Santa Crucian represen- 

Now represented only by the living Indian and African 
elephants the Proboscidea were formerly a some- 
deans b SC1 what extensive subordinal group of the ungulates, 

easily characterised by their peculiar molar teeth 
and tusks (the latter of which may be present either in the upper 
or lower jaw alone, or in both) ; their five-toed feet, in which the 
bones of the ankle and wrist are arranged on the linear plan, and 
the astragalus has a flat upper surface; and the presence of a 
trunk. While true elephants (Elephas) are totally unknown in 
South America, the genus Mastodon, distinguished by the low 
crowns and simpler structure of the molar teeth, is represented by 
two species in the Pampean of Buenos Aires, and is also stated to 
occur in the Monte Hermoso beds ; but (assuming the distinct- 
ness of Pyrotheriuni) the sub-order is unknown in the older deposits. 
Although at the present day Neogaea contains no existing fami- 
lies of either carnivores or ungulates absolutely pecu- 

Rodents. .. ....... 3 * 

liar to it, the case is widely different with regard to 
the rodents, or gnawing mammals. Existing rodents are divided 
into seventeen families, arranged under four sectional groups ; 
nine out of these seventeen families occurring in the Neogaeic 
realm, among which four are absolutely peculiar to it. A fifth 
(Octodontida) is mainly South American and West Indian although 
possessing a few representatives in Africa south of the Sahara, and 
a sixth (Hystricida) has two genera which are practically only 
South American. The significance of these facts will be apparent 
when it is stated that of the other zoological regions of the globe 
only two have any families of the order peculiar to them, and 
neither of these has more than two such families. In the Ethio- 
pian region, for instance, the only peculiar family is that of the 
African flying-squirrels (Anomalurid(E\ represented by two genera ; 
while the western division of the Holarctic region has the sewel- 


lels (Haplodontida), with one genus ; and the Sonoran region the 
pouched rats (Geomyidce) with two. It is further highly significant 
that all of these families which are mainly or chiefly Neotropical 
belong to one division of the order, the Hystricomorpha, and 
that certain of them are represented either in the Santa Cruz beds 
of Patagonia, or the Parana beds, where the whole of the other 
sections and families are totally unknown. This, however, is by 
no means all, for throughout the Tertiaries of North America 
below the Pliocene there are no Hystricomorpha at all, and at the 
present day there is but a single species the Canadian porcupine 
(Erethizott) inhabiting the whole of that country. On the other 
hand, both at the present day and in the Tertiaries, North America 
abounds in Sciuromorpha and Myomorpha. These facts clearly 
point to the existence of an impassable barrier between North 
and South America during a large portion of the Tertiary 
period. Moreover, even at the present day the Sciuromorpha 
are scarcely represented at all in Neogaea, while the Myomorpha 
are not very numerous. 

Of the squirrel-like rodents, constituting the section Sciuro- 
morpha, and including the four existing families of the African 
flying-squirrels (Anomalurid(z\ the squirrels and marmots (Sciur- 
idcK], the sewellels (Haplodontida), and the beavers (Castoridce), 
as well as the extinct American Castoroidida, the only living 
Neogaeic representatives are certain species of squirrels, none x>f 
which range south of Paraguay. The extinct Castoroididcz include 
large beaver-like rodents with complex molars like those of the 
viscacha, and are represented by the typical Castoroides from the 
Plistocene of the United States, and likewise by Amblyrhiza 
(Loxomylus) from caves in the West Indies, and, it is said, also 
from the later Tertiaries of Argentina. This family is accordingly 
a northern type. 

The second or murine group of rodents (Myomorpha) contains 
five families, namely the dormice (Myoxidce), the jumping-mice 
and jerboas (Dipodidce), the mice and rats (Muridce), the mole-rats 
(Spalarida), and the American pouched rats (Geomytda). Out of 
all these, practically the only one represented in the realm is the 
cosmopolitan Muridce, although among the Geomyidce the genus 
Heteromys enters the transitional Mexican sub-region. In the 


Muridce, however, as in North America, the true rats and mice 
(Mus) of the Old World are entirely wanting, except through 
human introduction ; their place being taken by the white-footed 
mice (Sttomys) common to the whole of the New World, and 
nearly related to the European hamsters. Being essentially a 
northern type, they are certainly recent immigrants into South 
America from the north ; and the same is doubtless true of certain 
genera of the family now peculiar to this realm, such as the fish- 
eating rats (Ichthyomys\ of Peru and Venezuela, the groove-toothed 
mice (Rhithrodori), one of which ranges as far south as Tierra del 
Fuego, and the Brazilian genus Holochilus, which is another form 
allied to the hamsters. Although the cotton-rat (Sigmodon) occurs 
in Central America, and occasionally wanders still further south, 
the whole tribe of voles and their kindred are absent from the 
entire realm. 

Represented by six families, all of which occur within its limits, 
the porcupine-like group, or Hystricomorpha, may be regarded as 
the characteristic rodents of Neogsea. From both the preceding 
sections of the order the members of this section may be readily 
distinguished by the angle (or hinder inferior projection) of the 
lower jaw taking its origin from a prominent ridge running along 
the side of the jaw, instead of from the inferior edge of the socket 
for the incisor teeth. Of the families confined to this realm, that 
of the cavies (Caviidcz], includes heavily-built rodents, with four 
front and three hind toes, rudimental or short tails, and the 
cheek-teeth divided by transverse folds of enamel into a number 
of thin parallel plates. The genera of this family include the true 
cavies (Cavia] so well known through the domestic guinea-pig 
all of which are small short-limbed forms ; the larger and taller 
Patagonian cavy (Dolichotis}; and the carpincho, or capivara 
(Hydrocha>rus\ which is the largest living member of the order, 
and is characterised by the large number of plates going to form 
the last molar tooth in each jaw. Although they do not appear to 
have been recorded from the Santa Cruz beds, remains of members 
of this family occur in the Parana 1 horizon, and also in that of 

1 There is some confusion with regard to the age of the Parana beds. They 
are overlain by marine strata which have been identified with the Patago- 
nian ; but it is more probable that they are newer than the Santa Cruz beds, 


Monte Hermoso. Of the former Plexochairus differs from Hydro- 
chorus only by the somewhat simpler structure of the last molar, 
Hydrochcerus itself occurring in the Monte Hermoso beds. Other 
forms from the Parana stage are Eucardiodon and Cardiotherium, 
apparently more nearly allied to Cavia. Here it should be 
mentioned that certain European Oligocene genera (Issiodoromys 
and Nesocerodori) have the molars so like those of the cavies that 
they are regarded by Dr Schlosser as nearly allied' to the family. 
By Prof. Zittel they are, however, included in the extinct family 
Theridomyidce, which is classed with the dormice and certain 
other families in a group regarded as intermediate between the 
Sciuromorpha and Hystricomorpha. If, as would seem probable, 
they are really allied to the latter, they are of the utmost import- 
ance as indicating a connection between the middle Tertiary 
rodent fauna of Europe with that of South America 1 . 

Nearly allied to the cavies are the agutis (Dasyprocta) and 
pacas (C&logenys), collectively constituting the family Dasy- 
proctidcE, and differing from the former in that the folds of enamel 
merely form notches on the sides of the crowns of the cheek-teeth. 
In a fossil state the family seems to be only known by remains of 
the existing genera from the Brazilian caves. The third peculiar 
family (Dinomyidce) is known merely by a single specimen from 
Peru. The only other family exclusively confined to the realm is 
that of the Lagostomatidcz (Chinchillidce], which includes not only 
the true chinchillas (Eriomys) and Cuvier's chinchilla (Lagidium), 
but likewise the viscacha (Lagostomus] of the Argentine pampas. 
All the members of this family have long bushy tails, elongated 
hind limbs, and the cheek-teeth divided by complete transverse 
folds of enamel into thin plates. Lagostomus occurs fossil not 
only in the Pampean, but likewise in the older Tertiaries of 
Parana; while it may be doubted whether Pliolagostomus and 
Prolagostomus of the Santa Cruz beds are really entitled to generic 
distinction. Other allied forms from the latter deposits are 

although their lowest portion is older than the Monte Hermoso stage. (See 
Ameghino, Bull. Ac. Cordoba, Vol. xiil. pp. 260, 261, 1894.) They may be 
partly made up of the remains of pre-existing beds. 

1 Dr Schlosser writes me to the effect that he is fully assured these forms 
are the ancestors of the Caviidce. 


described under the names of Perimys and Sphodromys, and thus 
indicate that the family is essentially South American. The 
largest known rodent is the extinct Megamys, from the Parana 
beds ; the typical species of the genus being described as equal in 
size to an ox. The porcupines (Hystricida), which form a 
practically cosmopolitan family, sufficiently distinguished by their 
spiny covering, are represented in South America by the two 
arboreal genera Synetheres and Chcztomys, differing from all their 
allies by their prehensile tails, although otherwise related to the 
North American Erethizon, with which they form a separate sub- 
family. Of Chcetomys an extinct representative occurs in the 
cavern-deposits of Brazil; and in the Santa Cruz beds the family 
is represented by apparently extinct generic types described under 
the names of Stiromys, Acaremys, and Sciamys. Since fossil 
remains of Hystrix date from the European Miocene or Oligocene, 
there is distinct evidence of a connection between the early South 
American rodents and those of the Old World ; while as Erethi- 
zon is first known from the Plistocene of North America, it may 
probably be regarded as a late immigrant into that country from 
the south. 

The largest of all the families of the Hystricomorpha is that of 
the Octodontidal, in which out of a total of nineteen genera 
upwards of fifteen belong to the realm under consideration, while 
the remaining four are African and mainly Ethiopian. In addition 
to other features, the rodents of this family have the crowns of 
the cheek-teeth marked by infoldings of enamel from both sides; 
there are usually five toes to each foot ; and the general form is 
more or less rat-like. Of the Neogaeic forms, the typical genus 
Octodon is represented by the degu of Chili and Peru, which is a 
large rat -like animal, with a brush-tipped tail; other species 
occurring in Bolivia. The latter country is likewise the home of 
the allied genus Habrocoma, the members of which vie with the 
chinchillas in the delicate softness of their fur. Nearly related are 
the burrowing South American tuco-tucos (Ctenomys\ characterised 

1 By some the family is divided into three; viz. Capromyidce, with the 
West Indian Capromys, the S. American Myopotamus, and. the African 
Triaulacodus ; Ctenodactylidcz, including the remaining African forms; and 
Octodontidce, comprising all the other American types. 


by the comb-like bristles on the hind feet, and their bell-like cry ; 
two Chilian species, constituting the genus Spalacopus, being 
distinguished by their rudimental ears. Schizodon, on the other 
hand, of which there is a single species from the Southern Andes, 
has the ears larger than in the tuco-tucos. The South African 
Petromys has been regarded as very closely allied to Spalacopus^ 
but it is more probable it should be classed with the other two 
African genera Ctenodactylus and Pectinator, to constitute a 
separate sub-family. The third sub-family includes all the other 
genera, one of which (Triaulacodus 1 }, as represented by the cane- 
rats, is Ethiopian, while the whole of the remainder are Neogaeic. 
Many of the species are of large size, some being arboreal and 
others aquatic. Of the latter the best known and largest is the 
coypu (Myopotamus), widely spread over South America ; while 
in the West Indies the group is represented by the equally large 
arboreal hutias (Capromys and Plagiodori). The other seven 
genera are South American, and include smaller rat-like forms, 
which in the case of the two genera Loncheres and Echinomys have 
flattened spines mingled with the fur; the others being known 
as Mesomys, Dactylomys, Cannabateomys, Cercomys, and Cartero- 
don. Several of the existing genera occur fossil in the caves of 
Brazil and the Pampean ; Myopotamus also occurring in the 
infra-pampean beds of Parana, together with the reputedly 
extinct forms described as Orthomys and Morenia. Other extinct 
genera, such as Neoremys, Scleromys, and Addphomys occur in the 
Santa Cruz deposits, and appear to be very closely allied to 
Myopotamus. It is noteworthy that an extinct Octodont (Pelle- 
grinia) allied to Ctenodactylus occurs in the Plistocene or Pliocene 
of Sicily; while Ruscinomys from the Pliocene of France is 
believed to belong to the same group. Finally, the genus 
Eocardia, together with certain other allied forms from the Santa 
Cruz beds, are regarded as indicating a separate family (Eocar- 
ditdce) of the section. 

With regard to the extinct family Theridomyida from the 
middle and upper Oligocene of Europe, which includes not only 
Theridomys and Archceomys, but probably also the above-men- 

1 The name Aulacodus being preoccupied, that of Triaulacodus is proposed 
in substitution. 


tioned Nesocerodon and Issiodoromys (p. 89), it seems highly 
probable that these are really the ancestral forms of the modern 
Hystricomorpha, although their lower jaws approximate to the 
general type of those of the more generalised Sciuromorpha and 

The last section of the order (Lagomorpha), which includes the 
hares and picas, and is essentially a northern one, is but poorly 
represented in Neogaea ; the picas (Lagomyidcz) being unknown 
there, while in the whole of the realm there are only two species of 
Leporidce, one of which is Brazilian. 

It has been usual in zoological systems to include under the 
title of Edentata not only the armadillos, anteaters, 
and sloths of South America, but likewise the Old 
World pangolins (Manida) and aard-varks (Orycteropodidaz). 
There can, however, be no doubt that there is little or no 
connection between the two groups, and the latter may accord- 
ingly be separated as a distinct order under the title of ErTodientia. 
In this restricted sense the edentates at the present day are, per- 
haps, the most characteristic and remarkable of all the Neogaeic 
mammals. Whereas, however, the sloths and anteaters are 
entirely Neogseic, a few of the armadillos have wandered at a 
comparatively modern date as far north as Texas ; but this does 
not detract from their essentially southern character, seeing that 
they are well represented in the Santa Cruz beds, and, if we 
exclude certain remains of doubtful affinity from the Oligocene 
of France, are quite unknown elsewhere. This, however, is by 
no means all, since there are two extinct families of the order, 
dating from the Santa Cruz beds, which were extremely abundant 
during the Pliocene and Plistocene epochs ; some few of these 
having managed to obtain an entrance into North America 
about the Miocene epoch. Central and South America may 
accordingly be considered as essentially the home of the edentates ; 
and are thus broadly demarcated from all other parts of the world. 
It would be superfluous to point out all the distinctive features of 
the order, but it may be mentioned that none of the living forms 
have teeth in the front of the jaws ; while in all those genera in 
which teeth are present, these are of comparatively simple 
structure, being unprovided with a coating of enamel, growing 


continuously without ever forming roots, and being mostly very 
similar throughout the series. 

The mailed, or loricate edentates are represented by the two 
families of the armadillos (Dasypodidce] and glypto- Armadillos 
donts (Glyptodontida], the latter of which died out and Glypto - 
at the close of the Plistocene or the commence- 
ment of the Recent epoch, whereas the former is still abundant. 
With the exception of the two species of pichiciagos (Chlamydo- 
phorus] from Mendoza and Bolivia, in which a solid coat of mail 
is confined to the hinder region of the body, the members of both 
families have their bodies protected by a bony armour, while 
their heads are guarded above by a shield of the same nature, 
and their tails are enclosed in a tubular sheath. Covered exter- 
nally with horny shields, like the shell of a tortoise, the carapaces 
of these animals are formed of a number of small plates of bone, 
either united everywhere by their edges into a continuous solid 
armour, or in the middle region of the body overlapping one 
another like the tiles on a roof. In the true armadillos (of which 
the living Argentine forms are all comparatively small creatures, 
although one Brazilian species reaches nearly a yard in length) the 
carapace consists of a nearly solid buckler in front and behind, while 
between the two are situated a variable number of movable over- 
lapping bands, which in some instances admit of the body being 
rolled up into the form of a ball. They have all long snouts, and 
simple, subcylindrical teeth. The true existing armadillos may 
be divided into the genera Dasypus, Xenurus, Priodon, Tolypeutes, 
and Tatusia. The first of these, in which there are six or seven 
movable bands to the carapace, is found throughout the Argentine 
Tertiaries to the Santa Cruz beds, one of the fossil species from 
the higher beds of the series having a skull of nearly a foot in 
length, and thus vastly exceeding all its living congeners in size. 
Tatusia, in which the carapace has from seven to nine movable 
bands, does not appear to be known below the Pampean beds, 
where it is represented by the large species of which the 
external skeleton is shown in the figure on p. 93. A third genus, 
Eutatus, which likewise comprises species of large size, and 
ranges from the Pampean to the Santa Cruz beds, is distinguished 
by having over thirty movable bands in the carapace. More 


remarkable than any is the extinct Peltephilus of the Santa Cruz 
deposits, in which the teeth form a continuous series up to the 
front of the jaws, while the skull has a very broad snout, and the 
humerus is of such a remarkable shape that it has been described 
as that of a monotreme. Indeed this genus seems to suggest 
that edentates are derived from animals with a fully developed 
series of teeth in the front of the jaws. The pichiciagos (Chlamy- 
dophorus], which are unknown before the Plistocene, form a 
sub-family by themselves ; and yet another sub-family group is 
indicated by the gigantic Chlamydotherium, of the Brazilian caves 
and the Pampean, which rivalled the largest glyptodonts in size, 
and had teeth of a more complex type than the true armadillos. 
Other species occur in the Catamarca and Monte Hermoso Ter- 
tiaries, although the genus is unknown in those of Patagonia. 

From the armadillos and their immediate kin the extinct 
glyptodonts differ in having the carapace in the form of a con- 
tinuous solid shield, without any movable bands in the middle 
region ; in addition to which the skull is characterised by its 
depth and shortness, while the teeth form long fluted prisms. 
The internal skeleton, as shown in figure 19, is characterised by 
the union of nearly the whole of the vertebrae of the back into 
a solid girder for the support of the massive carapace ; and the 
feet are furnished with much shorter claws than those of the 
burrowing armadillos, the hinder ones being almost nail-like in 
form. Most of the species from the Pampean formation are 
animals of gigantic dimensions, the length of the carapace being 
not unfrequently from six to eight feet ; and they are unquestion- 
ably some of the most extraordinary creatures that ever trod the 
earth. Although the majority are South American, some members 
of the genus wandered as far north as Texas, while from the 
upper division of the Loup Fork beds, corresponding to the 
lowest Pliocene, a North American form has been described under 
the name of Carioderma. 

In the typical genus Glyptodon, which ranges from the sand- 
dunes and Pampean formation to the Monte Hermoso beds, the 
tail-sheath, as shown in the annexed figure, is composed of a 
number of spiny rings, gradually decreasing in size from the root 
to the tip, and the polygonal plates of the carapace are each 


Cs. O 


ornamented with a distinct rosette-like sculpture. The allied 
genus Plohophorus, of which the remains occur alike in the 
Brazilian caves and the Catamarca and Monte Hermoso beds of 
Argentina, while agreeing with the last in the characters of the 
skull, has a carapace more like that of the undermentioned 
Panochthus, and a tail-sheath resembling that of the next genus. 
In addition to well-marked distinctive features of the skull, Loma- 
phorus is characterised by the great elongation and slender form 
of the carapace, which is produced on either side of the neck in 
the same manner as in the armadillos, while its margins lack the 
large bosses exhibited by the typical genus. The tail-sheath 
consists of a small number of rings at the base, followed by a 
long terminal tube ornamented with smooth, oval, bony plates, 
of which those along the sides and at the tip are larger than the 
rest. The genus, of which the species are much inferior in point 
of size to those of Glyptodon, has the same geological range as 

Another type of the family is represented by the animals 
constituting the genus Panochthus, in which the hexagonal bony 
plates of the carapace are arranged in more distinct rows on the 
sides of the body ; those of the back being ornamented either 
with a number of small granular tubercles (as in the species here 
figured), or with one circular central disc surrounded with several 
rows of much smaller discs. A more striking difference is dis- 
played in the structure of the sheath of the tail, which consists at 
the base of six or seven large smooth rings diminishing very 
rapidly in diameter, and terminates in a long and massive de- 
pressed tube, the sides of which are ornamented with large 
roughened bosses, probably surmounted during life with horny 
knobs, while the intervening spaces are covered with small bony 
ossicles. The species are of very large or medium size, and range 
from the Pampean to the Monte Hermoso beds in Argentina, 
while some occur in the Brazilian caves. Still more extraordinary 
is the gigantic Dadicurus of the Pampean, represented by a some- 
what smaller form from the Monte Hermoso beds. Having a 
total length in a straight line of close upon twelve feet, five of 
which are taken up by the ponderous tail, the Pampean repre- 
sentative of this genus has the outer surface of the plates of the 






carapace smooth ; each being perforated by three or four large 
holes for the passage of blood-vessels, and the whole being pro- 
bably invested with a continuous leathery skin, instead of each 
disc bearing a separate horny shield. Commencing with a small 
series of enormous, narrow, hoop-like rings, the tail-sheath termi- 
nates in a long, massive, depressed, and nearly smooth club- 
shaped tube, at the extremity of which are a number of rough 
disc-like surfaces, apparently for the attachment of large horns. 

None of the preceding forms, which include all the largest 
members of the family, range below the horizon of the Monte 
Hermoso, Catamarca, and Parana beds, but the group is also 
represented in the Santa Cruz deposits of Patagonia, although the 
single species found there is of much smaller dimensions than any 
of its later cousins, the whole length of the carapace not exceeding 
about two feet. It is noteworthy that this earliest known glypto- 
dont, on which the unwieldy name of Propalczohoplophorus has been 
conferred, presents certain indications of affinity to the armadillos 
in the structure of its carapace, in which incipient movable bands 
may be detected on the margins of the middle region. In this 
small size of their earliest definitely known representative, the 
glyptodonts resemble the under-mentioned ground-sloths. 

Unless the aforesaid remains from the Oligocene Phosphorites 
of France should prove to belong to the group, we are at present 
totally in the dark as to whence both the glyptodonts and the 
armadillos originally came ; and it is, indeed, quite probable 
that, like the other members of the order, they may have origi- 
nated in South America (if not in an Antarctic continent) from 
some at present quite unknown mammalian type. How such 
creatures, which seem absolutely unassailable, came to be extermi- 
nated, is one of those questions which it appears quite impossible 
to answer. 

Although they have not hitherto been discovered in a fossil 
state, the sloths, constituting the family Bradypodidce, 
are just as characteristic of Neogaea as the two pre- 
ceding groups. Their habits, however, necessarily restricting them 
to the tropical forest-districts, their absence in a fossil state 1 must 

1 A presumed fossil sloth was described from the Argentine, but the jaw on 
which it was founded proves to belong to the Megalotheriidce. 


not be taken as an indication that they did not exist during the 
Pampean epoch, since their remains are not likely to occur in 
the Argentine, although they might with more probability be 
looked for in Brazil. On the other hand, their specialised struc- 
ture makes it highly probable that they had not come into being 
at the date of deposition of the Santa Cruz beds. Of the dimen- 
sions of medium-sized monkeys, sloths are characterised by their 
short, rounded heads, and extremely long limbs, armed with very 
elongated curved claws ; in the genus Bradypus the latter being 
three in number on each foot, but in Cholcepus reduced to two 
in the fore feet. Their bodies are coated with very coarse ragged 
hair, and the tail is wanting. The teeth are oval prisms, some- 
what cupped in the middle of their grinding surfaces ; but in 
the last-named of the two genera the first pair in each jaw are 
larger than the rest, from which they are separated by an interval, 
and form tusks wearing against one another to oblique facets. 
Usually there are five upper, and four lower teeth on a side. 
The range of sloths extends from Mexico throughout the greater 
part of the forest-districts, although they do not appear to reach 
as far south as Paraguay. 

Likewise unknown in a fossil condition, the true anteaters, or 
Myrmecophagidce, constitute another exclusively Neo- 

.. ' . . i , Anteaters. 

gseic family, with nearly the same geographical range 
as the sloths, but represented in Paraguay. So unlike are these ani- 
mals to sloths, that it is at first difficult to believe that there is any 
close relationship between the two, and it is largely due to the evi- 
dence of the ground-sloths referred to below that it has been possible 
to discover how close the connection really is. In place of being 
rounded and shortened, the skull in the present family is more or 
less elongated and slender, with the jaws entirely devoid of all 
vestiges of teeth, and the tongue long, cylindrical, and extensile. 
An equally striking difference obtains in regard to the structure of 
the limbs, the fore foot of the great anteater having five toes, of 
which the middle one is much more powerful than the others, 
while all except the fifth are furnished with strong claws. In 
walking, the outer side and part of the upper surface of the fore 
foot is applied to the ground ; but in the hind limbs the sole 
forms the support in the ordinary manner. Whereas sloths are 


highly specialised as regards the structure of their limbs, in the 
present group the greatest degree of specialisation shows itself in 
the skull, in the majority of the species. There are but three 


members of the family, each representing a genus by itself, namely 
the great anteater (Myrmecophaga\ the tamandua (Tamandua], and 
the two-toed or little anteater ( Cycloturus] ; the latter being ex- 
clusively arboreal in its habits. 

The foregoing remarks on some of the structural features of 

the sloths and anteaters will the more easily enable 
si?th s Und " the rea der to understand the peculiarities of the 

extinct group of ground-sloths. They have been 
divided into a large number of genera and several families; but 
the former may be considerably reduced, and the whole of them 
included in the single family Megalotheriidce. Ranging in the 
Argentine from the Santa Crucian to the Pampean epoch and the 
overlying sand-dunes, the family has a geographical distributional 
area including North America as far as Kentucky. The South 
American forms vastly exceed those of N. America in point of 
number; and whereas the latter are found only in deposits of 
upper Pliocene and Plistocene age, the former, as we have seen, 
extend downwards to the Miocene. The members of this family 
may be denned as edentates without a carapace, the skull and 
dentition of the general type of those of the sloths, and the 


limb-bones and vertebrae like those of the anteaters. The skull is, 
however, somewhat more elongate than in the former, and in the 
case of the genus Scelidotherium approximates to that of the latter. 
The Plistocene forms include by far the largest representatives of 
the order, the Megalotherium 1 attaining a total length of about 
eighteen feet, with a bodily bulk fully as great as that of an 
elephant. Whereas all the members of the family whose remains 
occur in the Plistocene walked on the outer sides of their feet, in 
the small ancestral Patagonian forms this specialised character 
seems to have been less developed. 

The typical genus Megalotherium which includes several 
species, ranging in time from the Monte Hermoso and Cordoba 
beds to the Pampean, and in space from Argentina and Chili to 
South Carolina and Texas is sufficiently distinguished by having 
the teeth in the form of large quadrangular prisms, sometimes 
measuring as much as a foot in length, and wearing on their sum- 
mits into a pair of transverse ridges, owing to the presence of layers 
of unequal hardness. The allied genus Mylodon, including smaller 
forms which may be compared in size to rhinoceroses, differs from 
the preceding in the structure of the teeth, which are similar to 
those of the sloths ; the skull, as shown in the figure on 
p. 104, being comparatively short, with the teeth extending nearly 
up to the extremities of the jaws. In the skeleton of this genus 
the limbs are of moderate length and very powerful. The two 
outer toes of the fore feet are rudimental and clawless, but the 
three innermost provided with claws, of which the third is much 
larger than either of the others, this discrepancy being carried 
to a still greater extent in Megalotherium. It will be observed 
that the creature walked on the outer sides and part of the upper 
surfaces of the fore feet after the manner of a sloth ; but, unlike 
the latter, only the outer sides of the hind feet were applied to the 
ground ; the great middle toe, which in Megalotherium carried a 
gigantic claw, not touching the ground at all. In the structure of 
their feet these animals are thus more like anteaters than sloths, 
although the hinder pair are of a somewhat more specialised 
structure than in the latter. It may be mentioned that the 

1 The name Megatherium clearly requires amendment to Megalotherium. 




conformation of their teeth indicates that the ground-sloths were 
vegetable-feeders, and it is probable that they subsisted largely 
upon the young twigs and leaves of trees, which may have been 

FIG. 23. UNDER-SURFACE OF SKULL OF Mylodon. (Reduced.) 

brought within reach by the animals rearing themselves up 
against the trunks and pulling down the boughs with their fore 
paws. The present treeless condition of the Argentine pampas 
suggests that the ground-sloths were grazing rather than browsing 


animals, but their structure is not in favour of this view, and it 
must be remembered that their remains are likewise met with in 
Brazil, which was probably always as well wooded as at the present 
day. The disappearance of forests from the pampas cannot, in- 
deed, be regarded as more marvellous than the extinction of its 
Plistocene mammals. In the sand-dunes near the coast at Buenos 
Aires bones of some of the ground-sloths, as well as of glyptodonts, 
have been found in association with human remains, so that their 
extinction is an event of comparatively recent date. The genus is 
typically represented by Mylodon harlani of the Plistocene of 
Kentucky and other parts of North America, but is nevertheless 
essentially South American, ranging in Argentina from the Pam- 
pean beds to those of Parana and Monte Hermoso. The allied 
genus Megalonyx is exclusively restricted to the North American 
Plistocene and Upper Pliocene ; and here may be repeated the 
observation that the absence of remains of these ground-sloths 
from the Miocene of North America, coupled with their presence 
in the Santa Cruz beds of Patagonia, clearly indicates that they are 
late immigrants from the south into the northern half of the 
continent. , 

Nearly allied to Mylodon, the genus Glossotherium from the 
Plistocene of Argentina and Uruguay serves to connect it with 
another generic representative of the family known as Scelido- 
therium. In place of the comparatively short skulls of the 
mylodons, the species of this genus have the muzzle of the skull 
greatly elongated, so that there is a long toothless space in 
advance of the dental series ; and whereas the skulls of the species 
of Mylodon are essentially sloth-like, those of Scelidotherium show 
a marked approximation to the anteater-type. The species of 
Scelidotherium are of medium or rather small size ; and in space 
the genus ranges from Patagonia, through the Argentine, to Brazil, 
Bolivia, and Chili ; while in time it extends from the superficial 
sand-dunes and Pampean deposits to the lower Tertiaries of 
Parana, Monte Hermoso, Catamarca, and Santa Cruz, with a 
gradual decrease in the size of the species as we descend in the 
geological scale '. Nearly allied is the genus Catonyx, from the 

1 The Santa Cruz form has been quite unnecessarily separated under the 
name of Analcithermm. 





w ^ 

K^* - 


Brazilian caverns ; while Nothrotherium of the same deposits 
seems to have been another nearly related form with teeth of the 
Megalotherium type. The imperfectly known Nothropus of the 
Pampean and Ortotherium of the Parana beds seem, on the other 
hand, to be late survivals of another group typically represented 
by the genera Eucholaops and Pseudhapalops of the Santa Crucian 
epoch of Patagonia. These forms differ from all those noticed 
above in that the terminal joints of some of the toes have a 
median cleft as in the great anteater, and likewise in the 
elongation of the metatarsal bones ; and it seems probable that the 
hind foot was not so much everted as in the later representatives 
of the family. The skull is of the general type of that of Mylodon ; 
most of the molars being prismatic in form, and surmounted by a 
pair of transverse ridges, more or less closely connected at their 
extremities so as to produce an oval cavity on the grinding surface. 
The first tooth is, however, tusk-like, and separated by a gap from 
the others. In some of these early ground-sloths the skull did 
not exceed three inches in length ; but other species were con- 
siderably larger. They are evidently generalised types, and were 
probably nearly allied to the ancestral stock which gave rise to 
Mylodon and Megalonyx, if indeed they be not the actual pro- 
genitors of both. 

The last group for consideration is that of the marsupials, or 
pouched mammals, among which the family of the 
opossums (Didelphyidce), with the three genera 
Didelphys 1 , Dromiciops, and Chironectes, is now confined to the 
New World, the great majority of the numerous species being 
Neogaeic, although the common opossum (Didclphys marsupialis} 
ranges from Chili and Brazil to the United States. Although 
certain forms from the Santa Cruz beds described under the 
names of Eodidelphys and Prodidclphys were originally assigned to 
the present family, these have been subsequently identified t^y 
Dr Ameghino 2 with the under-mentioned family of the Microbio- 
theriidce. True opossums occur, however, in the Monte Hermoso 

1 This genus is divided into several sub-genera, regarded by some writers as 
entitled to generic rank. 

2 Bol. Ac. Cordoba, Vol. xm. p. 363 (1894). 




beds while, as mentioned in the last chapter, they were widely 
distributed in the northern hemisphere during the Oligocene. If 
the conclusions of Dr Ameghino are right as to the absence of 
these marsupials from the Santa Cruz beds, it is evident that 
opossums only reached South America from the north at the close 
of the Miocene or commencement of the Pliocene, and that they 
do not belong to the indigenous fauna. It has been generally 
considered that the common opossum of the United States is a 
direct survivor from the Oligocene forms of North America, but it 
is more probable that it is really a very recent immigrant from the 
south, seeing that fossil representatives of the genus are unknown 
from the North American Miocene and Pliocene. During the 
Miocene the group perhaps survived in the extreme south of 
North America. 

Although opossums are apparently wanting, the Santa Cruz 
beds have yielded remains of undoubted marsupials, but several 

FIG. 25. LEFT HALF OF LOWER JAW OF Prothylctcinus, 

nat. size.) 

of them are assigned by Dr Ameghino to a distinct group under 
the name of Sparassodonta. Foremost of these is the genus 
ProthylacinuS) already mentioned in the last chapter, which may 
be provisionally assigned to the Australian Dasyuridce. In having 
only three in place of four lower pairs of incisors this genus agrees 
with the latter family, and differs from the opossums ; while the 
whole character of the lower jaw and dentition is very similar to 
that of the Tasmanian Thylatinus, with the exception that the 
premolars are closer together. As in the Dasyuridce generally, 


there are four pairs of upper and three of lower incisor teeth in 
Prothylacinus, and the same is the case with the smaller Santa 
Crucian form described as Amphiproviverra, which appears to be 
of a distinctly dasyurid type, although not coming very near to 
any Australian genus. v 

With regard to the Microbiotheriida, as typified by the genus 
Microbiotherium, these, although they are not included by Dr 
Ameghino in the order, appear to be undoubted marsupials, since 
they have a dentition numerically the same as that of the opossums, 
vacuities in the palate, and an inflected angle to the lower jaw. 
From the opossums they differ by the non-production of the palate 
behind the last molars, and in the form of the lower jaw, in which 
the extremity is produced to a greater extent in advance of the 
canine. In all these respects they approximate to the Dasyurid 
genus Phascologale, from which they differ in having one pair less 
of incisors in each jaw. The ancestors of the Australian Dasyuridce. 
must, however, have originally had five pairs of upper and four of 
lower incisor teeth, as the former are retained in many of the ban- 
dicoots (Peramelida), while Myrmecobms occasionally develops four 
pairs of these teeth in the lower jaw. It seems therefore probable 
that the Microbiotheriidcz were minute polyprotodont marsupials 
of an Australian type. 

There is more difficulty in arriving at any satisfactory con- 
clusions as to the serial position of certain carnivorous mammals 
from the Santa Cruz beds, of which a large form described as 
Borhycena may be taken as an example. In these animals the 
dentition approximates to a certain extent to that of the primitive 
or creodont Carnivora of the earlier Tertiaries of the northern 
hemisphere, although retaining the marsupial feature of four pairs 
of molars and only three of premolars. The replacement of the 
teeth is also fuller than in the marsupials. Dr Ameghino has 
suggested that these animals were transitional between marsupial 
and eutherian carnivores, and that the latter group originated in 
South America ; but this idea is obviously untenable. A possible 
suggestion is that they may be specialised offshoots from the 
marsupial stock which died out without giving origin to any 

A small mouse-like mammal first described in 1863 upon the 


evidence of a single specimen from Ecuador under the name of 
Hyracodon fuliginosus, but whose affinities were not determined 
till 1895, when an example of a second and larger species was 
procured from Bogota, belongs to the sole surviving genus of a 
group of small marsupials which occur abundantly in the Santa 
Cruz beds, and were till quite recently regarded as extinct. Un- 
fortunately the name Hyracodon has been previously employed 
to designate an extinct ungulate, and it has accordingly been 
replaced by Ccenolestes. The essential characteristic of this group 
of marsupials, is that while their upper dentition is of a poly- 
protodont type, that of the lower jaw is very similar to the 
diprotodont modification. In the living species, for instance, 
there are four pairs of small upper incisors of a normal type, 


Acdestis oweni. (Much enlarged.) 

The first tooth on the right is the first incisor, and the one on the extreme 
left the first molar. 

followed by a large canine, while in the lower jaw, as shown in 
the accompanying figure of that of one of the extinct forms, there 
is a large pair of horizontally projecting incisors, succeeded by 
several minute functionless teeth, of which the first three represent 
the second and third incisors and the canine. In all the forms 
the molar teeth have quadrangular crowns, surmounted by four 
blunt tubercles, somewhat resembling in structure those of certain 
ungulates, and thus totally different from the triangular and 
sharply-cusped molars of the opossums and other polyprotodonts. 
In the living forms, as well as in certain fossil kinds (Epanorthus, 
Decastis and Acdestis] from the Santa Cruz beds, the last premolar 
tooth, as shown in the figure of the jaw of Acdestis, is of normal 
dimensions : and these forms may consequently be grouped in a 
single family, under the name of Epanorthidce. In another group, 


confined to the Santa Crucian horizon, where it is represented by 
the family Abderitidce, the last premolar in each jaw is much 
larger and taller than the other teeth, and has its crown in the 
form of a compressed cone, marked on the sides with vertical 
grooves, as exhibited in the figure of Abderites. A third family is 


known as the Garzoniidce. In all the skull is of an elongated 
form, with large vacuities both in the front and hinder part of the 
palate, and presents a considerable general resemblance to those 
of the Australian genera Peragale and Perameles. With the ex- 
ception of the retention of four pairs of upper incisors and the small 
size of all these teeth, the dentition exhibits, however, a remark- 
able approximation to that of the Australian diprotodont genus 
Dromicia. On the other hand, the feet are of normal structure, 
with five complete toes, none of which are united by integuments; 
the thumb and great toe being apparently slightly opposable to 
the other digits. Probably the rat-like tail is slightly prehensile 
at the extremity ; and a small pouch is present in the female. In 
the skeleton the lower jaw exhibits the usual inflection of the 
angle ; and the pelvis carries marsupial bones. 

Probably these Patagonian marsupials, which may be known 
as selvas, must be included in the diprotodont sub-order ; from 
the Australian representatives of which they differ by the small 
and numerous upper incisors and the non-syndactylous hind feet. 
Both these being generalised features, it is evident that if the 
selvas are true diprotodonts their ancestors must have originated 
from the polyprotodonts in Notogaea, for if they are of exclusively 
South American origin they must form a subordinal group by 


themselves. Assuming their affinities with the Australian type 
to be rightly determined, they constitute a most important link 
in the chain connecting the faunas of South America and Aus- 

In the last chapter it has been argued that, from the absence 
of allied forms in the Tertiaries of North America and Europe, as 
well as from their resemblance to the Australian dasyurids, it is 
difficult to come to any conclusion other than that the ancestors of 
the Santa Crucian polyprotodont marsupials reached the country 
from Australia, either by way of the Antarctic continent, or by a 
land-bridge in a more northern part of the Pacific. If this be 
correct, and likewise the supposition that the opossums origi- 
nated from the ancestral stock in South-eastern Asia, it will be 
evident that Didelphys and Ccenolestes met in South America 
after their ancestors had travelled half round the world in opposite 

It may be added that the alleged occurrence of monotremes 
in the Santa Cruz beds is due to bones of aberrant armadillos 
(Peltephilus) having been mistaken for those of that group 1 . 

Although in this volume the writer avoids laying much stress 
upon aquatic mammals, it may be mentioned that 


there are two genera ot dolphins belonging to the 
family Platanistidce, each represented by a single species, which are 
peculiar to the Neogaeic realm. These are Stenodelphis (Pontoporia] 
from the mouth of the Rio de la Plata, and Inia of the upper 
Amazon; the only other existing representative of the family 
being Platanista of the larger Indian rivers. 

After the foregoing survey of the chief features of the recent 

and fossil mammalian fauna of the Neogaeic realm, 
tinction ofThe ^ ts g enera l bearings on the relations of South America 

t o other parts of the world may be taken into con- 

sideration. It will, however, facilitate matters to 
give a tabular view of the orders, suborders, and families of non- 
volant land mammals represented in the realm. In the following 
table such groups as are either confined to Neogsea, or have only 
reached North America at a comparatively recent epoch are 

1 See Lydekker, An. Mus. La Plata Pal. Argent. Pt. III. p. 67 (1894). 


printed in italics ; the extinct types being indicated by the prefix 
of an *, while those dating from the Santa Crucian (or the earlier 
Patagonian) epoch are followed by the words Santa Cruz, and 
those from the Parana beds by the word Parana. 

I. PRIMATES. Santa Cruz. 

Cebida, Santa Cruz (* Hdmunculus). 


Phyllostomatida. One genus ranging to Cali- 

Emballonuridae. Seven peculiar genera. 
Vespertilionidae. Natalus, Thyroptera. 


Solenodontidce. West Indies. 


Felidae. No peculiar genera. 

Canidae. In addition to the cosmopolitan Canis, 

represented in Brazil by the peculiar 

Ursidae. No peculiar genera; *Arctotherium, 

common to N. America. 
Procyonidae Nasua, Cercoleptes, Bassaricyon, and 

* Cynonasua. 
Mustelidae. Galictis, and Conepatus, the latter 

extending into Texas. 



Cervidae. Cariacus, peculiar to New World. 
Dicotylidae. Peculiar to New World at present 


Camelidae. Lama. 
L. 8 



Tapiridae. Elsewhere only in Malaysia at the 
present day, but formerly widely distributed 
over the northern hemisphere. 

Equidae. Now unknown, except through intro- 
duction. In addition to the cosmopolitan 
Equus (including Tertiary forms), the pecu- 
liar Pampean genera * Hippidium and * Ono- 

3. * Litopterna. Santa Cruz. 

* Macraucheniidce. Santa Cruz. 

* Proter other iidtz. Santa Cruz. 

4. * Astrapotheria. Santa Cruz. 

* A strapotheriidce. Santa Cruz. 

* Homalodontotheriida. Santa Cruz. 

5. * Toxodontia. Santa Cruz. 

* Toxodontidce. Santa Cruz. 

* Typotheriida. Santa Cruz. 

* Pachyruchidce. Santa Cruz. 

6. * Pyrotheria. Patagonian beds lying below those of 

Santa Cruz. 

* Pyrotheriidte. Patagonian beds. 


Elephantidse. Represented by Mastodon in 
the Pampean and Monte Hermoso beds. 



Sciuridae. Represented by Sciurus as far south 
as Paraguay. 

* Castoroididse. Peculiar to New World. 


Muridae. Represented by species of the New 
World genus Sitomys, as well as by several 
peculiar types such as Rhithrodon, Ichthyomys 
of Peru, Holochilus of Brazil, etc. 



Caviidce. Paran a. 
Dasyproctidce . 

Lagostomatidcz. Santa Cruz. 
Hystricidae. Santa Cruz. 

Octodontidae. Mainly Neotropical, but also Ethi- 
opian. Santa Cruz. 

* Eocardiidce. Santa Cruz. 


Leporidae. Represented by two species of 

VII. EDENTATA. Santa Cruz. 

Dusypodida. Santa Cruz. A few forms range 
as far as Texas. 

* Glyptodontida. Santa Cruz. One Neogaeic genus 

ranges as far north as Texas, and a peculiar 
one has been described from further north. 

* Megalotheriidce. Santa Cruz. Mainly Neogaeic, 

but also ranging into North America. 


1. DIPROTODONTIA. Santa Cruz. (Elsewhere only in 


Epanorthidcz. Santa Cruz. (One existing genus, 

* Abderitidcz. Santa Cruz. 

* GarzoniidcB. Santa Cruz. 


Didelphyidae. Now mainly Neogaeic, where they 
date from the Monte Hermoso stage, but 
ranging into North America, and formerly 
widely spread over the northern hemisphere. 
Chironectes, several subgenera of Didelphys, 
and Dromiciops peculiar to the realm. 



VIII. MARSUPIALIA (continued}. 

Dasyuridae. Santa Cruz. Now confined to 
Notogaea, but apparently represented in the 
Santa Cruz beds by * Prothylacinus and 
* Amphiproviverra. 

* Microbiotheriidcz. Santa Cruz. 
Incertce, Seats. 

* Borhyanidce. Santa Cruz. 

Although the addition of the names of all the peculiar genera, 
both recent and extinct, would have rendered the distinctness of 
the Neogaeic mammalian fauna still more pronounced, yet the 
foregoing table as it stands is amply sufficient to show that 
Neogaea is entitled to form one of the three primary zoological 
realms of the world. Starting with the Santa Crucian epoch of 
Patagonia and the somewhat older Patagonian stage, which form 
the earliest date at which the history of the Tertiary land mammals 
of Neogaea can be taken up, there is evidence that at least the 
southern part of the area was populated by the following pecu- 
liar fauna. Firstly, monkeys of a type quite different from those 
of the Old World, but evidently allied to the existing Neogaeic 
forms, were abundant; while rodents, belonging to the same 
groups as those now inhabiting the continent, several of which 
were nearly allied to existing African forms, and more remotely to 
certain Oligocene European types, attained a great development. 
Probably Insectivora with V-shaped molars were also present. 
More peculiar are the extinct subordinal groups of ungulates 
described above, which appear to have been allied to the ancestors 
of the perissodactyles of the northern hemisphere, and may 
possibly be remotely connected with the African hyraces. At the 
same period flourished several families of edentates (a group 
which in its restricted sense was originally peculiar to Neogaea), 
such as armadillos, glyptodonts, and ground-sloths, most of the 
members of which were of comparatively small size ; but of the 
ancestry of this group nothing can be said with certainty. 
Among the marsupials, although opossums appear to have been 
wanting, there were several types seemingly allied to Notogaeic 
forms, while others which may be included in the order were more 




or less unlike any from other regions. In addition to true mar- 
supials, the only carnivorous types were the problematical 

Now, as stated in the earlier part of the chapter (p. 68), 
this fauna cannot be older than the lowest Miocene or highest 
Oligocene ; and among its deficiencies may be noted lemuroids, true 
carnivores, creodonts, artiodactyle and perissodactyle ungulates, 
and opossums, all of which were in existence during the Oligo- 
cene or Miocene in North America and Europe. Moreover, at those 
epochs the former country lacked the whole of the Neogaeic types. 

Clearly, then, there must have been a barrier between North 
and South America during the Oligocene and a 

Early Separ- 

portion or the whole of the Miocene; but before ationofN.and 
entering into the consideration of other evidence S ' Amenca - 
showing the nature of that barrier, it may be well to give a table of 
the mammaliferous Tertiary strata of North and South America, 
with their approximate European equivalents 1 . In descending 
order this runs as follows : 

Up. Pliocene 
Low. Pliocene 


Up. Oligocene 

Mid. Oligocene 

Low. Oligocene 
Up. Eocene 
Mid. Eocene 

Low. Eocene 
Lowest Eocene 

South America. 

Santa Cruz 

North America. 
Equus beds. 

Monte Hermoso f Palo Duro. 

(?) Parana Loup-Fork -j Nebraska 

tDeep River 

John Day 

/ Protoceras 

White River j reodon 


ium Beds 

Bridger -I Bridger. 

IWind River. 


Cave-deposits etc. 
? Crag. 
| Pikermi 


St Gerand-Le-Puy 




1 In compiling this table the writer is indebted to Prof. W. B. Scott. 
Many American geologists (among them Dr Scott) include the whole of the 


Regarding the geological evidence of a separation between the 
two Americas, Cretaceous marine strata occupy a large portion of 
Mexico; and in 1879 Dr Le Conte 1 wrote as follows. The shore 
line of the Gulf of Mexico "was much more extended both north- 
ward and westward than either now or in Tertiary times. From 
the Gulf there extended northwestward an immensely wide sea, 
covering the Plains region and the Rocky Mountain region as far 
westward as the Wahsatch range, and dividing the continent into 
two continents, an eastern or Appalachian, and a western or 
Basin-region continent. Probably also this sea connected across 
the region of Mexico with the Pacific, thus dividing the western 
continent into two, a northern and southern." Later observations 
have shown that the Cretaceous sea undoubtedly made a wide gap 
between North America and the southern portion of the con- 
tinent 2 ; while the existence of Oligocene or Miocene strata in the 
region of the isthmus of Parana shows that the separation, which 
probably continued through the Eocene, was in existence during 

Loup-Fork in the Miocene; while to the lower part of the same era they 
assign the John Day beds of America, and the St Gerand-le-Puy beds of 
Europe. Others (e.g. Prof. Osborn, Studies Biol. Labor. Columbia Coll. 
vol. i. pt. 2, p. 28, 1893) refer the Equus beds of North America to the 
Pliocene. The following quotation from a paper by Prof. Cope (American 
Naturalist, 1895, p. 599) conclusively proves their Plistocene age. There it 
is stated that "the Equus beds are found covering areas of various extent in 
Oregon, Nevada, California, the Staked Plains, Southern Texas, Chihuahua 
and the valley of Mexico. Their most eastern station is western Nebraska. 
They contain a fauna which includes one extinct species (Equus major Dek.) of 
the Megalonyx fauna, and the recent Castor fiber. They contain the extinct 
genus of sloths Mylodon, of a species different from that of the east, and four 
species of camels of the extinct genus ffolomeniscns, and a peccary. Recent 
species of Cam's and Thomomys occur, while two extinct horses (Equus occi- 
dentalis Leidy and E. tau Owen) are common. The hairy elephant (E. primi- 
genius) is abundant, while the Mastodon americanus is rare, if occurring at all. 
The proportion of recent to extinct species and genera in the Equus bed fauna 
is very similar to that occurring in the Megalonyx fauna, while they differ as to 

1 Elements of Geology, pp. 451, 452, New York (1879). 

2 This separation also existed in the Jurassic era, when, as shown by 
Neumayr (Erdgeschichte, 2nd ed. vol. II., p. 263), South America was united 
across the Atlantic area with Africa and Madagascar. 



the middle portion of the Tertiary epoch 1 . When the connection 
between North and South America was completed is not precisely 
fixed by geological evidence ; but the occurrence of a glyptodont 
in the Nebraska stage of the Loup-Fork group, shows that it must 
have been by the end of the Miocene 2 . The question of a con- 
nection between the two continents by way of the West Indies 
is discussed later in this chapter, where it is concluded that if 
such a connection existed at all, it must have been of a transient 

Having thus shown, both from palaeontological and geological 
evidence, that the early mammalian fauna of 

Incursion of 

Neogaea appears to have been totally isolated from Northern 
that of North America up to about the end of the 
Miocene, the question of the origin of that fauna may be deferred, 
and the irruption of northern types after the connection between 
North and South America had been established taken into con- 
sideration. It may be mentioned, however, that it was not till 
after this irruption of the northern forms that the essentially 
Neogaeic fauna attained its maximum development in respect to 
the bodily size of its constituents ; since a gradual increase in this 
respect maybe traced from the small glyptodonts and sloths of the 
Santa Cruz epoch, through the larger ones of the Monte Hermoso 
horizon, to the gigantic forms characteristic of the Pampean and 
the cavern deposits of Brazil. 

The presence of a glyptodont in the Nebraska stage of the 
Loup-Fork group in North America, and of northern forms in the 
Monte Hermoso horizon of South America, marks, then, the first 
commingling of the original faunas of the two halves of the New 
World. For the first time in the history of the southern continent 
this connection allowed of the immigration from the north of the 
true Carnivora, such as the existing cats (Felts) the extinct sabre- 
toothed tigers (Macharodus), dogs and foxes (Canida), bears 
(Ursus and Arctotherium\ raccoons (Procyonidtz), skunks and 
their allies (Mustetida), together with various ungulates belonging 
to sub-orders previously unknown in the realm. These latter 
include the guanaco and vicuna (Lama] of which ancestral forms 

1 See J. W. Gregory, Quart. Joum. GeoL Soc. Vol. LI. pp. 299, 300 (1895). 

2 As stated above, many refer the whole Loup-Fork group to the Miocene. 


are abundant in the North American Tertiaries New World deer 
(Cariacus], horses (Equidce) of various genera, tapirs (Tapirid&\ 
peccaries (Dicotylidte), and mastodons. Among the rodents, 
squirrels, the various genera of Muridce, and the hares, likewise at 
this epoch made their first, appearance on the scene. Opossums 
also at this time effected an entrance into the land which has now 
become their chief home. That this new fauna came in from 
North America, and not from any other part of the world, may be 
regarded as certain from the presence of such essentially New 
World types as raccoons and their allies, skunks, peccaries, 
Cariacus, and Camelidce (exclusive of the Old World genus 
Came/us, which is of late origin), coupled with the absence of 
true deer (Cervus), pigs (Sus), Old World monkeys, and lemurs. 

At the same time this union of the northern and southern 
halves of the New World allowed certain members of the original 
Neogaeic fauna to make their escape into North America, glypto- 
donts, as already said, making their appearance in the Nebraska 
stage of the Loup-Fork group of the United States, while the 
ground-sloth Megalonyx occurs in the Blanco Beds. 

Although it is not universally admitted 1 , there is some 
evidence to indicate that this land connection was of com- 
paratively brief duration, seeing that none of the characteristic 
extinct types of South American ungulates, nor any of the peculiar 
Neogaeic rodents, reached the northern half of the continent. 

During the whole time that the alluvial deposits of the Parana 
and Paraguay rivers were being laid down, and well on into the 
human period, the mammalian fauna of the Pampean epoch, formed 
by an admixture of southern and northern types, continued to 
flourish, until the time when there came a complete sweep of all 
the larger forms, clearing off the whole of the ground-sloths, 
glyptodonts, mastodons, toxodonts, macrauchenias, horses, sabre- 
toothed tigers, and the larger members of the camel tribe, and in 
the Argentine leaving only armadillos, guanacos, a few deer, a 
number of rodents, various cats and foxes, as well as skunks and 
certain other members of the weasel family, to represent the vast 
assemblage of strange and giant creatures that once roamed over 

1 See Gregory, op. cit. p. 300. 


its plains. With regard to this extraordinary, and apparently 
sudden disappearance of almost all the larger forms of animal life 
from South America, it may be pretty confidently asserted, from 
the organisation of the creatures themselves, that at the time when 
the ground-sloths flourished, extensive portions of what is now the 
open pampas of Argentina were covered with forest ; and why the 
whole district should have become practically treeless, seeing that 
trees like the Australian eucalypti grow, when introduced, with 
more vigour than in their native home, is exceedingly hard to 
understand. That the country even when thus denuded was 
unsuited to the needs of the larger forms of mammalian life, is, 
however, negatived by the circumstance that in many parts the 
horses and cattle introduced from Europe have run wild and 
increased to an almost unprecedented extent. Neither does it 
appear that the extermination can be attributed to a period of 
extreme cold, since a glaciation of the pampas would assuredly 
have left unmistakable evidence of its presence. It is likewise 
practically certain that the clean sweep of the forests of Argentina 
and the larger mammals of the whole of South America is not due 
to the hand of man. It has, indeed, been suggested that the vast 
herds of guanaco which formerly roamed the pampas may have 
cleared the forests by preventing the growth of the seedlings ; but 
when we recall the fact that numbers of this group of animals 
flourished during the period when the alluvial formation was in the 
course of being deposited, it scarcely looks as though this could 
have been a vera causa. Moreover, if the forests were by some 
means or other actually destroyed, and the extermination of their 
animal denizens thus encompassed, there would still remain the 
disappearance of plain-dwelling forms, like horses, to be accounted 
for. Some have thought that pumas, by preying on the colts, 
were the active agents in this instance ; but even if such were 
really the case, it gives no help with regard to the disappearance 
of the ground-sloths and glyptodonts; the latter being such 
strongly armoured creatures that it is absolutely certain they were 
not killed off by any animal foes. The problem is further com- 
plicated by the circumstance that the fossil remains of nearly all 
the larger animals which formerly inhabited the pampas are also 
found in the caverns of Brazil, where the climate is now, and 


probably always has been, tropical. Up to the present, it is, 
accordingly, impossible to account satisfactorily for the disappear- 
ance of all the larger forms from among the mammalian fauna of 
South America. 

Returning to the fauna itself, it may be asserted that before 
the great intrusion of northern forms the mammals of Neogaea 
were far more distinct from those of the rest of the world than 
is the case with the fauna of any other region, with the exception 
of Australasia; and that consequently there can be no hesita- 
tion in allowing this part of the earth's surface to take rank as a 
primary realm. At the time when the Santa Cruz fauna was so 
decidedly marked off there was a much more general similarity 
between the faunas of all the other regions of the world (exclusive 
of Notogaea) than is the case at the present day, and it is this 
antiquity of the differentiation of the Neogaeic fauna that supports 
so strongly its claim to distinctness. 

It has been suggested that the first land-connection between 
South and North America was probably of limited extent or short 
duration ; and some evidence of a later separation between the 
two areas is afforded by the beds of marine shells already mentioned 
as occurring in the upper part of the Pampean deposits ; these 
beds marking an epoch of submergence of a considerable portion 
of the area 1 . Subsequently to this final submergence of portions 
of Argentina and Uruguay there was a great upheaval of the 
country, indicated not only by the upraising of the aforesaid marine 
beds, but likewise by that of the sand-dunes fringing most parts of 
the Argentine coast. This upheaval, which has taken place within 
the human period, certainly resulted in the final union of the two 
Americas ; and since its date there has probably continued to be 
a greater and greater admixture of the two originally distinct 
faunas, so far as climatic conditions have permitted. It is, how- 
ever, not a little curious that some of the original northern types, 
such as the vicunas and guanacos, have entirely died out in their 
original habitat, to flourish only in the southern half of the con- 

1 There is some evidence to show that the isthmus of Panama was never 
completely submerged after the Pliocene, but it may have been so narrow as 
not to allow of much migration of the fauna. 


Hitherto especial stress has been laid on the fossil mammals 
of Neogaea as entitling the tract to form a primary Distinctness 
realm, on account of the distinctness of its fauna of the existing 
from that of the rest of the world during a consider- 
able portion of the Tertiary epoch. Even at the present day, 
however, when, as already shown, its mammalian fauna contains a 
very large admixture of types which have immigrated from the 
north at a comparatively recent epoch, it still stands widely apart 
from other regions. On this point may be quoted the admirable 
summary given in "Island Life" 1 by Dr Wallace, who writes that 
among the peculiar mammals we have " the prehensile-tailed 
monkeys and the marmosets, the blood-sucking bats, the coati- 
mundis, the peccaries, the llamas and alpacas [vicunas and 
guanacos], chinchillas, the agutis, the sloths, the armadillos, and 
the ant-eaters ; a series of types more varied and more distinct 
from those of the rest of the world than any other continent can 
boast of. Among birds we have the charming sugar-birds, forming 
the family C&rebidce, the immense and wonderfully varied group 
of tanagers (Tanagridce), the exquisite little manakins and the 
gorgeously-coloured chatterers ( Cotingidce) ; the host of tree- 
creepers of the family Dendrocolaptidce, the wonderful toucans 
(Rhamphastid<z\ the puff-birds (Bucconidcz), jacamars (Galbu- 
lid<z), todies ( Todidce), and motmots (Momotida) ; the marvellous 
assemblage of four hundred distinct kinds of humming-birds 
(Trochilidce\ the gorgeous macaws (Ara), the curassows (Cracida), 
the trumpeters (Psophiidcf), and the sun-bitterns (Eurypygida). 
Here again there is no other continent or region that can produce 
such an assemblage of remarkable and perfectly distinct groups of 
birds ; and no less wonderful is its richness in species, since these 
fully equal, if they do not surpass, those of the two great tropical 
regions of the Eastern Hemisphere (the Ethiopian and the 
Oriental) combined." Not less noteworthy among the birds are 
the screamers (PalctTiiedeidcz) ; the tinamus ( Tinamida), which 
while outwardly resembling game-birds, agree with the struthious 
birds in the structure of their skulls ; and the rheas (Rheida), or 
South American ostriches, whose nearest allies are the true 

1 Pages 50, 51. In this quotation the scientific names of some of the groups 
have been added. 


ostriches of the Old World. The hoatzin (Opisthocomus\ the oil- 
bird (Steatornis\ and the boat-bill (Cancroma) are likewise 
peculiar Neogaeic types. Still more remarkable is the solitary 
Andean survival (Ccenolestes] of the diprotodont marsupials of 
the Santa Cruz epoch. A curious feature of the Neogaean 
forest-mammals whether they belong to the old fauna or the 
new is the frequency of prehensile power in the tail. Not only 
does this occur in several genera of monkeys, but also in Cerco- 
leptes, Synetheres, Chcetomys, Capromys prehensilis, Cydoturus, 
Didelphys, and Ccenolestes. A parallel is to be found elsewhere 
only in Australia. 

After referring to the deficiency of the many types of 
mammals alluded to in an earlier paragraph of the present 
chapter, the author adds that "Among birds we have to notice the 
absence of tits, true flycatchers, shrikes, sun-birds, starlings, larks 
(except a solitary species in the Andes), rollers, bee-eaters, and 
pheasants, while warblers are very scarce, and the almost cosmo- 
politan wagtails are represented by a single species of pipit 

Whether, therefore, we consider its richness in peculiar forms of 
animal life, its enormous variety of species, its numerous defici- 
encies as compared with other parts of the world, or the prevalence 
of a low type of organisation among its higher animals, the 
Neotropical region stands out as undoubtedly the most remark- 
able of the great zoological regions of the earth." 

The distinctness is, however, by no means confined to 
mammals and birds. Of the land molluscs, Mr A. H. Cooke 1 
writes that they present a marked contrast to those of North 
America. "Instead of being scanty, they are exceedingly 
abundant; instead of being small and obscure, they are among 
the largest in size, most brilliant in colour, and most singular 
in shape that are known to exist. At the same time they 
are, as a whole, isolated in type, and exhibit but little relation 
with the Mollusca of any other region." 

Having arrived at the conclusion that the original Neogseic 
mammalian fauna, exemplified in the Santa Cruz 

Origin of the 

Santa Cruz beds, has not been derived from North America, 
it remains to endeavour to account for its origin. 
1 The Cambridge Natural History Mollusca, p. 342 (1895). 


This, however, is a difficult and perplexing subject which it is 
scarcely possible to explain fully in the present imperfect state of 
palaeontological knowledge. 

With regard to the marsupials of an Australian type, it has 
already been stated in the preceding chapter 1 that these appear to 
have been derived from Notogaea by means of a southern land- 
bridge. The hypothesis there suggested is that the immigration 
has taken place via the Antarctic continent, probably across the 
southern Pacific 2 . It is known that shallow water extends from 
southern Patagonia andTierra del Fuego to South Shetland; while 
between Australia and the Antarctic land there are no depths ex- 
ceeding 2000 fathoms. On the other hand it is just possible that 
the connection may have been by way of Polynesia. 

In this place reference may be made to certain very remark- 
able resemblances existing between animals of groups other than 
mammals respectively inhabiting Neogsea and Notogsea. The first 
instance is that of two peculiar families of freshwater fishes, known 
as the Haplochitonida and Galaxiidce. Of these the former has 
one Australian and a second South American genus, while the 
latter is represented by a single genus (Galaxias), common to 
New Zealand, Australia, and the extremity of South America. 
This, however, is by no means all, since one species of the last- 
mentioned genus (G. attenuatus) is found in regions as remote from 
one another as New Zealand and Tasmania on the one hand, and the 
Falkland Islands and the extremity of Patagonia on the other. 
Commenting on this, Dr Wallace remarks 3 , it is impossible to 
believe that a land connection between South America and 
Notogaea could have existed " within the period of existence of 
this one species of fish, not only on account of what we know of 
the permanency of continents and deep oceans ; but because such 
a connection must have led to much more numerous and im- 
portant cases of similarity of natural productions than we actually 
find. Rather must we look to the transport of the ova across the 

1 Supra, p. 55. 

2 Possibly, as suggested below, the connection may have been nearer the 

3 Geographical Distribution of Animals, Vol. I. pp. 401, 402; see also Vol. 
II. pp. 82, 83. 


southern seas, aided perhaps by the Antarctic ice, and a former 
greater extension of South America towards the pole." After 
remarking how such transmission might take place with but little 
extension of the present Antarctic lands, Dr Wallace adds that 
" there is evidently some means by which ova or young fishes are 
carried moderate distances, from the fact that remote Alpine lakes 
and distinct river-systems often have the same species. Glaciers 
and icebergs generally have pools of fresh water on their surfaces ; 
and whatever cause transmits fish to an isolated pond might 
occasionally stock these pools, and by this means introduce the 
fishes of one southern island into another." 

Allowing all due weight to these objections to a land con- 
nection between Notogaea and Neogsea, it seems almost im- 
possible to believe that the transit has taken place in the manner 
suggested by Dr Wallace. 

Another piece of evidence is afforded by some observations of 
Mr F. E. Beddard l in regard to the intimate relationship existing 
between the earth-worms of New Zealand and Eastern Australia 
on the one hand, and those of Patagonia on the other. Without 
committing himself to any theory as to how the communication 
took place, the author is content to say that "the facts seem to 
point to a more recent communication between Patagonia and 
New Zealand than between either of those countries and the Cape 
of Good Hope." 

Assuming a land connection, earth-worms would suggest that 
it was probably not in very high latitudes. Now I have been 
informed on verbal authority that there is a curious similarity 
between the slugs of Patagonia and those of Polynesia. And it is 
probable that the latter tract indicates a subsiding area which 
was formerly connected with Patagonia. Possibly, therefore, there 
may have been a land connection between Patagonia and Australia 
via Polynesia ; and this may have been the line through 
which Neogsea received the Notogseic elements in its fauna. 
Whether it could have existed at a date sufficiently late for the 
passage of the marsupials, it is impossible to say. If it existed, 
it probably allowed only a limited communication between the 

1 Appendix, No. 5, pp. 170, 171. 


Notogaeic and Neogaeic mammals ; and it is easy to imagine that 
the Polynesian mammals (if they existed) were drowned out by 
submergence, as has undoubtedly been the case with many of 
those of the West Indies. In dismissing this part of the subject, 
it may be observed that it appears impossible to adequately 
explain the presence of a Notogaeic element in the fauna of 
Neogaea without the aid of some form of southern land connection; 
although there is not sufficient evidence to show in what latitude 
such connection (or connections) existed. 

Attention must now be directed to the Santa Crucian mammals 
other than marsupials. With the exception of the edentates, which 
probably originated in Neogsea, or possibly in some still more 
southern land, all the evidence points to the whole of these 
being originally of northern derivation ; the ungulates having 
affinities with the ancestral types from which the earlier Tertiary 
perissodactyles were descended, while the rodents have certain 
relationships with the early European members of the order. The 
monkeys again were probably descended from lemuroids ; and 
the solenodons are evidently related to the Old World insectivores. 
It has been shown that this portion of the fauna did not come 
from North America; and it is certainly not derived from Notogaea. 
Accordingly, the only route by which it could have entered is by 
way of Africa. The only marked community between the Ethio- 
pian and Neogaeic faunas as regards mammals relates to the hystri- 
comorphous rodents ; but this community is a very marked one, 
and difficult to explain on any other hypothesis than that of a 
connection between the two areas. The possibility of a close 
community of origin between the toxodonts and the hyraces has 
already been mentioned ; and if it be substantiated, it will be 
highly important. Of course, on the supposition of an African 
origin for the eutherian mammalian fauna of Tertiary Neogaea, 
it must be taken for granted that the ancestral types entered 
Africa long before the progenitors of its modern fauna ; 
although probably not before the ancestors of the Malagasy 
fauna. It may be objected that we ought to find Neogaeic 
Tertiary types of ungulates in Africa ; but we are unacquainted 
with the Tertiary palaeontology of that country, and it is 
quite probable that the peculiar subordinal Neogaeic types of 


ungulates were only developed as such in America itself. Even 
if they ever existed in Africa there is no more reason why they 
should have survived there than in America. As the evidence 
for the presence of Insectivora in the Santa Cruz deposits is 
not very strong, the case of the West Indian solenodons must 
not be pressed too strongly, but their affinity to the tenrecs of 
Madagascar, and the absence of allied types in the North American 
Tertiaries, both point to their having reached Neogsea with the 
other eutherians. 

Regarding a possible connection between Africa and South 
America by way of the Antarctic continent, Dr Blanford 1 writes 
as follows: "Singularly enough, so far as our present information 
as to the depths of the southern oceans goes, there would appear 
at first sight to be less difficulty in supposing a former extension 
of the southern continent to Australia and South America than to 
Africa, the depth as shown on the ' Challenger' charts south of the 
former continents nowhere exceeding 2000 fathoms, whereas to the 
south of Africa there is represented a considerable belt of greater 
depth. But on an Admiralty chart on which all the known deep 
soundings are marked, none are shown south of the southern 
extremity of Africa.... So far as our present information goes, 
the ocean south of the Cape of Good Hope may be no deeper 
than the Mozambique channel, though probably the depth is 
greater in the former case." 

Before discussing certain relationships between the Ethiopian 
fauna and that of Neogaea, it seems advisable to 

"Antarctica" . . . _ 

and the South refer to some recent views as to the existence of 

a S reat southern circumpolar continent in Tertiary- 
Ethiopian times, extending into comparatively low latitudes, 

and connected, at all events temporarily, with 
America, Africa, and Australia. For this continent the name 
Antarctica has been suggested by Dr H. O. Forbes 2 , who urges 
that many of the types of animal life now confined to the southern 
hemisphere have originated there. It is chiefly to show the fallacy 
of these latter views that the subject is referred to here ; palaeon- 
tological evidence clearly proving that several of the groups of 

1 Appendix, No. 8, p. 100. 

2 Appendix, No. 15. 


animals assumed to be essentially southern, really had a northern 
origin. It may be premised that, according to the view of 
Dr Forbes, " Antarctica " followed nearly the 2000 fathom line, 
extending northwards from a circumpolar area by broad expan- 
sions, one to join an old New Zealand continental island (includ- 
ing the Antipodes, Macquarries, New Zealand, and Chatham, Lord 
Howe, Norfolk, and the Kermadec and Fiji Islands); a second to 
East Australia and Tasmania ; a third to the Mascarene and 
adjacent islands ; perhaps one to South Africa ; and finally one 
to South America. 

As regards the marsupials, which are among those considered 
to be southern types, the evidence of the northern Jurassic and 
Cretaceous kinds alluded to in the preceding chapter, coupled 
with the presence of opossums in the Oligocene of the northern 
hemisphere, renders it practically certain that the group did not 
originate in the southern hemisphere. 

Among other groups cited by Dr Forbes as being mainly or 
exclusively southern in their distribution and origin are the parrots 
(Psitiaci) and trogons (Trogonidce). But both these are represented 
in a fossil state in the Oligocene strata of France, and are thus 
shown to have been originally denizens of the temperate regions 
of the northern hemisphere. Take, again, the case of the struthious 
birds, or Ratitse, which although cited as a southern group, are 
represented by an ostrich (Struthio) in the Pliocene of Northern 
India and the Crimea, while the former deposits have yielded 
remains of a three-toed genus allied probably to the emeus and 
cassowaries. Much the same may be said in regard to the giant 
land-tortoises (Testudo\ which although now confined to the 
Galapagos and Mascarene islands, were abundant in Northern 
India, Greece, France, and the United States during the Pliocene, 
and also occur in the French Miocene and Oligocene, as well as 
in the Plistocene deposits of the Maltese caves. The group was 
thus evidently a northern one originally, and as it is unknown in 
the southern hemisphere before the Pampean epoch of Argentina 
and the superficial deposits of Madagascar, its southern migration 
probably did not take place till the Miocene, or even the 
Pliocene. Indeed, the separation of North and South America 
indicates that, if the Galapagos tortoises came from the former 
L. 9 


country, they could not have reached their present habitat till the 
end of the Miocene. 

On the question of the southern or northern origin of some of 
the above-mentioned birds, Professor Huxley 1 , so far back as 1868, 
wrote as follows: "I watch the progress of M. Alphonse Milne- 
Edwards's researches with great interest, to know whether parrots, 
pigeons, Dromceidce (Casuariidas) and Rhtzidce occur in force, or 
at all, among the Miocene birds. If they are absent from the 
Miocene fauna of Arctogaea, it will be necessary to suppose that 
these groups of birds are of sufficiently ancient origin to have been 
separated, even before the Miocene epoch in Austro-Columbia 
(Neogaea) and Australasia, whence they have subsequently 
colonised part of Arctogsea ; while, on the other hand, their 
presence in European Miocene formations will render it possible 
that the colonisation has taken place the other way, and that these 
birds have attained their wonderful multiplicity and diversity of 
forms in Austro-Columbia and Australasia simply in consequence 
of the very favourable nature of the conditions to which they have 
been exposed in that country. 

"I confess I incline to the latter supposition. The distribution 
of Psittacula, for instance, is quite unintelligible to me upon any 
other supposition than that this genus existed in the Miocene 
epoch, or earlier, in Northern Arctogaea, and has thence spread 
into Austro-Columbia, South Africa, India, and the Papuan 
Islands, where it is now found." 

Although the term Psittacula has now been restricted so as to 
include only the Neogaeic forms, this passage is almost prophetic ; 
both parrots and pigeons having, as already stated, been dis- 
covered in the French Oligocene, while the Australian and 
probably the South American ratite birds appear to have had an 
Indian forerunner. And here it may be mentioned that the 
South American ostriches (Rhea] which are primitive types allied 
to the ostrich, would seem to have made their way into Neogaea 
via Africa, as there are no traces of ancestral forms in the North 
American Tertiaries. 

On the other hand, there is considerable probability that the 

1 Appendix, No. 18, p. 319. 


penguins (Spheniscidcz), which present a relation to other birds 
somewhat analogous to that exhibited by the edentates to other 
mammals, having no apparent affinity with any group -may prove 
to be an exception to the rule of the northern origin of most of the 
existing southern types of terrestrial vertebrates, since they are 
quite unknown in the north, and occur fossil both in New Zealand 
and Patagonia. ', / 

Another marked instance of the northern origin of southern 
types is afforded by the side-necked, or pleurodiran Chelonia, 
which although now restricted to the more southern parts of the 
globe, were abundant during Secondary and early Tertiary times 
throughout the northern hemisphere. A striking example of this 
is shown in the family Pelomednsida, whose existing representatives 
are confined to Africa, Madagascar, and South America. Among 
these, two out of three genera, namely Sternothoerus and Pelomedusa 
are found in Ethiopian Africa and Madagascar, one of them also 
ranging into the Sinai tic peninsula ; while the third (Podocnemis} 
has five species in South America and a sixth in Madagascar. 
There occurs, however, in the upper Cretaceous of the United 
States the allied extinct genus Bothremys, and Podocnemis itself is 
represented in the London Clay and the Eocene of the Punjab. 
Here the inference would seem to be that the latter genus 
originated in the northern half of the Old World, passed by way of 
India into Madagascar and Africa, and thence by a southern route 
into Neoggea. Even if this particular genus occurred in the early 
Eocene of North America, which it does not, it could scarcely have 
crossed the sea into South America ; and the migration can hardly 
have taken place since the union of the two continents. In 
commenting on the distribution of Podocnemis Dr Blanford 1 
observes that as the incursion of more modern types into Africa 
appears to have driven out many of the older, it is in Madagascar 
that traces of the relationship of the modern fauna to that of 
Neogaea should be looked for. One such instance is the occur- 
rence there of Podocnemis^ and a second that of the Centetidcz in 
that island. Perhaps the occurrence of sucker-footed bats only in 
Brazil where they are represented by the single species of 

1 Appendix, No. 8, p. 101. 



Thyropoda, and in Madagascar, where there is the sole member of 
the allied genus Myxopoda^ may be an analogous instance. 

In the second family of the pleurodiran Chelonia, the Chelyidcz, 
which are now restricted to South America and Australia (the 
genera in the two areas being distinct), there is at present no 
evidence of the derivation of the Australian forms, but of the 
Neogaeic types Platcmys is represented in the Cretaceous of the 
United States, and Hydraspis in the Eocene of Bombay. Although 
the northern origin of the family is thus proved, the explanation 
of how the existing forms attained their present distribution is 
very difficult. Possibly Hydraspis may have reached South 
America by way of Africa; but it is difficult to believe, in the 
absence of its remains, that Platemys survived in North America 
till the late Miocene communication with Neogaea was estab- 

Among snakes, the boas of the genera Corallus and Boa are 
confined to South America and Madagascar, and thus have 
precisely the same distribution as Podocnemis. Now although true 
boas are unknown as fossils, the allied extinct genus Paleryx 
occurs in the European Oligocene, thus pointing to the northern 
origin of the group, which has probably reached South America by 
way of Madagascar and Africa. 

Another remarkable case is afforded by the limbless lizards of 
the family Amphisbcenidcz, which are now almost equally divided 
between South America and South Africa, although one genus 
extends into the Mediterranean area, and two are found in North 
America ; the two genera Amphisb&na and Anops being common 
to South America and Africa, while the northern ones are different. 
The northern origin of the family is, however, indicated by the re- 
cent discovery of fossil forms 1 in the White River Oligocene of the 
United States. Here the evidence strongly points to a southern 
connection between Neogaea and Africa; Tertiary forms having 
probably existed in Europe or Asia as well as in North America. 
A second instance that may be cited among lizards is the family 
of the Iguanidcz, which while now mainly Neogaeic, has represen- 
tatives in the warmer parts of North America, and also includes 

1 Baur, American Naturalist, Vol. xxvii. p. 998 (1893). 


two outlying genera in Madagascar, and a third in the Fiji 
and Friendly Islands. But fossil iguanas occur in the French 
Oligocene, and it may hence be suggested that the group may 
have reached Neogaea via Madagascar and Africa; while if 
the connection between Patagonia and Polynesia alluded to above 
were substantiated, the origin of the Polynesian forms could be 
accounted for. 

During the middle portion of the Secondary period a very 
curious resemblance between the fauna of Ethiopia and Neogaea 
is exhibited by the occurrence in both of certain very peculiar 
reptiles known as Mesosaurns (Stereosternum), which have been 
referred to the Sauropterygia. Remains of these reptiles have 
been obtained at San Paolo in Brazil, and in Griqualand West 
and other parts of South Africa, but nowhere else ; and, although 
the type may be of northern origin, this curious distribution 
apparently points strongly to a connection between Africa and 
South America as far back as the Secondary epoch. This con- 
nection, as pointed out by Neumayr 1 , was, however, probably by 
way of the Atlantic. 

Somewhat similar relationships to those of living reptiles are 
exhibited by fishes, among which the ffaplochitonidce and 
Galaxiida have been already mentioned. Very remarkable is the 
case of the lung-fishes Lepidosirenidce, where there is a very close 
relationship between the West African Protopterus and Lepidosiren 
of Brazil and Paraguay ; the Australian Ceratodus being markedly 
distinct from both. Although the two former are unknown as 
fossils, teeth of the latter are abundant in the Trias and Jurassic of 
Europe, India, South Africa, and the United States ; while during 
the Palaeozoic era extinct families of the subclass (Dipnoi) were 
abundant in the northern hemisphere. Clearly, then, the group was 
originally northern in origin; and Ceratodus apparently migrated 
south both into Africa and Australia. Taking into account the 
Cretaceous separation of North and South America, and the close 
alliance between Lepidosiren and Protopterus, it is, however, 
difficult to see how the latter reached its present habitat except by 
way of Africa. If this be so, and the connection between South 

1 Vide supra, p. IT 8, note. 


Africa and South America in Tertiary times was only in high 
latitudes, a warm epoch in the southern hemisphere must have 
been necessary for the passage of such tropical forms. It might 
be urged that as Ceratodus dates from the Trias, the other two 
genera might have reached their present habitats at a very 
distant epoch ; but their specialisation is against their antiquity. 
Another family which is essentially southern is that of the Osteo- 
glossidce, represented by Arapaima of the Brazilian rivers, and 
Osteoglossum, with one species from Brazil and the Guianas, a 
second from Sumatra, and two others from Australia. But the 
northern origin of the family is indicated by the occurrence of the 
extinct genus Dapedoglossus in the Eocene of Wyoming. Here 
there is a presumption that Osteoglossum originated in Asia, from 
which it passed in one direction by way of Malaysia to Australia, 
and in another through Africa to South America. Two other 
families of freshwater fishes have a somewhat similar distribution ; 
the first being the Chromidid<z y which includes spiny fishes mainly 
characteristic of tropical America and Africa, but extending 
eastwards into Syria, and sparingly represented in Southern India 
and Ceylon. In a fossil state they occur in the Cretaceous of 
Syria ; and, although none of the genera are common to the two 
continents, they are highly suggestive of a connection between 
Africa and South America. The second family is that of the 
Characiniidce, comprising fish more nearly allied to the carps, and 
now exclusively confined to tropical America and Africa. Although 
the palaeontological record is a blank, this can scarcely be taken 
as a sufficient indication that the family has always been a 
southern one. 

From the foregoing facts it may be considered that the as- 
sumption of an Antarctic continent is unnecessary 
to explain the origin of the many forms of vertebrate 
life which are now exclusively or mainly southern ; nearly all of 
these, with the exception of the edentates and penguins, being of 
northern derivation, and thus apparently showing a southern 
migration of the older forms of life. The Cretaceous and Tertiary 
break between North and South America appears, however, to 
have prevented the occurrence of such migration in the western 
hemisphere till the close of the Miocene : and it is accordingly 


necessary to look elsewhere for the origin of the Neogaeic fauna. 
That Africa has been the great feeder appears the most probable 
explanation ; although in the case of the marsupials it seems 
necessary to look to Notogaea as the point of origin. Clearly, 
however, the presumed connections between Neogaea, Notogsea, 
and Africa have not been very continuous or- very extensive 
in Tertiary times, or the faunas of these areas would have been 
more alike than they are ; and this suggests that the northern 
extension of Antarctica has not been so great as has been 
supposed. Whether the presumed connection between Notogaea 
and Neogaea has taken place by way of Antarctica or Polynesia 
may be left an open question. With regard to Africa, the recent 
researches of Dr Gregory 1 on the West Indian corals, in the 
course of which it is urged that a shallow-water connection "ex- 
tended across the Central Atlantic in at latest Miocene times," 
while the southward extension of the Atlantic is a comparatively 
recent feature, indicate the possibility that the land-connection 
which existed in Jurassic times between Brazil and Western 
Africa may have persisted till the Tertiary era. 

As already mentioned, the Neogaeic realm includes but a 
single region the Neotropical : and in this four 

,..,,..,, , Sub-regions. 

sub-regions have been defined, and are named as 
follows. Firstly we have the Brazilian sub-region, which includes 
not only Brazil, but likewise the Guianas, Venezuela, Colombia, 
Ecuador, Paraguay, and those portions of Peru and Bolivia lying 
on the Brazilian side of the Andes, together with the eastern 
slopes of that portion of the great mountain-chain itself. This is 
essentially an area of dense tropical forests, locally interspersed 
with open pastures, or "campos." The second is the Chilian 
sub-region, comprising Chili, Argentina proper, Uruguay, Pata- 
gonia, and such portions of Peru and Bolivia as are not included 
in the preceding. It is chiefly an area of open plains and 
pampas, although including the high Andes. Thirdly, there is 
the Mexican sub-region, which embraces the isthmus of Panamk, 
Central America, and Southern Mexico, and may be regarded to 
a great extent as a transitional tract between the typical Neo- 

1 Quart. Journ. GcoL Soc. Vol. LI. pp. 306 307 (1895). 


tropical and the Sonoran regions. Lastly, the Antillean sub- 
region includes the West Indian Islands, exclusive of Trinidad, 
which for zoological purposes may be regarded as part and parcel 
of the South American continent. 

From the survey of the fossil forms it has been shown that 
during the Plistocene epoch the mammalian fauna of the Chilian 
and Brazilian sub-regions was similar, and it may consequently be 
inferred that the present differentiation of the two areas in this 
respect is a comparatively modern feature, probably due to 
the disappearance of the forests from the Argentine. At the 
present time the mammalian fauna of the Brazilian sub-region is 
essentially that of the Neotropical region as a whole, nearly all the 
characteristic groups being present within its limits, while several 
are almost or quite peculiar to it. Among the latter, the great 
ant-eater (Myrmecophaga) is practically confined to this sub-region \ 
while most of the sloths and marmosets are limited to it, although 
a few extend northwards into or through the isthmus. The pacas 
(Ccelogenys), and the giant armadillo (Priodon} the sole repre- 
sentative of its genus are likewise restricted to this tract ; as 
is the bush-dog (Icticyon), and also one genus of tree-porcupines 
( Chcetomys], while most of the spiny rats (Echinomys and Loncheres) 
are confined to it. The carpincho (Ifydroc/icerus) the largest 
of living rodents likewise chiefly pertains to the Brazilian 
region, although extending southwards into Uruguay. The 
American monkeys are also very abundantly represented here ; 
the genera Lagothrix, Pithetia, Brachyurus, Brachy teles (Eriodes), 
and Callithrix being restricted to it. Among the forms that are 
unrepresented, may be mentioned guanacos and vicunas (Lama), 
viscachas (Lagostomus), the Patagonian cavy (Dolichotis), and 
chinchillas (Eriomys and Lagidium). 

The Mexican or Central American sub-region differs chiefly 
from the last by the paucity of the essentially Neotropical forms, 
and the large mingling of Arctogaeic types ; among the latter, 
the shrews (Soriridce), a pouched rat (Heteromys}^ and the caxo- 
mistle (Bassariscus) being noticeable. In the Chilian sub-region 
marmosets, monkeys, sloths, tapirs, and peccaries are wanting; 

1 According to Senor Figueira it just enters Uruguay. 


while the carpincho, as already mentioned, only borders on it in 
Uruguay. Among the characteristic types are prominent the 
vicunas of the Andes, the guanacos of the Argentine pampas and 
Patagonia, the spectacled bear of the Andes, the chinchillas of the 
same elevated regions, together with the aquatic coypu (Myopo- 
tamus}, the burrowing viscacha, and the cursorial Patagonian 
cavy ; all the three latter being plain-dwelling forms. Armadillos 
are abundant; and among these the sub-family represented by the 
beautiful little pichiciagos, or fairy-armadillos (Chlamydophorus) 
is peculiar, one of the two species inhabiting open plains near 
Mendoza, in the Argentine, while the second (regarded by some 
as a distinct genus) is found in the Bolivian highlands. 

The Antillean, or West Indian sub-region, which comprises the 
West Indian Islands (exclusive of Trinidad, Tobago, and some of 
the adjacent islets, which are zoologically a part of continental 
South America), differs widely from the other three by the extreme 
poverty of its mammalian fauna; monkeys, marmosets, carnivores, 
and edentates being wanting, and the class mainly represented by 
bats, insectivores, and rodents, although a species of aguti (Dasy- 
procta antilliensis) is found in the islands of St Vincent and Santa 
Lucia, in the Lesser Antilles group, as also in Tobago. In 
addition to a single species of white-footed mouse (Sitomys) said to 
inhabit Hayti and Martinique, and which may also occur in some 
of the other islands, the West Indian sub-region is especially 
characterised by the large arboreal rodents known as hutias, which, 
while belonging to the family Octodontidce, represent two genera 
totally unknown elsewhere. Of these, the genus Plagiodon has 
but a single species, confined to Hayti and Jamaica; although in 
the allied Capromys three existing species are found in Cuba, and 
the fourth in Jamaica, the extinct kind occurring in the former 
island 1 . The nearest relative of the hutias appears to be the 
South American coypu, but the group seems also to show affini- 
ties with the porcupines. From caves in the small island of 
Anguilla, at the northern extremity of the Lesser Antilles group 
have been obtained remains of a large extinct beaver-like rodent 
known as Amblyrhiza (Loxomylus), which has also been recorded 

1 Chapman, Bull. Amer. Mus. Vol. IV. p. 314 (1892). 


from the Pliocene of Argentina a fact of importance as serving 
to connect the Antillean fauna with that of the mainland. As 
mentioned above, this genus belongs to a family (Castoroididce) 
typified by the extinct Castoroides of the Plistocene of Ohio and 
Georgia with species which rivalled a bear in point of size. The 
other Antillean native mammals (exclusive of the bats, to which it 
will be unnecessary to refer) are the two species of the genus 
Solenodon respectively inhabiting Cuba and Hayti, and constitut- 
ing by themselves a separate family among the Insectivora. It 
has been already mentioned that the nearest allies of these strange 
creatures are the tenrecs (Centetidce) of Madagascar; and thus both 
are probably derived from unknown extinct insectivores formerly 
inhabiting the northern hemisphere. As Jamaica and probably 
several other of the West Indian islands contain large masses of 
sedimentary deposits of Tertiary age, it is probable that they 
come under the denomination of continental islands; and there 
seems little doubt, from the evidence of their mammals alone, that 
they have been connected with the mainland 1 . Dr Wallace is of 
opinion that "originally they probably formed part of Central 
America, and may have been united with Yucatan and Honduras 
in one extensive tropical land. But their separation from the 
continent took place at a remote period, and they have since been 
broken up into numerous islands, which have probably undergone 
much submergence in recent times. This has led to that poverty 
of the higher forms of life, combined with the remarkable similarity 
which now characterises them ; while their fauna still preserves a 
sufficient resemblance to that of Central America to indicate its 
origin." Recently, the connection of the West Indies with the 
mainland has been worked out more fully by Mr J. W. Spencer 2 , 
who, from observations made on the buried river channels so 
numerous in some of the islands, concludes that there have been 
several epochs of connection with the continent, one of which was 
so late in date as the Plistocene epoch. The extinction in the 
islands of the great majority of the mammals of the continent 
is attributed to drowning. 

1 This is not the opinion of Dr A. Agassiz, who regards them as oceanic 

2 Geological Magazine, 1894, pp. 448 451. 


Here a presumed connection between North and South 
America by way of the West Indies must be referred to, the 
evidence in favour of which has been summarised by Dr J. W. 
Gregory 1 as follows: "It is not at all certain that when the 
isthmus of Panama was submerged there was a free communica- 
tion between the Atlantic and Pacific Oceans. The Caribbean 
Sea may then have been a gulf from the Pacific, separated from 
the Atlantic by the land area of the hypothetical 'Antillia.' That 
there was once a connection between North and South America 
along the chain of the Windward Islands, Cuba, the Bahamas, and 
Florida is not improbable. Evidence for this, either in whole or 
in part, has been advanced by De Castro and others. Further 
evidence could be adduced from the study of the land-shells, and 
also from the remarkable distribution of Peripatus. That Cuba 
was once connected with Yucatan and Florida is almost certain ; 
that this connection was in existence in the Pliocene, and probably 
also in the Plistocene, is shown by the evidence collected by De 
Castro. That the area of the Windward Islands was occupied by 
land in the lower Tertiary is also most probable. But this was all 
submerged at the period when the Oceanic [Miocene] deposits of 
Barbados were laid down. There is no adequate evidence to 
show that at any time after this was there more land in this region 
than there is at present." With regard to the land-shells, Mr 
A. H. Cooke 2 writes "that a certain number of the characteristic 
North American genera are found in the Antillean sub-region, 
indicating a former connection, more or less intimate, between the 
West Indies and the mainland.... A small amount of South Ameri- 
can influence is perceptible throughout the Antilles, chiefly in the 
occurrence of a few species of Bulimulus and Simpulopsis. The 
South American element may have strayed into the sub-region by 
three distinct routes : (i) by way of Trinidad, Tobago, and the 
islands northward ; (2) by a north-westerly extension of Honduras 
towards Jamaica, forming a series of islands, of which the Rosalind 
and Pedro banks are perhaps the remains; (3) by a similar 
approximation of the peninsula of Yucatan and the western 
extremity of Cuba." This seems to indicate that such Antillean 

1 Quart. Journ. Geol. Soc. Vol. LI. p. 305 (1895). 

2 Cambridge Natural History Mollusca, pp. 345, 346 (1895). 


connection as may have existed between North and South 
America was of a very incomplete and transitory nature ; and that 
before the end of the Miocene there was never any route in this 
direction by which mammals passed from the one continent to the 

In this connection it may be mentioned that Dr Hart Merriam l 
has proposed to unite Central America with the West Indies to 
form a separate zoological region the Tropical of equal rank 
with the Sonoran ; but however much may be urged in favour of 
this view, the multiplication of regions is much to be deprecated. 

The practical or entire absence of non-volant native mammals, 
both recent and fossil, from all the other South 

Other Islands. . .....,- . . 

American islands, with the exception of 1 icrra del 
Fuego and the Falklands, properly excludes their consideration 
from this volume, although it is almost essential that a few words 
should be said with regard to the Galapagos group, more especially 
since conflicting views have been expressed concerning their rela- 
tions to the mainland. In respect to Tierra del Fuego and some 
of the adjacent islets, these may really be regarded as a part of 
the continent, since the main dividing channel is extremely 
narrow, and species like the guanaco are common both to the 
islands and the mainland. And although the Falkland Islands 
lie about 350 miles to the eastward of southern Patagonia, yet 
they are separated by a comparatively shallow sea (less than a 
hundred fathoms in depth), and it is thus evident that they were 
connected at no very distant date with the mainland. Of the two 
indigenous mammals, the most remarkable is the Falkland Island 
wolf (Cam's antarcticus], which differs markedly from all the Canidce 
of the mainland, and is apparently closely allied to the North 
American coyote (C. latrans). The other is a species of groove- 
toothed mouse (Rhithrodon). With regard to Fernando Noronha, 
the poverty of its fauna induces Mr Beddard 2 to class it among 
oceanic islands, although there is some affinity between its fauna 
and flora and those of South America and the West Indies. 

Of far more interest are the Galapagos islands, situated on the 
equator, at a distance of about five hundred miles westward of the 

1 Appendix, No. 19, p. 33. 

2 Ibid. No. 5, pp. 190, 207. 


coast of Ecuador. Entirely volcanic in structure, they are sur- 
rounded by a sea of great depth ; and according to the view both 
of Wallace and Darwin, they have never been connected with the 
mainland, while the latter observer is also of opinion that for 
countless ages they have been separated from one another. The 
known mammals include a bat (Atalapha], a rat (Afus), doubtless 
introduced, and a peculiar species of white-footed mouse (Sitomys 
bauri). Of reptiles the islands contain two peculiar genera of 
iguanoid reptiles, and no less than five species of giant tortoises 
belonging to the genus Testudo. The iguanoids are nearly related 
to South American types ; but there are no tortoises now living 
on the mainland, although a large species flourished in Argentina 
during the Pampean epoch, while, as already stated, others are 
now found living in the Mascarene islands, and extinct species 
occur in the middle and later Tertiary deposits of the United 
States, Europe, and Northern India. It may accordingly be 
taken for granted that both the iguanoid lizards and the giant 
tortoises reached the island from the South American mainland ; 
but the question is how did they arrive ? Dr Wallace is of opinion 
that both were transported across the sea, although by what means 
is unknown. This view is, however, disputed by Dr G. Baur 1 , 
who believes that the Galapagos islands were formerly connected 
not only with one another, but likewise with the mainland. He 
observes that if the Galapagos be oceanic islands, their inhabitants 
could only have been introduced by accident from other regions ; 
" but on such a supposition we are absolutely unable to explain 
the harmonious distribution, we cannot explain why every, or 
nearly every, island has its peculiar race or species, not repre- 
sented on any other island. If some animals could be carried 
over hundreds of miles to the islands, why are they not carried 
from one island to the other? But besides that, how could we 
make plain the presence of such peculiar forms as the gigantic 
land-tortoises? According to the elevation theory, we can only 
think of an accidental importation of these tortoises by some 
current, because they are unable to swim. After the islands had 
been elevated out of the sea, it happened once, by a peculiar 

1 Proceedings American Antiquarian Society, Oct. 1891. 


accident, that a land-tortoise was carried over. Alone it could 
not propagate. This was only possible after a similar accident 
imported another specimen of the same species, of the other sex, 
to the same island. Or we could imagine that at the same time 
animals of both sexes were thus accidentally introduced. By this 
we could at least explain the population of a single island. But 
how did all the other islands become populated? To explain 
this we should have to invoke a thousand accidents. The most 
simple explanation is given by the theory of subsidence. All the 
islands were formerly connected with one another, forming a 
single large island ; subsidence kept on, and the single island was 
divided up into several islands. Every island developed, in the 
course of long ages, its peculiar races, because the conditions on 
these different islands were not absolutely identical." 

Further evidence in favour of the same view is adduced by 
Mr W. B. Hemsley 1 , who draws attention to the marked simi- 
larity of the flora of the Galapagos Islands to that of the South 
American mainland. 

The difficulty of accounting for the transport of reptiles like 
land-tortoises across five hundred miles of sea is undoubtedly very 
great ; yet, in the face of their volcanic nature and the depth of 
the surrounding ocean, it is somewhat difficult to accept the view 
that the Galapagos Islands were connected with South America. 
In the paragraph quoted Dr Baur appears to forget that the first 
tortoise carried to these islands may have been a gravid female ; 
and also that if an ancestral species were established on one of 
the islands, there would be nothing very wonderful in individuals 
being carried to some of the others, where they might eventually 
differentiate into distinct specific types. Moreover, it seems quite 
within the bounds of possibility that the original introduction may 
have been effected by means of eggs transported on natural rafts. 
That the Galapagos tortoises were derived originally from equally 
gigantic continental forms, may be taken for granted ; and the 
existence at the present day of such creatures only in these and 
the Mascarene islands is one more instance of the survival in the 
southern hemisphere of ancient types which were formerly abun- 

1 Nature, vol. lii. p. 623 (1895). 


dant on the opposite side of the equator. The practical absence 
of mammals from the Galapagos Islands is of little import one 
way or the other, as they might have been drowned out during the 
subsidence; but perhaps, on the whole, a suspension of judgment 
as to the relation of these islands to the mainland is the wisest 
course to adopt at present. 

Finally, whether the hypotheses that have been advanced in 
the present chapter to explain the origin of the peculiar mammalian 
fauna of Neogaea be substantiated or the reverse, there can be 
little doubt that Dr Wallace has been misled in his statement 
that this area, so " far as we can judge from the remarkable 
characteristics of its fauna and the vast depths of the ocean east 
and west of it, has not during Tertiary, and probably not even 
during Secondary times, been united with any other continent, 
except through the intervention of North America." 



Features of the Arctogceic Fauna Community of earliest Fauna Evidence of 
Secondary Reptiles Puerco Fauna Lemuroids Insectivora Carnivores 
Rodents Ungulates Summary of the characteristics of the Mamma- 
lian Fauna of Arctogsea. 

ARCTOGSEA, or the Arctogaeic realm, includes the whole of the 
countries of the globe which do not come within the limits of 
either the Neogaeic or Notogseic realms, and thus embraces by far 
the greater portion of the land-surface. Nearly the whole of this 
vast tract lies to the northward of the equator ; the only portions 
lying below that line being the southern half of Africa, together 
with Madagascar and some of the Malayan islands. As stated 
in the introductory chapter, the term Arctogaea was originally 
proposed by Professor Huxley 1 , but, although subsequently used 
in one case by Dr Sclater 2 , and at a still later date adopted by 
Dr Blanford 3 , has failed to obtain general recognition. There 
can, however, be no doubt that, so far as mammals at least 
are concerned, the whole of this vast tract is entitled to hold only 
the same relative rank as each of the realms treated of in the two 
preceding chapters; and that if we regard each of the regions 
into which the area under consideration is divided by Sclater and 
Wallace as equivalent to each of those two realms, we have an 
exceedingly unequal series of divisions. Not only have the 
Neogseic and Notogseic realms no species of mammal common to 
one another, but if we eliminate the genus Cants, which is of 
comparatively recent introduction into these areas, we shall find 

1 Appendix, No. 18. 

2 Ibid., No. 27, p. 214. 

3 Ibid., No. 8, pp. 76, 77. 

[CHAP, iv.] ITS UNITY. 145 

that all the genera, and likewise most of the families, together 
with some of the subordinal or even ordinal groups of mammals 
are likewise perfectly different. Were it not also for the compara- 
tively recent union between South and North America, to which 
allusion has been made in the preceding chapter, we should 
likewise find just as well marked a distinction between the 
mammalian fauna of these two countries ; and, as a matter of" 
fact, when we go back to the middle portion of the Tertiary 
epoch, we find such a distinction actually existing. Again, were 
it not for the intermediate connecting Austro-Malayan region, 
which forms, as we have said, a kind of zoological No-man's-land, 
there would be an equally stringent line of division between the 
Notogasic realm and Asia. If, on the other hand, we take the 
different regions of Arctogsea, we find not only a certain number 
of species of mammals common to two or more regions, but 
when we pass back into the Tertiary epoch, the whole faunas of 
several of such regions merge more or less completely into one 
another, instead of becoming more distinct than they are at the 
present day. The lion and the leopard, for instance, are common 
to India and Ethiopian Africa, and during the Plistocene epoch 
ranged over a considerable portion of Europe ; while the range of 
the tiger includes not only India and Ceylon, but likewise a 
considerable portion of Central Asia and China. The caracal 
and the hunting-leopard are also common to India and Africa ; 
the British fox ranges not only over Europe and a large portion of 
Asia north of the Himalaya, but likewise over a part of North 
America; and the common otter is found alike in India and 
Europe. Still more numerous are the species of mammals com- 
mon to Europe, Northern Asia, and North America. 

Recapitulating some of the details given in the introductory 
chapter, it may be observed that by Messrs Sclater and Wallace 
the area here included in the Arctogaeic realm was divided into 
the Nearctic, Palaearctic, and Oriental regions. Professor Newton, 
who was subsequently followed by Dr Heilprin, proposed to 
brigade the first two of these together under the name of the 
Holarctic region. At a still later date Dr Blanford 1 , who as 

1 Appendix, No. 8, p. 76. 
L. IO 


already stated, takes Arctogaea as one of the three primary 
divisions of the globe, proposed to subdivide it as follows, viz.: 

1 . Madagascar. 

2. Africa, south of the tropic of Cancer. 

3. Oriental, South-eastern Asia, and Malayan islands to 

Wallace's line. 

4. Aquilonian, Europe, Asia north of the Himalaya, Africa 

north of the tropic of Cancer, and America north of 
about 45. 

5. Media-Columbian, America, between about 25 and 45 

north latitude. 

The importance of this division was, firstly, the recognition of 
the right of Madagascar and the adjacent islands to form a region by 
themselves ; and, secondly, the separation of the Medio-Columbian 
region from the rest of North America. And it will be noted that, 
if we take away that area, the Aquilonian region corresponds to 
the Holarctic of Newton and Heilprin. A further modification was 
proposed in 1892 by Dr C. H. Merriam 1 , who gave the name of 
Sonoran region to the area corresponding approximately with the 
Medio-Columbian of Dr Blanford, and suggested that the southern 
portion of the Eastern Holarctic region should form a region by 
itself; the name of Boreal region being adopted for what remained 
of the Holarctic after the subtraction of the Sonoran region and 
a corresponding area in the Eastern Hemisphere. 

Although from many points of view the retention of such 
well-known terms as Palsearctic and Nearctic would be a great 
convenience, the close resemblance of the existing mammalian 
fauna of the whole of northern Arctogaea compels us to adopt 
the view that the area forms but a single zoological region. For 
this region the name Holarctic may be retained ; while for the 
southern portion of the old Nearctic region, the term Sonoran is 
the most appropriate. In the Eastern hemisphere the whole of 
that portion of Arctogsea not included in either the Malagasy, 
Ethiopian, or Oriental regions is provisionally included in the 
Holarctic, although when our knowledge of distribution is less 

1 Appendix, No. 19. 


imperfect it may subsequently be found practicable to separate a 
distinct Mediterranean region. 

In an area of such vast extent as the Arctogseic realm, which 
embraces countries from the equator to the most 
northern habitable lands, it is, of course, perfectly the Arctogseic 
unnecessary to say anything as regards climate 
and physical features, and we may accordingly proceed forth- 
with to discuss the leading features of the mammalian fauna as a 

From the Notogaeic realm, in its typical form, Arctogaea is 
distinguished by the absence of monotremes and diprotodont 
marsupials, not only at the present epoch, but so far as we know, 
in past times also. From the Neogaeic realm it is equally well 
differentiated by the absence at the present day of all the peculiar 
Neogaeic types of edentates, and likewise at all epochs of Neo- 
tropical monkeys and marmosets. Such of the former as are 
found in North America are indeed, as we have seen in the last 
chapter, only intruders from the south since the epoch of the 
earlier Pliocene; and in the Miocene the Arctogaeic fauna was 
further distinguished from that of Neogaea by the absence of the 
peculiar subordinal groups of ungulates characteristic of the latter. 
The Insectivora, which with the exception of the solenodons are 
practically wanting in Neogaea, and are unknown in Notogaea 
proper, are abundant in all the regions of this realm. 

We might almost go one step further than this, and say that 
Arctogaea previous to the Pliocene epoch was characterised by 
being the sole habitat of almost all the families of Eutherian 
terrestrial mammals, with the exception of those characteristic of 
the Santa Crucian epoch of Neogaea. But although this would be 
practically true, it would land us in the difficulty that the Ethio- 
pian region would probably have to be excluded from Arctogaea, 
seeing that the higher mammals of the former region are but 
comparatively recent immigrants. Still, it may be stated that 
northern Arctogaea is the original habitat of all the modern types 
of the higher Eutherian mammals. 

Another feature of Arctogaea is the absence at the present day 
of all marsupials except opossums, while these are only sparingly 
represented in its western half. Moreover, with the exception of 



the same family group, which are only known from strata of the 
Oligocene and Miocene epochs, marsupials appear to have been 
absent from a large part of the realm during the Tertiary period, 
although there is reason to believe that during the Eocene they 
must have survived in south-eastern Asia 1 . Among volant 
mammals, bats of the Neogaeic family PhyllostomatidGR* are now 
absent from the whole of the realm, with the exception of a part 
of the Pacific side of North America. Again, to revert to the 
non-volant forms, the Lemuroid suborder of the Primates seems to 
have been absolutely restricted to Arctogaea, at least since the 
Miocene epoch, although it may turn out that the monkeys and 
marmosets of South America may be descended from ancestral 
lemuroids which inhabited that country at an epoch previous 
to the deposition of the Santa Crucian beds. 

To take a comprehensive survey of the whole Secondary and 

Communit Tertiary mammalian faunas of Arctogaea would entail 

of earliest such a mass of palaeontological and anatomical 

detail that it would only weary the majority of our 

readers, and we must accordingly limit ourselves to noticing some 

of the most striking features in the earlier faunas, and then pass 

on to the consideration of some of the more widely spread modern 


In the chapter devoted to the Notogaeic realm, it has been 
already pointed out (p. 51) that during the Jurassic period Europe 
and North America were populated with a fauna of polyprotodont 
marsupials of small size, some of which appear to have been the 
ancestral types from which those now inhabiting the Notogseic 
and Neogaeic realms were derived, while others have disappeared 
entirely. It will be unnecessary to recapitulate the names of the 
more representative of these forms, but it may be stated that while 
the fauna of the lower Jurassic Stonesfield Slate has no equivalent 
in North America, that of the upper Jurassic Purbeck beds of Dor- 
setshire is paralleled in the latter area. Although some difference 
of opinion prevails among palaeontologists as to the identity of the 
American with the European genera, there can be no doubt that 

1 Vide suprd, p. 55. 

2 The genus Necromantis, from the French Oligocene, has been assigned 
to this family. 


many of them are very closely allied indeed, while some are 
probably inseparable. Contemporary with these early marsupials 
were members of the group known as Multituberculata, which are 
probably more or less closely related to the existing monotremes, 
or egg-laying mammals, and form with them the subclass Proto- 
theria. An essential feature of these multituberculates is that 
the molar teeth were divided by one or more grooves into longi- 
tudinal ridges, covered with numerous blunt tubercles ; such 
grooves being very generally two in number in the upper molars, 
while the lower teeth have but a single one. Apparently in all 
cases the extremities of the jaws were armed with a pair of chisel- 
like incisor teeth, behind which in the upper jaw there may have 
been a pair of smaller teeth. Very generally the last premolar 
tooth, as in the English Purbeck genus Plagiaulax, was com- 
pressed and trenchant in shape, with its upper edge regularly 

FIG. 28. RIGHT SIDE OF LOWER JAW OF Plagiaulax. Enlarged. 
p. premolars, m. molars. 

convex, and its sides marked by oblique grooves; but in other 
forms (Polymastodoti) this tooth was of a more tubercular type. 
Without going into disputed questions, it may be stated that this 
group was represented by closely allied forms in the Jurassic of 
both Europe and North America; while it is also known, from 
the evidence of a single genus (Tritylodon)^ to have extended its 
range to South Africa ; this genus also occurring in the European 
Trias, and thus affording another instance of the wide range of the 
earlier faunas. 

Although in Europe the only known traces of mammalian life 
during the succeeding Cretaceous period occur in the Wealden 
beds (which are the immediate successors of the Jurassic Pur- 
becks), in North America a well-developed fauna of polyprotodont 


marsupials and multituberculates is met with in rocks of Cretaceous 
age ; and it is most probable that if suitable freshwater beds were 

FIG. 29. UPPER SURFACE OF SKULL OF Tritylodon. Somewhat reduced. 

extant, the same fauna would be found in Europe. By the 
commencement of the Tertiary epoch, most of this old fauna seems 
to have disappeared; but in the Puerco beds of the United States, 

Natural size and enlarged. 

and the equivalent deposits of Cernays, in the south of France, 
which seem to form a transition between the Secondary and 
Tertiary, the Multituberculata still persisted, and it is noteworthy 
that one genus (Neoplagiaulax] at least was common to the 
northern half of the eastern and western hemispheres. 

It thus appears, so far as the available evidence permits of our 
forming a judgment, that during both the Jurassic and Cretaceous 


epochs a single mammalian fauna was spread over Europe and 
North America. This being so, it is a fair inference that a similar 
fauna characterised a considerable portion of Asia ; while the 
occurrence of the above-mentioned genus Tritylodon points to the 
conclusion that it likewise ranged over Africa. Accordingly, it 
would appear that not only did the whole of Arctogaea then form 
a single zoological realm, but that this realm was indivisible into 

The evidence for this unity is, however, by no means restricted 
to mammals, but is supplemented and extended by vid 
the extinct reptiles of the Secondary epoch of the Secondary 
earth's history. During the Triassic and early 
Jurassic periods there flourished an extinct ordinal group of rep- 
tiles known as the Anomodontia, remarkable for many structural 
resemblances to mammals, and likewise for the peculiarities of 
their dentition. As a well-known example of one section of this 
group may be cited the dicynodonts, in which the teeth were 
reduced, at most, to a single pair of tusks in the upper jaw, the 
remainder of the jaws being ensheathed in horn to form a beak ; 
whereas Galesaurus represents a second section in which the teeth 
simulate those of the carnivorous mammals. These anomodonts 
are known to have been spread over Europe, India, Africa, and 
North America ; the dicynodont types from the three areas first 
named being so alike that there is little question that some of 
them were generically identical. The North American forms, 
which mostly or exclusively come from beds assigned to the 
Permian epoch, do not include dicynodonts, but are allied to 
certain other African families, and are also closely related to their 
European contemporaries. 

If we turn to another order of the same class, namely the 
Dinosauria, as represented by the Iguanodon of Europe and the 
Atlantosaurus of the United States, we find not less well- 
marked similarities in the Jurassic and Cretaceous fauna of the 
whole of Arctogaea, this group being represented by closely allied, 
and in many cases generically identical forms, not only in Europe, 
India, and South Africa, but likewise in Madagascar and South 
America. For instance, in that section of the order known as 
the Sauropoda, which includes the most gigantic forms, and is 


characterised by the presence of large chambers in the sides of the 
vertebrae of the neck and trunk, we find not only that several 
genera, such as Morosaurus, are common to the upper Jurassic 
and lower Cretaceous strata of Europe and the United States ; but 
we also find, one genus (Titanosaurus) in India, Europe, and 
Patagonia, while a second (Bothriospondylus) occurs in countries 
as far apart as England and Madagascar 1 . Again, in the carni- 
vorous or Theropodous section of the order, as typified by the 
English Megalosaurns, we find certain closely allied or identical 
generic types common to Europe and South Africa. Further 
evidence in the same direction is afforded by the discovery in the 
Jurassic of Madagascar of a genus of extinct crocodiles (Steneo- 
saurus) which were abundantly represented during the same epoch 
in Europe. Among the class of fishes we have also the genus 
Ceratodus, now living in Queensland, represented in the Secondary 
rocks of Europe, India, Africa, and North America. 

With regard to the land-fauna of Australia at the same epoch 
we have less evidence ; anomodonts, and, we believe, dinosaurs, 
being unknown from that country. Among the amphibians, how- 
ever, we find in the extinct order of Labyrinthodontia certain 
genera, such as Bothriceps and Micropholis, common to Australia 
and South Africa, both of these being closely allied to the Indian 

This reptilian evidence thus clearly points to the conclusion 
that during the greater part of the Secondary period not only 
had Arctogaea a single widely-spread fauna, but that the same 
fauna was represented in South America, and at least partially in 
Australia. Hence at this date no zoological realms can be distin- 
guished, and it was probably not till late in Cretaceous times that 
Arctogaea was differentiated from the rest of the world as a realm. 
Needless to say, the great continents and islands during the 
epochs in question must have had free communication with one 
another, and it is highly probable, as Dr Blanford 2 suggests, that 
Madagascar then formed a line of connection between Africa and 
India. It is possible that even at the early part of the Secondary era, 

1 Possibly future discoveries may show differences worthy of generic distinc- 
tion between these forms, but this would not affect the general question. 

2 Appendix, No. 8, pp. 88, et seq. 


"when South Africa was united to India via Madagascar on one 
side, and to South America on the other, especially if the Indo- 
Malay continent was also connected with the Australian, there 
may have been a girdle of land, chiefly in low latitudes, round 
nearly three-quarters of the earth's circumference from Peru to 
New Zealand and the Fiji Islands." The vertebrate testimony 
does not, however, countenance the idea that such southern land 
was cut off from Europe and northern Asia by sea. It is true 
that the evidence in favour of such an isolation is afforded by the 
identity of the Carboniferous (Damuda-Talchir) floras of Australia, 
South Africa, Peninsular India, and Central Argentina 1 , and their 
total dissimilarity from those of Europe, Northern Asia, and North 
America ; and it is suggested that the same conditions may have 
prevailed during the Jurassic 2 . This, however, the vertebrate 
evidence certainly does not support ; and hence, while admitting 
the isolation of a great southern (subtropical) continent during 
the Palaeozoic era, it appears probable that since that epoch most 
of the southern lands have been from time to time more or less 
closely connected with those to the north 3 . 

Leaving these difficult problems with the foregoing remarks, 
we pass on to notice briefly the Puerco mammalian 

Puerco Fauna. 

fauna of the United States, which, together with the 
approximately equivalent Cernaysian fauna of Europe, is of especial 
interest as showing a transition between the Cretaceous and 
Tertiary. As we have already said, this fauna includes several 
representatives of the multituberculates, which are essentially a 
Secondary group, and one of which is common to the Cernaysian 
fauna. In addition to these, four orders of eutherian mammals are 
represented, namely the Primates, the Carnivora, the Ungulata, 
and the extinct group known as the Tillodontia. It is, however, 
very noteworthy that in all these orders it is only the lowest 
sections that were in existence during the Puerco epoch. Thus 

1 See F. Kurtz, Rev. Mus. La Plata, Vol. vi. p. 117, and Rec. Geol. Surv. 
India, Vol. xxvni. p. in (1895). 

2 Appendix, No. 8, p. 96. 

3 Dr Blanford writes to me that he believes the Palaeozoic connection be- 
tween South America and South Africa was tropical or subtropical, rather than 
antarctic, and hence that "the evidence for an antarctic continent in upper 
Mesozoic or Tertiary times is very slight indeed." 


all the Primates belong to the lemuroid section, and include no 
monkeys or apes ; and the carnivores are represented solely by 
the extinct creodont group, which differs from the existing members 
of the order by the simpler and more primitive structure of the 
limbs and teeth. The ungulates, again, belong exclusively to two 
extinct suborders, respectively termed the Condylarthra and the 
Amblypoda, both of which are very primitive types, with five-toed 
limbs of simple structure, the former still retaining evidences of 
affinity with the early carnivores. The tillodonts are quite unlike 
any existing forms, having a pair of incisor teeth similar to those 
of rodents in the front of the jaws, while their cheek-teeth recall 
those of the ungulates. 

All the Puerco mammals are characterised by the lowness of 
the crowns of their molar teeth, which carry simple tubercles, 
generally arranged in a triangle ; this type of tooth being known 
as the tritubercular, and occurring in all the orders found in the 
Puerco. It will be unnecessary to mention the names of the 
genera occurring in this horizon, and it will suffice to state that 
while peculiar to these beds, many of them belong to families ' 
characteristic of the overlying Tertiaries. Thus we have the 
Anaptomorphida among the lemuroids, the Arctocyonida, Mesony- 
chidce, Proviverridce, and Miacida in the carnivores, and the 
PhenacodontidcR among the condylarthrous ungulates. The Cernay- 
sian fauna is mostly represented by such fragmentary specimens 
that the determination of the affinities of its members is a matter 
of considerable difficulty; but the forms were all more or less 
nearly allied to those of the Puerco, and the creodont genus 
Dissacus is common to the two formations ; while Arctocyon, 
which is met with in the Cernaysian, also occurs in higher horizons 
both in Europe and America. 

A very remarkable fact connected with the Puerco fauna is 
that out of the 39 generic types by which it is represented, only 
eight are followed by analogous forms in the overlying Wahsatch 
beds, three of which became extinct in the still higher Bridger 
deposits. This leads Messrs Osborn and Earle to the conclusion 
that this early mammalian fauna was a kind of failure as regards 
development, and that only a few of its less specialised members 
persisted to give rise to the mammals of later periods. 


With the Puerco and Cernaysian faunas we take leave of the 
Secondary multituberculates, and as we ascend in 

. . ... . Lemuroids. 

the Tertiary series we find a gradual and progressive 
modification of the eutherian mammals towards the modern types. 
In the ungulates especially the modification displays itself in the 
more complex structure of the molar teeth, and in the reduction of 
the number of toes ; the culmination of the latter line of develop- 
ment being reached in the modern horses among the perissodactyle 
section of the order, and in the ruminants among the artiodactyles. 
As regards the molar teeth, the chief features are a lengthening of 
the crowns in the more specialised later forms, accompanied by 
complex infoldings of the surface of the crown and sides, whereby 
the short-crowned, or brachydont type, as exemplified in the tapirs, 
has developed into the tall, or hypsodont type characteristic of the 
horses. Instead, however, of tracing the succession of the various 
faunas, it will suit our present purpose better to refer to the 
distribution of the more widely spread groups which are either 
characteristic of Arctogsea as a whole, or which were common to 
that realm together with Neogsea during the Plistocene period. 

The lemuroids, which are at present unknown beyond the 
limits of this realm, are first met with in the Puerco beds, where 
they are represented by Indrodon, and it is probable that the 
Cernaysian mammal described as Plesiadapis (of which the upper 
cheek-teeth are shown in the annexed figure) belongs to the same 


p. premolars, m. molars. 

group. It will be observed that in the latter genus the molars are 
of the tritubercular type; the same being the case in Anapto- 
morphus of the lower or Wahsatch Eocene of America. In other 
forms, however, as in Hyopsodus and Pelycodus of the lower Eocene 
of North America, and probably also of the European Eocene, as 
well as in Microchcerus of the Oligocene of France and England, 
the upper molar teeth have squared crowns, and thus approximate 


to those of modern lemurs. These early forms differ however from 
the latter in that the first of the three lower premolar teeth does 
not assume the form and functions of a canine. Another well- 


GENUS Microchtxrus. Nat. size and enlarged. 

known European lemuroid is Adapts, of the European Oligocene, 
differing from all living forms in having four pairs of premolar 

With the Oligocene, lemuroids seem to have disappeared from 
western Europe, and they apparently ceased to exist about the 
same date in North America, after which the entire order of the 
Primates is unrepresented in the latter country. At the present 
day, as we shall see, lemuroids are confined to the Malagasy, 
Ethiopian, and Oriental regions ; but at what epoch the southern 
migration took place cannot yet be determined. 

Omitting mention of the bats, we pass on to the Insectivora, 
among which we have the mole family (Talpidce) 
distributed over the whole of the Holarctic as well as 
the Sonoran region, although all the genera but one are distinct 
on the two sides of the Atlantic ; the single common type being 
the shrew-moles (Urotrichus\ which have one species in Japan 
and another in the United States, thus affording an instance of the 
near affinity of the fauna of eastern Asia to that of North America. 
The earliest known fossil forms which have been assigned to the 
typical genus Talpa occur in the upper Oligocene strata of Europe, 
while in the middle Oligocene the family is represented by the 
allied Amphidozotherium (Protalpa). The shrews (Soriddce)^ 
which likewise date from the Oligocene of the Continent, range 
over the whole realm, and also enter the Austro-Malayan region 


as well as the Mexican subregion, although they are represented 
in Madagascar only by a single species of the widely spread genus 
Crocidura. Unknown in America, the latter genus, which includes 
the well-known musk-shrews, is widely spread over Europe, Asia, 
and Africa, extending as far east as Amurland ; but the typical 
genus Sorex is practically confined to the Holarctic region 1 , while 
other genera have a more local distribution. 

With the exception of the civet tribe ( Viverridce] and hyaenas 
(Hyanidce), which are unknown in the New World, 

....... . Carnivora. 

the majority of the existing families of the Carnivora 
have, if we except Notogaea, a cosmopolitan distribution, while in 
many cases this extensive distribution also holds good with regard 
to genera. In Europe the Felidce and Canidce, together with the 
Mustelida seem to have made their first appearance in the lower 
Oligocene, when they were accompanied by the extinct creodonts ; 
while in America the two former are known from the John Day 
group, corresponding to the European Miocene. The bears 
( Ursidce) are, however, a later group, being unknown before the 
Pliocene. Although the whole of the families mentioned above 
are represented in South America at the present day and during 
the Plistocene, they are, as we have seen in the last chapter, 
unknown in the presumably Miocene Tertiaries of Patagonia, and 
they are therefore originally of Arctogaeic origin. Although most 
of the extinct American Tertiary genera of cats are distinct from 
those of Europe, the sabre-toothed tigers (Machcerodus) were 
common to both areas, and likewise ranged into Notogaea; and 
the existing Felis has a similar cosmopolitan distribution. A more 
.specialised sabre-toothed genus (Eusmilus) is likewise common 
to North America and Europe. As examples of extinct American 
cats, we may name Nimravus and Archcelurus ; while the Oligocene 
^Elurictis may be cited as an Old World form. The aforesaid 
distinction between the Oligocene and Miocene Felidcz of North 
America and Europe is, however, an indication that by this date 
the mammalian faunas of Western and Eastern Arctogaea had 
become differentiated to a certain extent, although, as now, there 
were many types common to the two areas. 

1 It has one species in the Sonoran. 


Much the same story is told by the fossil dogs ( Canidce) of the 
two areas. In the Miocene of North America we meet with the 
genus Temnocyon, characterised by the cutting heel of the lower 
carnassial tooth; while the more civet-like Cynodictis appears to be 
confined to the Tertiaries of Europe. The bear-like genus Am- 
phicyon, differing from modern dogs by its plantigrade feet, is 
confined to the European Oligocene and Miocene and lower 
Pliocene, but is represented in the Miocene of America by the 
nearly allied Daphcemis. Through the intervention of the still 
larger Dinocyon of the European Miocene, the foregoing groups are 
intimately connected with the bears (Ursidcz) by means of the 
genus Hycenarctus, which is common to the Miocene and Pliocene 
of Europe and the Pliocene of India; and this suggests that the 
bears are originally an Old World group, which have subsequently 
migrated into America. As to whether true dogs (Cam's) and cats 
(Felis) originated in America or Europe, we have no means of 
deciding 1 . The large weasel family (Mustelidce) calls for no special 
mention here, although its comparative poverty in South America 
proclaims its Arctogaeic origin. 

The community between the mammalian faunas of Eastern 
and Western Arctogaea is perhaps better exemplified by the ex- 
tinct creodont carnivores, since none of the families occurring there 
have been definitely recognised in the South American area. 
Differing from modern carnivores by the absence of a pair of 
differentiated carnassial teeth in each jaw, as well as by the 
scaphoid and lunar bones of the wrist generally remaining 
separate, and by the nearly flat upper surfaces of the astragalus in 
the ankle, these creodonts make their first appearance in the 
Puerco, and mostly died out in the Oligocene, although a few seem 
to have survived till the Miocene. In the typical family Hyczno- 
dontida we find the genus Hycenodon common to the Oligocene of 
both sides of the Atlantic, while the European Pterodon seems to 
have a transatlantic representative in the so-called Hemipsalodon of 

1 Scott, Trans. Amer. Phil. Soc. xvn. p. 75, concludes that the evolution 
of Cam's took place in North America, the ancestry being traced through 
Cynodesmtts of the John Day Beds to Daphcemis of the White River, and 
thence to the creodont Miacis of the Bridger ; Cynodictis forming a lateral 

IV.] RODENTS. 159 

Canada. Oxycena, again, is found both in America and Europe, 
although in a lower horizon in the former than in the latter. In a 
second family (ProviverridcR) the typical genus Proviverra is met 



with in the Bridger Eocene of America, and the French Oligocene; 
while in the Arctocyonidce, in which the upper molar teeth are 
bluntly tritubercular, Arctocyon of the lowest Eocene of Europe is 
represented by two allied genera in the American Puerco, one of 
which has been described as Clcenodon. 

In the rodents there are three more or less widely distributed 
existing families restricted to Arctogaea. Of these, 
the jerboas and their allies (DipodidtzY occur in all 
the regions with the exception of the Malagasy, Oriental, and 
Sonoran, although the genera from the different areas are more 
or less markedly distinct. The other two, namely the picas or 
tailless hares (Lagomyida) and the beavers (Castoridce) are now 
severally represented by a single genus confined to the Holarctic 
region. The picas date from the Oligocene of Europe, and the 
family not improbably originated in eastern Arctogaea ; while the 
beavers have fossil representatives in both hemispheres, with the 
Miocene and Pliocene genus Chalicomys common to the two. 

Although members of the typical genus range into the Neogaeic 
realm as far south as Paraguay, the squirrel family (Sciuridce) may 
be regarded as a typical Arctogaeic one, the ground-squirrels 
(Tamias), marmots (Arctomys) and susliks (Spermophilus} being 
restricted to the Holarctic region, though others range over the 

1 It has been suggested by Dobson that the Dipodidcz are Hystricomorpha. 
This, however, is disproved by Dr Scott (P. Ac. Philad. 1895, pp. 269 286), 
who finds in the Uinta Oligocene genus Protoptychus an ancestral type of the 
family, which thus appears to be of N. American origin. From this family are 
probably descended the Geomyidce. 


whole of the tropical and temperate parts of the realm with the 
exception of Madagascar. Remains of Sperm ophilus and Sciurus 
are met with in the later Tertiaries of Europe'; and the extinct 
Plesiarctomys, which is common to the Oligocene and Miocene of 
Europe and North America, seems to be a connecting form 
between the squirrels and marmots, having upper molar teeth of 
the tritubercular type. 

As regards the cosmopolitan family Muridcz, including the 
rats, voles, lemmings, etc., it will suffice to say that originally it 
was undoubtedly Arctogaeic ; the forms respectively inhabiting the 
Neogaeic and Notogaeic realms being comparatively recent immi- 
grants. Both the subfamilies of the voles (Microtince) and the 
CricetincB are common to the entire Holarctic region ; the latter 
being represented in the eastern half by the hamsters (Cricetus) 
and in the western by the white-footed mice (Sitomys], while they 
are the sole rodents inhabiting Madagascar, and have one species 
in Ethiopian Africa, where there is also the closely allied Deomys, 
forming a subfamily by itself. The cricetines are indeed evidently a 
primitive type, which in the Old World have been largely supplanted 
or driven south by the more specialised. Murince (true rats and mice) ; 
but as these are represented in the Middle Tertiaries of both 
eastern and western Arctogaea, it is difficult to decide which was 
their original habitat. Little need be said in regard to the hares 
(Leporidce), except that they range over the whole of Arctogasa, 
and have two outlying representatives in Neogaea, which are 
doubtless comparatively recent immigrants, although one is known 
to have inhabited Brazil since the Plistocene. 

A not less marked feature of Arctogaea is the absence of most 
of the Neogaeic rodent families noticed in the preceding chapter. 
The existence of members of one of these (Octodontida) in Africa 
is mentioned in the same place 1 , where a reference to the occur- 
rence of allied forms (Theridomyidce] in the European Tertiaries 
will also be found 2 . 

One of the most important features in connection with the 
Arctogaeic ungulates is the total absence of the 

Ungulates. 5 6 

peculiar subordmal groups characteristic of the 
1 See also the chapter on the Ethiopian region. 2 Supra, p. 86. 


South American Tertiaries ; although, as stated in the last chapter, 
there are indications of a distant affinity between these and some 
of the primitive early European perissodactyles. So far as our 
present knowledge goes, both the perissodactyle and artiodactyle 
suborders made their appearance in North America during the 
lower or Wahsatch Eocene ; the former group at least also dating 
from the same epoch in Europe, where the genus Hyracotherium, 
which is common to the Bridger and Wahsatch Eocene, and is one 
of the earliest ancestors of the horses, occurs in the London Clay. 
Commencing with the Artiodactyla, or even-toed group 
characterised by the toes corresponding to the third and fourth 
of the typical pentedactyle limb being symmetrical to a line drawn 
between them and taking into consideration only such families as 
have a wide distribution, we have first to do with certain extinct 
types which serve to connect the pigs with the ruminants. The 
most pig-like of these are the animals forming the family Chcero- 
potamidce, characterised by their broad upper molar teeth carrying 
five blunt tubercles, three of which are on the front half of the 
crown. Although the typical genus Chosropotamus appears to have 
been confined to the lower Oligocene of Europe, the much larger 
animals known as Elotherium (fig. 35) were common to the upper 
Oligocene of both hemispheres. Nearly allied is the family of the 
Anthracotheriidce, in which the low tubercles of the molars assume 


a more or less crescentic, or selenodont structure, thus foreshadow- 
ing those of the ruminants. Here, again, the typical genus is 
restricted to the Old World, but the nearly allied Ancodus (Hyo- 





potamus] has the same approximate distribution as Elotherium, 
although it also occurs in the Miocene of northern India, together 
with Anthracotherium and a genus (Tetraconodon] closely allied to 

A group of smaller Eocene and Oligocene mammals, con- 
stituting the family Ccenotheriidce, differ from the preceding in 
that their upper molars have two cusps on the front, and three on 
the hinder half of the crown. Here none of the genera are com- 
mon to the two sides of the Atlantic, Ccenotherium and Dichobumis 
being European, while the latter is represented in the Bridger 
Eocene by the closely allied Homacodon. It has been suggested 
that this group includes the ancestors of the camel tribe. 

The latter group (Camelidce), now represented only by the 
camels (Camelus) in the Old World, and the guanacos, vicunas, 
and their domesticated allies the llamas in South America, was 
formerly widely distributed in Arctogaea, which was doubtless its 
original home, since, as we have seen in the preceding chapter, 
the llamas and their allies are but comparatively recent immigrants 
into Neogaea. Camelus itself is represented in a fossil state in the 
Pliocene of northern India and the Plistocene of Algeria ; but no 
other Old World representatives of the family are known. Very 
different, however, is the case with North America, where, from 
the Plistocene downwards, we meet with a host of extinct types, 
such as Pliauchenia, Procamelus, Protolabis, etc., gradually con- 
necting the existing forms with a small animal from the middle 
Oligocene known as Poebrotherium, which exhibits many very 
generalised characters. A still earlier representative of the family 
in Leptotragulus, of the lower or Uinta Oligocene, which itself may 
be sprung from the aforesaid Homacodon of the underlying Bridger 
beds. It is thus perfectly evident that the cameloids were ori- 
ginally a N. American group. One branch crossed the area now 
occupied by Bering Strait to found the camels of the Old World ; 
while, probably at a later date, a second branch passed over the 
isthmus of Panama to persist in the guanacos, vicunas, and 
llamas of South America. The disappearance of the whole group 
from the northern half of the New World is a very remarkable fact, 
but is paralleled by that of the elephants, rhinoceroses, lemurs, and 
several other groups. 

II 2 


The Tragulidce, or chevrotains, which form a group distinct 
both from the cameloids and the true ruminants, are now confined 
to the Oriental and Ethiopian regions, being represented in the 
former area by the true chevrotains (Tragulus), and in the latter 
by the single existing species of water-chevrotain (Dorcatheriuni). 
Representatives of the latter genus occur, however, in the Miocene 
and Pliocene strata of Europe ; while in the Oligocene we meet 
with the more generalised extinct genera Prodremotherium and 
Bachitheriuwi. In North America the group is but poorly repre- 
sented, and apparently confined to the White River Oligocene, 
where we find two types described under the names of Leptomeryx 
and Hypertragulus ; the latter differing from the existing forms by 
having both the third and fourth metacarpals and the correspond- 
ing metatarsals separate, instead of being fused together to form a 
cannon-bone. It is difficult to decide whether the group was 
originally an Old or a New World one ; but, on the whole, it is 
probable that it originated in the former area. 

We now come to the true ruminants, or Pecora, forming the 
most specialised group of all the artiodactyle section of the ungu- 
lates, and characterised by the completely crescentic, or selenodont 
conformation of the columns of their molar teeth, which are 
frequently of great height ; and likewise by the fusion of the third 
and fourth metacarpals and metatarsals into a cannon-bone, and 
by the imperfect development or disappearance of the lateral 
metacarpals and metatarsals. In the family of the Cervida the 
typical deer, or those included in the genus Cervus, are almost 
exclusively Arctogseic 1 , being found in all the regions of the realm, 
except the Sonoran, Ethiopian, and Malagasy. The reindeer 
(Rangifer] and elk (Alces] are also solely Arctogseic forms, but 
have a more restricted range, being confined to the more northern 
portions of the Holarctic region. A more striking case is afforded 
by the hollow-horned ruminants, or Bovidce, the whole of the 
numerous members of which are confined to the realm under con- 
sideration, with the sole exception of the anoa (Bos depressicornis] 
of Celebes ; and even the latter, as we have seen in an earlier 
chapter, is very closely related to certain extinct Indian forms. It 

1 The only exception is Cervus timoriensis, which may have been introduced 
into its present habitat. 


may be noticed, however, that Bovidce are much more numerously 
represented in Eastern than in Western Arctogaea ; the latter area 
having only the genera Bos, Ovibos, Ovis, and Haploceros, each 
with a single species confined to the more northern portions of 
the continent ; the last genus being peculiar to this area. 

We now pass to the perissodactyle, or odd-toed ungulates, 
which while agreeing with the artiodactyle section in the inter- 
locking arrangement of the bones of the wrist and ankle joints, 
and the pulley-like upper surface of the astragalus bone in the 
latter, differ by the third or middle toe and its supporting meta- 
carpal or metatarsal bone being symmetrical in itself, and larger 
than the lateral ones, when such are retained. In this suborder 
the family of the tapirs (Tapiridce), although now mainly Neogaeic, 
with one outlying Malayan species, was formerly widely spread in 
northern Arctogaea, fossil remains belonging to the single existing 
genus Tapirus being abundant in the Pliocene of Europe, although 
none appear to have been recorded from North America. In 
both Europe and America there occurs, however, the ancestral 
genus Protapirus ; its remains having been obtained in the former 
area from the upper Oligocene phosphorites of France, and in the 
latter from the nearly equivalent Uinta beds. Possibly a doubtful 
form {Palceotapirus) from the middle Eocene of France should 
also be included in the same family. The Uinta and Bridger 
deposits have also yielded a more or less nearly allied form known 
as Isectolophus, which apparently also occurs in the European 
Eocene, where it has been described as Lophiodon. Indeed, in our 
view, both this genus, and the still earlier American Systemodon, 
which appears to have been the earliest known representative of 
the tapiroid stock, may be included in the family Lophiodontidce, 
where we should also place the ancestral types of the horses. In 
this family the genus Lophiodon, as now restricted, seems to have 
been confined to the Eocene of Europe, where it died out without 
giving rise to descendants, the same being the case with the 
allied Eocene genus Helaletes^. The well-known Hyracotherium, 
which was an animal of the size of a fox first described from 
the London Clay but subsequently recorded from the North 

1 Recorded from Europe by Osborn and Wortman, Bull. Amer. Mus. 
Vol. vii. p. 360 (1895). 


American Eocene, is, however, the proximate ancestor of the 
horse-family (Equid<z)\ and we have thus evidence that the fore- 
runners of both the horses and tapirs were widely spread over the 
whole of northern Arctogaea. Hyracotherium, it may be observed, 
had the typical forty-four teeth characteristic of the earlier euther- 
ians, and four toes to the front, and three to the hind feet; but in 
the still earlier Phenacodus, which seems to be the ultimate 
ancestor of the horses, each foot was furnished with five complete 
toes. As other instances of the community between the early 
Tertiary mammalian fauna of the northern halves of the two 
hemispheres, we may cite the genus Pachynolophus of the middle 
and upper Eocene of Europe and the Bridger Eocene of the United 
States, which connects Hyracotherium with the under-mentioned 
horse-like animals ; and also Hyrachyus, typically from the 
Bridger, but probably occurring in the French phosphorites ; the 
latter being more nearly related to the rhinoceroses. 


Passing by several less important genera confined to one or 
the other hemispheres, we come to the family Pal&otheriidce, 
which is an ill-defined one including forms connecting the pre- 
ceding with the undoubted Equidce. Here the typical Oligocene 
genus Pal&otheriuni) which includes large tapir-like animals with 
three toes to each foot, is exclusively European. The same is the 
case with the contemporary Anchilophus, represented by smaller 
forms with more decidedly horse-like affinities; but with the 
Miocene and upper Oligocene Anchitherium, used in its wider 
sense, we once more come to a genus common to Western and 
Eastern Arctogaea. In this genus the jaws are provided with the 
typical forty-four teeth; but the last lower molar has generally lost 
the third lobe found in the preceding forms ; the fifth metacarpal 
bone being still represented by a splint. In the small A. 



I6 7 

(Mesohippus) bairdi of the White River Oligocene of the United 
States, the incisor teeth lack the infoldings of the summit of the 
crown characteristic of the horses, and the lateral digits are 
relatively large ; the whole size of the creature being comparable 


to that of a Newfoundland dog. On the other hand, in the typical 
A. aurelianense, from the European Miocene, the summits of the 
incisors were infolded, as in the horses. In spite of this resem- 
blance, Professor Scott, from the structure of the limbs, is of 
opinion that the latter species was not an ancestor of the modern 
horses, but that this position was occupied by A. bairdi. 

Restricting the term Equidce to those members of the suborder 
in which the crowns of the cheek-teeth are very tall (Jiypsodonf), 
with complicated infoldings of their enamel, and the hollows thus 
formed completely filled with cement, we have in the lower 
Pliocene of North America the three-toed genus Protohippus, 


distinguished from the modern horses by the shorter crowns of the 
cheek-teeth. The widely-spread genus Hipparion differs in having 
the anterior inner column of the upper molars completely 

1 68 



detached 1 ; the feet being generally three-toed, although in one 
Indian species the lateral digits are wanting. These three-toed 
horses are met with in the Pliocene of Europe, Asia, and North 


America ; and it is suggested by Professor Cope that while in the 
Old World the intermediate stage between Anchitherium and the 
modern horses was occupied by this genus, in the New World this 
gap was filled by Protohippus. The true horses (Equus\ charac- 
terised by the one-toed feet and the union of the anterior inner 
column of the upper cheek-teeth with the adjacent middle column,, 
although now confined to the Old World, where they date from 
the Pliocene, were formerly abundant in North America during the 
Plistocene, and, as we have seen, were likewise represented during 
the same epoch in South America. The oldest forms appear to 
be those from the Siwalik Hills of northern India ; and it is thus 
evident that the group was originally an Arctogseic one 2 . The 
genus is now represented in all the regions of eastern Arctogaea, 
with the exception of Madagascar, and its extinction in the New 
World cannot be satisfactorily explained. 

1 This pillar forms the lowest part of the unshaded area in figure 38. 

2 Professor Scott, to whose views we have already alluded, in a paper pub- 
lished in Tr. Amer. Phil. Soc. Vol. xvn. pp. in 112 (1894), is of opinion 
that the genus Equus was evolved in North America, and that Anchitherium, 
in its restricted sense, was off the direct line. He arranges the direct ancestral 
forms of the upper Tertiary, from above downwards, in the order Protohipptis , 
Desmatippus, Miohippus, and Mesohippus ; Anchitherium branching off from 
Miohippus, and Hipparion from Protohippus. 


The rhinoceroses (Rhinocerotidce) originally had a distribution 
very similar to that of the horses, with the exception that they 
never entered Neogaea; and they also agree with the latter in their 
present extinction in North America, where they are unknown 
after the Pliocene. On both sides of the Atlantic the true 
rhinoceroses appear to have commenced in the lower Oligocene ; 
and in both areas the earliest forms were hornless. Early species 
with a pair of horns placed transversely on the nose are likewise 
met with both in Europe and North America; but the modem 
two-horned rhinoceroses appear to be restricted to the Old World, 
where one extinct species (Rhinoceros antiquitatis] ranged as far 
north as the Arctic circle during the Plistocene period. On the 
whole, it seems preferable to include all the living and most of the 
extinct species in the typical genus Rhinoceros ; the existing forms 


A. median valley ; D. anterior, and E. posterior crests ; F. posterior valley ; 

H. crochet. 

being confined to the Oriental and Ethiopian regions. Rhino- 
ceroses, it may be observed, differ from all the preceding families 
of the suborder by the upper cheek-teeth having a continuous 
outer wall, instead of being divided by a vertical ridge into two 


distinct lobes ; and while all the living forms have but three toes 
to each foot, in some of the extinct hornless species the front limb 
was four-toed. In the typical genus the number of the front teeth 
is more or less reduced, but in the extinct Hyracodon and 
Amynodon of the upper Eocene of North America the full forty- 
four teeth were developed ; and as allied forms also occur in the 
Oligocene, it would seem highly probable that the group originated 
in North America, whence it migrated westwards by way of what 
is now Bering Strait to attain its most specialised development in 
the Old World. It is, however, noteworthy that the genus 
Cadurcotherium, from the French Oligocene, which should ap- 
parently find a place in this family, and is distinguished by the 
narrowness of its upper molars, makes a curious approximation 
in the structure of these teeth to the Neogaeic Homalodontothe- 
rium 1 . The most specialised representative of the family is the 
huge Elasmotherium, of the Siberian Plistocene, whose molars shew 
a resemblance to those of the Equidce. 

Another family of perissodactyles (Titanotheriidce) is typically 
represented by certain huge, somewhat rhinoceros-like, mammals, 
generally having a pair of transversely-placed tuberosities on the 
nasal region of the skull, and characterised by a peculiar arrange- 
ment of the tubercles on their upper molars. These teeth, which 
have very short crowns, also differ from those of the rhinoceroses 
in having the outer wall divided into two lobes by a vertical ridge; 
while the last lower molar is distinguished from the corresponding 
tooth of the latter by having a third lobe. The typical genus 
Titanotherium is mainly North American, where it ranges from the 
lower Oligocene to the upper Eocene, but is also represented in the 
Tertiaries of the Balkans, although unknown in those of western 
Europe. An allied genus (Brachydiastematotherium) has also been 
recorded from eastern Europe, but all the other members of the 
family, such as Palceosyops, are North American. This family 
accordingly appears to have been mainly an American one, but 
was probably represented in Asia as well as in eastern Europe. 

The remarkable genus Chalicotherium, whose geological range 
in the Old World extends from the Oligocene of France to the 

1 Supra, p. 82. 





lower Pliocene of India, while species also occur in the Miocene of 
the United States, has molars strikingly like those of the Titano- 
theriida, but its feet differ from those of all existing ungulates in 
terminating in huge curved claws much resembling those of 
the South American edentates. Indeed, one genus of the family 
(Macrotheriuni] was long regarded as belonging to the latter 
group. Of the two genera which occur in the European Miocene, 
Macrotherium has the fore limb much longer than the hinder ; 
whereas in Chalicotherium (Ancylotherium) the two are more nearly 
equal in length. It is to the former genus that the North American 
forms are assigned. 

The proboscidean ungulates, which differ markedly from the 
two preceding groups in the structure both of their teeth and 
limbs, and are now represented only by the Indian and African 
elephant, form a comparatively small assemblage, most of whose 
members may be included in the family Elephantida. Among 
these, the most specialised types constitute the genus Elephas, 
characterised by the complexity of the structure of the cheek-teeth, 
which generally assume the form of more or less elevated parallel 
plates, with the intervals filled with cement. In certain of the 
earlier species from the Pliocene of Asia the plates of these teeth 
are, however, comparatively low and less numerous, with the 
intervening valleys almost devoid of cement; so that these 
stegodont elephants, as they are called, form a complete transition 
towards the mastodons. Commencing in the Indian Pliocene, 
elephants ranged over the whole of Europe and Asia during the 
Plistocene, while we have also evidence of their occurrence during 
the same epoch in northern Africa; and they were likewise repre- 
sented by two species in North America, one of which ranged as far 
south as Texas. One of these American species was identical 
with the European mammoth (E. primigenius], while the second 
(E. columbianus] was nearly allied; both being near relatives of the 
existing Indian elephant. The extinct stegodont elephants being 
confined to south-eastern Asia, it is interesting to note that the 
species of Mastodon making the nearest approximation to Elephas 
are met with in this region alone; and from this it may be inferred 
that the evolution of the latter genus took place in that part of* the 
world. All mastodons, it may be mentioned, have comparatively 




simple molar teeth, in which the crowns are surmounted by low 
transverse ridges, frequently separated into more or less distinct 
tubercles, and with the intervening valleys open, such ridges 
varying from three to five in number in the majority of the teeth, 


although more numerous in the last of the series. Both in Europe 
and North America mastodons make their appearance in the 
Miocene, although in the latter area they are unknown before the 
Deep River beds forming the upper portion of that stage; and 
whereas they disappeared in the Old World with the Pliocene, 
they persisted in the New till the succeeding Plistocene age. 
During the Pliocene, as we have seen in the last chapter, they 
obtained an entrance into South America, so that they cannot be 
regarded as absolutely characteristic of Arctogaea. On the whole, 
it is probable that the group originated in the Old World, although 
we are still in the dark as to its relationship to other ungulates. 

The only other Arctogaeic genus of proboscideans is Dino- 
therium, which constitutes a family by itself, and is known from the 
Miocene and Pliocene of Europe and India, but is unrepresented 
in America. In these animals only one of the true molars has 
three ridges, the others having but two; and tusks were present in 
the lower jaw alone. 

A fourth subordinal group of the ungulates, which is more 
primitive than any of the foregoing, and has been designated by 
Professor Cope the Amblypoda, or short-footed group, has one 


family (Coryphodontidce] common to the lower Eocene of both 
hemispheres, while a second family ( Uintatheriid<z), of later age is 
strictly North American. All these animals had limbs of an 
elephantine type, each foot being furnished with five toes, and the 
bones of the wrist and ankle joints arranged on the linear plan. 
The genus Coryphodon, which includes species varying in size 
from a tapir to a rhinoceros, had forty-four teeth, with the canines 

FIG. 43. LEFT UPPER CHEEK-TEETH OF Coryphodon ; reduced. 

well developed, and the molars bearing prominent oblique ridges. 
First discovered in the London Clay, and subsequently in the 
lower Eocene of France, this genus has since been recognised in 
the lower Eocene of the United States, where nearly perfect 
skeletons have been obtained. 

In a still more primitive group known as the Condylarthra, 
which apparently contains the ancestral stock of both the artio- 
dactyles and perissodactyles, it is believed that the genus Phena- 
codus (the ultimate parent of the horses), typically occurring in the 
Wahsatch Eocene of the United States, is represented in the Swiss 
Eocene; and the same has been stated to be the case with 

Summarising the results of the foregoing survey, it may be 
stated that as regards its mammalian fauna Arctogaea 

Summary of . 

the character- as a whole is characterised by the following features, 
mammalian Absence of monotremes and diprotodont marsu- 
faunaof Arcto- pials. No existing polyprotodont marsupials except 
opossums, and these only in its western half. No 
Tertiary marsupials known except opossums, although other types 
probably existed in South-eastern Asia. No existing edentates 1 , 
and fossil ones only in North American Plistocene and Pliocene. 
No marmosets (Hapalida), and no monkeys of the family Cebidce. 
Bats of the family Phyllostomatida wanting, save for a few on the 
1 The aard-varks and pangolins are here excluded from the Edentata. 


Pacific side of North America. Insectivores abundant. In the 
following list of more or less widely distributed families the ma- 
jority are for the most part confined to this realm, while the others 
contain exclusively Arctogaeic genera. Such groups as are prac- 
tically confined to Arctogaea are printed in italics, those which are 
extinct having an * prefixed ; while the letter H. denotes such of 
the existing ones as are restricted to the Holarctic region : 


Talpida. H. 

Soricidce. Enters Austro-Malayan region and Mexican 


* Hycenodoriidce. 

* Proviverridce. 

* ArctocyonidcE. 


Sciuridae. Mainly Arctogseic, although species of Sciurus 

extend in Neogaea as far south as Paraguay. 
Castorida. H. 

* Theridomyida. H. 
Lagomyidce. H. 


* Chceropotamidtz. 

* A nthracotheriida. 

* Ccenotheriidce. 

TraguhdcE. Extinct in N. America. 

Cervidae. The genus Cervus exclusively Arctogaeic. 

Bovida. Represented by Bos depressicornis in Celebes. 

* Lophiodontidce. 

* Palaotheriidce. 

Rhinocerotida. Extinct in W. Arctogaea. 

* Chalicotheriidce. 

* Titanotheriida*. Mainly W. Arctogaeic but occurring 

in the Balkans. 
Elephantidae. Elephas. 

* CoryphodontidcK. 




* Plagiaulacidce. 

* Bolodontidce. Only Secondary in Eastern Arctogsea. 

The following table exhibits some of the better known Tertiary 
mammalian genera common to both halves of Arctogaea, together 
with allied types restricted to the western and eastern hemispheres ; 
such as are still existing being indicated by a f : 







Neogaea in Plistocene). 










Procamelus, etc. 












t Camelus. 
f Dorcatherium. 




UNGULATA (continued). 







Helaletes. Lophiodon. 




Anchitherium. Anchilophus. 


t Equus (also Neogaeic). 
t Rhinoceros. 


Titan otherium. 

Chalicotherium. Macrotherium. 
t Elephas. 
Mastodon (also Neogaeic). 


( Typically N. American, 
Phenacodus IV* J , 

i but stated also to 
Protogoma i . ^ 

I occur in Europe. 

In the foregoing table, only some of the better known types 
have been selected, but these suffice to show that throughout the 
Tertiary epoch a certain number of genera were common to 
western and eastern Arctogaea. It is true that, with the exception 
of those still existing, we have no evidence that any of these ever 
reached the Ethiopian region ; and it is quite probable that many 
or the whole of them never did so. By its present fauna that 
region is, however, so closely connected with the Pliocene and 
modern mammals of Asia and Europe, that there can be no ques- 
tion of its right to inclusion in the same realm ; and this being so, 
L. 12 


Madagascar cannot be excluded. Still, it is quite probable that 
during the later Tertiary epoch the Ethiopian and Malagasy 
regions were almost as distinct from the rest of Arctogaea as are 
Neogaea and Notogaea at the present day, and if such conditions 
had continued, the former areas would have been entitled to con- 
stitute a realm by themselves. 

In the sequel we shall discuss the whole number of existing 
genera of non-volant terrestrial mammals common to the eastern 
and western halves of the Holarctic region, and likewise such 
living and extinct types as are respectively peculiar to Eastern and 
Western Arctogsea. These, taken in conjunction with the fore- 
going tables, will show that, in spite of the forms common to the 
two latter areas, there has always been a large number of types 
restricted to one side or the other of the Atlantic basin. And 
this leads to the conclusion that, although during a considerable 
portion, or the whole of the Tertiary period, there was a free land- 
communication between North America and Eastern Asia by way 
of Bering Strait, yet that this connecting land must have been 
comparatively narrow, so that the faunas of the more southern 
portions of both areas developed to a great extent independently 
of one another. 

Not the least curious feature in connection with the com- 
munity of types on the two sides of the Atlantic is the precise 
parallelism in the development of many of the groups in both 
areas. In both, for instance, the phyla of the horses and rhino- 
ceroses were practically similar, although it is thought that the 
stage occupied in the one area by Hipparion was held in the other 
by Protohippus. If this particular suggestion should prove to be 
well founded, it will be self-evident that the true horses have been 
independently evolved in the two areas ; and it almost seems as 
if the same had been the case with the rhinoceroses and certain 
other groups. Had the culminating forms been devolved in only 
one of the two areas, we should not expect to find the whole of 
the ancestral links present in both. 



Mammalian groups peculiar to Eastern Arctogsea Tertiary Mammalian 
Faunas of Eastern Arctogaea Oligocene Fauna Miocene Fauna Older 
Pliocene Fauna Pikermi and allied Faunas Siwalik Fauna Higher 
Pliocene Faunas. 

ALTHOUGH northern Europe and Asia forms but one zoological 
region with the corresponding part of North America, 
yet there are numerous groups of mammals con- 

fined respectively to its eastern and western portions, ^u> Eastern 
which clearly show that the communication between 
the two areas was always more or less limited. In this chapter 
attention will be first directed to some of the most striking 
peculiarities of the mammalian fauna of Eastern Arctogaea, after 
which the whole fossil fauna may be taken into consideration. 

In addition to the total absence of existing opossums (Didel- 
phyid(E\ and the presence in its warmer portions of fruit-bats 
(Pteropodidce), which, however, are common to Notogaea, Eastern 
Arctogaea is especially characterised as being the home of all the 
higher Primates; namely the family Stmiidce, which includes the 
man-like apes and gibbons, and the Cercopithecidcz, embracing all 
the other Old World monkeys. From the South American monkeys 
(Cebidcz) both these families are broadly distinguished by having 
two pairs of premolar and three of molar teeth, whereas in the 
former group there are three pairs of both premolar and molar 
teeth. Not only are the two families in question confined at the 
present day to the Eastern hemisphere, but the same appears to 
have been the case at all epochs, since no trace of a fossil monkey 
has ever been recorded from North America. This remarkable 
isolation of the distributional areas of the Simiidcz and Cercopi- 
thecidce on the one hand, and of the Cebidce. (and Hapalidcz) on 

12 2 


the other, points unmistakeably, in spite of their external similarity, 
to the dual origin of the monkeys of the Old and New Worlds. 
Both may, however, have originated from different groups of 
lemuroids, which, as indicated in an earlier chapter, ranged over 
the whole of Northern Arctogaea during the earlier part of the 
Tertiary epoch. 

At the present day the man -like apes, which are few in 
number, have an extremely limited distribution. The chim- 
panzees (Anthropopithecus) are restricted to Equatorial Africa, the 
gorilla (Gorilla) is found only in the hottest regions of Western 
Africa, the orangs (Simla} are confined to the islands of Borneo 
and Sumatra, and the smaller gibbons (Hylobates) are inhabitants 
of South-eastern Asia, from Assam and Burma to Hainan. Ex- 
tinct species of chimpanzees and orangs occur, however, in the 
Pliocene of Northern India; while gibbons are met with in the 
Miocene of France and Baden, although there is some difference 
of opinion whether these are generically identical with the Asiatic 
forms, or should be assigned to a genus apart (Pliopithecus}. The 
former deposits have also yielded remains of a large extinct ape 
(Dryopithecus)) apparently of a somewhat more generalised type 
than all the existing forms. 

The ordinary monkeys and apes (Cercopithecidce) have a wider 
distribution, ranging over most of the warmer parts of Eastern 
Arctogaea, and being represented by a single species at Gibraltar, 
and by two others in Moupin, in Eastern Tibet, and a fourth in 
Japan. This family, by the way, is not exclusively Arctogaeic, 
since, as we have seen, one species of a peculiar genus (Cynopi- 
thecus) inhabits Celebes. Apart from the latter, the family is 
represented by eight living genera, among which the Ethiopian 
Papio has extinct representatives in the Pliocene and Plistocene 
of India, as have also the Asiatic Macacus and Semnopithecus. The 
two latter genera also occur in the Pliocene of France and Italy; 
and a tooth of a species of Macacus has been obtained from the 
Plistocene brick-earths of Essex. In addition to these, there are 
certain extinct genera from the European Tertiaries; among 
which Mesopithecus from the lower Pliocene of Greece agrees with 
Macacus in its short and stout limbs, but approximates to 
Semnopithecus in the character of its skull and dentition. Dolicho- 


pithecus, from the French Pliocene, has a longer muzzle; while 
Oreopithecus, from the Italian Miocene, seems to connect the 
CercopitheddcB with the Simiida. 

It is thus evident that during the latter portion of the Tertiary 
epoch monkeys and apes were spread over the greater part of 
Eastern Arctogaea; and their extensive diffusion is a proof that 
this half of the realm could only have been connected with North 
America by land situated so far north that it formed an impass- 
able barrier to these animals. Although the smaller extinct 
European monkeys do not necessarily indicate a very high 
temperature in the regions they inhabited, there can be little doubt 
that at the era when Dryopithecus flourished southern Europe at 
least enjoyed a moist tropical climate. 

Not less characteristic of Eastern Arctogaea are the existing 
lemuroids, of which there are three families ; the largest (Lemurida) 
ranging over the Oriental, Ethiopian, and Malagasy regions, the 
Tarsiidce, with a single genus, being exclusively Oriental, while 
the sole representative of the Ckiromyida is Malagasy. The 
numerous existing members of the Lemuridce. are all characterised 
by the first of the three lower premolar teeth assuming the form 
and functions of a canine; and as this feature is unknown in 
any of the Tertiary representatives of the sub-order, this family 
appears to be exclusively confined to the area under consideration. 
For the most part, the Oligocene lemuroids of Europe seem 
likewise to have been markedly distinct from those of North 
America. Microchcerus^ , for instance, which is represented both 
in France and England, indicates a family characterised, among 
other features, by the general presence of only three pairs of 
premolar teeth in each jaw ; and Adapts, which is likewise 
common to the same two countries, has four such teeth. 

Although there are several families of Insectivora peculiar to 
the eastern hemisphere, the only one of these with a wide distri- 
bution is that of the hedgehogs and their allies (Erinaceida), which 
has representatives in the eastern Holarctic, Oriental, and Ethiopian 
regions. Although none are known from America, extinct repre- 
sentatives of this family are common in the European Oligocene. 

1 Teeth figured on p. 156. 


Of these Pal&oerinaceus appears to be allied to the true hedgehogs 
(Erinaceus) ; whereas other genera, such as Necrogymnurus, 
connect the former with the existing long-tailed and spineless 
Gymnura of the Malayan islands. This group is thus essentially 
characteristic of Eastern Arctogaea as a whole. 

Turning to the Carnivora, we have, in addition to the Prote- 
leidce, of which the sole representative is the African aard-wolf, two 
important families practically confined to this half of the realm. 
The first of these is the extensive group of the civets, mungooses, 
and their allies ( Vtverridce), which has no representatives in the 
New World, although a single species in two genera ranges into 
the Austro-Malayan region. Out of a total of twenty-three genera 
included in this family only one mungoose (Herpestes) and the 
common genet (Genetta) range into Europe, most of the other 
forms being confined to the Oriental, Ethiopian, and Malagasy 
regions. Civets ( Viverra) and ichneumons, together with several 
remarkable extinct genera, such as Stenoplesictis, were, however, 
common in the European Tertiaries, from the lower Oligocene 
upwards. And from the circumstance that the last-named genus 
presents characters connecting the Viverridce with the weasel 
tribe (Mustelida), it would seem probable that the latter family 
was originally evolved in Eastern Arctogaea, although it has now a 
considerable number of American representatives. 

By means of certain extinct forms from the lower Pliocene of 
Southern Europe and Northern India known as Ictitherium, the 
civets are very closely connected indeed with the hyaenas 
(ffy&nidce) ; the three living representatives of which may be 
included in the single genus Hyana. Although the striped hyaena 
(H. striata) is now confined to Southern Asia and Northern 
Africa, it occurred in the Pliocene epoch in France and England; 
while the larger spotted hyaena (ff. crocutd] of Southern Africa 
ranged during the Plistocene era over the greater part of temperate 
Europe, and likewise extended eastwards as far as India. 
Numerous extinct species of the same genus are found in the 
Pliocene of Europe and India; and two extinct genera from the 
same deposits Hyanictis and Palhyizna connect the living 
forms with the aforesaid Ictitherium. Although one extinct 
Tertiary North American genus has been tentatively assigned to 

V.] RODENTS. 183 

it, the family is thus essentially an Eastern Arctogaeic one; and it 
may be assumed that, as its living representatives are inhabitants of 
hot climates, the extinct forms were unable to exist sufficiently far 
north to permit them to cross by the bridge of land via Bering 

Among the rodents, in addition to the two widely spread 
families Myoxidcz and Spaladdce. confined to Eastern Arctogaea, 
we find in the Muridce the sub-family of the gerbils (Gerbillina) 
similarly restricted, their range including the whole of Eastern 
Arctogaea with the exception of the Malagasy region. 

The subfamily Murtna, which includes the true rats and mice 
(Afus) is likewise restricted to the Old World. These rodents 
differ from the hamsters (Cricetus) and the New World white- 
footed mice (Sitomys), which, with other forms, constitute the 
sub-family Cricetince, by the molar teeth in the upper jaw having 
their tubercles arranged in three longitudinal rows ; whereas in the 
latter they form only a double series. Distributed over all the 
regions of Eastern Arctogaea, with the exception of Madagascar, 
this group is also represented in the Australian region. Of the 
two extensively distributed families restricted to the area under 
consideration, the first is that of the dormice (Myoxidce) whose range 
includes the Eastern Holarctic and Ethiopian regions. Distin- 
guished from all other rodents by the absence of a caecum or 
blind appendage to the intestine, and further characterised by 
the complicated infoldings of the enamel on the crowns of their 
molar teeth, these beautiful little creatures now attain their 
maximum development in Africa. In a fossil state they are first 
known from the lower Oligocene of Europe, and are likewise 
common in the Miocene. Another family which does not range 
beyond the limits of Eastern Arctogaea is that of the Spalacida, 
typically represented by the great mole-rat (Spalax typhlus] a 
blind creature, ranging over south-eastern Europe, Persia, Meso- 
potamia, Syria, and Egypt. The allied genus (Rhizomys), in which 
the eyes, although minute, are not covered with skin, includes 
several species, whose distributional area embraces the north of 
India, Tibet, China, Burma, Malaysia, and Abyssinia ; but the 
three remaining genera are restricted to the Ethiopian region. 
The whole of the foregoing belong to the mouse-like, or Myo- 


morphous section of the order, but among the Hystricomorphous 
rodents we have the typical porcupines (Hystricince), in which the 
tail is never prehensile, practically confined to Eastern Arctogsea, 
where they range over south-eastern Europe, and the Ethiopian 
and Oriental regions. The Javan species of the typical genus 
(Hystrix javanicd] is, however, found in the island of Timor, 
in the Austro-Malayan region, although it is doubtless of late 
introduction there, and may not improbably have been transported 
by human agency. In a fossil state porcupines of this sub-family 
are common in the European Tertiary as far down as the lower 

Turning to the ungulates, we have in the artiodactyle section 
two closely allied families, which if we except certain pigs from the 
Austro-Malayan region and Papua, which may have been originally 
introduced by man are restricted to the area under considera- 
tion. Both these families, moreover, have representatives, either 
living or extinct, in all the regions of Eastern Arctogsea, inclusive 
even of Madagascar, so that they may be reckoned among its most 
characteristic mammals. The Hippopotamidtz now restricted to 
the Ethiopian region, where they are represented by the widely- 
spread common Hippopotamus amphibius, and the much smaller 
terrestrial H. liberiensis of the West Coast ranged during the 
Plistocene and upper Pliocene epochs over the greater part of 
Europe as far north as England ; one species from these deposits 
being apparently indistinguishable from the common African form. 
Extinct species are met with in the Plistocene of Algeria, the 
Plistocene and Pliocene of India, the Pliocene of Burma, and the 
superficial deposits of Madagascar; some of these differing from 
the common hippopotamus by the presence of three, in place of 
two, pairs of incisor teeth in each jaw. Whatever may have been 
the case with the swine, it is evident that the hippopotami were 
never able to exist sufficiently far north to cross by way of Bering 
Strait into the New World. In the pigs (Suidez) which among 
other features differ from the Hippopotamidce by the nostrils being 
perforated in a fleshy disc at the extremity of the muzzle, and like- 
wise by the structure of the teeth the typical group of the genus 
Sus ranges over most of the Eastern Holarctic and the whole 
of the Oriental region, being replaced in the Ethiopian and 


Malagasy regions by the potamochcerine group, frequently reckoned 
as a distinct genus. The Ethiopian region is, however, the sole 
habitat of the wart-hogs (Phacochoerus}. 

Among extinct artiodactyles we have two well-marked families, 
distinguished from the foregoing by the crescentic columns of 
their short-crowned cheek teeth the Anoplotheriidce and Dicho- 
dontidce likewise confined to Eastern Arctogaea, although their 
remains have hitherto been obtained only from the eastern section 
of the Holarctic region. The first of these includes several genera 
from Oligocene strata, characterised by having three columns on 
the front half, and two on the hinder half of their upper molars. 
In the typical Anoplotherium there were forty-four teeth, arranged 


in a continuous even series, and the feet were provided with either 
three or two toes. Dacrytherium differs by the molars being more 
like those of Ancodus, and also by the deep cavity for the 
reception of a gland on each side of the face in front of the eye ; 
while the small and elegantly formed animals described as Xiph- 
odon have the crowns of the first three premolar teeth elongated 
and trenchant, the feet being two-toed. In the Dichodontida, of 
which there are likewise several genera, the cheek-teeth are more 
completely selenodont, with only four columns on the crowns of 
the upper molars ; and it is not improbable that in this family we 
have the ancestral types of both the chevrotains and the deer. 

In the Camelidce, as we have already seen, the typical genus 
Camelus, which is found living (although not in a wild state) in the 
Eastern Holarctic, Oriental, and Ethiopian regions, and fossil in 
the Plistocene of Algeria and the Pliocene of India, is likewise 
confined to Eastern Arctogsea. And the same is true, both in the 


recent and fossil state, with the genera Tragulus and Dorcatherium, 
which at the present day alone represent the Tragnlida. 

The family of giraffes (Giraffidce), of which the Ethiopian 
Giraffa cameiopardalis is the sole existing survivor, was formerly 
extensively distributed over the area under consideration, to which 
it appears to have been always restricted, albeit represented by 
a considerable number of generic types. True giraffes (Giraffd) 
ranged during the Pliocene epoch over Greece, Persia, India, 
and China, and allied types are to be found in Visnutherium 
of the Pliocene of India and Burma, and Helladotherium from 
the corresponding formation of Greece. Still more gigantic than 
the latter were the huge Hydaspitherium, Bramatherium, and 
Sivatherium, of the Indian Pliocene, in all of which the simple 
horns of the giraffes were replaced by large antler-like appendages, 
differing considerably in their arrangement in the different genera. 
Other members of the family are Samotherium, of the Pliocene of 
the Isle of Samos, and Palceotragus from the equivalent deposits 
of Attica, in both of which the females appear to have been horn- 
less, although the males had a pair of simple, compressed, and 
nearly upright horn-cores. The former is represented by a species 
rivalling the giraffe in the size of its skull, but the latter was a 
much smaller animal. This group likewise extended to Northern 
Africa, where a large species from the Algerian Pliocene has been 
described under the name of Libytherium. 

Although the extensive family of the Bovidce, including the 
oxen, sheep, goats, antelopes, etc., is now represented in the 
northern part of the western Holarctic region by the American 
bison (Bos americanus\ the bighorn (Ovis canadensis], the musk- 
ox (Ovibos moschatus], and the so-called Rocky Mountain goat 
(Haploceros montanus), together with a few extinct forms from the 
superficial deposits, while the anoa (Bos depressicornis) is peculiar 
to Celebes, the greater number of its representatives belong to 
Eastern Arctogsea. The whole of the numerous genera of ante- 
lopes, together with the true goats, as well as the great majority of 
the sheep, are, for instance, restricted to this area. Moreover, 
whereas the musk-ox is now solely North American, it was common 
in Europe during the Plistocene ; while the bighorn is closely 
allied to the Kamschatkan wild sheep (O. nivicola), and the 


American bison not far removed from its Caucasian cousin (Bos 
bison], so that all these forms are probably descended from 
ancestors inhabiting Eastern Arctogaea. 

In the perissodactyle section of the ungulates, if we take fossil 
forms into account, there are no families peculiar to this area; but 
among extinct forms we have the large Oligocene genus Palceo- 
therium 1 , and the Eocene Lophiodon absolutely restricted to it. 

Although occurring only in Syria and the Ethiopian region, and 
at present unknown in a fossil state, the peculiar subordinal group 
of ungulates represented solely by the hyraces (Procaviidce) per- 
haps deserves mention among the types characteristic of Eastern 
Arctogaea. A nearly similar observation applies to the extinct 
proboscidean family Dinotheriidce, in which the single known 
genus (Dinotherium) ranges from the Miocene and Pliocene of 
Europe to the Pliocene of Northern India. 

Of the edentates, with the exception of certain very doubtful 
forms from the French phosphorites, which may prove to be 
reptilian, we have no evidence of the existence of any representa- 
tives in the Old World. There are, however, in Eastern Arcto- 
g?ea two very peculiar families commonly assigned to the same 
order as the latter, although it seems preferable to regard them as 
indicating an ordinal group (Effodientia) by themselves. Of these 
the pangolins (Manidce), which are distinguished from all other 
mammals by their covering of overlapping horny scales, are now 
confined to the Oriental and Ethiopian regions, to which the one 
living genus Manis is common ; but they appear to have been 
represented in the Oligocene phosphorites of France by smaller 
extinct forms, to which the names Necromanis and Leptomanis* 
have been given. The second family, Orycteropodidce, of which 
the only living members are the Ethiopian aard-varks (Orycteropus), 
differ very widely from the last, the body being nearly naked, and 
the molar teeth characterised by a peculiarly complex structure 
which is unique in the whole mammalian class. A fossil species 
of the existing genus has been discovered in the lower Pliocene of 
Samos and Maraga, in Persia ; while the extinct genus Pal&orycte- 

1 Teeth figured on p. 166. 

2 The so-called Palceomanis, from the Pliocene of Samos, turns out to have 
been founded on remains of an ungulate. 


ropus, from the French phosphorites, is believed to have belonged 
to the same family, so that both groups appear formerly to have 
been widely distributed. 

Summarising the results of the foregoing survey, we may put 
in a tabular form the leading features of the mammalian fauna of 
Eastern as distinct from that of Western Arctogsea. In the sub- 
joined table the letters E., M., O., H. respectively indicate the 
Ethiopian, Malagasy, Oriental, and Eastern Holarctic regions ; 
and when a family is represented in any of such regions only in a 
fossil state, a f is added to the denoting letter. The names of such 
families or groups as are practically peculiar to the area 'under 
consideration are printed in italic type ; while extinct groups have 
an * prefixed. 


Simiidce. O. E. H.f 

Cercopithecidcz. O. E. H. ; also extending into the Austro- 

Malayan region. 
Lemuridtz. O. E. M. 
* Microchoeridce. H. 
* Adapida. H. 


Pteropodidse. O. E. ; also common to Notogaea. 

Erinaceidce. O. E. H. 


Viverridce. O. E. H. M.; two species extending their 

range into the Austro- Malayan region. 
Hycenidcz. O. E. H.f 


Muridae ; the sub-family Gerbillince, O. E. H., is restricted 
to Eastern Arctogaea, while the Murinae are exclu- 
sively confined to the Old World, but range into 

Myoxidce. H. E. 

Spalacida. H. O. E. 


Hystricidae ; in this family the Hystricina are practically 
restricted to Eastern Arctogaea, although a Javan 
species is found in Timor. 


Hippopotamidce. E. O.f H.f M.f 

Suidce. H. O. E. M., also extending into the Austro- 
Malayan region. 

* Anoplotheriida. H. 

* Dichodontidce. H. 

Camelidae ; in this group the genus Camelus, H. E. O., is 

peculiar to Eastern Arctogaea. 
Tragulidae ; the existing genera Tragulus, O., and Dorca- 

therium, E. H.t, as well as several extinct ones, 

are restricted to this area. 
Giraffida. E. O.f H.f 
Bovidae; the whole of the true antelopes and goats, as 

well as most of the sheep and oxen, are restricted to 

Eastern Arctogaea, North America now preserving 

only one species of Bos, Ovis, Ovibos, and Haplo- 


* Palaeotheriidae ; Palceotherium. 

* Lophiodontidae ; Lophiodon. 
Procaviida. E. H. 

* Dinotheriida. H. O. 


Manida. E. O. H.f 
Orycteropodida. E. H.f 

To these may be added the absence of living opossums (Didel- 
phyida}. It will be observed that in this list such existing families 
as are confined to one of the regions of the area considered are 
for the most part omitted. Tertiary families which are at present 
unknown beyond the Eastern Holarctic region have, however, been 
included, since the limitation is probably in great part due to our 
want of knowledge of the early Tertiary faunas of the other 

Although the differences indicated between the faunas of 


Eastern and Western Arctogaea, which will be still more apparent 
after the consideration of the mammals of North America, seem at 
first sight to indicate that these two areas should form distinct 
divisions of the realm, yet the community of the fauna of the 
northern portion of the two hemispheres forbids this view. This 
question may, however, be more fully discussed in the chapter 
devoted to the Holarctic region. 

Before entering upon the consideration of the different zoolo- 
gical regions into which the realm is divided, it is 
Mlmma r Han essential to take a brief survey of the Tertiary mam- 
Faunas of malian faunas of Eastern Arctogaea. By this alone 

Eastern . 

Arctogeea. it is possible to understand the true relations of the 

existing faunas to one another ; while such a survey 
also serves to demonstrate that the regions in question are but 
features of the present epoch of the earth's history ; and that even 
as late as the Pliocene portion of the Tertiary epoch the dis- 
tinctions now obtaining between the Holarctic, Oriental, and 
Ethiopian regions had no existence. In our survey we may omit 
the Eocene period, and commence with the lower Oligocene ; and 
it will simplify matters to give lists of some of the more important 
and better known generic types characterising the faunas of the 
different horizons. The leading affinities of many of the genera 
mentioned have been already alluded to in the present or preced- 
ing chapters ; but it would vastly exceed the limits of our space 
to attempt to point out the distinctive features of the others. 
Accordingly, the reader must either take them on trust, and treat 
them practically as abstract terms, or he must refer to some 
palaeontological treatise in order to find the real nature of the 
animals indicated by such generic names. 

Although it is essential to our purpose to notice the Oligocene 

faunas of Eastern Arctogaea, it is important to 

FauIS* 06116 observe that our knowledge of these is practically 

limited to Western Europe. We are consequently 

quite unable to say how far the geographical range of such 

faunas extended, although it is probable that this embraced a 

large portion of the Eastern Holarctic region. Whether, however, 

at this epoch Ethiopian Africa had received a large mammalian 

fauna must be left for future discoveries to determine. 


Under the term of lower Oligocene (the upper Eocene of 
many writers) are included a large series of strata, such as the 
freshwater beds of Bembridge and Hordwell in the south of 
England, the gypsum of Montmartre near Paris, and the corre- 
sponding black lignite beds of De'bruge in Vaucluse 1 . A consider- 
able part of the fauna of the Quercy phosphorites of Central France 
likewise comes under the same category, only we have here a 
mixture of Middle and Upper Oligocene forms. And in the case 
of the siderolites, or bone-earths of Switzerland, this admixture is 
carried to a still greater degree, undoubted Eocene types occurring 
with those properly characteristic of the Oligocene. In the 
following list such genera as are found only in the phosphorites 
have the letter P. after them; while after those peculiar to the 
siderolites the letter S. is added; both letters being given when 
the genera are common to the two formations. As already said, 
only some of the better-known forms are selected. 

Among the lemuroid Primates, we have the genera Adapts 
and Microch&rus, both of which occur in the English beds as well 
as in the phosphorites ; these being the last European representa- 
tives of the group. The Insectivora include Necrogymnurus (P.S.), 
allied to the Malayan Gymnura, Amphidozotherium, together with 
the existing genera Sorex and Talpa. More remarkable is the 
occurrence in the phosphorites of an insectivore described as 
Pseudorhynchocyon, which is believed to be a member of the family 
of jumping shrews (Macroscelidida), now confined to the Ethiopian 

The true Carnivora are represented by Eusmilus (P.), a 
highly specialised ally of the sabre-toothed tigers, as well as by 
the more cat-like sElurictis, and the generalised Pseudcelurus. 
In addition to species of true civets, referred to the living genus 
Viverra, the Viverrida include Amphictis (P.), Stenoplesictis (P.), 
and Palaoprionodon (P.); the two latter being generalised forms 
closely connecting the family with the Mustelidce, which is 
represented by Plesictis (P.). To the Canida may be assigned 
the genera Cynodon (P.S.), Cephalogale (P.), and Cynodictis, 
together with a species which may be included in Amphicyon 

1 See table on p. 117. 


(P.); while the creodont division of the order is represented by 
Hyanodon, Pterodon, Oxycena (P.), and Proviverra. Among the 
rodents we may note the squirrel-like Sciuroides (P.S.) and Pseudo- 
sciurus (S.), the existing genus Sa'urus, and the extinct Plesiarctomys 
and Plesiospermophilus, which likewise belong to the same family. 
Among the Muridce, Cricetus includes ancestral types of the 
hamsters ; while the dormice are represented by the existing genus 
Myoxus., As noticed previously, Theridomys, Nesocerodon, and 
Protechinomys seem to be ancestral forms allied to some of the 
existing South American rodents. 

The ungulates are very strongly represented; the pig-like 
group including Cebochoerus, Cheer opotamus and Elotherium (P.) 
among the Chcsropotamida^ Acotherulum, and Anthracotherium (P.) 
and Ancodus in the Anthracotheriidce. The anoplotheroids com- 
prise Anoplotherium, D aery t her ium, and Xiphodon ; Dichobunus 
and Ccenotherium (P.) are the characteristic forms among the 
Ccenotheriidce; and Dichodon, Gelocus, and Lophiomeryx among the 
DichodontidcR\ while the chevrotains are represented by Prodremo- 
therium (P.) and Bachitherium (P.). In the perissodactyle section 
of the same order, we have Pachynolophus (P.) to represent the 
Lophiodontidce, Palceotherium and Anchilophus in the Palceo- 
theriida, Protapirus (P.) as an ancestral form of the tapirs, and 
Rhinoceros (P.), Cadurcotherium (P.), and Hyrachyus (P.) as 
representatives of the rhinoceroses. The aberrant Chalicotherium 
has also one species from the phosphorites. The Effodientia in- 
clude Leptomanis (P.), Necromanis (P.), and Palceorycteropus (P.); 
while the existing genus Didelphys alone represents the mar- 

It will be seen that in this fauna the existing generic types are 
very few, and if the whole of the extinct ones had been given, their 
relative proportion would have been still less. The ungulates were 
abundant, and among these the perissodactyles proportionately 
more numerous than at the present day ; while the anoplotheres 
are in some respect transitional between the latter and the typi- 
cal artiodactyles. All the ungulates had brachydont teeth, and 
annectant types between the modern pigs and ruminants were 
abundant ; the traguloids being the highest development among 
the artiodactyle section of the order. Creodont carnivores still 


persisted, although more modern types had already made their 
appearance on the scene ; and opossums flourished. 

The middle stage of the Oligocene is represented in Europe 
by the freshwater marls and clays of Hempsted in the Isle of 
Wight and the corresponding beds of Ronzon, near Puy-en-Velay, 
the lignitiferous strata of Cadibona in Liguria, the deposits of 
Fontainebleau and Ferte-Alais in France, and likewise by certain 
beds in Hungary and at Monte Promina in Dalmatia. Among 
the small fauna of this stage we may notice the following. In the 
Insectivora, Tetracus, an ally of the hedgehogs; Cynodon, Amphi- 
cynodon, Plesictis, and Hycznodon among the carnivores; Anthra- 
cotherium, Ancodus, Elotherium, Ccenotherium, Gelocus, and Rhino- 
ceros in the ungulates; and opossums (Didelphys). While this 
fauna is closely related to the preceding, it has lost a number of 
early ungulate types, such as Anoplotherium and Xiphodon among 
the artiodactyles, and Pal&otherium and Anchilophus in the 
perissodactyles. On the other hand, the pig-like forms, such as 
Ancodus, Anthracotherium, and Elotherium, attained an extra- 
ordinary degree of development. Among the creodont carnivores, 
we may note the final disappearance of the genera Pterodon and 
Proviverra, although Hycenodon still survived. 

The upper Oligocene (lower Miocene) fauna is a large and 
characteristic one, well represented in the freshwater beds of 
St Gerand le Puy, in the Allier, as well as in those of Weisenau 
and other localities in the neighbourhood of Mayence. Among 
the mammals may be mentioned the following ; existing genera 
being denoted by the prefix of a f. 

INSECTIVORA. fTalpa. fSorex. 

Geotrypus. Dimylus. 

t Myogale. Palseoerinaceus. 

Plesiosorex. t Erinaceus. 

CARNIVORA. Cephalogale. fViverra. 

Amphicyon. t Herpestes. 
Plesictis. Proaelurus. 

Potamotherium. Hyaenodon. 


L. 13 


RODENTIA. Theridomys. fSpermophilus. 

Archaeomys. f Sciurus. 

Issiodoromys. Chalicomys. 

t Myoxus. Titanomys. 

t Cricetus. 

UNGULATA. Anthracotherium. Csenotherium. 

Hyotherium. f Tapirus. 

Amphitragulus. t Rhinoceros. 

MARSUPIALIA. f Didelphys. 

Of this fauna, Professor von Zittel writes that it seems at first 
sight closely akin to those of the middle and lower Oligocene ; the 
same ordinal and subordinal groups, and in many instances the 
same genera characterising the whole three horizons. In the lack 
of lemuroids, the reduced number or final disappearance of 
opossums, creodonts, and anoplotherioids, in the greater abundance 
of forms like Anthracotherium, Hyotherium, and Dremotherium, 
which were but poorly represented in the lower Oligocene, and in 
the number of new types, such as Tapirus, Amphitragulus (an 
ancestral chevrotain), Chalicomys (an early beaver), Titanomys (an 
ally of the picas), Erinaceus, Dimylus (a form connecting the 
shrews with the hedgehogs), Potamotherium (a generalised otter), 
Herpestes (mungooses), Procelurus (a primitive type of cat), we 
notice, however, the marked difference of this fauna from its fore- 
runners. Among the incoming genera it is noteworthy that there 
is none for which an ancestral type cannot be found in the lower 
Oligocene ; the main difference occurring in the more specialised 
characters of the members of the later fauna. With the exception 
of certain bats, insectivores, rodents, and the opossums (such as 
Vespertilio, Erinaceus, Sorex, Myogale, Talpa, Sciurus, Spermo- 
philus, Cricetus, Myoxus, and Didelphys], the majority of the 
genera are, however, still extinct. 

It is probable that the beds in the Balkans which have yielded 
remains of the North American Tertiary genus Titanotherium 
belong to some portion of the Oligocene epoch. 

We now come to the Miocene epoch, which, as at present re- 
stricted, forms in Europe but a small section of the Tertiary era. 


It includes the well-known freshwater strata of Sansan in Gers (the 
middle Miocene of the older geological classifica- 
tions), together with the corresponding beds of F ^n a cene 
Steinheim in Styria, and likewise the somewhat 
newer (upper Miocene) deposits of CEningen, in Baden. Grive-St- 
Alban, in the valley of the Rhone, is likewise another well-known 
locality where mammaliferous strata of this age are developed; 
and, among other places, we may also mention Monte Bamboli 
in Italy, San Isidro in Spain, and Oran in Algeria. 

For the first time in Europe we meet with remains of true 
Primates, of which there are three genera belonging to the Simiidce, 
two of which, Dryopithecus and Oreopithecus *, are extinct, but the 
third seems scarcely separable from the existing Oriental Hylobates. 
In the Insectivora we meet with the existing European genera 
Talpa, Myogale (desmans), Erinaceus, Soreoc, and Crocidura ; while 
the extinct Lanth another turn seems to be allied to the tree-shrews 
( Tupaid] of the Oriental region, and Galerix intermediate between 
the latter and the j Limping-shrews (Macroscelidtdce) of Ethiopian 
Africa. Among the Carnivora, where the creodonts have disap- 
peared, the cats are represented by the sabre-toothed tigers 
(Machcerodus] and Pseudcelurus. In addition to the existing 
genera Viverra and Herpestes, we have among the civet tribe the 
extinct Progemtta. The dogs include the existing Cam's, together 
with the extinct Hemicyon and Pseudocyon ; while the larger forms 
described as Dinocyon and Hycznarctus connect the former with 
the bears. The Mustelidce. are represented by species of the 
typical living genus Mustela, together with certain more or less 
closely allied extinct types ; and Enhydriodon filled the place 
of the modern otters. 

From among the rodents the generalised types allied to those 
now characteristic of Neogsea have all disappeared, nearly all the 
recorded forms apparently pertaining to existing genera. In the 
Sciuridce not only have we true squirrels (Sdurus\ but the Ethio- 
pian spiny squirrels (Xerus) are likewise represented, as are also 
the more widely distributed flying-squirrels of the genus Stiuro- 
pterus, which now inhabit both Eastern and Western Arctogaea 

1 Some of the characters of these genera have been already mentioned on 
pages 1 80, 18 r. 



Chalicomys, Cricetus, and Myoxus are survivors from the Oligocene; 
but porcupines (Hystrix] are new comers. Picas of the existing 
genus Lagomys are likewise to the fore ; and it is a question 
whether those distinguished by the name of Myolagus might not 
be included under the same title. 

A marked approximation to the modern type is likewise the 
characteristic feature of the ungulates of the European Miocene ; 
although in this group living genera still remain in the minority. 
The pigs (Sutda) include, for instance, the genus Hyotherium, in 
which the molar teeth are tuberculated and of the general type of 
those of some of the living members of the family ; and also the 
more aberrant Listriodon, characterised by the presence of a pair 
of transverse ridges on each of the teeth of the same series. A 
species of the existing West African genus Dorcatherium alone 
represents the chevrotains (Tragulidte) ; while we have forerunners 
of the deer (Cervidce) in the extinct Pal&omeryx and Dicroceros, 
both characterised by the simple structure of their antlers ; and 
Protragoceros a generalised type of antelope marks the first 
appearance of the hollow-horned ruminants (Bovida], which now 
form such a numerous and characteristic group in the fauna of 
Eastern Arctogaea. Perissodactyle ungulates are less numerous. 
Anchitheriiim, to some of whose distinctive characters allusion has 
been made above, constitutes the representative of the equine line 
at this stage ; and tapirs and rhinoceroses belonging to the 
existing genera were likewise common. Some of the latter were, 
however, still hornless, and in none was more than a single horn 
developed. The aberrant ChalicotheriidcB^ forming the last family 
of this section of the order, and characterised by the extraordinary 
resemblance presented by their claws and toes to those of eden- 
tates, are here represented by the gigantic Macrotherium. Finally 
the Miocene is notable as being the stage at which proboscideans 
first made their appearance on the scene in Europe. In this group 
we have species of Mastodon, which, as already explained, includes 
the ancestors of the modern elephants ; and likewise one of the 
more aberrant Dinotherium. 

Compared with the Oligocene, the loss of so many antiquated 
types, coupled with the appearance of proboscideans and man-like 
apes, and the general modern facies of all the mammals of the: 


Miocene, indicates the lapse of a considerable interval of time 
between the deposition of the two series of strata. And that this 
is really the case, is demonstrated by the fact that there occurs 
between the two a considerable thickness of marine deposits which 
have not hitherto yielded remains of land mammals. It may be 
noticed that while many of the insectivores and rodents from this 
horizon belong to genera now inhabiting the Eastern Holarctic 
region, among other forms we have marked instances of Oriental 
(Lanihanotherium. Hylobates] or Ethiopian ( Galerix, Dorcatherium, 
and Xerus) affinities in this assemblage; and it is thus evident that 
at the epoch in question there was no trace of the differentiation 
of Eastern Arctogaea into regions. 

Still more markedly are the same features displayed by the 
older Pliocene fauna of Europe and Southern Asia. 
This fauna, which was formerly regarded as of upper cen^Faun'a 
Miocene age until shewn by Dr Blanford to be 
unquestionably referable to the succeeding era of geological his- 
tory, had a very wide distribution ; and it is represented at certain 
localities, mostly at long distances from one another, by an extra- 
ordinary profusion of remains. One of these charnel-holes occurs 
at the village of Pikermi, near Athens, a second in the Isle of 
Samos, in the Turkish Archipelago, and a third at Mont Leberon, 
in Provence. This fauna is also met with locally in the valley of the 
Rhone, at the foot of the Pyrenees, in Spain, Asia Minor, and at 
Maraga in Persia. It is likewise represented in the regions lying 
to the north of the Alps, only here the number of forms is less, 
and the antelopes and giraffe-like ruminants, fitted for roaming 
over the open plains of the south, are conspicuous for their 
absence; their place being taken by forest-haunting deer. The 
sand-beds of Eppelsheim in Hessen-Darmstadt, together with 
strata in the neighbourhood of Vienna, and others in Hungary 
and Rumania, may be cited as localities where the northern 
section of this fauna is preserved. 

Taking first the European and Western Asiatic portion of this 
fauna, and leaving its Oriental members for subse- 
quent consideration, we find the Primates represented 
solely by an extinct genus of monkey, taking its 
name of Mesopithecus on account of presenting certain features 


intermediate between the existing Semnopithecus and Macacus. 
The insectivores are likewise known only by a solitary form, a 
shrew (Sorex) ; but this is probably due to the nature of the strata 
being unfitted for the preservation of the remains of such small 
creatures. The Carnivora, on the other hand, were abundant, the 
Felidce being represented not only by the sabre-tooths (Machter- 
odus), but true cats (Felis) likewise making their appearance on 
the scene. Hyaenas display a great variety of development, there 
being one species of the typical genus Ifycena, with certain 
resemblances to the existing Cape form, while the more generalised 
types known as Lycyana and Hycenictis were likewise present, as 
were also species of Ictitherium^ and the allied Palhyana, which, 
as already mentioned, formed a connecting link between the 
hyaenas and the civets. True dogs seem to have been absent from 
this assemblage ; but Amphicyon still survived from the Miocene, 
and an aberrant form known as Simocyon made its appearance. 
Hycznarctus was likewise another survivor from the Miocene, and 
may be regarded as a forerunner of the true bears. Finally, in the 
weasel tribe (MustelidcB) we have representatives of the existing 
genus Mustela, as well as the extinct Pal&omephitis and Promeles, 
the latter being an ancestral type of the badgers. 

Rodents make but a poor show, as we have only the extinct 
beaver-like Chalicomys, a species of porcupine (ffystrix), and a 
representative of the curious little spiny mice (Acomys\ now 
characteristic of Syria, Palestine, and north-eastern Africa. 

A remarkable advance over their Miocene forerunners is 
displayed by the ungulates, especially those from Pikermi and 
Maraga. Here, in the artiodactyle section, we meet for the first 
time with true pigs of the genus Sus, which at this period ranged 
over the greater portion of Europe, and some of which attained 
very large dimensions. Water-chevrotains (Dorcatherium) serve 
to connect the Miocene representatives of their genus with the 
existing West African form; while muntjacs (Cervulus), now 
confined to the Oriental region, filled the place of the stags. 
Giraffe-like creatures were numerous, for not only have we true 
giraffes belonging to the existing Ethiopian genus Giraffa, but the 
gigantic hornless Helladotherium stalked over the plains of Greece, 
and the allied but horned Samotherium inhabited the area now 



occupied by the Turkish Archipelago, and extended eastwards as far 
as Persia; Palceotragus being a smaller but allied form. The Bovidce 
are represented by antelopes, most of which present a marked 
Ethiopian facies, although Tragoceros (probably the direct descen- 
dant of the Miocene Protragoceros} is an aberrant form, with 
compressed horn-cores like those of the goats. And it may be 
remarked that most of the Pikermi antelopes have short-crowned 
molar teeth, in which respect they resemble the existing eland, 
kudu, and their allies. Of the Pliocene. forms, Palczorias, which is 
common to Southern Europe and Algeria, seems to be inter- 

FlG. 45. SKULL OF Palczorias. 

mediate between the kudus (Strepsiceros) and elands (Orias}\ 
while the so-called Protragelaphus is so closely allied to the 
existing Ethiopian harnessed antelopes (Tragelaphus) as to be 
included by some in the same genus. On the other hand, 
Palaoryx is nearly related to the gemsbok and its allies (Oryx), 
although with certain resemblances to the sable antelope group 
(Hippotragus). Gazelles (Gazella}, which are essentially inhabi- 


tants of open plains, were likewise abundant ; one being considered 
a near relative of the South African springbok. The genus 
Helicophora, on the other hand, closely resembles the water-buck 
group (Cobus), which is exclusively Ethiopian. In the perisso- 
dactyle division, the three-toed horses (Hipparion) seem to have 
approximated in general structure to the Ethiopian zebras, and, 
like those animals, may have been ornamented with dark and light 
stripes. While some of the Pliocene rhinoceroses were hornless, 
another was a two-horned species closely allied to the common 
African Rhinoceros bicornis, of which it may be regarded as the 
parent form. There is also an extinct genus (Leptodon), of some- 
what uncertain affinity ; while tapirs are found in the Eppelsheim 
beds, although not apparently in the southern area. The Chalico- 
theriidft were represented by the typical genus Chalicotherium 
(Ancylotheriuni), which, as we have seen, was a near ally of the 
Miocene Ma cr other him, and also occurs in the Oligocene phos- 
phorites. As in the Miocene, the proboscideans include only 
Mastodon and Dinotherium; the one species of the former ranging 
from Greece to Persia, but being different from all the Indian forms 
of the same epoch. Finally, the occurrence of an aard-vark 
(Orycteropus] both in Samos and Persia serves to accentuate the 
Ethiopian affinities of the southern section of this fauna. 

We have thus evidence that one and the same fauna extended 
from Spain and Algeria across Southern Europe to Asia Minor 
and Persia; and we may infer from the deposits at Samos, that 
what is now the y-Egean sea formed a tract of land connecting 
Greece with Turkey. It is further evident that there must have 
been free communication across the Mediterranean basin (which 
in Cretaceous times is known to have been a mare clausnm, in the 
physical, and not the political sense of the term) between Europe 
and Africa. This communication may have existed both by way 
of Gibraltar, and also between Italy, Sicily, and Malta on the 
one hand, and Tunis on the other; since the Plistocene mammals 
of the islands in question clearly indicate continental connection. 
While the antelopes and hipparions of this fauna prove the exist- 
ence of open plains during the lower Pliocene epoch, the host of 
individuals of Mesopithecus as unmistakeably point to the presence 
of extensive forest-tracts. In the northern section of the fauna, as 


displayed at Eppelsheim, the Ethiopian affinities are much less 
apparent, aard-varks and the whole of the giraffe-group being 
absent, while tapirs and deer were abundant. That there was a 
more or less marked separation between the two areas thus seems 
evident; and the tapirs and muntjac-like deer, both of which 
seem wanting in the Siwalik fauna, are indicative, so far as they 
go, of Malayan affinities. 

Nearly related to that of Pikermi, Samos, and Persia, the 
celebrated Siwalik fauna of India and the adjacent 
countries presents certain well-marked differences; F a^ hk 
this being specially shown by the occurrence of several 
essentially modern types quite unknown in the former. More- 
over, there are a considerable number of peculiar genera which do 
not occur in the western fauna; while we also come across certain 
Miocene, and even Oligocene types, which are equally strange to 
the latter. Although in some cases these occur in beds which are 
not improbably of upper Miocene age, in others they appear 
mingled with the later forms ; but, in any case, they indicate a 
survival in this area of archaic types which at that time had com- 
pletely disappeared from Europe. 

Originally discovered in the outer ranges of the typical Hima- 
layan area, the Siwalik fauna has been traced towards the 
north-west into the Punjab, Kach, Sind, and the north-eastern 
frontier of Baluchistan ; the beds from the two latter areas being 
lower in the series than those from the typical Siwalik hills, and 
containing an older assemblage of forms, although several are 
common to all. An outlier of the same fauna occurs in Perim 
Island in the gulf of Cambay. Eastwards the Siwalik fauna 
ranged through Sylhet and Assam to Burma, whence it has been 
traced at intervals, as in Java, Sumatra, and the Philippines, into 
China and Japan. In China it extended from Yunnan in the 
south-west northwards through Szechuen to Kansu, and thence 
eastwards through Shensi to Shansi, its extreme eastern limit being 
indicated by the discovery of a Siwalik elephant's tooth at 
Shanghai. Northward of Kansu the fauna ranged into Mongolia, 
probably by way of the gap formed by the course of the Hwang-ho 
through the Ala-shan mountains if such mountains existed at the 
time. And it is not a little remarkable that of the few Mongolian 


forms at present known, two (Hycena macrostoma and Equus 
sivalensis) are identical with species from the Siwalik Hills 1 . 

As regards the fauna itself, we find, in the first place, the 
Primates much more fully represented than at Pikermi, and all by 
existing generic types. Of the man-like apes (Simiid&\ there is 
a chimpanzee (Anthropopithecus) presenting a more human type 
of dentition than its living Ethiopian cousins; while a single tusk 
indicates the former existence of an orang (Simla) allied to the living 
Bornean and Sumatran species. The other three generic types 
belong to the Cercopithecida, and include baboons of the Ethiopian 
genus Papio ( Cynocephalus), together with species of Semnopithecus 
and Macacus, the former genus being exclusively, and the latter 
mainly, Oriental at the present day, although both occur in the 
later Pliocene of Europe. Doubtless owing to the unsuitability 
of the strata for the preservation of small specimens, no remains 
of insectivores have hitherto been obtained. The Carnivora are, 
on the other hand, well represented; the Felicia including large 
and small species of the typical genus Felis, and apparently one 
of the allied Cynahirus (hunting-leopard), now exclusively Oriental 
and Ethiopian. Mach&rodus had two species ; and another 
form has been identified with the European Oligocene genus 
SEhiridis. Civets include species of Viverra larger than any now 
existing; this genus being also one now confined to the Ethiopian 
and Oriental regions, although more abundant in the latter than in 
the former. The CanidcE, in addition to a survivor of the Miocene 
Amphicyon, were represented by wolves and jackals (Cam's), as 
well as by a species apparently allied to the long-eared fox (Otocyon} 
of Africa. While in the Ursidce the generalised Hycenarctus still 
survived, true bears (Ursus) make their appearance for the first 
time, the single known Siwalik species presenting, however, a 
marked approximation in the characters of its skull and dentition 
to the Indian sloth-bear (Melursus). Among the few known 
representatives of the Mustelida, we have a large marten (Mustela), 
probably allied to the living yellow-throated Indian species ; a 
ratel, belonging to a genus (Mellivora) now restricted to India 
and Africa; and likewise an otter (Lutra) whose nearest affinities 
are with an existing Sumatran species. The same family also 

1 Lydekker, Rec, Geol. Sw~v. India, Vol. xxiv. pp. 207 211 (1891). 


includes a member of the otter-like genus Enhydriodon, the other 
species being from the Italian Miocene. Among the most 
remarkable features of the Siwalik Carnivora is the survival of 
a species of Hycenodon, of which the remains have been discovered 
in the Punjab. 

The Rodentia are but very imperfectly known. They include 
a representative of the bamboo-rats (Rhizomys), which are now 
exclusively Oriental, and belong to the family Spalacida; and, 
among the Muridce, a species of Nesocia, which genus is likewise 
confined to the Oriental region. The other forms are a porcupine 
(Hystrix] and a hare (Lepus). 

A very long list is presented by the ungulates, which are 
numerous not only in generic, but likewise in specific types. The 
pig-like artiodactyles include, among the family Suidce. several 
representatives of the true pigs (Sus), some of which attained 
gigantic dimensions, while others are remarkable for the complex 
structure of their molar teeth, which show a marked resemblance 
to those of the existing Ethiopian wart-hogs (Phacochczrus). A 
still more elaborate structure is displayed by the corresponding 
teeth of the allied genus Hippohyus, which is peculiar to this 
fauna ; and the family is also represented by species of the 
European Miocene genera Hyotherium and Listriodon, the remains 
of the two latter being mostly obtained from the Punjab and 
districts to the west. The same areas are mainly those which 
have yielded remains of Anthracotheriidcz, although some of these 
have been discovered in Sylhet. In this family we have species of 
the European genera Anthracotherium and Ancodus, the former of 


which is elsewhere unknown above the Middle Oligocene; and 
there are also three peculiar types, respectively known as Meryco- 
potamus, Hemimeryx, and Chctromeryx, differing from all the rest 


in having only four columns on the crowns of the molars, as 
shown in the annexed figure, and thus presenting a marked 
approximation to the ruminants. The earlier Tertiary Chczropo- 
tamidcz likewise had a survivor in the genus Tetraconodon, which 
was represented by a large pig-like creature remarkable for the 
enormous size of its simple conical premolar teeth. The pig- 
like group closes with Hippopotamus, which makes its appearance 
on the scene for the first time in this formation, where it is 
represented by a generalised species with three pairs of incisor 
teeth in each jaw. Turning to the groups with fully-developed 
selenodont molars, we have first to notice the occurrence of fossil 
camels of the existing genus Camelus, which are unknown else- 
where except in the Algerian Plistocene. As we have seen that 
the Camelidce. were originally a New World group, it is interesting 
to note that these earliest Old World representatives occur in Asia 
instead of Europe; and it is further noteworthy that in the 
structure of their molar teeth the Siwalik camels retain evidences of 
affinity with the South American guanacos and vicunas which are 
lost in their living descendants. The Tragulidce contain repre- 
sentatives of the true chevrotains ( Tragulus) and water-chevrotains 
(Dorcatherium\ now respectively characteristic of the Oriental and 
Ethiopian regions, while among the deer ( Cervidce) we have species 
of the Oligocene European genus Palaomeryx, together with 
others belonging to Cervus, the representatives of the latter being 
all closely allied to existing Oriental types. Not improbably also 
a musk-deer (Moschus) should be included among the Siwalik 
Cervidce. Among the Giraffidce, in addition to true giraffes 
(Giro/a), which are common to the Pikermi beds, and extended 
eastwards into China, we have the peculiar gigantic antlered types re- 
spectively known as Vishnutherium, Sivatherium, Hydaspotherium, 
and Bramatherium, of which the first seems common to the Siwaliks 
of Burma and the Punjab, while the second is confined to the 
more easterly Himalaya, the third to the Punjab, and the fourth to 
Perim Island. They include the most gigantic of all ruminants, 
Sivatherium almost rivalling an elephant in bulk. 

Not one of the least curious features in this marvellous fauna is 
that while, as we have seen, deer of Oriental types were abundant, 
antelopes closely allied to those now inhabiting the Ethiopian 





region where deer are totally absent were likewise extraordinarily 
numerous. Of the African genera we have a species of Bubalis 
intermediate between the hartebeests and the blesbok, a member 
of the sable antelope group (Hippotragus\ a kudu (Strepsiceros), an 
eland (Orias), and probably a representative of the water-buck 
group (Cobus). On the other hand, Oriental forms are not 
wanting, as proved by the occurrence of a nilgai (Boselaphus), and 
probably of a four-horned antelope (Tetraceros] ; while the widely- 
spread gazelles (Gazelld) were likewise present. Goats and oxen 
for the first time made their appearance ; the former group being 
represented not only by species belonging to the typical Capra, 
but likewise to the shorter-horned genus Hemitragus, now confined 
to India and Arabia. The oxen (Bos) included members of all the 
existing groups, that is to say typical oxen, bison, buffalo, and 
smaller forms with upright triangular horns nearly allied to the 
anoa of Celebes. 

The perissodactyle ungulates, so numerous in the earlier 
Tertiary formations, have now become proportionately much fewer 
as compared with the artiodactyles. While typical forms of 
Hipparion were present, one species differs from the rest by the 
loss of the lateral toes, and thus resembles the modern horses 
(Equus\ which here make their appearance for the first time. 
Rhinoceroses include not only hornless forms, but likewise one 
species allied to the existing Oriental Rhinoceros unicornis and R. 
sondaicus, and a third as closely related to the African Burchell's 
rhinoceros (7?. simus]. In the same group Chalicotherium is a 
survivor from older formations. 

Finally, the proboscideans exhibit a development unparalleled 
in any other formation or epoch. Dinotherium appears for the 
last time in the Siwaliks of Perim Island, Kach, Sind, and the 
Punjab ; while the mastodons include a large number of species, 
some of which present such a close approximation to the so-called 
stegodont elephants (which, as already mentioned 1 , are peculiar to 
this fauna) as to render it impossible to draw any well-defined 
demarcation between the genera Mastodon and Elephas. Not 
only does the Siwalak fauna include the aforesaid stegodont, or 

1 Stipra, p. 172. 


transitional elephants, but likewise one which may well have been 
the ancestor of the species now inhabiting India. Eastwards these 
transitional elephants and mastodons have been traced into Java, 
Borneo, China, and Japan ; and, as stated in an earlier chapter, 
there can be no doubt that the modern elephants were evolved in 
this area. 

Although, as shown in the foregoing survey, the Siwalik fauna 
differs in certain respects from that of Pikermi, Samos, Leberon, 
etc., yet there can be no hesitation in regarding the whole lower 
Pliocene fauna of Europe, North Africa, Asia Minor, and South 
and East Asia as essentially one ; and consequently at this epoch 
there was no possibility of distinguishing between the Palaearctic 
and Oriental regions. Whence Ethiopian Africa had by this 
time received the forerunners of its present higher mammalian 
fauna, we have, unfortunately, no decisive evidence. Writing 
some years ago, Dr Blanford 1 seems to suggest that the irruption 
of the modem African fauna was anterior to the Pliocene. After 
referring to certain peculiarities connected with the existing 
mammalian fauna of India and the Malayan area, he observes that 
" these cases of isolation probably indicate that the animals belong 
to an older fauna, now partly replaced by newer types, and that 
the older fauna was common to India and Africa. It is very 
probable that these animals are descended from the ancient 
tropical fauna of the early Tertiary times. But, so far as it is 
possible to judge, the process of variation would have caused a 
greater distinction between forms so widely separated and exposed 
to such different conditions, if the period of isolation were great ; 
and it is difficult to suppose that the lands inhabited by the 
ancestors of the Simiidce, Lemuridce, Tragulida, and Manidce of 
the Oriental and Ethiopian regions can have been separated prior 
to the early part of the Miocene period." 

This is perfectly true so far as it goes, but since, as we have 
seen, genera like Hippopotamus, Bos, Capra, Equus, and Elephas 
are unknown previous to the Siwalik epoch, and some of them 
at least were evolved at or about that time in the Indian area, it 
seems necessary to assume the existence of a free land communi- 

1 Manual of Geology of India, ist ed. p. Ixviii. (1879). 


cation between the Ethiopian and Oriental regions at least as late 
as the lower Pliocene epoch. With regard to where this connec- 
tion was situated, we may note, in the first place, that Dr Wallace : 
was of opinion that even the Pikermi fauna made its way into 
Africa chiefly through Syria, although a brief connection of Europe 
with Tunis is admitted. When the passage in question was 
written, little or nothing was, however, known as to the Pliocene 
fauna of Algeria. And although this undoubtedly indicates a 
western connection between Europe and Africa, yet even in the 
Pliocene the Sahara probably formed, as now, a barrier 2 across 
which the fauna of northern Africa could not pass south. Accord- 
ingly, even the Pikermi fauna may have come round byway of Egypt. 
Be this as it may, it seems clear that the Siwalik fauna entered 
Africa by way of Syria or Arabia, or possibly by both. The most 
direct line of communication would be via the Gulfs of Oman and 
Aden ; and some indication that such a line of connection may 
have existed is afforded by the distribution of the goats of the 
genus Hemitragus. As already stated, fossil species of this genus 
occur in the Siwaliks of Perim Island and the Himalaya, while of 
the three existing forms, one is Himalayan, a second confined to 
the Nilgiri and certain other South Indian ranges, and the third 
inhabits Oman. So far as it goes, the evidence of these goats is 
strongly suggestive of the former existence of a land-bridge across 
the mouth of the Persian Gulf, as otherwise we should expect to 
find living species in Persia and other parts of western Asia. If 
the existence of such a bridge be admitted, we only require another 
across the narrow strait of Bab-el-Mandeb to give a free line of 
communication between India and Africa in this direction. 

Whether, however, the migration from India to Africa took 
place at the north or south end of the Red Sea, or at both ends, 
it is certain that the connecting land must have been of consider- 
able width, and suited to the passage of mammals of all kinds. 
In referring to the nature of the connection, Dr Wallace 3 remarks 
that " we may now perhaps see the reason of the singular absence 

1 Geographical D^strib^ttwn of Animals, Vol. I. p. 288. 

2 The idea that there was a Tertiary sea in the Sahara is incorrect; see 
Blanford, Quart, Journ. GeoL Soc. Vol. XLVI. p. 90 (1890). 

3 Op. cii. p. 291. 


from tropical Africa of deer and bears ; for these are both groups 
which live in fertile or well-wooded countries, whereas the line of 
immigration from Europe to Africa was probably always, as now, 
to a great extent a dry and desert tract, suited to antelopes and 
large felines, but almost impassable to deer and bears." The 
Siwalik chimpanzee, however, indicates most unmistakably that 
the communication by way of Arabia or Syria between the Ethio- 
pian and Oriental regions must have embraced a forest-area, and 
accordingly have been of considerable width. 

With regard to the question why so many genera which existed 
in India and southern Europe during the Pliocene should have 
disappeared from those areas to live on in Africa, all we can say is 
that it is quite evident that a southern migration of the fauna has 
certainly taken place, and that this was probably induced by the 
cold heralding the approach of the glacial period. Although we 
have few, if any, decisive physical evidences of a cold period in 
India, yet the existence of a goat (Hemitragus) nearly allied to a 
Himalayan species in the ranges of southern India seems to indi- 
cate that such must have occurred, as it would be quite impossible 
for the ancestral form to have crossed the intervening plains under 
present conditions of temperature. It is further noteworthy that 
many of the animals which have disappeared from India, such 
as chimpanzees, hippopotami, giraifes, water-chevrotains, and 
ostriches, are precisely those which are now restricted to very hot 
climates ; whereas the lion, tiger, rhinoceroses, elephants, and 
monkeys, which both now or during the Plistocene are known 
to be capable of existing in cold climates, have persisted. 

Leaving these exceedingly difficult questions, two other points 
may be noticed in connection with the Siwalik fauna. In the 
first place, since the Siwalik hills themselves form ranges of con- 
siderable height on its southern flank, it is evident that the 
Himalaya was much lower during the lower Pliocene epoch than 
it is at present ; Dr Blanford l stating that the movement which 
led to its elevation "has been distributed over the Tertiary and 
post-Tertiary period, and a great portion in post-Plistocene." 
This will account for the community between the lower Pliocene 

1 See Geol. Mag. Decade 3, Vol. ix. p. 166, note (1892). 
L. 14 


fauna of the Himalayan area and Mongolia, the Himalaya at that 
epoch not forming, as now, an impassable barrier to the north of 
the Oriental region. The second point relates to the survival in 
the Siwalik fauna of archaic forms, which had disappeared at that 
date from Europe. This fact, especially since old types such as 
lemurs and gymnuras are even now met with in the Oriental 
region, lends support to the view advanced in an earlier chapter 1 
that marsupials may have lived on in south-eastern Asia long after 
they had completely disappeared from Europe. 

Our knowledge of the later Pliocene faunas of Eastern 

Arctogaea is mainly confined to Europe, where at 
cene Kaunas. " this period the general distribution of land and sea 

was apparently very much the same as at the 
present day. Spain was, however, connected with Africa, as was 
probably also Italy by way of Sardinia and Malta. A portion of 
Italy was, however, submerged, while in Belgium, Holland, and 
the south-east of England the sea intruded upon what is now land ; 
but, on the other hand, Britain was joined to the Continent. 
Few mammaliferous deposits of this age have been preserved to 
us, but among these are the Crags of the east coast of England 
(which contain numerous fossils derived from earlier formations), 
the fresh-water beds of the Val d' Arno in Italy, as well as others in 
the Auvergne, in the Rhone valley, at Roussillon, and in the 
neighbourhood of Montpellier. The following genera are in- 
cluded in this fauna, those which are extinct having an asterisk 

PRIMATES. Semnopithecus. 

* Dolichopithecus. 

INSECTIVORA. Sorex (Shrews). 
CARNIVORA. *Machaerodus (Sabre-tooths). 

Felis (Cats). 

Viverra (Civets). 


Canis (Wolves and Foxes). 

* Hyaenarctus. 

1 Supra, p. 57. 




CARNIVORA (cont.). 

Ursus (Bears). 

^Elurus (Cat-bears). 

Mustek (Martens and Weasels). 

Lutra (Otters). 
RODENTIA. Arctomys (Marmots). 

* Chalicomys. 
Castor (Beaver). 

*Trogontherium (Giant Beaver). 

Cricetus (Hamsters). 

Microtus (Voles). 

Mus (Rats and Mice). 

Hystrix (Porcupines). 

Lagomys J 

Lepus (Hares) 

Sus (Pigs). 


Cervus (Deer). 

Alces (Elk). 

Cervulus (Muntjacs). 

* Palaeoryx. 
Gazella (Gazelles). 
Bos (Oxen). 
Tapirus (Tapirs). 
Equus (Horses). 

*Hipparion very rare. 

* Mastodon! 


Elephas J 


In this list by far the greater number of the genera are living 
ones, and if we removed from it types like Hyana, Hippopotamus, 
Rhinoceros and Elephas, which were spread during the Pliocene 
and Plistocene epochs over the greater part of Eastern Arctogaea, 
its Ethiopian resemblances are by no means strongly marked. 
Although the larger forms (as in the succeeding Plistocene epoch) 



include a considerable number of genera now mainly confined to 
tropical or subtropical countries, the rodent fauna exhibits a 
marked Palsearctic facies, thus indicating an approximation to the 
existing state of things. Among the extinct rodents, Pellegrinia, 
from the Sardinian Pliocene, belongs, however, to the Octodontida, 
and is probably allied to the existing African Ctenodactylus. Tro- 
gontherium is a gigantic extinct type of beaver, which also persisted 
into the Plistocene. The deer include northern types unknown 
in the lower Pliocene. 

One of the most remarkable features of this fauna is the 
occurrence of a large species of sElurus, a genus represented 
elsewhere only by the cat-bear or panda (^E. fulgens) of the 
eastern Himalaya, which, although formerly regarded as the type 
of a family by itself, is now included in the American Procyonidce 
(raccoons). The fossil species has been hitherto detected only in 
the English Crag ; the genus may, however, be expected to occur 
in the Siwaliks, since it is quite clear that it must have been 
originally connected with the American representatives of the 
family by forms inhabiting Eastern Asia. 

With the end of the Pliocene epoch this brief survey of the 
Tertiary mammalian faunas of Eastern Arctogaea may be brought 
to a close, since the Plistocene mammals can be more con- 
veniently considered under the headings of the different regions of 
this great province. While throughout the Oligocene, Miocene, 
and lower Pliocene epochs no trace of the present zoological 
regions of this half of the Arctogseic realm is shown, when the 
Upper Pliocene is reached there are faint indications of the 
demarcation of the Eastern Holarctic. At the time of the Plisto- 
cene, as will be shown in a later chapter, the Eastern Holarctic, 
Oriental and Ethiopian regions appear to have assumed a still 
more marked distinction, although this is to a great extent 
obscured by the wide range even at that epoch of genera like 
Hippopotamus, Rhinoceros, Elephas, Macacus, etc. Moreover, 
several species which are now confined to one of the three regions 
in question had then a more extensive distribution, so that it is 
only during the recent epoch that the Holarctic, Oriental, and 
Ethiopian regions attained the full faunistic peculiarities by which 
they are now characterised. 



Limits Mammalian Fauna Relations of Madagascar to the Mainland. 

INCLUDED by Drs Sclater and Wallace within the Ethiopian 
region, Madagascar and the adjacent groups of 
islands were referred to a region apart by Dr Blan- 
ford 1 ; this separation being justified not only by the mammalian 
fauna, but likewise by many other groups of animals. To quote 
Dr Wallace, this region "comprises, besides Madagascar, the 
islands of Mauritius, Bourbon, and Rodriguez, the Seychelles, 
and Comoro Islands. Madagascar itself is an island of the 
first class, being a thousand miles long, and about two 
hundred and fifty miles in average width. It lies parallel to 
the coast of Africa, near the southern tropic, and is separated 
by 230 miles of sea from the nearest part of the continent, 
although a bank of soundings projecting from its western 
coast reduces this distance to about 160 miles. Madagascar is a 
mountainous island, and the greater part of the interior consists of 
open elevated plateaus ; but between these and the coast there 
intervene broad belts of luxuriant tropical forests." It is this 
forest-district which forms the home of most of its peculiar fauna. 
As regards geological structure, it [appears from the researches of 
Messrs Cortese and Baron that, roughly speaking, a line drawn 
from north to south so as to divide the island into two longitudinal 
halves, gives an area of granitic and volcanic rocks on the right or 
eastern side, and on the left or western side one of sedimentary 
deposits, containing beds belonging to the Jurassic, Cretaceous, 
Eocene and recent epochs. Blown sand occurs in abundance 

1 Appendix, No. 8, p. 76. 


around the coast, and numerous old lake-basins or marshes, some 
of very large dimensions, form receptacles where remains of the 
later faunas have been preserved. With the exception of the 
Comoro group, which contain a few species, the non-volant mam- 
malian fauna is confined to Madagascar, so that the other islands 
do not properly come within the province of the present work. 
It is, however, important to observe that the Seychelles differ from 
almost all oceanic islands in consisting largely of granitic and 
other crystalline rocks. 

In an island lying so close to the African continent as Mada- 
gascar, the natural assumption would be that, if it 
fa^ia" 1 " possessed a mammalian fauna at all, such fauna 

would be closely allied to that of the mainland. As 
a matter of fact, precisely the reverse is the case, and out of a 
total of fully 28 genera of non-volant mammals now or recently 
inhabiting the island, only three are common to Africa. This, 
however, is by no means all, for out of these three genera two 
{Hippopotamus and Sus] are such as have probably crossed the 
intervening channel, although at a time when it was narrower than 
at present, while it is quite possible that the third (Crocidura) may 
have been introduced by human agency. Even this, however, 
scarcely gives a true idea of the case. In the first place, not only 
are the peculiar genera unknown in Africa, but they are equally 
strange to all the other regions of the world. In the second place, 
these genera belong to groups which form only a very small por- 
tion of the existing mammalian fauna of the Ethiopian region. 
At the present day, as will be more fully indicated in the following 
chapter, Ethiopian Africa is especially characterised by its nume- 
rous antelopes, as well by giraffes, zebras, rhinoceroses, elephants, 
hippopotami, wart-hogs, bush-pigs, lions, leopards and various 
other large cats, baboons, anthropoid apes, aard-varks, and 
ostriches. But, with the exception of the aforesaid bush-pig and 
extinct hippopotamus, not a single representative of any one of 
these groups is found in Madagascar. In place of such animals, 
Madagascar is populated by a host of lemurs, so numerous that 
the number of their species considerably exceeds that of all the 
other non-volant mammalian inhabitants of the island. Civet- 
and mungoose-like species, all pertaining to peculiar genera, alone 

VI.] THE FAUNA. 21$ 

represent the numerous Garni vora of the mainland; the Insectivora, 
in addition to the aforesaid Crocidura, or musk-shrew, include only 
the peculiar family of the tenrecs (Centettdce), which is confined to the 
island, and a representative of the Ethiopian family Potamogalida ; 
while the rodents comprise five genera of the cosmopolitan 
mouse-family (Muridce], more or less closely allied to one another, 
but different from any found elsewhere. 

The following is a list of the genera of non-volant Malagasy 
mammals ; those which are extinct being indicated by an asterisk, 
and the names of all the groups peculiar to the island printed in 


Chirogale (Mouse-lemurs) ; 4 species. jieiroAd.le<*s 

Microcebus (Dwarf Lemurs) ; 5 species. * 

Opolemur (Fat-tailed Lemurs) ; 2 species. ^ 

Lemur (True Lemurs) ; 8 species. 

Mixocebus (Hattock) ; i species. ? ^ 

Hapalemur (Gentle Lemurs) ; 2 species. ' y ( \ 

Lepidokmur (Sportive Lemurs) ; 8 species. 7.' $c~*V\ 

Avahis (Avahi) ; i species. ? 

Propithecus (Sifakas) ; 4 species.^ 7 - 

/^m'(Endrina) ; i species. 


* Megaladapis (Giant Lemur); i species. K^l*< 


Chiromys (Aye-aye) ; i species. 

Crocidura (Musk-shrews) ; i species. 

Centetes (Tenrec) ; i species. 

Hemicentetes \ 2 species. 

Ericulus (Hedgehog-tenrec) ; i species. 

Echinops-j i species. 

Microgale (Long- tailed tenrecs) ; 3 species. 

Oryzorictes (Rice-tenrecs) : 2 species. 




Geogale; i species. 



Cryptoprocta -, i species. 


Fossa ; i species. 

Galidictis (Striped Mungooses) ; 2 species. 

Galidia (Ring-tailed Mungoose) ; i species. 

Hemigalidia (Brown-tailed Mungoose) ; i species. 

Eupleres (Small-toothed Mungoose) ; i species. 



Hypogeomys ; i species. 
Nesomys ; 2 species. 
Brachytarsomys ; i species. 
Hallomys; i species. 
Eliurus; 2 species. 


Sus (Potamochcerus) ; t species. 


Hippopotamus ; i species (extinct). 

Considering the fauna in more detail, it may be first mentioned 
that the lemurs (Lemuroidea) differ from the higher, or Anthropoid 
Primates by their generally lower grade of organisation, as well as 
by certain features of the skull and internal anatomy which need 
not be more fully noticed here. They all have fox-like, expression- 
less faces ; and, with the exception of the aye-aye and the Asiatic 
tarsiers, they are characterised by the innermost pair of upper 
incisor teeth being separated from one another in the middle line. 


At the present day lemuroids are represented elsewhere only 
in the Ethiopian and Oriental regions ; the African forms being 
more nearly allied to the Malagasy types than are those of Asia. 
As stated in an earlier chapter, the group was, however, well 
represented in the lower Oligocene of western Europe, where 
certain forms (Mtcroc/ioerus) distinctly approximate some of the 
living kinds, although differing in the conformation of the first 
lower premolar tooth, which in the existing Lemuridce, assumes 


tic. upper canine ; k. lower canine ; pm. premolars ; m. molars. 

the form and function of a canine or tusk. In the latter family (of 
which the distribution is coextensive with that of the suborder) the 
first three genera in the foregoing list belong to a subfamily 
(Galagina) distinguished by the elongation of the bones of the 
tarsus, and represented by an allied genus (Galago) on the African 
mainland. The next four genera constitute the typical subfamily 
(Lemurincz\ which is absolutely confined to Madagascar and some 
of the islands of the Comoro group, and of which the ring-tailed 
lemur (Lemur cattd) is one of the most familiar examples in 
European menageries. All these lemurs, which have long, 
although non- prehensile tails, differ from the first subfamily by the 
normal structure of the bones of the ankle. The third subfamily 
(Indrisin<z\ which is likewise peculiar to this region, includes the 
avahi, sifakas, and the endrina, all of which differ from the two 
preceding groups by having only thirty, in place of thirty-six teeth ; 
while the endrina is peculiar in having the tail rudimentary. The 
group includes the largest living lemurs ; the sifakas and endrina 




differing from other members of the suborder by their diurnal 
habits. They form a characteristic feature in every wooded Mala- 
gasy landscape, there being scarcely a copse in the island which is 
not tenanted by one or more of these strange creatures ; and when 

FIG. 49. RING-TAILED LEMUR (Lemur catta). 

walking from covert to covert, they do so in an erect posture, with 
their hands clasped behind their necks. 

Whereas the endrina (the largest living lemur) is only two feet 
in length exclusive of the rudimentary tail, the extinct Megala- 
dapis, whose remains have been obtained from the Ambolisatra 
marsh, had a skull three times the size of that of the latter, so that 


the whole animal might be compared in size to a mandrill. The 
skull of this species is characterised by the great elongation of 
the face, and in several respects shows resemblances to that 
of the European Oligocene genus Adapts] although the upper 
molar teeth are peculiar in having tritubercular crowns, whereas 
those of all modern lemurs are quadrangular. There is consider- 
able reason to believe that the giant lemur was actually living in 
the middle of the seventeenth century, an otherwise unknown 
animal being described by De Flacourt in 1658 under the name of 
tretretretre, or tratratratra, which accords fairly well with the fossil 
remains. The giant lemur is, however, not the sole extinct 
member of the group from Madagascar, since the hinder part of a 


skull indicates another, but at present unnamed genus, apparently 
allied to Hapalemur. The last of the Malagasy lemurs is the 
singular aye-aye (Chiromys); a creature representing by itself a 
separate family, broadly distinguished from all other members of 
the suborder by the curious resemblance of the dentition to that of 
the rodents, to say nothing of the extreme elongation and slender- 
ness of the middle finger of the hand. 

Apart from the single musk-shrew, the Malagasy insectivores 
all belong to the group with tritubercular upper molar teeth, 
which, as already mentioned, is now confined to the more southern 
portions of the world, and is evidently a very primitive one. The 
small mouse-like creature (Geogale auritd] representing the Pota- 


mogalidce differs from its Ethiopian cousin, not only in its inferior 
dimensions, but likewise in having but thirty-four in place of forty 
teeth ; and it is possible that, when more fully known, it will have 
to be assigned to a family by itself. As stated in a previous 
chapter 1 , the tenrecs (Centetida) appear to have their nearest allies 
in the West Indian solenodons, although the relationship is now 
believed to be somewhat less close than was formerly supposed. 
The common tenrec ( Centetes), which is the largest member of its 
order, measuring from a foot to sixteen inches in length, is a tail- 
less creature, remarkable for the possession of four pairs of upper 
molar teeth, as in marsupials. Much smaller are the two species 
of Hemicentetes , which, in addition to differences in the dentition, 
are distinguished by having rows of spines along the back at all 
ages, instead of merely in the young condition. The hedgehog- 
tenrecs, forming the genera Ericulus and Echinops, are small 
forms having the whole of the back and tail covered with close-set 
spines. The two other genera are spineless at all ages ; Microgale 
being readily distinguished by the inordinate length of the tail 
which is equal to twice that of the head and body, while 
Oryzorictes has this appendage relatively short. 

The largest, and at the same time one of the most peculiar of 
the Malagasy carnivores is the fossa ( Cryptoprocta), which, although 
usually included in the Viverridce, is so different from all other 
members of that group that it has been regarded as constituting a 
family by itself, specially characterised by the feline type of denti- 
tion. On the other hand, Daubenton's civet (Fossa), although 
representing a genus by itself, has its nearest relative in the widely 
distributed Oriental rasse ( Viverra malaccensts). The latter species, 
although now found both in Madagascar and the Comoro group, 
has in all probability been introduced there. Of the four remain- 
ing genera, Galidictis, Galidia, and Hemigalidia are more or less 
closely allied to the mungooses, although presenting certain 
structural differences from other genera ; but the fourth (Eupleres) 
is so markedly distinct as to constitute a subfamily by itself. 

The five genera of murine rodents call for but little remark, 
although it is noteworthy that they are all more or less closely 

1 Supra, p. 70. 




allied, and belong to the cricetine section, which contains the 
oldest members of the family. Nothing need be said in regard to 
the two ungulates, except that they both belong to Ethiopian 
types. Although bats are not taken much into account in the 
present volume, it is important to notice a peculiar distribution of 
the fruit-bats or Pteropodida ; more especially as this coincides with 
that of many Malagasy birds. On this point Dr Blanford writes 
that " the only African genus belonging to the family is Epomo- 

FIG. 51. THE FOSSA ( Cryptoprocta ferox] . 

^^ which is confined to the continent, whilst throughout the 
Mascarene archipelago, and even in the Comoro islands in the 
Mozambique channel, the typically Oriental genus Pteropus occurs, 
and is represented in various islands by five species, one or two of 
them only distinguished by critical characters from the common 
' flying-fox ' of the Indian peninsula." 

In groups other than mammals, certain common features 
between the reptiles of Madagascar and South America have been 

1 A second genus, Scotonycteris, has been described from the Cameruns 
since this passage was written ; and the author has omitted mention of 
Trygenycferis (Megaloglossus}. 


mentioned in an earlier chapter 1 , where it was attempted to show 
that although these instances of discontinuous distribution might 
be explained by parallel migration from a common northern 
centre, yet that the Tertiary mammalian evidence indicated that 
the American forms had reached their present habitat by way of 
Madagascar and Africa. It will suffice to add here that giant 
land-tortoises, which existed in the Mascarenes during the present 
epoch, are represented by extinct species from the superficial 
deposits of Madagascar; and that the latter have also yielded 
remains of gigantic flightless birds (dEpyornis) markedly distinct 
from any other known type. And here it may be mentioned that 
the chamseleons (Chamaleontida) present a certain similarity in 
their distribution to the lemuroids, the Malagasy region including 
23 out of the 49 species, while nearly all the others are Ethiopian. 
As a whole, the Malagasy reptiles, with the exception of the 
snakes, are stated to be more nearly allied to those of the main- 
land than are either the mammals or the birds ; but the 
amphibians exhibit more decided traces of Oriental affinities. 

Concentrating our attention mainly on the mammals alone, 
their distinctness from those of all other parts of 

Relations of .... 

Madagascar to the world are quite sufficient to indicate the right of 

the Mainland. _ _ , r -\ r ~\ 

Madagascar to form the centre of a separate zoologi- 
cal region. In the survey of the lower Oligocene fauna of Europe 
it has been shown that both lemuroids and civet-like carnivores 
were common, one of the latter having been referred to the existing 
genus Viverra. Hence it is probable that to this fauna we must 
look for the ancestors of the Malagasy mammals. The only 
lemuroids closely allied to those of Madagascar are the African 
galagos, and as the civet-family ( Viverridcz] is better represented 
in Africa than elsewhere, it may be taken for granted that Mada- 
gascar received its mammalian fauna from the mainland. Putting 
aside the hippopotamus and bush-pig, which doubtless arrived 
later, the Malagasy fauna can, however, have been derived from 
Africa only at a time anterior to the introduction of the modern 
types of ungulates into that continent, when it was chiefly popu- 
lated by lemuroids and civet-like carnivores 2 . The question then 

1 Supra, p. 131. 

' 2 It is of course probable that some of the Oligocene primitive ungulates 


narrows itself as to the probable date of the connection between 
the island and the continent. Now, so far as can be determined, 
none of the European Oligocene lemuroids are referable to the 
family Lemuridce and since both the Ethiopian and Malagasy 
representatives of the subfamily Galagince resemble one another in 
the peculiar structure of the ankle, or tarsus, it is pretty evident 
that not only was the family, but likewise the subfamily differenti- 
ated before the separation of Madagascar. Allowing time for the 
southward migration of the Oligocene lemuroids and civets, arid 
the modification of the former into the Galagince, it seems 
impossible to put the separation at an earlier date than the Upper 
Oligocene, while it might well be Miocene 1 . Confirmation of 
this comparatively late separation of the island is afforded by some 
observations of Dr Blanford with regard to the passage of the 
bush-pig across the intervening strait, for it is evident that both 
that animal and the hippopotamus must have reached Madagascar 
by swimming, as otherwise more ungulates would assuredly have 

migrated into Africa with the lemuroids and Viverridce ; but if so, all have 
died out. The Tertiary palseontological history of Africa or Madagascar can 
alone decide this point ; but if ancestors of the South American extinct ungu- 
lates reached their home by way of Africa, it is certain that primitive members 
of that order must have first passed into that continent. 

1 It must be remembered that we are here dealing with the mammalian 
evidence alone. In regard to the molluscan Mr A. H. Cooke (Conchologist, 
1893, p. 131) states that this region possesses sufficient individuality from that 
of the mainland to entitle it to separation. The Helicida are peculiar, not 
being found in the Mascarenes, Seychelles, or Comoros. They seem to be 
related to certain Cingalese and Australian types. Upwards of fifty-four species 
of Cyclostoma are known, distributed over Madagascar, the Comoros, Seychel- 
les, Mauritius, and Bourbon. The African Bulimi are represented by two 
species, but Achatina (so common there) is scarce; and groups of Bulimi are 
peculiar. A single species of the genus Caliella is identified with an Indian 
form ; and unmistakable indications of Oriental affinities are afforded by the 
freshwater molluscs. There are two species of Paludomus, Bithynia occurs, 
and while several of the Melania are of a type common in the Indo- Malayan 
countries, the Melanatria, which are peculiar to Madagascar, have their nearest 
allies in Ceylon or India. Although not a single African freshwater bivalve 
has yet been recorded from Madagascar, yet several Ethiopian genera of 
gastropods occur there, and, in common with the land-molluscs, indicate a 
former connection between Madagascar and Africa, and this, in Mr Cooke's 
opinion, occurred at an immeasurably remote epoch. 


been found in the island. After remarking that bush-pigs are 
stated to be more aquatic in their habits than ordinary swine, Dr 
Blanford 1 asks "how far could Potamochxrus swim? Surely it 
is not likely that it could cross the Straits of Dover. I think we 
are justified in assuming about ten miles as a probable limit of its 
power of crossing the sea, but, to be safe, let us suppose double as 
much. Then, in Pliocene or Plistocene times, quite as probably 
the latter as the former, when Potamochczmis reached South Africa, 
Madagascar was separated by a channel not more than twenty 
miles broad. The conclusion is inevitable, that nearly the whole 
depression of upwards of a thousand fathoms is of Pliocene or 
Post-pliocene date. Of course it must not be assumed that this 
date is proved. What we may consider, however, as beyond any 
doubt is that the depression cannot be older than the Middle 
Tertiary." This view may be taken as practically identical with 
the one here advanced, namely that Africa and Madagascar were 
united till the period of the upper Oligocene or Miocene. 

With the exception of the fruit-bats and Daubenton's civet, 
which, as already mentioned, is more nearly allied to the Oriental 
rasse than to the Ethiopian Viverridce, the Malagasy mammals do 
not exhibit any well-marked alliance with those of India. But the 
case is different with the birds, molluscs, and certain other groups ; 
while we have no evidence that giant land-tortoises ever inhabited 
the African mainland, although an extinct species is known from 
the Indian Pliocene. 

Basing his conclusion on evidence drawn from several sources, 
Dr Blanford, in the communication last cited, is of opinion that 
there was formerly a direct land-connection between India and 
South Africa, and that this connection "included the Archaean 
masses of the Seychelles and Madagascar, that it continued 
throughout upper Cretaceous times, and was broken up into 
islands at an early Tertiary date. Great depression must have 
taken place, and the last remnants of the islands are now doubt- 
less marked by the coral atolls of the Laccadives, Maldives, and 
Chagos, and by the Saya de Malha bank. It is immaterial 
whether Bourbon, Mauritius, and Rodriguez ever formed part of 

1 Appendix, No. 8, p. 88. 


the Mascarene land or not." It is added that if future soundings 
should indicate the absence of a bank extending the whole way 
from India to Africa, it may be a question whether the whole of 
the ocean-bed between those two countries has not sunk to its 
present depth since the Cretaceous era 1 . 

This presumed connection satisfactorily explains much in 
regard to the distribution of the molluscs. It is, however, certain 
that fruit-bats did not exist in the early Tertiary, and the Pteropus 
must accordingly have made the journey across the sea from 
India, aided by what remained of the chain of islands, which may 
have been more extensive during the Pliocene. The same 
explanation also holds good with regard to most of the Oriental 
types of birds. The case of the land-tortoises is, however, more 
difficult. Nearly allied forms have been found in Mauritius, 
Rodriguez, Madagascar, and Aldabra; and since this group is 
unknown, even in Europe, before the Oligocene, it is evident that 
they could not have travelled from India by means of the con- 
necting land-bridge, which is considered to have been broken up 
at the commencement of the Tertiary epoch. This being so, the 
probability is that they originally came from Africa ; but whether 
they entered Madagascar with the ancestral lemurs, or whether 
they, or their eggs, were transported across the channel when 
narrower than at present, there is no evidence to show. Be this 
as it may, it is probable that they reached Rodriguez and Mauritius 
across the intervening sea, since even if these islands ever joined 
Madagascar, such union must apparently have been at a date 
anterior to the existence of true tortoises. That none of these 
tortoises could have been transported by sea from India is proved 
by an observation of Dr Blanford to the effect that on this line the 
currents invariably set from the Seychelles to India. It may be 
added that some writers have considered it probable that the giant 
tortoises of the Malagasy region, like those of the Galapagos 
Islands, attained their large dimensions after they had reached 
the islands they respectively inhabit. The existence of gigantic 

1 Neumayr (Erdgeschichte, 2nd ed. vol. n. p. 262, 1895) considers that when 
India was connected with Madagascar during the Jurassic era, only the southern 
extremity of that island was joined to South Africa. 

L. 15 


species on nearly all the great continents during the Tertiary 
epoch seems, however, an insuperable objection to this view. 

In the absence of any evidence as to the Tertiary vertebrate 
palaeontology of eastern Africa, I have no suggestion to offer as to 
the origin of the gigantic birds of the genus SEpyornis, which 
during the late Plistocene or recent epoch formed such a marked 
feature in the Malagasy avifauna. 

NOTE. The Author has reason to believe that several new 
Malagasy mammals have been discovered by Dr C. I. Forsyth-Major; 
but as no description of these had appeared when this chapter was 
passed for press, they could not be noticed. 



Extent Characteristics of the Mammalian Fauna Birds Past History of 
Ethiopia Subregions. 

IT would be difficult to find a much greater contrast to the 
mammalian fauna of Madagascar than is presented by that of 
Africa south of the tropic of Cancer ; the one area, as shown in the 
last chapter, being characterised by the number of lemurs, together 
with its peculiar Viverridce. and insectivores, while the other is 
distinguished from all other parts of the world by the extraordinary 
number (both as regards genera, species, and individuals) of large 
ungulates which roamed through its plains and forests until deci- 
mated or exterminated by the hand of man. As regards the 
number of individuals of large animals inhabiting equal areas, it is 
quite probable that at the date when the bison flourished in its 
millions on the North American prairies, the balance in this 
respect may have been in favour of the New World ; but whereas 
the prairies had but a single species, Ethiopian Africa was popu- 
lated (for it is unfortunately necessary to write in the past tense) 
with a host of species of antelopes, together with buffaloes, giraffes, 
hippopotami, zebras, rhinoceroses, and elephants. Such a fauna 
has existed during the recent epoch in no other part of the world, 
and in past times has only been paralleled by the lower Pliocene 
fauna of southern Europe and Asia, although even this, as regards 
the number of generic and specific types of antelopes, is by no 
means its equal. 

Separating Madagascar and the associated islands as a distinct 
division, the Ethiopian region may be taken to include such 
portions of Africa and Arabia as lie to the south of the tropic of 



Cancer ; northern Africa, as it did in the Pliocene, clearly forming 
a part of the Holarctic region. The greater part of 
the Sahara, as well as the northern portion of 
the Nubian desert, although included in the Holarctic, will form 
a kind of transition zone towards that region, as is also the case 
with Syria, where a considerable number of Ethiopian types of 
mammals are met with, while western Arabia shows a decided 
approximation to the Oriental region, as is well exemplified by 
the occurrence there of a species of the short-horned goats con- 
stituting the genus Hemitragus. As has been stated in an earlier 
chapter, the Sahara and Nubian deserts, although they have 
apparently never been submerged since the Cretaceous epoch, seem 
always to have formed a more or less complete barrier to the passage 
of the mammals of Algeria and the adjacent countries into the 
Ethiopian region ; and the main migration from the north and east 
has thus taken place along the north-eastern side of the continent. 
With the exception of its southern extremity, the whole of this 
vast area lies within the tropics. As regards its physical features, 
Dr Heilprin writes that " it presents several well-marked physical 
peculiarities. In the first place, we have the vast expanse of 
desert, which in the north occupies a transverse band varying in 
width from about four to nearly ten degrees of latitude. This is 
succeeded by what may not improperly be termed the open 
pasture-lands, which as a narrow belt bounds the Sahara on the 
south, curves southwards at about the position of Kordofan, and 
occupies the greater portion of the continent lying east of the 
thirtieth parallel of east longitude and south of the fifth parallel of 
south latitude. A very considerable portion of this pasture-tract 
forms a plateau of from four thousand to five thousand feet eleva- 
tion. Included within it, and bounded on the west by the Atlantic 
Ocean, is the region of the great equatorial forests, to the present 
day a terra incognita in great part both to geographers and 
naturalists. That portion of the African continent lying south of the 
tropic of Capricorn differs in many respects, both as to its physical 
configuration and its vegetable products, from the region to the 
northward, and is characterised by a vegetation which is one of 
the richest and most remarkable on the globe. With this marked 
peculiarity in its vegetable development there is of necessity a 


certain amount of faunal peculiarity superadded as well, but this is 
not sufficiently pronounced to permit of the separation of this 
tract from the tract lying immediately to the north. We have thus 
on the continent three strictly defined faunal sub-regions : (i) the 
pasture-lands already described, constituting the East Central 
African sub-region, through whose vast expanse there is manifest 
a strong identity in the character of the animal products, the same 
or very closely related forms being in many instances found at the 
extreme points of this sub-region ; (2) the forest-tract, constituting 
the West African sub-region, whose animal products naturally 
differ very essentially from those of the last ; and (3) the desert or 
Saharan sub-region, containing a comparatively limited fauna, 
which, with almost insensible gradations, merges into the fauna of 
the Mediterranean tract. To the same division belong in great 
measure the desert tracts of Arabia, or that portion of the 
peninsula lying to the south of the tropic of Cancer." 

Although Dr Wallace had previously divided continental 
Ethiopia into an East African, West African, and South African 
sub -region, the foregoing arrangement seems, on the whole, 
preferable. There are, however, considerable reasons for regard- 
ing Somaliland as a sub-region by itself, and South Arabia should 
perhaps constitute another. Although the precise determination 
of such areas does not come within the province of this work, it is 
most important to notice that the West African or Equatorial 
forest tract continues right across the continent as far eastwards as 
the Congo- Nile watershed, that is to say, close up to Wadelai, 
where all traces of the West African fauna are suddenly lost. On 
this point Mr O. Thomas 1 writes that "the abruptness with which 
the change of fauna occurs on the watershed is, considering the 
insignificant nature of the physical barriers, very remarkable, and 
almost unequalled in the distribution of the mammals of any part 
of the world. The reason of the change is, however, clear enough, 
being not the occurrence of such barriers to migration as moun- 
tains or rivers, but the abrupt ending of the great West African 
forest, which, as we know from the travels of Schweinfurth and 
ethers, extends quite into this region, but abruptly ceases before 
the slopes of the upper Nile basin are reached." 
1 Proc. Zool. Sec. 1888, p. 17. 


Before considering the leading characteristics of the Ethiopian 

mammal fauna and its relations to that of other 

characteris- regions, both in the present and the past, it is 

tics of Mam- . & . . 

maiian Fauna, desirable to make reference to certain deficiencies, 
which are very difficult, if not impossible to explain 
adequately with our present knowledge. 

Although deer (Cervus), typical pigs (the genus Sus in its 
restricted sense), and bears are met with in northern Africa, no 
member of any one of these genera with the single exception of 
a pig (Sus sennaarensis) from the Sennaar district of Upper Nubia, 
inhabits Ethiopia. Even the entire family of the Cervtdce is 
unrepresented. These deficiencies form a most marked contrast 
between the Ethiopian region on the one hand, and both the 
Oriental and the Holarctic on the other. Almost equally con- 
spicuous is the absence of goats and sheep ; the only exceptions 
being the occurrence of a species of Capra in the highlands of 
Abyssinia, and one of Hemitragus in Oman, in south-eastern 
Arabia. The absence of sheep and goats is, however, by no means 
so remarkable as that of the other groups above mentioned, since 
the former are exclusively mountain animals, and probably need 
some general lowering of the temperature to enable them to pass 
from one chain to another, and of the existence of such cold 
period there seems no evidence in Ethiopian Africa. A somewhat 
similar explanation will probably apply to the total absence of 
marmots (Arctomys\ susliks (Spermophilus], chipmunks (Tamias), 
beavers (Castoridce], voles (Microtince), and picas (Lagomys), since 
all these are inhabitants of elevated or northern areas. More diffi- 
cult to explain is the absence of all shrews (Soricidce), with the 
exception of one genus peculiar to the region ; but the deficiency 
of moles (TalpidcR) may perhaps be accounted for by the slow 
travelling powers of these animals, which did not allow them 
time to pass into Ethiopia during the (probably short) period 
when its connection with other regions was of such a nature as to 
permit their living in the intermediate lands. Possibly also the 
absence of moles from peninsular India has something to do with 
this deficiency. In this connection it is worth remark that the 
place held in the Holarctic region by moles is by no means 
unoccupied in the Ethiopian, both the golden moles ( Chrysochloris), 


and the so-called Cape mole (Bathyergus), with its allies, having 
similar subterranean habits. 

Together with the Oriental, the Ethiopian region shews a 
marked distinction from all others as the sole habitat of the 
man-like apes (Simiidce). The Ethiopian forms comprise the 
chimpanzees (Anthropopithecus) and gorilla (Gorilla), both of 
which are restricted to the equatorial forest-region, where the 
former ranges as far east as Uganda, although the latter has a 
more circumscribed distribution. The occurrence of a fossil 
chimpanzee in the Indian Pliocene affords the most convincing 
evidence of the derivation of a large part of the Ethiopian fauna 
from what is now the Oriental region. Among the ordinary 
monkeys and baboons (Cercopithetida) there are five genera con- 
fined to this region. Of these, Colobus differs from the Oriental 
langurs (Semnopithecus] by the absence or rudimentary condition of 
the thumb, which frequently has lost all trace of a nail. On the 
other hand, the large genus Cercopithecus^ which is most fully 
represented in the forest region, is as nearly related to the Oriental 
Macacus, from which it differs in the less prominent muzzle, and 
the absence of a projecting heel, or hinder lobe to the last lower 
molar tooth. This heel is, however, present in the mangabeys, or 
white-eyelid monkeys (Cercocebus), all of which are exclusively 
confined to the forest tract. Although the dog-faced baboons 
(Papio 1 ) have a wider distribution, ranging from the Cape to 
Arabia, some of the largest and most peculiar forms, such as the 
mandrill and drill, are confined to West Africa. This genus is one 
of those common to the Ethiopian region and the Indian Pliocene. 
The nearly-allied gelada baboons (Theropithecus], of which there 
are two representatives, are, on the other hand, exclusively north- 
eastern types, one being confined to Abyssinia. Among the 
lemuroids, the galagos (Galago*) which, as stated in the last 
chapter, belong to a sub-family which attains its maximum develop- 
ment in Madagascar extend right across the equatorial portion of 
the continent, descending somewhat on the east coast, where they 
are very numerously represented. The pottos (Perodicticus], which 
are nearly related to the lorises of the Oriental region, are, how- 

1 Syn. Cynocephahis. * Including Otogale. 




ever, exclusively confined to West Africa, where they are known 
by two species, regarded by some as representing as many genera. 

It will be unnecessary to say more with regard to the Chiro- 
ptera than that the fruit-bats (Pteropodidce) are represented solely by 
three peculiar genera in the Ethiopian region, of which Epomo- 
phorus has the greater number and the most peculiar of its species 
confined to the western forest tract, while the single species of 
Scotonycteris is solely found in this part of this continent, as is 
also the case with that of Trygenycteris. 

Of great importance from a distributional point of view are the 
Ethiopian Insectivora, since of the five families found within the 

FIG. 52. AFRICAN JUMPING-SHREW (Macroscelides tetradactyhis]. 

area under consideration, two are almost or exclusively confined 
to it, while the third has only one aberrant representative in Mada- 
gascar, and even this may prove entitled to constitute a family by 
itself when its structure is more fully known than is at present the 


case. It is further noticeable that two of the families belong to 
the primitive group characterised by their tritubercular upper molar 
teeth, and were accordingly in all probability very early immigrants 
into the country. The first family almost peculiar to the region is 
that of the jumping-shrews (Macroscelididce), the members of which 
are easily recognised by their elongated hind limbs, long snout, 
and leaping habits. While the typical genus Macroscelides has 
representatives throughout the region and also extends into 
Holarctic Africa, the four species of Rhynchocyon, in which the 
legs are shorter and the snout longer, are restricted to East Africa. 
To the latter is closely allied the European Oligocene genus 
Pseudorhynchocyon, and it would thus seem that the family, 
although unrepresented in Madagascar, arrived at a relatively 
early date in Ethiopia. On the other hand, since the hedgehogs 
are represented only by the widely-spread typical genus Erinaceus, 
together with a West African species which has been separated as 

FIG. 53. WEST AFRICAN POTAMOGALE (Potamogale velox]. 

Proechinus, it is probable that they did not make their appearance 
on the scene till a later epoch. The Soricidce, or shrews, are 
represented in Ethiopia only by three species belonging to the 
peculiar genus Myosorex, differing from all the other genera in 


which the teeth are white by the absence of long hairs on the tail 1 . 
With the West African genus Potamogale, we come to the first of 
the two families with tritubercular upper molars ; the present one 
(Potamogalidce] being represented elsewhere only by the Malagasy 
Microgale, of which, as already said, the systematic position is 
doubtful. The potamogales, which attain a couple of feet in 
length, are thoroughly aquatic in their habits, swimming by the aid 
of the highly compressed tail. It has generally been considered 
that there is only a single species, but it has recently been 
suggested that there may be two. The family is probably an 
ancient one, although we have no fossil evidence to this effect, 
Microgale, even if it belong to another family, indicating that the 
group was among the earlier mammalian colonists of Ethiopia. 
The golden moles (Chrysochloridcz], which take their name from 
the brilliant metallic lustre of the fur in the majority of the species, 
are blind, earless, fossorial insectivores, having the middle toe of 
the fore foot furnished with an enormously powerful claw. As the 
moles (Talpida] form a group nearly related to the shrews, so the 
golden moles are equally nearly related to the tenrecs ( Centetidcz) 
of Madagascar, from which they may be regarded as a highly 
specialised offshoot. Accordingly, it may be taken for granted 
that their ancestors obtained an entry into Ethiopia with the 
ancestral lemurs. It is not improbable that the prevalence of 
higher types of mammalian life has been the cause of the assump- 
tion of mole-like habits in the Chrysochloridce ; the tenrecs, which 
live in an island where the competition is much less severe, having 
retained the original primitive type. The golden moles, which 
may all be included in the single genus Chrysochloris, are mainly 
confined to South Africa, although one species extends on the 
east coast as far north as Ugogo. 

Turning to the Carnivora, it is unnecessary to say anything 
with regard to the Felida, except that three species of felts are 
common to Ethiopia and India; while the single species of 
hunting-leopard (Cyncelurus] is likewise found in both countries, 
the genus being apparently also represented in the Indian Pliocene. 
In the civet family ( Viverridce) the true civets ( Viverra] and 

1 See Dobson, Proc. Zool. Soc. 1887, p. 575. 


mungooses (Herpestes] are common to the Ethiopian and Oriental 
regions ; but the whole group attains a far greater development 
within the former area than elsewhere. Although the common 
genet is an inhabitant of the southern portion of the eastern 
Holarctic region, all the other species of Genetta are Ethiopian. 
The West African linsang (Poland] is the Ethiopian representative 
of the beautiful linsangs (Linsangd) of the eastern portion of the 
Oriental region ; the distribution of this group being a well-marked 
instance of the close alliance of the fauna of the Malayan 
countries to that of West Africa, to which reference will again be 
made. The Oriental palm-civets (Paradoxurus) are represented 
by the nearly-allied Ethiopian genus Nandinia, of which one 
species is West African, while the other comes from Nyasaland. 
The small-toothed mungoose (Helogale] is common to West and 
East Africa ; and the allied genus Bdeogale has representatives 
on both sides of the continent. Two other peculiar Ethiopian 
genera, Cynictis and Rhynchogale, have each but a single species : the 
former being South African and the latter East African. The 
cusimanses (Crossarchus), although mainly characteristic of the 
forest tract, have one representative in Abyssinia. Lastly, the 
meerkat, the sole representative of the genus Suricata, is an 
exclusively South African form. A peculiar family (Proteleidce) is 
constituted by the aard-wolf (Proteles], ranging from the Cape to 
Somaliland, and a near ally of the hyaenas, from which it is 
distinguished by the extremely feeble development of the denti- 
tion. Both the spotted hyaena (Hyczna crocuta) and the brown 
hyaena (H. fused) are now confined to Ethiopia, but the former 
ranged over a large portion of Europe as well as southern India 
during the Plistocene epoch ; and as all the three living species 
are included in the same genus, there is no generic type in this 
family restricted to the Ethiopian region. 

In the Canidcz wolves are absent, but the jackals are repre- 
sented by species allied to Cam's aureus, which occurs in North 
Africa; wild dogs (sub-genus Cyon) are, however, wanting. 
Although the long-eared foxes or fennecs, such as Cants chama, 
are common in Ethiopia, they are by no means characteristic, 
since they range into North Africa, Syria, Persia, and Afghanistan ; 
being, in fact, like the gazelles, desert-haunting forms. There are, 

2 3 6 



however, two genera of the family, each with a single species, now 
confined to this region ; the first being represented by the Cape 
hunting-dog {Lycaon pictus], which is a large, somewhat hy?ena-like 
animal, easily recognised by its spotted coloration and long bushy 
tail, and distinguished from the other genera by having only four 
toes on both the fore and hind feet. That the genus is of northern 

FIG. 54. CAPE HUNTING-DOG (Lycaon pictits). 

origin is proved by the occurrence of remains of an extinct species 
in the Glamorganshire caves. The small Lalande's fox (Otocyon 
megalotis) of South Africa, in addition to the enormous size of its 
ears, is peculiar in having four pairs of molar teeth in the lower 
jaw, and either three or four in the upper. Possibly a fossil species 
from the Indian Pliocene may be an allied type. 

Passing by the Ursidce. and Procyonidce as unrepresented in 
this region, we find the Mustelida very poorly developed, martens 
(Mustela) being absent, and true weasels very scarce. The striped 
Gape weasel (Poedlogale] constitutes, however, a genus by itself; 
while the similarly coloured Cape polecat (fctonyx), is one of two 
representatives of a small genus, the second of which ranges from 




Sennaar to Egypt, and is also stated to occur in Asia Minor. A 
fossil species from the European Miocene may perhaps belong to 
this genus. The ratels (Mellivora) which are now represented by 
one Ethiopian and a second Indian species, are proved to be com- 
paratively late immigrants from the north into this region by the 
occurrence of a fossil species in the lower Pliocene of Northern 
India. As the animal described under the name of Galeriscus is 
at present known only by a skin, it is not even certain that it 
belongs to the Mustelidce. at all. 

Among the squirrel-like rodents, the most striking feature is 
the absence of the true flying-squirrels and their replacement by a 


distinct family (Anomaluridce.}, characterised, among other features, 
by the presence of scales on the under surface of the root of the 
tail. Mainly characteristic of the forest area, the typical genus 
Anomalurus has, nevertheless, representatives on the eastern side 


of the continent; although the mouse-like long-tailed flying- squirrel 
(Idiurus) is exclusively West African. While true squirrels (Sci- 
urus] are common to this and the other regions of Arctogaea, the 
pigmy squirrels constituting the genus Nannosciurus are repre- 
sented by one species (N. minutus) in West Africa, while all the 
others are Malayan. The spiny squirrels of the genus Xerus are 
now, on the other hand, exclusively Ethiopian, although their 
northern origin is proclaimed by the occurrence of an extinct 
species in the French Miocene. Dormice (Myoxidce) are exceed- 
ingly abundant in Ethiopia, and if it is considered desirable to 
split up the family into more than two genera, the genus Graphi- 
urus, characterised by the short, cylindrical, and tufted tail, and 
the simple structure of the molar teeth, will be peculiar to this 
region, as will also be the single West African form described as 
Claviglis. Dormice, as mentioned earlier, date in Europe from 
the lower Oligocene, and therefore they might well have entered 
Ethiopia with the ancestral lemuroids, although, so far as it goes, 
their absence from Madagascar is against this view. In the 
mouse-family (Muridce) five sub-families are met with in Ethiopia, 
two of which are peculiar to the region. Curiously enough, the 
cricetine sub-family, which is the oldest of all, and the only one 
met with in Madagascar, is represented only by a single highly 
specialised genus (Trilophomys). The inference from this would 
seem to be that the ancestral cricetines and lemuroids entered 
Ethiopia together, whence some migrated to Madagascar ; the 
former group, with the exception of the one peculiar genus, having 
completely died out on the continent, where the remaining 
murines are more recent immigrants from the north. In this 
family the Eastern Arctogaeic group of the gerbils (Gerbillince) 
includes six genera, out of which no less than five, namely 
Pachyuromys, Mystromys, Otomys, Dasymys, and Malacomys, are 
exclusively Ethiopian, the last being West African. The elongated 
hind limbs and the transverse laminae of the molars readily serve 
to distinguish the gerbils from the exclusively Ethiopian sub- 
family Dendromyin&i in which the molars are rooted and tuber- 
culated and the ears remarkably hairy. The typical genus 
Dendromys includes two dormouse-like forms, one from South, 
and the other from East Africa; the other genera being Lima- 


comys, Steatomys, and Lophuromys, the last having the fur replaced 
by fine flattened bristles. Although there is no palaeontological 
history of either of the preceding sub-families, the Cricetince have 
already been shown to date from the lower Oligocene of Europe. 
Their sole Ethiopian representative is the single species of the 
genus Trilophomys^, from Upper Nubia and the Red Sea littoral 
in the neighbourhood of Suakin, and perhaps ranging into southern 
Arabia. The crested rat, as the creature is called, takes its name 
from the prominent crest of stiff hair running down the back ; 
while it is specially characterised by the roofing over of the whole 
upper surface of the skull with bone, on which account it has 
(quite unnecessarily) been made the type of a family by itself. 
Perhaps, however, the most interesting member of the family 
inhabiting Ethiopia is a species of mouse known as Deomys, repre- 
senting both a genus and sub-family by itself, and characterised by 
having its upper molars intermediate between those of the crice- 
tines and murines. Doubtless, therefore, this rodent is a somewhat 
modified descendant of the true cricetines which entered Africa 
while it was still united to Madagascar; its habitat being that 
refuge for ancient types, the lower Congo valley. The two other 
generic representatives of the family, Cricetomys and Saccostomus, 
although resembling the hamsters in the presence of cheek- 
pouches, have molars like other Murina, and are accordingly 
referred to that sub-family. While there are two species of the 
second genus, there is but one of the first. By some zoologists 
the striped mice, as typically represented by the Barbary mouse, 
are separated from Mus to form a genus Arvicanthis* , which is 
mainly Ethiopian. 

The small family of the Spalacidce, or burrowing-rats, has four 
genera out of six exclusively Ethiopian, while a fifth (Rhizomys\ 
which is mainly Oriental, enters Abyssinia. Of the four Ethiopian 
genera, which constitute a sub-family by themselves, Bathyergus 
includes only the great Cape mole-rat of South Africa; and 
Myoscalops has also but a single species, although there are several 
of Georhychus. Closely allied to the latter are two tiny little 
burrowing and nearly naked creatures (Heterocephalus) from 

1 Syn. Lophiomys. '* Syn. Isomys. 


Somaliland, which may be regarded as degraded descendants from 
that type. The family is unknown either in Madagascar or in a 
fossil state in Europe, although an extinct species of Rhizomys 
occurs in the Indian Pliocene, and it is, therefore, in all proba- 
bility a comparatively late immigrant into Ethiopia. In addition 
to one genus ranging from Nubia to Siberia, the jerboa-family 
(DipodidcE) has one peculiar Ethiopian genus, represented solely 
by the Cape jumping-hare (Pedetes) ; a form so different from all 
the others that it must constitute a sub-family apart. With this 
genus we leave the mouse-like, and come to the porcupine-like 
group of the rodent order, in which the family Octodontidce. has 
nearly all of its representatives which are not Neogaeic confined to 
this region, although the gundi (Ctenodactylus) of North Africa, in 
the neighbourhood of Tripoli, is south Holarctic. This genus and 
an allied type (Pectinator) from Somaliland, together with the next 
form, alone represent a sub-family typically characterised by the 
presence of a horny comb-like appendage and stiff bristles on each 
of the hind feet ; Pectinator thus being Ethiopian. The South 
African rock-rat (Petromys), although now included in the same 
group, approximates to a sub-family of which all the members are 
South American ; the resemblance between one of the latter and 
the Ethiopian species being curiously close. The two species of 
cane-rat (Triaulacodus 1 ) constitute the sole African representatives 
of a sub-family containing a large number of Neogseic genera: 
Probably, as there has already been occasion to remark, both the 
Neogaeic and Ethiopian representatives of this family trace their 
origin to the extinct Theridomyidce of the Oligocene of the Hoi- 
arctic region or some nearly allied forms ; and as certain forms 
occur in the Santa Cruz beds of Patagonia, it is probable that the 
migration into Ethiopia was at least as early as that of the early 
lemuroids, the extinct Pellegrinia of the Sicilian Pliocene being 
the last survivor of the group in the northern half of the Old 
World 2 . Although there is no generic type of porcupine (Hystri- 

1 This name is proposed to replace Aulacodus, Temm. (1827), which was 
preoccupied in 1822 by Eschscholtz for a genus of Coleoptera. 

2 There is difficulty in this respect on account of the absence of hystrico- 
morphous rodents from Madagascar; but perhaps the early forms entered the 
west side of the continent, while the lemurs travelled in along the east. 


cidai) peculiar to Ethiopia, it is not improbable from the wide dis- 
tribution and antiquity of the group, that these rodents also entered 
tropical Africa at a comparatively early epoch. The deductions 
drawn from these rodents as to a connection between Africa and 
South America have been mentioned in Chapter III. 

Among all the striking features of the mammalian fauna of 
Ethiopian Africa, none is more remarkable than the enormous 
preponderance of ungulates, many of which are of great corporeal 
bulk. Of these a large number of genera and two families are 
absolutely peculiar to this region. As may be gathered both from 
their absence at the present day in Madagascar, and the late 
epoch at which their remains are found in the Tertiaries of the 
Holarctic and Oriental regions, all these creatures have reached 
the Ethiopian region but recently. The Hippopotamida is one 
of the two families now practically peculiar to the region, the 
common species {Hippopotamus amphibius) having ranged over a 
considerable portion of Europe during the Plistocene and upper 
Pliocene ages, while even in the beginning of this century it 

FIG. 56. HEAD OF WART HOG {Phacochtxrus cethiopicus). 

frequented lower Egypt. The pigmy hippopotamus (H. liberi- 

ensis) of western Africa, which is referred by many writers to a 

genus apart, and more resembles the pigs in its mode of life, 

I, 1 6 


appears to be more nearly allied to a small species from the 
Sicilian and Maltese Pliocene. As already mentioned, with the 
exception of a single species (Sus sennaariensis], true pigs are 
unknown in Ethiopia, their place being taken by the two species 
of bush-pigs, forming the potamochoerine group of the same genus, 
and distinguished by the simpler structure of the molar teeth, as 
well as by the tendency of the front premolars to fall out in the 
adult. The reason for the occurrence of a third species of the 
group in Madagascar has been already sufficiently discussed 1 . 
Still more distinctive of the region are the hideous wart-hogs 
(Phacocharus\ specially characterised by the facial warts from 
which they take their name, the huge tusks, and the great com- 
plexity of the last molar tooth in each jaw ; the tusks and these 
molars being frequently the only teeth remaining in aged animals. 
It is, however, very noteworthy that certain extinct species of Sus 
from the Pliocene of India and Algeria have their last molars of a 
type which could easily be developed into those of the wart-hogs ; 
and it would accordingly seem that the latter are comparatively 
recent descendants of ordinary pigs. Although wild camels are 
unknown in the region, the Tragulida are represented in West 
Africa by the water-chevrotain, which is now the only existing 
species of the genus Dorcatherium, although fossil forms occur in 
the Pliocene of India and the European Miocene ; the ancestral 
forms having probably entered the region from India. The second 
ungulate family now confined to Ethiopia is the Giraffidce, of 
which there appears to be only a single living species, although the 
North African form shows a decided difference in coloration from 
its southern brother. The occurrence of species belonging to the 
existing genus Giraffa in the lower Pliocene of Greece, Persia, 
India, and China, shows that giraffes came into Africa with the 
other ruminants ; the African species being very probably the 
direct descendant of the extinct Indian one. 

Abounding as it does in ungulates in general, Ethiopian Africa 
is the especial home of the antelope group, which here takes 
the place of the sheep and goats so characteristic of the elevated 
districts of the eastern half of the Holarctic region. Regarding 

1 Supra, p. 223. 




their distribution in the Ethiopian region, Dr Sclater writes 1 that 
although " antelopes are to be met with in every part of Africa, 
they are most numerous where the country is comparatively open, 
and where there are grassy plains interspersed with sheltering 
bushes. South of the tropic of Capricorn this condition generally 
prevails, and throughout the Cape Colony and its adjoining terri- 
tory they are or, at all events, before the advent of a European 

FIG. 57. WATER-CHEVROTAIN (Dorcatherium aqiiaticum}. 

population were, everywhere abundant. The early settlers at the 
Cape describe antelopes as to be met with in herds of thousands 
on the veldt, and in parts of Africa where the white man and his 
destructive firearms have not yet penetrated a similar condition 
prevails even at the present day. When we advance further north 
and meet with the dense forests of the Niger and Congo basins, 
we find the mass of antelopes holding rather to the more open 
lands on the eastern coast, throughout which they are to be met 

Natural Science, vol. I. p. -255 (1892). 

1 6 2 


with in great abundance up to Cape Guardafui. The vast plains 
traversed by the Upper Nile and its tributaries are likewise well 
stocked with antelope life ; but in the great Sahara only some of 
the more desert-loving forms are to be found. In Senegambia 
again, and in the more open districts on the West Coast, many 
forms of antelopes occur, but they cannot rival the numbers and 
varieties of those of Eastern and Southern Africa." Although 
most of the genera of Ethiopian antelopes are peculiar to the 
continent, a few desert-haunting types range into southern Arabia, 
and hence northwards into Syria, where they enter the Holarctic 
region. Many of the groups have extinct representatives in the 
lower Pliocene of southern Europe and India, and since existing 
Ethiopian genera are more common in the latter than in the 
former area, it seems probable that the great migration into 
Ethiopian Africa has taken place from the east, by way of Syria or 
Arabia. As such extinct genera or species have been already 
noticed 1 , it will suffice to take a very brief survey of the genera 
now mainly or exclusively confined to Ethiopian Africa. 

The first section includes the hartebeests and their allies the 
bontebok and blesbok, all of which may well be included in the 
genus Bubalis, although the two latter are often separated as 
Damaliscus. One species of the typical group ranges into Syria, 
while a second is an inhabitant of Tunis. To the same section 
also belong the wildebeests, or gnus ( Connochcetes). On the other 
hand the numerous species of duikerboks (Cephalophus) consti- 
tute, so far as Africa is concerned, a section to themselves. They 
are, however, allied to the Indian four-horned antelope (Tetraceros\ 
and it is not improbable that they are represented in the Pliocene 
of the Siwalik Hills. While many species of the genus are found 
in East and South Africa, the largest kinds are confined to the 
forest-districts of the West Coast. The small African antelopes 
classed by Sir V. Brooke in the Ceruicaprince and included in 
the genera Neotragus and Nanotragus are now referred by Messrs 
Sclater and Thomas to a section apart, under the name of 
Nanotragin<z, and are classed in six genera. Of these, Madoqua ~, 
with six species, includes Salt's antelope (M. saltiand)\ Nanotragus 

1 Supra, pp. 197 206. 2 Syn. Neotragtis. 


is represented only by the minute royal antelope (JV. pygmaus} 
of Guinea; Nesotragus is typified by the Zanzibar steinbok 
(JV. moschatus] ; the true steinbok (Raphiceros campestris'} forms 
the fourth genus ; the oribi of South Africa ( Oribia scoparia) is 
still more distinct; while the well-known klipspringer (Oreotragus 
saltator}, which ranges from the Cape to Abyssinia, differs from 
all the rest by its coarse brittle fur. 

The Cervicaprince, include larger forms. Foremost among 
these is the South African rheebok (Pelea) ; while the water-buck 
and its allies (Cobtts) are some of the largest of all antelopes. 
The last representative of this section (Cervicaprd) is typified by 
the South African rietbok, but there are also other species in West 
and East Africa. The fine South African antelope known as the 
pala (sEpyceros], together with an allied species from the West 
Coast, form the first representatives of another section. Here 
belong the true gazelles (Gazella), more characteristic of the desert 
tracts of the eastern half of the Holarctic region, although repre- 
sented in South Africa by the somewhat aberrant springbok, as 
well as by several more typical species on the East Coast. Clarke's 
gazelle (Ammodorcas) of Somaliland is, however, the sole species 
of an exclusively Ethiopian genus ; and the same is the case with 
Waller's gazelle (Lithocranius\ of which the range extends on the 
East Coast from Somaliland to Kilimanjaro. The single species 
of the East African genus Dorcatragus seems to be an aberrant 
gazelle, with the trunk-like muzzle of Madoqua, but retaining 
the small gland-pits of the type. Yet another section of 
antelopes is typified by the beautiful sable antelope and its 
allies (Hippotragus), in which the horns sweep backwards in a 
graceful curve, and are ringed nearly to their tips. This genus is 
exclusively Ethiopian, but in the one typified by the gemsbok 
(Oryx), and characterised by the long and slender horns being 
either straight or but slightly curved, and ringed only at the base, 
the range includes all the desert regions of Africa, Arabia, and 
Syria, although the majority of the species are Ethiopian. To the 
same section belongs the addax antelope (Addax), but although 
this animal occurs as far south as latitude 18 N., it is mainly an 
inhabitant of the deserts of North Africa, Arabia, and Syria. The 
last section includes some of the largest and at the same time the 




most beautiful of all antelopes, the three Ethiopian genera being 
all characterised by the more or less strongly-marked spiral twisting 
of their horns, and the short crowns of their molar teeth ; the last 
feature distinguishing them sharply from the sable antelope and 
gemsbok group. The first of the genera in question includes the 

FIG. 58. HEAD OF GEMSBOK ( Oryx gazdla) . 

harnessed antelopes (Tragelaphus), in which the spiral twisting of 
the horns is less marked than in the other two ; these antelopes 
frequenting forest or jungle, and being most numerous in western 
Africa. The kudus (Strepsiceros] agree with the last in that the 
females are hornless, but the horns of the males form a more 
corkscrew-like spiral than is generally the case with the harnessed 


antelopes; while in the elands (Orias) 1 the horns are present in 
both sexes and form a close spiral. 

Turning to the perissodactyle section of the order, we find 
Ethiopian Africa the home of several species differing markedly 
from any now living in other parts of the world, although, accord- 
ing to the system here adopted, these do not constitute generic 
groups by themselves. There are two, or possibly three species of 
Rhinoceros, both of which are two-horned, and differ from those of 
the Oriental region in the absence of canine and incisor teeth. Of 
these the common African rhinoceros (R. bicornis) is closely allied to 
an extinct species from the Pikermi beds of Attica; while Burchell's 
rhinoceros (R. simus), which is now nearly exterminated, has its 
nearest allies in the extinct R. platyrhinus of the Siwalik Hills, and 
the woolly rhinoceros (R. antiquitatis) of the Plistocene of Europe 
and northern Asia. In addition to being the habitat of the parent 
form of the domestic ass, which is confined to Somaliland and the 
adjacent regions, Ethiopian Africa is also characterised by contain- 
ing all the striped horses, or zebras, separated by some authorities 
as a distinct genus, under the name of Hippotigris. The best 
known representatives of this group are the true or mountain 
zebra (Equus zebra], Burchell's zebra (E. burcheili\ Grevy's zebra 
(E. grevyi] of the Galla country, and the quagga (E. quagga). The 
latter, although formerly abundant in the southern extremity of the 
continent, is now fast verging on extinction, and serves to connect 
the more typical members of the group with the true asses. In 
spite of the difference in their coloration, the zebras are indeed 
indistinguishable in their osteology and dentition from the latter, 
and it is quite possible that they are represented in a fossil state in 
the later Tertiaries of Europe, while they are not improbably the 
direct descendants of the ancestral genus Hipparion. The absence 
of tapirs from the Ethiopian region is as remarkable as the want 
of deer ; but it is noteworthy that the former are unknown among 
both the Pikermi and Siwalik faunas. 

Although the African elephant (Elephas africanus) is markedly 
distinct from the Oriental species, yet the two are so closely con- 
nected by intermediate extinct forms that it is impossible to regard 

1 The name is usually spelt Oreas, but as it is derived from <5/>etds, the 
proper orthography is Orias. 




them as the representatives of separate genera. The existing 
species is now confined to the Ethiopian region, but since its 
fossilised remains occur in deposits of Plistocene age in Algeria, 
Spain, and Sicily, it is evident that, like the spotted hyaena and 
lion, it formerly enjoyed a much more extensive range. 

With the exception that a single species is found in Syria, the 
small rodent-like ungulates, known as hyraces, which constitute 
not only a family (Procavtidce) but likewise a sub-order (Hyracoidea) 
by themselves, are especially characteristic of the Ethiopian region, 
where they are represented by a large number of species, more 

FIG. 59. CAPE HYRAX (Procavia capensis]. 

particularly in the southern portion of the continent. Although 
these animals closely resemble the rhinoceroses in the structure of 
their molar teeth, they differ markedly from all the perissodactyles 
in having the carpus constructed on the linear type 1 , and 
from all other living forms of the order in that their single pair 
of upper incisor teeth grow continuously throughout life, as in 
the rodents. They were formerly divided into at least two 
generic groups, but both the terrestrial and arboreal forms are now 
included in the single genus Procavia. Nothing definite is 

1 See figure 12 on p. 78. 


known as to the past history of the group 1 ; but it has been 
suggested (p. 85) that they may be allied to certain extinct South 
American ungulates. 

The list of peculiar Ethiopian mammals is brought to a close 
by the aard-varks (Orycteropodida), which although generally in- 
cluded in the Edentata have nothing to do with the typical South 
American representatives of that order, and are here, together with 
the pangolins, regarded as forming an ordinal group Effodientia 
by themselves. Only a single genus (Orycteropus] now exists, of 
which there are two living Ethiopian species, and there are 
extinct species in the Pliocene of Persia and Samos. A skull 
from the Plistocene of Madagascar has been described as Plesi- 
orycteropus, and another genus occurs in the French Oligocene. 
Not improbably some members of the family entered Africa and 
Madagascar with the ancestral lemuroids and civets, but the dis- 
covery of the Pliocene forms renders it probable that the existing 
genus is a later immigrant. 

Finally, it may be mentioned that among the more widely- 
spread genera a few species of mammals are either now common 
to the Ethiopian region and India, or were so during the Plistocene 
age. In the Felida the lion (Felts leo), leopard (F. pardus), jungle- 
cat (F. chaus], caracal (F. caracal), and hunting-leopard (Cynce- 
lurus jubatus] still range over the two areas ; fossilised remains of 
the first three of these also occurring in the European Plistocene 
deposits. On the other hand, the spotted hyaena (Hycena crocutd), 
which lived in Southern India (as well as in Europe) during the 
Plistocene era, is now restricted to Ethiopian Africa; and the 
same is the case with the giant pangolin (Mains gigantea) of West 
Africa, fossilised remains of which have been discovered, in 
company with those of the spotted hyaena, in a cavern in 

The following table shows the genera and family of mammals 
now more or less exclusively restricted to the Ethiopian region ; 
the names of such as are practically peculiar to this area being 
printed in italic type. 

1 I am informed that a skull belonging to an extinct member of this group 
has been discovered in the Pliocene of Samos, but no description has been 


I. Primates. 



Anthropopithecus. Equatorial Africa. Fossil in 

Indian Pliocene. 
Gorilla. W. African. 

Co lob us. 

Cercopithecus. Largely W. African. 
Cercocebus. W. African. 
Theropithecus. N. E. African. 
Papio. Fossil in Indian Plistocene and Pliocene. 



Galago. Equatorial and E. African. 
Perodicticiis. W. African. 

II Insectivora. 

MACROSCELIDID&. Fossil in European Oligocene. 
Macroscelides (including Petrodromus}. One N. 

African species. 
Rhynchocyon. E. African. 


Proechinus. W. African. 


POTAMOGALID^:. Elsewhere only in Madagascar. 

Potamogale. W. African. 

Chrysochloris (including Chalcochloris). S. and 
E. African. 

III. Carnivora. 

Cynaelurus. Elsewhere only in India, the same species 
being common to the two regions. 


III. Carnivora (cont.}. 


Genetta. One species in South Holarctic region. 
Poiana. W. African. 
Nandinia. W. and E. African. 
Helogale. W. and E. African. 
Bdeogale. W. and E. African. 
Cynictis. S. African. 
Rhynchogale. E. African. 
. Crossarchus. 
Suricata. S. Africa. 

Proteles. S. and E. African. 

Lycaon. S. and E. African. Fossil in European 

Plistocene. f 

Otocyon. S. African. 


Mellivora. One African and one Indian species ; and 

another from the Indian Pliocene. 
Ictonyx. Ranges into Egypt, and perhaps Asia 


Pcecilogale. S. African. 
Galeriscus. E. African. 

IV. Rodentia. 


Anomalurus. W. and E. African. 
Idinrus. W. African. 


Nannosciurus. Elsewhere in Malaysia. 
Xerus. Fossil in European Miocene. 



Claviglis. W. African. The right of the one species 
to generic distinction is doubtful. 


IV. Rodentia (cont.\ 

Mystromys. S. African. 
Otomys. S. E. and W. African. 
Dasymys. S. African. 
Malacomys. W. African. 
Dendromys. \ 

Limacomys. These represent a peculiar Ethiopian 
Steatomys. sub-family the Dendromyina. 

Lophuromys. j 

Trilophomys. N. E. African, and (?) S. Arabian. 
Deomys. W. African. Alone represents a sub-family. 

Saccostomns. W. African 1 . 

Bathyergus. \ Constitute a sub-family the Ba- 
Georychus. thyergina ; the last of the four 

Myoscalops. being confined to Somaliland, 

Heterocephalus. ) while the first is S. African. 

Pedetes. The sole representative of a sub-family. 
OCTODONTID^:. Elsewhere at the present time only in 

the Neogaeic realm and N. Africa. 
Pectinator. N. E. African. 
Petromys. S. African. 
Triaulacodus. W., S. and E. African. 

V. Ungulata. 

n Eur P ean and Asiatic 


AK1. UACiYLA. Plistocene and Pliocene, and 

HIPPOPOTAMI DAI. 4 . ,, , -r. , 

also in Madagascar. Formerly 

Hippopotamus. in lower Egypt. 

Sus ; the Potamochczrine group, frequently regarded as 
a distinct genus, is peculiar to the Ethiopian and 
Malagasy regions. 

1 If Arvicanthis be accepted as a genus it should come here. 


V. Ungulata (cont.\ 

Dor cat her ium. W. African ; fossil in European Miocene, 

and also in the Pliocene of India. 

GIRAFFID/E. ( Fossil in lower Pliocene of Europe and 
GiraffaA Asia. 

Bubalis (including Damaliscus}. Ranges into Syria 

and Tunis ; fossil in Indian Pliocene. 

Madoqua. N. E. and E. African. 
Nanotragus. W. African. 
Oribia. S. African. 
Oreotragus. S. and E. African. 
Dorcatragus. Somaliland. 
Pelea. S. African. 
Cobus. Fossil in Indian Pliocene. 
Cervicapra. S. W. and E. African. 
sEpyceros. S. and W. African. 
Ammodorcas. Somaliland. 
Lithocranius. E. African. 
Hippotragus. Fossil in Indian Pliocene. 
Oryx. Ranges into Syria. 
Tragelaphus. Perhaps fossil in European Pliocene. 

Strepsiceros. ) 

-, . f Fossil in Indian Pliocene. 

Onas. I 


Rhinoceros ; the species without front teeth are now 
peculiar to the Ethiopian region, although allied 
forms occurred in the European and Asiatic 
Pliocene and Plistocene. 

Equus ; all the striped species confined to this region. 


V. Ungulata (cont.). 

PROCAVIID/E. I Range into Syria. 

VL Effbdientia. Ethiopian and Oriental. 

ORYCTEROPODIDJS. Fossil in French Oligocene. 

Orycteropus. Fossil in Pliocene of Samos and Persia. 

Among groups other than mammals, attention may be directed 
to the remarkable difference between the birds of 
Ethiopia and those of Madagascar. On this point 
Dr Blanford 1 writes that "the most characteristic African families, 
such as plantain-eaters (Musophagidce), colies (Collidce), and wood- 
hoopoes (Irrisoridce), barbets, hornbills, secretary-birds (Serpen- 
tarius], and a number of genera, such as Lamprotornis (glossy 
starlings), Buphaga (ox-peckers), Laniarius, and Telephonus, that 
are the common and familiar birds of every part of Africa south of 
the Sahara, are entirely wanting in the Mascarene Islands, in- 
cluding the Seychelles, Mauritius, etc., while no fewer than four 
peculiar families and a number of genera confined to the archi- 
pelago replace them. Amongst the Mascarene birds, too, are 
found several representatives of Oriental genera, or genera closely 
allied to Oriental types, and without any near Ethiopian relations. 
Foremost among these are certain bulbuls, forming the genera 
Ixocincla and Tylas, the former composed of species which have 
been usually referred to the typically Oriental genus Hypsipetes, 
and the latter nearly affined. In fact, as was shown by Geoffrey 
St Hilaire, and as Hartlaub has since pointed out, there is in the 
Mascarene avifauna a more marked connexion with Indian than 
with Ethiopian types. In the Seychelles, especially, out of the 
seven Passerine genera represented by peculiar species, three, 
Nectarinia, Zosterops, and Tchitrea, are Indian and African, one, 
Foudia, is Ethiopian, but not Indian, and two, Copsychns and 
Hypsipetes, or Ixocincla, are Indian but not African." 

All this is confirmatory, not only of the right of Madagascar 
and the Mascarenes to form a region by themselves, but likewise 

1 Appendix, No. 8, p. 89. In this quotation the English or Latin names 
have in some cases been added to the original. 


of -the distinction between the Ethiopian and Oriental regions, 
which some have proposed to unite. It is, however, somewhat 
remarkable that secretary-birds (Serpentarius) are unknown in 
Madagascar, seeing that they are represented in the upper Oligo- 
cene of France, and may therefore be presumed to have entered 
Ethiopia with the ancestral lemuroids and civets. Finally, the 
ostriches (Struthio), which are now mainly confined to Africa and 
Syria, are evidently recent immigrants into the region, the genus 
being represented in a fossil state in the Indian Pliocene. 

With the exception of the occurrence of remains of certain 
existing species, such as Rhinoceros simus, Phaco- 
charus, etc., in the superficial deposits of southern 
Africa, nothing is known of the mammalian Tertiary 
palaeontology of the Ethiopian region. Fortunately, however, the 
clue given by the existing fauna of Madagascar and the Tertiary 
faunas of Europe and southern Asia enables a considerable portion 
of the past history of the population of the region to be given with 
a fair degree of completeness. And here, with one important 
exception, Dr Wallace's explanation, as given in the Geographical 
Distribution of Animals 1 > may be accepted almost in its 
entirety; the one exception being, as mentioned in an earlier 
chapter, that the Sahara was never a sea during Tertiary times ; 
although it appears always to have formed a barrier between 
northern Africa and Ethiopia. As already mentioned, the an- 
cestral types of the existing mammalian fauna of Madagascar ob- 
tained an entrance into Ethiopia some time during the Oligocene 
period, and soon after ranged over the whole of what are now the 
Ethiopian and Malagasy regions, which were then united and 
possessed a common fauna. During the Pliocene age, when 
Madagascar had become isolated, came the great irruption into 
Ethiopia, of the higher and larger mammals, such as apes, monkeys, 
ungulates, etc., which were then flourishing all along southern 
Europe and Asia. Finding the country unoccupied and eminently 
suited to their existence, these rapidly attained a development now 
unequalled in any other part of the world ; many new genera being 
apparently evolved within the Ethiopian area, although a large 

1 Vol. i. p. 285 292. 


number were already in existence at the time of the southern 
migration. Several of these existing genera are met with in the 
Pikermi deposits of Greece, but more were confined, at this epoch, 
to the Pliocene of Persia, Samos, and India ; and it may there- 
fore be assumed that the great migration was by way of Syria or 
Arabia. Dr Wallace has indeed expressed the opinion that a 
certain number of types among them the elephants and rhinoce- 
roses obtained an entrance to the westward of Tunis ; but there 
are no true elephants in the Pikermi deposits, and apparently 
none in those of Persia, whereas their remains abound in the 
Siwalik Hills. As to the rhinoceroses, although the Pikermi 
species is closely allied to the African Rhinoceros bicornis, the 
Siwalik R. platyrhinus is equally close to R. simus ; and in the 
Siwaliks we meet with chimpanzees (Anthropopithecus), baboons 
(Papio), ratels (Mellivora), hippopotami, water-chevrotains (Dorca- 
therium\ and several genera of Ethiopian antelopes, all of which 
are totally unknown in the Pikermi beds. Ostriches, too, are 
first known in the Siwaliks ; while aard-varks occur in the Persian 
and Samos beds. All the evidence accordingly points to the 
great immigration having taken place along the eastern side of the 
continent ; and the existence of certain species of mammals which 
are either still common to India and Africa, or which were so 
during the Plistocene epoch, lends support to this view. Further 
testimony in this direction is aiforded by the occurrence of closely- 
allied generic types in the Ethiopian and Oriental regions. 
Among the lemuroids, for instance, the Oriental lorises (Nycti- 
cebus and Loris) are replaced in Western Africa by the potto and 
awantibo (Perodicticus) ; while in the Viverrida the true linsangs 
(Linsanga) of the eastern half of the Oriental region are repre- 
sented in Fernando Po by the allied Poiana, and the Oriental 
palm-civets (Paradoxurus} have very close allies in the two species 
of the Ethiopian genus Nandinia. A less marked instance is 
afforded by the occurrence of the water-chevrotain (Dorcatheriuni} 
in West Africa and of the true chevrotains in southern India and 
the eastern half of the Oriental region. 

And here it may be remarked that especial stress has been laid 
upon the much greater resemblance that exists between the fauna 
of the eastern, or Malayan, division of the Oriental region and 


Western Africa, than between that of peninsular India and 
Eastern and South Africa ; large man-like apes and linsangs being 
confined in the Oriental region to its eastern half, while palm- 
civets, lorises, and chevrotains are more abundant there than in 
other parts of the same region. This, however, appears to be 
mainly or entirely due to similarity of climatic conditions, and 
not to original distributional distinctions. And, it may be re- 
marked, increased acquaintance with the fauna of Ethiopia tends 
to show that types formerly thought to be confined to the western 
half of Africa really extend far to the eastward ; chimpanzees, for 
instance, being now known to range as far east as Ugogo, while 
the genus Nandinia, which was originally known solely by a West 
African species, is now proved to have an eastern representative in 

The similarity between the fauna of the Malayan sub-region 
and that of Western Africa naturally leads on to the consideration 
of the former land-connection between India and Africa. Writing 
on this subject, Dr Wallace 1 observes that "we may now perhaps 
see the reason of the singular absence from tropical Africa of deer 
and bears ; for these are both groups which live in fertile and well- 
wooded countries, whereas the line of immigration from Europe to 
Africa was probably always, as now, to a great extent a dry and 
desert tract, suited to antelopes and large felines, but almost 
impassable to deer and bears. We find, too, that whereas remains 
of antelopes and giraffes abound in the Miocene 2 deposits of 
Greece, there were no deer (which are perhaps a somewhat later 
development), neither were there any bears, but numerous forms 
of Felidce, Viverrida, Mustdida, and ancestral forms of Hyczna, 
exactly suited to be the progenitors of the most prevalent types of 
modern African zoology." 

As mentioned in an earlier chapter, since this passage was 
written the discovery of the remains of a species of chimpanzee 
in the Indian Siwaliks has shown quite clearly that the line 
of communication between India and Africa must have included 
a wooded tract comparable to the existing equatorial African 
forest region ; and this would be true even if the migration 

1 Op. dt. p. 291. 

2 The Pikermi beds were formerly universally held to be of Miocene age. 

L. I 


had taken place from Africa to India, which was not the 
case. Evidence of such a tract is, I believe, afforded by the 
occurrence of fossilised tree-stems in many districts which are now 
desert. And along this tract there can be little doubt that the 
ancestors of the mammalian types now common to the West 
African and Malayan sub-regions originally wandered from their 
common Indian home. Subsequently the whole of the countries 
lying between eastern Africa and India have become deforested, 
while in Africa itself the forest-area has shrunk away from the 
eastern side of the continent. 

This leaves the question of the absence of bears and deer from 
Africa without any adequate explanation. Bears are, however, in 
the main, mountain animals, some of which, like the isabelline 
bear of the Himalaya, inhabit districts where there is but little 
forest ; and it is noteworthy that, with the exception of the sloth- 
bear, which forms a genus apart (Melursus)^ there are no bears in 
India proper, although a fossil species allied to the sloth-bear 
occurs in the Siwaliks. This being so, when we take into account 
the absence of ursine remains from the Pikermi and Persian beds, 
there is nothing very wonderful in the fact that none of these 
animals entered Ethiopia during the great Pliocene migration. 
The absence of all Cervidce is more difficult to explain, seeing 
that deer of an Oriental type are abundant in the Siwaliks, while 
they are also sparingly represented in the Pikerrni beds. Typical 
deer of the red-deer group are, however, totally wanting in the 
Siwaliks, as they are in the Oriental region at the present day ; 
and we are, therefore, perfectly able to account for their absence 
from the Ethiopian region, although they occur in Africa north of 
the Sahara. With regard to the absence of Oriental types of deer 
in Ethiopia, it can only be said that it is as difficult to see any 
reason why these should have continued to flourish since the 
Pliocene in the Oriental region without ever having entered Africa, 
as it is to explain why giraffes, hippopotami, and ostriches should 
have disappeared from the former area to survive in the latter. 

Another difficulty is presented by the case of the pigs, bat here 
it may be suggested that the absence of the typical group of the 
genus Sus from the whole of Ethiopia, with the exception of 
Sennaar, may perhaps be accounted for by all the other species of 


that genus which originally entered the country having been 
developed into the more specialised Phacochxrus. Attention has 
already been called to the fact that the molars of some of the 
Indian Tertiary species of Sus show a distinct approximation to 
those of Phacochairus, and further evolution might easily lead to 
the development of the latter. Since this genus is unknown in a 
fossil state from other regions, is it improbable that it has originated 
from Sus within the limits of its present habitat ? 

Summarising the results of the foregoing survey of the mam- 
malian fauna of Ethiopia, it would appear that the Sahara has for 
a very long period formed a barrier between Ethiopian Africa and 
the northern part of the continent. When first populated by 
Tertiary mammals, Ethiopia and Madagascar were in union, and 
formed but a single zoological province, which would seem to 
have been to a great extent isolated from the rest of the Old 
World during the Miocene epoch. Had such conditions persisted 
|his province would have been entitled to form a primary 
zoological realm by itself. During the Pliocene epoch, however, 
Madagascar became separated, while a more complete union of 
the continent with Asia by way of Syria or Arabia permitted the 
influx of larger and higher mammals from the eastward. Hence 
there is a most intimate relationship between the Pliocene fauna 
of India and the one now inhabiting Ethiopia ; but the distinc- 
tion between the two areas at the present day is fully sufficient 
to justify the separation of the Ethiopian from the Oriental region. 
Of all the zoological regions of the world, the Ethiopian may be 
regarded as the one which has been most recently evolved ; and 
it may be shortly characterised by the sole possession of the 
gorilla and chimpanzees ; the absence of bears and deer ; and the 
presence of the African elephant, hyraces, rhinoceroses devoid of 
front teeth, zebras, hippopotami, wart-hogs, giraffes, numerous 
genera of antelopes, and aard-varks, as well as by the great de- 
velopment of its large ungulates in general. It shares with the 
Oriental region the distinction of being the sole habitat at the 
present day of man-like apes, true civets ( Viverra), linsangs, palm- 
civets 1 , ratels (Mellivora), elephants, rhinoceroses, and pangolins 

1 Also in the Austro-Malayan region. 



(Mam's), together with the rodent genera Nannosciurus, Golunda 
and Atherura. The probable relationship between Ethiopia and 
Neogaea has been sufficiently discussed in the third chapter. 

Although, from the mammalian point of view, Ethiopia is a 
very modern region, as a continent it is one of the oldest, the 
greater portion of its area having been land since the Palaeozoic 
epoch. As has been shown in an earlier chapter, in Palaeozoic 
times southern Africa formed a portion of the great southern or 
equatorial continent distinguished from more northern lands by 
the peculiar characteristics of its flora; and it is probable that 
it remained connected with India until late in the Secondary 
epoch 1 ; as is proved by the identity of the flora and reptiles of the 
two areas. Early, however, in this epoch there must also have 
been free communication with Europe, as shewn by the close 
alliance of the anomodont reptiles of the two continents, and 
likewise by the occurrence in both of the mammalian genus 
Tritylodon. In the Cretaceous, so far as vertebrates are conf 
cerned, our knowledge of the Ethiopian region is practically a 

In conformity with the plan adopted in other cases, the sub- 
regions into which Ethiopian Africa may be divided 

Sub-regions. . . 

will be treated very briefly. 

Writing of the desert-tracts of the Saharan sub-region, where 
the necessary conditions for existence are largely wanting, Dr 
Heilprin observes that " there is a marked impoverishment of the 
fauna. The more formidable carnivores, such as the lion and the 
leopard, are absent from most districts, leaving their places to be 
filled by some minor cats, the hyaena, jackal, fox, and fennec. 
The hoofed animals are represented (in some parts) by the buffalo, 
and a limited number of antelopes (Gazella, Oryx, and Addax). 
Among rodents the families of rats (Muridce) and jumping-mice 
(Dipodidcd) are fairly represented, in addition to which we have 
the porcupine and hare (Lepus mediterraneus}. The ostrich is 
sufficiently abundant throughout most of the sub-region." 

In marked contrast to the poverty of the Saharan districts, 
is the remarkable richness of the great equatorial forest-tract, 

1 Vide siipra, p. 224. 


which is the exclusive home of all the man-like apes and of the 
lemurs of the genus Perodicticus. To this sub-region also belong 
PotamogaUi the African linsang (Poiana), as well as several other 
genera of Viverridcz, such as Nandinia and Helogale. Among the 
rodents the flying-squirrels of the family Anomalurida are very 
characteristic of the forest tract, and the peculiar murine genus 
Deomys, as also Saccostomus, is restricted to it. The water-chevro- 
tain (Dorcatheriuni) is solely West African, as is the small Liberian 
hippopotamus ; while certain genera of antelopes, such as the 
duikers (Cephalophus), harnessed antelopes (Tragelaphns\ and 
elands (Orias), have larger and finer representatives here than 
elsewhere. The giant pangolin (Manis gigantea), the largest 
member of its genus, is likewise a West African form. 

Although South Africa has a certain number of mammalian 
genera, such as Cynictis, Suricata, Otocyon, Pcecilogale, Bathyergus, 
Pedetes, Petromys, and Pelea, peculiar to it, others, such as the 
golden moles (Chrysochloris) and aard-wolf (Proteles], have been 
proved to extend far up the east coast, the latter occurring in 
Somaliland. Hence it seems advisable to unite Dr Wallace's 
South African sub-region with that portion of his East Central 
sub-region which is not included in the equatorial forest-tract. Of 
this East Central sub-region, as it may be collectively called, it 
will suffice to say that it is the home of the greater number of the 
characteristic Ethiopian mammals exclusive of those restricted to 
the forest-tract. Here antelopes attain their greatest numerical 
development, both as regards genera and species; and here is 
also the true home of the lion and the spotted hyaena ; while to 
this sub-region are confined the Cape hunting-dog (Lycaon) and 
the aard-wolf (Proteles). The distribution of other genera is 
sufficiently indicated in the table already given. 

The light which has recently been thrown upon the mammals 
of northern Somaliland has shown that, as regards antelopes at 
least, these are so peculiar that it may be questioned whether this 
tract is not entitled to be separated as a sub-region by itself. 
According to the lists given by Dr Sclater 1 and Capt. Swayne 2 , 

1 Natural Science, vol. I. p. 264 (1892). 

2 Seventeen Trips to Somaliland, London (1895). 




there are no less than sixteen 
northern Somaliland, of which the 

1. Bubalis swaynei. 

2. Madoqua swaynei. 

3. phillipsi. 

4. ,, guentheri. 

5. Oreotragus saltator. 

6. Dorcatragus megalotis. 

7. Cobus ellipsiprymnus. 

8. Gazella pelzelni. 

species of antelopes found in 
names are as follows, viz. : 

9. Gazella spekei. 

10. ,, soemmerringi. 

11. Ammodorcas clarkei. 

12. Lithocranius walleri. 

13. Oryx beisa. 

14. Tragelaphus decula. 

15. Strepsiceros kudu. 

1 6. imbubis. 

Among these Nos. i, 2, 3, 4, 6, 9, n are quite peculiar to this 
district, while No. 12, which, like Nos. 6 and n, is the sole repre- 
sentative of its genus, is only found elsewhere along the east coast 
as far south as the Tana river. Another generic form peculiar to 
Somaliland is Heterocephalus, including two small naked rodents 
with burrowing habits ; while in the same order the single species 
of Pectinator is restricted to this district. Among species may be 
mentioned two musk-shrews (Crocidura smithi and C. somalica), a 
hedgehog (Erinaceus sclateri), as well as a banded mungoose 
(Crossarchus somalicus). The Somali ostrich seems likewise to 
represent a species by itself. On the other hand, in addition to 
those mentioned in the foregoing list of antelopes, there are 
several East or South African mammals, such as the aard-wolf, 
which range into Somaliland, and further evidence is perhaps 
desirable before the right of that country to form a separate sub- 
region can be admitted. 

The case is more clear with regard to south-eastern Arabia, 
whence Mr O. Thomas 1 records the following fifteen species of 
land-mammals, viz. : Xantharpyia amplexicaitdata, Taphozous nudi- 
ventriS) Rhinopoma microphyllum, Erinaceus niger, Crocidura 
murina, Herpestes albicauda, Cants pallipes, C. leucopus, Gerbillus 
dasyurus, Mus rattus, Lepus omanensis, Gazella muscatensis, Oryx 
beatrix, Hemitragus jayakari, and Procavia syriaca. Of these Mr 
Thomas remarks that their geographical relationships " are, as 
might be expected, about equally with Africa and India, three of 

1 Proc. Zool. Soc. 1894, p. 449. In one case the generic title has been 
altered, in order to bring it into harmony with the system followed in this work. 


the species being distinctly African in affinities, three Indian, and 
the remainder either peculiar or widely-spread, and of no special 
significance." The association of a goat belonging to the Oriental 
genus Hemitragus with such an essentially Ethiopian animal as a 
hyrax {Procavia}, shows that we are here truly on the border-land 
between the two regions in question. 

In conclusion, from whatever aspect it be regarded, Ethiopia 
is one of the most interesting of the regions of Arctogaea; and if, 
as is suggested in an earlier chapter, it has been the feeder by 
means of which South America received its earliest mammalian 
Tertiary fauna, it is entitled to an importance above all the other 
zoological regions of the realm to which it pertains. 



Sub-regions Characteristics of the Mammalian Fauna Past History of the 
Region Malayan sub-region Nicobars, Mentawi, and Christmas Islands 
Philippine sub-region. 

FAR inferior in extent to the Ethiopian, the Oriental region is 
taken to include those portions of the continent of Asia lying 
south of the Holarctic region (with the exception of southern 
Arabia, which is Ethiopian), together with the islands of Ceylon, 
Formosa, the Philippines, Sumatra, Java, Borneo, and numerous 
smaller ones. In India the northern limits of the region are 
formed by the higher ranges of the Himalaya, while " Wallace's 
line" constitutes the eastern boundary denning it from the Austro- 
Malayan region of the Notogaeic realm. In a region so diversified 
as the Oriental, it would not be natural to expect a homogeneous 
fauna; and, as a matter of fact, there are in this respect great 
diversities between the different portions of the region, many of 
the peculiar genera having a very restricted distribution. Never- 
theless, the positive and negative features of the mammalian fauna 
of the region as a whole are sufficient to indicate its zoological 
unity, and also to differentiate it from the Ethiopian region, to 
which it is now most closely allied. In the Himalaya there is a 
gradual transition towards the Holarctic fauna ; and it is probable 
that in this portion of the area the differentiation between the 
Oriental and Holarctic faunas has been largely due to the eleva- 
tion of the Himalaya itself, which has taken place entirely since 
the early part of the Tertiary period, and is to a considerable 
extent of Post-tertiary date. It has already been shown that the 
older Pliocene fauna of northern India and Burma contained a 


remarkable admixture of mammalian genera now respectively 
confined to the Ethiopian and Oriental regions, together with 
some of a Holarctic facies ; and the completion of the elevation 
of the Himalayan chain was probably an important factor in the 
dispersal and differentiation of this common fauna. Holarctic 
types are again met with in force in the open desert regions on 
the north-western frontier of India. On the other hand, the 
fauna of the Philippines exhibits an approximation to that of the 
Austro-Malayan region, and thus shows a blending between the 
Arctogaeic and Notogaeic realms. In physical features the Oriental 
region displays great variation, a large portion of peninsular India 
consisting of open dry grassy plains, whereas the slopes of the 
eastern Himalaya, together with the greater part of Assam, Burma, 
and the Malayan countries are clad with luxuriant forests ; these 
tropical or sub-tropical forest-regions being those where the fauna 
attains its fullest and richest development. 

The poorest part of the region is, as Dr Wallace 1 observes, the 
great triangular plateau forming the Indian peninsula; this area 
differing remarkably from the Himalaya in its geological features, 
and having been land since an extremely ancient date, whereas 
the Himalayan area consists very largely of marine formations. 
Since it is stated in the passage cited that peninsular India 
during the Tertiary period existed as an island entirely discon- 
nected from the Himalaya and Burma, it may be well to quote 
the later and more authentic views of the authors of the Manual 
of the Geology of India 2 on this subject. After reference to the 
extent of Eocene rocks in northern India, it is there stated that 
" the Peninsula of India in Eocene times was part of a tract 
of land, perhaps of a great continent united to Africa ; that there 
was a sea to the eastward, extending far to the north-east, in the 
region now occupied by the Assam hills, and another sea to the 
north-west, covering great part, if not the whole, of Persia, Baluch- 
istan, the Indus plain, and a portion of the upper Ganges plain. 
An arm of this sea extended from the north-west up the Indus 
valley in Ladak. The Himalaya, and perhaps Tibet, wholly or 
in part, were raised above the sea ; but formed in all probability 

1 Geographical Distribution of Animals, vol. I. p. 314. 

2 First edition, pt. I. p. liii. 


land of moderate elevation. Whether the Himalayan land was 
united to the Peninsula is, of course, uncertain but very pro- 
bably it was ; for there is no evidence of marine conditions having 
existed in the Ganges plain to the east of the Dehra Dun ; and if 
the ferruginous bands of the Subathu group be laterite, as they 
appear to be, the trappean detritus composing them must have 
been derived, in all probability, from the peninsular area; and the 
latter must consequently have extended northward to the base of 
the Himalayas, in the neighbourhood of Umballa.... In Miocene 
times, although marine conditions prevailed throughout western 
Sind, the area of the sea was very much smaller than in the Eocene 
period ; for all the marine beds of the Punjab and Sub-Himalayas 
are destitute of marine fossils, and are probably fluviatile de- 

From this it would appear probable that during the whole of 
later Tertiary times, at least, there has been no isolation of 
peninsular India from the eastern Himalaya and the Burmese 
countries ; and consequently that the differences between the 
faunas of these areas are mainly or solely due to their differences 
of physical features and climate. 

By Dr Wallace the Oriental region is divided into four sub- 
reerions ; namely (i) the Indian, comprising the whole 

Sub-regions. J ^ ' 

of upper India, (2) the Ceylonese, including southern 
India and Ceylon, (3) the Indo-Chinese, embracing Assam, Burma, 
and such portion of China as lies within the limits of the region, 
and (4) the Indo-Malayan, which includes not only the Malayan 
archipelago and islands, but likewise the Philippines. 

So far as India and its dependencies are concerned, the follow- 
ing amended scheme has been proposed by Dr Blanford l , viz. : 
i. Himalayan. The southern slopes of the Himalaya, from the 

base to about the limit of trees. 

ii. Indian. India from the base of the Himalaya to Cape 
Comorin, with the exception of the Malabar coast, but with 
the addition of northern Ceylon. 

iii. Malabar or Ceylonese. The Malabar coast and the neigh- 
bouring hills as far north as the Tapti river, together with 
southern Ceylon. 

1 Fatina of British India Mammalia, p. v. 


iv. Burmese. All Burma except south Tenasserim, and with the 

addition of Assam and the intervening countries, 
v. Malayan. South Tenasserim, the Malay Peninsula, and the 

Malayan Islands as far as Wallace's line. 
Whether the Philippine Islands should be included in this 

sub-region, or should form one by themselves, may be 

vi. Indo-Chinese. Although not free from objection, this term 

may be employed for the sub-region indicated by that 

portion of China coming within the limits of the Oriental 


Regarding these sub-regions in general, Dr Blanford observes 
that some " may require further subdivision. Thus the fauna of 
the North-west Provinces and Punjab differs considerably from 
that of southern India, and both areas exhibit zoological dis- 
tinctions from the forest-clad tracts of south-western Bengal. 
There is also much difference between the animals of Pegu and 
Arakan, on the one hand, and those of the drier regions of upper 
Burma on the other ; and even greater distinctions may be traced 
between those found in the sub-tropical and those inhabiting the 
temperate regions of the Himalaya. On the other hand, the sub- 
tropical Himalayas were united with the Burmese sub-region by 
Wallace, and the two are, perhaps, zoologically more allied to 
each other than to any other sub-region." 

Recent discoveries clearly indicate that the Philippine Islands, 
exclusive of Palawan and the Calamianes, should form a sub-region 
by themselves. 

Taking the Oriental region as a whole, it may be stated that 
the number of peculiar generic types of mammals is 
less than in the case of the Ethiopian; and that 
there are but two families absolutely confined to it, 
although a third is very nearly so. While sharing 
with the Ethiopian region the want of several groups of insectivores 
and rodents, such as the typical shrews (Sorex), marmots (Arcto- 
mys), and voles (Microtus), it lacks some of the other deficiencies 
of that region, true pigs (Sus) and deer being abundant, although 
the latter belong to groups distinct from those of the Holarctic 
region, while bears, belonging to two genera, are likewise met 


with. Wart-hogs (Phacochcerus), aard-varks (Orycteropus\ hyraces 
(Procaviid(B\ and jumping-shrews (Macroscelididce) are among 
some of the more characteristic Ethiopian animals which are 
wanting in the Oriental region at the present day, and have not 
hitherto been obtained there in a fossil state. Giraffes, a number 
of genera of antelopes, and hippopotami, are equally conspicuous 
by their absence, although this, as we have seen, is but a com- 
paratively recent feature of the region, since most of these forms 
are represented in the Pliocene of India and Burma. A notable 
feature of the Oriental as distinct from the Ethiopian region is the 
circumstance that the great majority of its fruit-bats (like those of 
Madagascar) belong to the typical genus Pteropus, which is 
wanting in Ethiopia, while the three genera of the family found in 
the latter area are absent from the present region. Indeed there 
is a very curious dissimilarity between the flying-mammals of the 
two areas, Ethiopia possessing the flying-squirrels of the family 
Anomaluridce, while the Oriental flying-squirrels all belong to the 
SriuridiE, the genus Pteromys being peculiar to the region. Among 
the Insectivora, the so-called flying -lemur (Galeopithecus) is a 
peculiar Oriental type which has no Ethiopian representative. A 
somewhat similar instance to that of the flying-squirrels is afforded 
by the tree-shrews (Tupaiid&) of the Oriental region, which are 
represented in Ethiopia by the jumping-shrews (Macroscelidida). 
From the Holarctic region, the Oriental is distinctly differentiated 
by the presence of apes and lemurs and the abundance of 
monkeys, together with the presence of the groups mentioned 
above as being now restricted to this and the Ethiopian region, 
and likewise by the absence of the typical elaphine deer, 
marmots, susliks, voles, etc., and the scarcity of sheep and true 
goats, which, indeed, enter the region only in the north-western 
frontier of India. 

Commencing with the man-like apes of the family Simiidce, we 
find the Oriental region destitute of chimpanzees and gorillas, 
whose place is taken by the orangs (Simla] of Borneo and 
Sumatra, characterised by the reddish, instead of blackish, colora- 
tion of their hair, and their more wide departure from the human 
type. Orangs appear, however, to have inhabited northern India 
during the Pliocene, and as chimpanzees were then also in exist- 




ence there, it would seem that the region must be regarded as the 
original home of the larger man-like apes. The smaller long- 
armed apes known as gibbons (Hylobates) are likewise characteristic 
of the Oriental region, where they range from Assam through the 
Burmese and Malayan countries to Hainan and Cambodia. In 
the upper Miocene these apes occurred in Central Europe, but 
perhaps on account of their small size their remains have not 
hitherto been obtained from the Indian Pliocene. Among the 
Cercopithecidcz the Oriental genera of monkeys are now entirely 
distinct from those of Ethiopia, although, as we have seen, the 

FIG. 60. SLOW LORIS (Nycticebus tardigradus). 

African genus Papio occurs in the Indian Pliocene and in Madras 
survived till the Plistocene. And here it may be noticed that this 
is the only one of the Ethiopian genera of monkeys that is found 
in Arabia. Of the other genera, Macacus is mainly Oriental, 
although with representatives in northern Africa, Kashmir, Tibet, 
and Japan ; while the langurs (Semnopithecus] are likewise almost 
wholly confined to this region, although they range into Kashmir 
and eastern Tibet. As both these genera are found in the 
European and Indian Pliocene, they are evidently ancient types 
which were formerly widely spread in the eastern half of the Old 


World. Each represents a sub-family by itself; and as both these 
sub-families have Ethiopian genera, which are unknown in a fossil 
state, it may be suggested that the latter (like the wart-hogs) have 
been evolved within the limits of the Ethiopian region from the 
Oriental types. To the same sub-family as the langurs belongs the 
singular proboscis monkey (Nasalis) of Borneo. Among the 
lemuroid Primates there are Oriental representatives of two 
families. Of these the Lemurida include the lorises of the genera 
Nycticebus and Loris, the former ranging over the Burmese, 
Malayan and Indo-Chinese sub-regions, while the latter is con- 
fined to southern India and Ceylon. Although these animals are 
nearly allied to the pottos (Perodicticus) of western Africa, nothing 
is known as to their past history. The tarsiers (Tarsiidce\ of 
which there is but a single genus (Tarsius\ although several 
specific forms have been recognised, are almost confined to the 
Malayan sub-region, but are represented in Celebes, as they are in 
the Philippines. 

Of the Insectivora, the most aberrant and remarkable forms are 
the flying-lemurs (Galeopithecus), constituting a sub-order by them- 
selves, and ranging from south Tenasserim through the Malay 
peninsula and islands to the Philippines. As with the tarsiers, we 
have no palaeontological history of these creatures, which are 
probably comparatively modern types. The most characteristic 
Oriental family of this group is that of the tree-shrews (Tupaiidce], 
whose members have the form and habits of squirrels, with the 
structure of shrews. The typical genus Tupaia ' ranges from India 
throughout the Burmese and Malayan regions, but is unknown in 
Ceylon ; while the single representative of the pen-tailed tree- 
shrews (Ptilocercus], characterised by the pen-like extremity of the 
exceedingly long tail, is confined to Borneo and some of the adja- 
cent islands. As mentioned in an earlier chapter, the European 
Miocene genera Lanthanotherium and Galerix appear to be an- 
cestral types of this family ; and this distribution of the family is a 
well-marked instance of the curious affinity existing between 
certain mammalian genera of the middle Tertiaries of Europe and 

1 Two species (T. murina and 7". frenata) are often separated as Dendro- 
gale, but as there is an annectent form (see Thomas, Proc. Zool. Soc. 1892, p. 
-225), this appears unnecessary. 




those of the Malayan sub-region, the absence of Tupaia from 
Ceylon probably indicating that the genus is essentially a Malayan 
one which has immigrated but recently into India. The hedgehog 
family (JErwaceidce), which, as already shown is an ancient one, 
has a very remarkable distribution in the Oriental region ; true 
hedgehogs (Erinaceus] ranging into India, but apparently not oc- 
curring in Ceylon, and being unknown to the west of the Bay of 
Bengal. In the latter districts their place is taken by the spineless 
and more rat-like animals forming the genus Gymnura and the 

FIG. 61. TREE-SHREW {Tupaia tana). 

allied Hylomys ; and here, again, we have to note the occurrence 
of an allied type in the European Oligocene, which has been 
described under the name of Necrogymnurus. In passing, it may 
be remarked that the survival of these early Tertiary insectivorous 
types in the Malayan sub-region serves to lend support to the 
suggestion made in a previous chapter 1 that opossums may also 
have survived in the same area long after they had ceased to exist 
in western Europe. 

Passing over the moles (Talpida), of which the typical genus 
Talpa only just impinges on the region in the frontiers of India> 

1 Supra, pp. 51, 57- 


we come to the Soricidce or shrews. Here the typical shrews 
(Sorex) are wanting, while the section of the family to which that 
genus belongs (characterised by the reddish-brown tips to the 
teeth) is represented only by the genus Soriculus, ranging from the 
southern slopes of the Himalaya to China. Of the widely-spread 
musk-shrews (Croddura) it is unnecessary to speak; but it may 
be mentioned that of the two almost tailless and scaly-footed 
species forming the genus Annrosorex one is from Assam and the 
other from eastern Tibet and Pekin. Chimarrogale includes two 
aquatic shrews, one of which is found in the eastern Himalaya, 
the hills north of Burma, and M 4 Kina Balu in Borneo, while 
the second is Japanese. 

Among the Carnivora the region is especially rich in Felidce, 
containing more species than any other part of the world. The 
tiger (Felis tigris) is usually regarded as one of the most character- 
istic mammals of India, but as its range extends northwards to 
Siberia, while its fossilised remains have been found within the 
Arctic circle, and it is unknown in Ceylon, there is a great proba- 
bility that this feline is a comparatively recent immigrant into 
India from the north-east. The range of the lion (F. leo) in this 
region is limited to India, not extending to the eastward of the 
Bay of Bengal, and as this animal was widely distributed during 
the Plistocene in Europe, while it ranges all over Africa, it may 
be regarded as essentially a western type, or exactly the opposite 
of the tiger. Possibly certain remains from the Indian Plistocene 
which have been assigned to the latter animal may really belong to 
the former. As noticed on p. 234, there are other species of 
Felis, as well as the hunting-leopard (CynteJurus), which are com- 
mon to India and Africa, some of these occurring in the European 
Plistocene, while only the jungle-cat (F. chaus) is found to the 
eastward of the Bay of Bengal, and there not further east than 
Burma. On the other hand, there are certain species, like the 
clouded leopard (F. nebulosa) and the marbled cat (F. marmorata), 
which are essentially eastern forms, their range including the 
Malayan sub-region and India, but not Ceylon. The rusty- 
spotted cat (F. rubiginosa) and the Indian desert-cat (F. ornatd] 
are species whose range is limited in one case to India and 
Ceylon, and in the other to India alone. 


In the civets and their allies ( Viverridtz] the Oriental region 
approaches the Ethiopian in richness, and thereby stands in 
marked contrast to the Holarctic, which contains only a single 
species of Genetta and another of Herpestes in southern Europe, 
although the latter genus ranges into Kashmir. Of the true civets 
( Viverrd] the whole of the species, with the exception of one from 
the Ethiopian region, are Oriental, and some are confined to the 
countries to the east of the Bay of Bengal; one small species being 
separated by many zoologists as Viverricula. The beautifully- 
coloured linsangs (Linsangd) are exclusively Oriental, and are con- 
fined to the eastern Himalayan and Malayan sub-regions, although 
represented in West Africa by the nearly allied Poiana. Equally 
characteristic of the region are the two species of Hemigale, which 
are, however, exclusively Malayan, H. hosei being limited to the 
mountains of North Borneo. The palm-civets of the genus Para- 
doxurus range throughout the region, and have also representatives 
in Celebes : their place being taken in the Ethiopian region by 
the allied genus Nandinia. On the other hand, the two species 
of small-toothed palm-civets constituting the genus Arctogale are 
restricted to the Burmese and Malayan sub-regions ; the same 
being also the case with the binturong, which is the sole repre- 
sentative of the genus Arctitis, distinguished by its pencilled ears 
and prehensile tail. Still more circumscribed is the range of the 
peculiar genus Cynogale, of which the single species is confined to 
the Malayan sub-region. All the foregoing forms belong to the 
sub-family Viverrina \ the Herpestincz, which are so numerous in 
the Ethiopian region, being represented only by species of the 
large and widely-spread genus Herpestes. Both this genus and 
Viverra date from the European Oligocene, the latter also 
occurring in the Pliocene of France and India ; but none of the 
others are known in a fossil state. It is, however, probable that 
most of the other genera are comparatively modern derivatives 
from the original stock ; and the high development in the Malayan 
sub-region of a group first known from the Oligocene of Europe 
is another instance of the relationship of the faunas of these 

Although the striped hyaena (Hycena striata) is by no means 
confined to India, its range extending through south-western Asia 
L. 18 


to northern Africa, it is unknown in Ceylon or in the countries on 
the eastern side of the Bay of Bengal, which forms, indeed, the 
present limits of the range of the genus in this direction. As 
remains of the existing African spotted hyaena (H. crocutd) have 
been met with in a cave in Madras, while they are common in the 
Plistocene of southern and central Europe, it is manifest that both 
these animals are as essentially western types as is the lion. And 
it is a curious circumstance that nearly all these western types of 
mammals ranging into India (of which a list is given in the sequel) 
belong to genera which date only from the Miocene or Pliocene, 
whereas very many of the Malayan or eastern types date from the 
Oligocene. During the Pliocene a single species of hyaena ranged 
as far eastwards as China, and species were exceedingly abundant 
in India at the same epoch. 

As regards its Canidcz, the Oriental region is inferior to 
the Ethiopian in lacking any peculiar generic type, although it 
possesses a true wolf (Cants pallipes), and three species of wild 
dog (C. rutilans, etc.), the latter, on account of the absence of the 
last tooth in the lower jaw and other differences, being frequently 
referred to a distinct genus, under the title of Cyan. Whereas, 
however, the Indian wolf, which ranges into southern Arabia, is 
unknown either in Ceylon or in the countries to the east of the 
Bay of Bengal, the wild dogs are found throughout the region, and 
have also a representative beyond it in the mountains of Central 
Asia, and they are likewise known by fossil species from the 
European Plistocene. The wolf, which is very closely allied to 
the European species, may be the descendant of a fossil form 
found in the Siwaliks, but its absence from Ceylon would seem to 
indicate that it has only reached southern India at a comparatively 
modern date. No foxes are known to the east of the Bay of 
Bengal, and the jackal does not range east of Burma. 

The Oriental region is the home of three well-marked species 
of bears, and thereby presents a decided contrast to the Ethiopian. 
Of these the Himalayan black bear (Ursus torquatus) ranges from 
the forest districts of the Himalaya to Burma, and thence to the 
Indo-Chinese sub-region. The small Malayan bear (U. malay- 
anus] is restricted to the Burmese and Malayan sub-regions ; and 
the great Indian sloth bear (Melursus ursinus], which is the sole 




representative of a separate genus, is confined to India and 
Ceylon, and is known to have been an inhabitant of Madras since 
the Plistocene era. Not improbably it may be the descendant of 
a Siwalik species ( U. theobaldi), which is the earliest known repre- 
sentative of the true bears; and the Malayan species may be 

FIG. 62. INDIAN SLOTH-BEAR (Melursus ursinus). 

derived from a small extinct bear whose remains occur in the 
Plistocene of the Narbada valley. 

One of the most remarkable of Oriental carnivores is the 
panda, or cat-bear (sElurus fulgens), which ranges from the East- 
ern Himalaya to Yunnan, and is the single existing Old World 
representative of the Procyonida. Curiously enough, the remains 
of a much larger species of the same genus have been discovered 
in the English Pliocene ; and it is thus evident that sElurus 
formerly enjoyed a wide range. From the restriction of all the 
other known members of the family to the New World, fossil 
types may be looked for in eastern Asia, as it is quite clear that 
the distributional area of the group must once have been con- 

In the Mustelidce there are four generic types very character- 

18 2 

2 7 6 



istic of the region, although two of them are not confined to it. 
The first of these comprises the three species of sand-badger 
(Arctonyx), two of which are found in the Himalayan and Burmese 
sub-regions, while the third is Tibetan. The single Oriental 
species of ratel (Mellivora) is restricted to India, exclusive of 
Ceylon ; a fossil species occurring in the Siwaliks. The only other 
living form is Ethiopian. Its distribution would thus seem to 
indicate that the genus originated in northern India, whence it 

FIG. 63. INDIAN RATEL (Mellivora ratel). 

migrated into Africa while there was a free communication between 
the two continents, and that it only reached southern India (where 
it is unknown on the Malabar coast) at a comparatively recent 
epoch. The third genus, Helictis, comprising badger-like animals 
with long bushy tails, is represented by four species, ranging from 
India to China, but unknown in Ceylon. Lastly, the teledu, or 
small burrowing badger (Mydaus meliceps) of the Malayan sub- 
region, is the sole representative of its genus, and is found at con- 


siderable elevations in Java, as well as in Sumatra and Borneo. 
No fossil representatives of either of these two genera are at 
present known. 

Among the rodents, the grooved-toothed squirrel (Rhithro- 
scturus) is a peculiar type confined to the island of Borneo; 
and the pigmy squirrels (Nannosciurus) are represented by four 
Malayan species, the only other member of the genus being West 
African. The true squirrels (Sciurus), as mentioned above, attain 
their maximum development in the Malayan sub-region. The 
flying - squirrels are represented by the genera Pteromys and 
Sciuropterus, the former being exclusively Oriental, and the latter 
having one species in the eastern, and a second in the western 
half of the Holarctic region, in addition to being represented in a 
fossil state in the French Miocene. In the Muridcz there are no 
less than eleven genera in most cases respectively represented 
by only a single species peculiar to this region, while another is 
Oriental and Ethiopian only. Of the peculiar types, one of the 
most remarkable is Chrotomys, from the mountains of Luzon, in 
the Philippines, belonging to the sub-family Hydromyince, of which 
the typical forms are Australian 1 . From other members of the 
sub-family the single species of this genus differs in having three 
(in place of only two) pairs of molar teeth, thereby forming a 
link with ordinary murines. This animal, which is about the 
size of a rat, is easily recognised by the presence of an orange or 
buff line running down the middle of the back. Luzon has also 
yielded another rat, provisionally referred to the Australian genus 
Xeromys*. Another unique Oriental type is found in the long- 
tailed dormouse-like form from the Malabar coast known as 
Platacanthomys, which constitutes a sub-family by itself. Phlao- 
mys, likewise representing a separate group of the same rank, is 
restricted to the Philippines, and is characterised by the molar 
teeth being divided into three transverse lobes. The one species 
is of very large size. Nearly allied is a huge, rough-haired, grey 
or blackish rat, from the mountains of Luzon, which may be 
compared in size to a small marmot, and for which the name 
Crateromys has been suggested. This differs from Phlceomys by 

1 Vide suprh, p. 40. 

2 Appendix, No. 31 


the smaller claws and more bushy tail, and also by the completely 
tuberculate molars. The single species of the Burmese rat-like 
Hapalomys differs from all other members of the sub-family 
Murincz in possessing three longitudinal rows of tubercles on 
the lower as well as on the upper molar teeth. The one repre- 
sentative of the allied genus Vandeleuria ranges from India and 
Ceylon to Yunnan. The pencil-tailed tree-mice (Chiropodomys), 
of which there are three species, are restricted to the Burmese and 
Malayan sub-regions; and the small red rat representing the genus 
Pithechirus is known only from Sumatra and Java. With the 
shrew-rat (Rhynchomys] we revert to several peculiar forms from 
the mountains of Luzon, in the Philippines. This rodent, which 
is about the size of an ordinary rat, has the muzzle extraordinarily 
slender and elongated, with very feeble incisors, and it is pro- 
bable that it lives on animal substances, possibly caterpillars. 
The two other Philippine types form the genera Carpomys and 
Batomys ; the former with two, and the latter with a single 
species. They are more or less dormouse-like forms, with blunt 
muzzles, thick woolly fur. and long and well-haired tails. Lastly, 
the bush-rats (Golunda) have one Indian and another Ethiopian 

An interesting instance of how the present distribution of a 
genus is explained by palaeontological discoveries is afforded by 
the brush-tailed porcupines (Atherura], now represented by one 
species from the Malayan, and a second from the West African 
sub-region ; the connecting form being one of which fossil teeth 
have been found in the Karnul district of Madras. From this 
it may be inferred that the genus was probably also represented 
in the Siwaliks. To the same family (Hystricida) belongs a 
peculiar porcupine from Borneo, constituting the genus Trichys. 

Passing on to the ungulates, we have first to notice a peculiar 
group of oxen forming a section of the genus Bos, which is con- 
fined to this region, and characterised, in addition to certain features 
of the skull and horns, by the dark colour of the males, or of 
both males and females. Of these, the gaur (B. gaurus) inhabits 
both India and the Malayan countries, but appears never to have 
reached Ceylon ; while the banteng (B. sondaicus) is confined to 
the countries on the east of the Bay of Bengal. Fossil representa- 




tives of this group occur in the Indian Plistocene ; and certain 
generalised oxen from the Siwalik Hills and the Pliocene of 
southern Europe, in which the females were generally or always 
hornless, may have been the ancestral type. The Indian buffalo 
(B. bubalus] is markedly distinct from the Ethiopian forms, and 
has ancestral representatives in the Indian Pliocene and Plisto- 
cene. While abundant in Ceylon, it is probably unknown in a 
truly wild state to the east of the Bay of Bengal. The Philippine 
buffalo, or tamarao (B. mindorensis) is regarded by some as a 

FIG. 64. JAPANESE SEROW (Nemorhadiis crispus}. 

cross between the last and the anoa of Celebes ; ancestral types of 
the latter occurring, as already mentioned, in the Siwaliks. In the 
same family the short-horned goats of the genus Hemitragus are 
represented by two Indian species, one inhabiting the Himalaya 
and the other the Nilgiris ; the third living species being south 
Arabian. One extinct species occurs in the Siwaliks and a second 
one in Perim Island, so that the group is essentially an Indian 
one ; and, as already mentioned, its present distribution can only 


be accounted for by a lowering of the temperature during a past 
epoch. Of the goat-like genera Nemorhcedtis and Cemas, the 
former has a wide range in the region and also extends into 
northern China and Japan, while the latter is represented solely 
by the Himalayan goral ; no fossil types of either being known. 
In its poverty of antelopes (exclusive of the widely-spread gazelles) 
the Oriental presents a most remarkable contrast to the Ethiopian 
region, although this poverty is largely a feature of the present 
epoch, African types being common in the Siwaliks. The sole 
existing forms are the four-horned antelope (Tetraceros quadri- 
cornis], the black-buck (Antilope cervicapra)^ and the nilgai (Bos- 
elaphus tragocamelus], each of which forms a genus by itself, and 
all of which are restricted to India, exclusive of Ceylon. Indeed, 
it is a remarkable feature that true antelopes and gazelles are 
unknown to the eastward of the Bay of Bengal ; although this may 
be chiefly or entirely due to the countries to the eastward being 
unsuited to their habits. The nilgai, which has fossil representa- 
tives in the Indian Plistocene and Pliocene, is allied to the kudu 
group of Africa, while the four-horned antelope is a near relative 
of the duikers. It will be unnecessary to say anything with regard 
to the true goats (Caprd) and sheep (Ovis) inhabiting the region, 
since these are found only on the north-western frontier of India, 
and are obviously intruders from the Holarctic region. It is, 
however, important to mention that extinct representatives of one, 
if not of both groups, occur in the Siwalik Hills. 

The abundance of Cervidce is one of the most noticeable 
features distinguishing the Oriental from the Ethiopian region ; 
there being an equally marked difference in this respect between 
the former and the Holarctic area. Although the majority of the 
Oriental deer are now included in the genus Cervus, the typical, 
or elaphine group, as represented by the red deer and the wapiti, 
is entirely wanting, its place being taken by the sambur and its 
allies (C. unicolor), forming the rusine group; the swamp-deer 
(C. duvaucelt), which with another species constitutes the rucervine 
group, and the Indian spotted deer (C. axis), alone representing 
the axine group. Rusine deer are abundant in the Indian 
Siwaliks, but appear to be unknown in the Pikermi beds. 
Although they have one Tibetan representative, the smaller deer 


known as muntjacs (Cervulus) which are characterised by the 
length of the pedicles of the antlers and the shortness of the antlers 
themselves form a very characteristic Oriental group, ranging over 
the entire region. Not improbably they are represented in the 
Pliocene of Europe. 

The chevrotains, or TragulidcK^ which have already been shown 
to be abundant in the European Oligocene and Miocene remains 
of the West African genus occurring in the latter deposits and 
the Indian Pliocene are represented in the Oriental region by 
Tragulus, which dates from the Siwalik epoch, and ranges from 
India and Ceylon to the Philippines. Although wild camels are 
now everywhere unknown, it is probable that India and the 
Holarctic region was their original home, remains of the genus 
Camelus being found in the Pliocene of the Siwalik Hills. 

The large number of species of true pigs (Sus} characterising 
the Oriental region is a notable feature, India itself being in- 
habited by a species (Sus cristatus] nearly allied to the European 
wild boar, while the Malayan sub-region is the home of a consider- 
able number of species differing more or less markedly from the 
latter. The genus is well represented both in the Pliocene and 
Plistocene of India, but in neither of these formations are there 
any of the Ethiopian types of the family. 

With the exception of the Ethiopian, the Oriental region is now 
the sole one where the family Rhino cerotida still exists ; but there 
is a remarkable difference between the species inhabiting the two 
areas, all the three living Asiatic forms being furnished with teeth 
in the front of the jaws, which, as we have seen in the last chapter, 
are wanting in the African species. While one of the Oriental 
rhinoceroses (R. sondaicus} ranges from eastern Bengal to the 
Malayan islands, and a second (R. sumatrensis) from Assam to 
the same, the great Indian species (R. unicornis) is unknown to 
the eastward of Assam, as it is in Ceylon. Fossil remains of the 
latter are found in the Plistocene of the Narbada valley, while 
ancestral types both of this species and of R. sondaicus are met 
with in the Pliocene of the Siwalik Hills. It is, however, very 
remarkable that Ethiopian types of the genus occur not only in 
the last-named deposits, but likewise in the Plistocene of Madras ; 
the total extinction of this group in India being, as in the case of 


other Ethiopian types, almost impossible to account for. One of 
the two-horned extinct Indian rhinoceroses (R. platyrhinus] ap- 
pears to have been the ancestor both of the existing R. simus of 
Africa and R. antiquitatis of the Plistocene of northern Asia and 
Europe ; the evolution of the latter species having not improbably 
taken place in the countries lying between India and China, 
whence the creature wandered northwards and westwards with the 
mammoth to the Arctic tundras. With regard to the Equidce, it 
will suffice to mention that species of Equus occur in the Plisto- 
cene of Central India and Madras, and that wild asses (of a type 
markedly different from the African wild ass) occur in Sind and 
Kach. The genus, like the antelopes, is, however, totally un- 
known in the countries to the east of the Bay of Bengal, as it is in 
Ceylon. In the Tapiridcz, the Malayan tapir (Tapirus indicus) 
inhabits the Malay Peninsula as far north as Mergui, and also the 
islands of Sumatra and Borneo. It is important to notice that 
although fossil remains of tapirs are unknown from the Pliocene of 
the Siwaliks, they are met with in caverns in China. 

Distinguished, among other features, from its African cousin by 
the thinner and more numerous enamel-plates of its molar teeth, 
the Indian elephant (Elephas indicus) ranges over the greater part 
of the region, being found in suitable districts in India, Ceylon, 
Burma, the Malay Peninsula, Cochin China, and Sumatra. This 
species is a near ally of the mammoth (E. primigenius) ; and it 
may prove that both are descendants of a Siwalik species (E. 
hysudricus], which has molar teeth of the type we should expect to 
find in such an ancestral form. If this view be correct, the 
mammoth has probably wandered to northern Europe and Siberia 
from the countries lying just to the east of India. It has been 
mentioned in an earlier chapter that the extinct so-called stego- 
dont elephants (such as E. clifti and E. insignis] are mainly 
confined to this region, although some of the species are found in 
north China and Japan. As these elephants form the transition 
between Elephas and Mastodon, and also since the species of the 
latter genus which may be regarded as the original stock of the 
elephants is confined to the Indian and Malayan Pliocene, it may 
be taken for granted that the elephants have been developed from 
the mastodons within the limits of the Oriental region. In the 


Plistocene of the Narbada Valley in central India there occurs a 
species (E. namadicus] closely allied to the contemporary Euro- 
pean E. antiqims, in both of which the molars are intermediate in 
structure between those of the living Indian and African species. 
Elephas planifrons of the Siwaliks, which has molars of a still more 
generalised type, is equally closely allied to E. meridionalis of the 
upper Pliocene of Europe; and it is quite probable that the 
former may be the original ancestral stock of the African elephant. 
It is worth mentioning that the stegodont elephants survived till 
the Plistocene ; and also that some of the species of this group 
inhabiting India, as well as certain mastodons, ranged as far east- 
wards as Java, Borneo, China, and Japan. 

FlG. 65. WHITE-BELLIED PANGOLIN (Manis tricuspis). 

The last mammals that we have to mention are the pangolins 
(Mamdcz), which are now common to the Oriental and Ethiopian 
regions, and appear to be represented by an extinct genus in the 
European upper Oligocene. The presence of horny scales in- 
vesting the whole of the body and tail serves to distinguish the 


pangolins from all other mammals whatsoever ; and the Oriental 
species are further characterised by having the median series of 
scales on the body continued to the tip of the tail, and likewise by 
the presence of numerous isolated hairs between the scales of the 
back, as well as by the presence of small ears. Of the three 
Oriental species, Manis javanica ranges from Burma through the 
Malay Peninsula to Java and Borneo ; M. aurita extends from 
Nipal to the Indo-Chinese sub-region ; while M. pentedactyla is 
restricted to India and Ceylon. The most remarkable feature 
connected with the distribution of the group is, however, the 
circumstance that claw-bones indistinguishable from those of the 
giant pangolin (M. giganted] of West Africa have been discovered 
in a cavern in the Karnul district of Madras. 

In the following list the leading results of the foregoing survey 
are put in tabular form, the italics indicating groups or species 
peculiar to the region. 

I. Primates. 

Simia. Borneo and Sumatra ; fossil in India. 
Hylobates. Burmese and Malayan ; fossil in European 


Macacus. Now mainly Oriental, but occurring on 

the southern borders of the Holarctic region ; 

fossil in the European and Indian Pliocene. 
Semnopithecus. An outlying species in Eastern Tibet 

and one in Kashmir ; fossil in Pliocene of Europe 

and India. 
Nasalis. Borneo. 

Nycticebus. Burmese, Malayan, and Indo-Chinese. 
Loris. S. India and Ceylon. 

Elsewhere only in Celebes. 
Tarsius. Malayan, extending into Celebes. 


II. Insectivora. 


Galeopithecus. Malayan. 


Tupaia. Indian and Malayan. 

Ptilocercus. Borneo and some adjacent islands. 


Gymnura. Burmese and Malayan. 
Hylomys. Burmese and Malayan. 


Soriculus. Himalayan and Indo-Chinese. 

Anurosorex. Known by one species from Assam and 
a second from Tibet and Pekin. 

Chimarrogale. Represented by one species from the 
eastern Himalaya, hills north of Burma, and Mt 
Kina Balu, Borneo, and a second from Japan. 

III. Carnivora. 

FELID^:. Very numerous in the region. 

Cynaelurus. Indian and Ethiopian ; the one species 
being common to the two areas ; fossil in Indian 


Viverra. All the species, except a single Ethiopian 
one, are Oriental, one of these being frequently 
regarded as the representative of a distinct genus 
( Viverricula) ; fossil in European Oligocene and 
European and Indian Pliocene. 

Hemigale. Malayan. 

Linsanga. Malayan and E. Himalayan. 

Paradoxurus. An outlying species in Celebes. 

Arctogale. Burmese and Malayan. 

Arctictis. Burmese and Malayan. 

Cynogale. Malayan. 


Melursus. India and Ceylon. 


III. Carnivora. (Cont.) 


sElurus. Eastern Himalayan and Burmese ; fossil in 

English Pliocene. 

Arctonyx. Two E. Himalayan and Burmese species, 

and probably a third from Tibet. 
Mellivora. One Indian and another Ethiopian 

species ; fossil in Indian Pliocene. 
Helictis. India to China. 
Mydaus. Malayan. 

IV. Rodentia. 


Rhithrosciurus. Borneo. 

Nannosciurus. Represented elsewhere by a single 

West African species. 
Sciurus. This almost cosmopolitan genus appears to 

attain its maximum development in the Malayan 


Chrotomys. Philippines (Luzon). 
Xeromys. Philippines (Luzon), and Australia. 
Phlceomys. Philippines. 
Crateromys. Philippines (Luzon). 
Hapalomys. Burma. 
Vandeleuria. India and Burma. 
Chiropodomys. Burmese and Malayan. 
Pithechirus. Sumatra and Java. 
Rhynchomys. \ 

Carpomys. > Philippines (Luzon). 
Batomys. / 

Golunda. One Indian and one Ethiopian species. 

Atherura. One Malayan and one West African 

species ; fossil in Indian Plistocene. 
Trichys. Borneo. 


V. Ungulata. 

Bos. The Bibovine group exclusively Oriental. 
Hemitragus. Two Indian species, and a third in the 

South Arabian sub-region of Ethiopia; fossil in 

Indian Pliocene. 
Nemorhaedus. Largely Oriental (Himalayan, Bur- 

mese, and Malayan), but extending into northern 

China and Japan. 
Cemas. Himalayan. 
Tetraceros. Indian. 

Antilope. Indian ; fossil in Plistocene. 
Boselaphus. Indian ; fossil in Plistocene and Pliocene. 

Cervus. The Rusine, Ruce.rvine, and Axine groups 
of this genus are characteristic of the" region, 
although the first is also represented in the Austro- 

Cervulus. Mainly Oriental, but with one Tibetan 


Tragulus. India, Ceylon, and Malayan sub-region ; 
fossil in Indian Pliocene. 

Sus. Attains its maximum specific development in 
the Malayan sub-region. 


Rhinoceros. The three existing Oriental species 
differ from the Ethiopian forms in having front 


Tapirus. Malayan : elsewhere living only in the 
Neogaeic realm, but widely distributed in a fossil 
state, although absent from the Siwaliks. 


V. Ungulata (cont.). 


Elephas. The existing Oriental elephant is widely 
different from the Ethiopian, although nearly 
allied to the Holarctic mammoth ; the extinct 
Stegodont group is mainly Oriental, although 
extending into north China and Japan. 

VI. Effbdientia. 


Manis. Elsewhere only in Ethiopian region ; fossil 
in Indian Plistocene. 

The relations of peninsular India to the Himalayan area have 
been already discussed at the commencement of 
of^hV Region y this cna P ter \ wm ' le tne land-connection which 
appears to have existed between India and Mada- 
gascar, and thus with Africa, has been alluded to in an earlier 
one. The latter connection must have ceased to exist before 
the Pliocene era ; and, as we have seen, the descendants of the 
Siwalik mammals would appear to have made their way into 
Ethiopia across Syria or Arabia. During the Pliocene, India, at 
least, could not have been distinguished as a region from Ethiopia 
as it exists at the present day ; and even in the Plistocene the 
connection between the faunas of the two areas was much more 
intimate than it is now. The full reason for this gradual dis- 
appearance of the modern Ethiopian types from the Indian area 
will probably never be known ; but there can be little doubt that 
the gradual refrigeration of the northern hemisphere with the 
advent of the glacial period has been largely instrumental ; the 
present distribution of Hemitragus being only explicable on the 
hypothesis of a marked lowering of the temperature over India. 

The more peculiar mammals now inhabiting the Oriental 
region may be roughly arranged under five headings. The first 
will include those that are common to India and some of the 
countries to the west or south-west, but are, for the most part, 
unknown in either Ceylon or the countries to the eastward of the 
Bay of Bengal. Under this category may be included the follow- 
ing, viz. : 




INSECTIVORA. Erinaceus. 

CARNIVORA. Felis leo. Ethiopian, and European Plisto- 

Felis chaus. Ethiopian, and European Plistocene ; 

ranges eastward into Burma. 
Felis caracal. Ethiopian. 
Cynaelurus jubatus. Ethiopian. 
Hyaena striata. Western Asia and North Africa. 
Hyaena crocuta. Ethiopian ; occurs both in India 

and Europe during the Plistocene. No 

representative of the genus known in the 

Burmese or Malayan countries. 
Canis aureus. S. W. Asia ; also ranges into 


Canis pallipes. South Arabian. 
Mellivora. The Indian and Ethiopian species 

very closely allied. 

RODENTIA. Golunda. Ethiopian ; the Indian species ranges 
into Ceylon. 

UNGULATA. Hemitragus. South Arabian. 

Antelopes. ) None known east of the Bay of 
Equus. j Bengal. 

The second group includes such genera and species as are 
common to India and Ceylon, but are unknown elsewhere. Here 
we have : 




Felis rubiginosa. 

Golunda ellioti. 

Bos bubalus. 
Cervus axis. 
Tragulus memimna. 

EFFODIENTIA. Manis pentedactyla. 




The third group, which is a small one, comprises types which 
are confined to India ; it includes 
Felis ornata. 
Rhinoceros unicornis. 

In the fourth group we have generic or specific forms common 
to India and the countries to the eastward of the Bay of Bengal, 
but unknown in Ceylon or in the countries to the west or south- 
west. This list comprises the following, viz. : 
PRIMATES. Hylobates. 
CARNIVORA. Felis tigris. 

Felis nebulosa. 

Felis marmorata. 




UNGULATA. Bos gaums. 

Bos frontalis. 


Cervus porcinus. 

Finally, we have (among others) the following group of genera 
and species confined to the countries lying immediately to the 
eastward of the Bay of Bengal, viz. : 




INSECTIVORA. Galeopithecus. 



RODENTIA. Rhithrosciurus. 



UNGULATA. Bos sondaicus. 

Tragulus javanicus. 

EFFODIENTIA. Manis javanica. 

Other forms might be added to several of these lists, but 
those included are sufficient for the purpose of showing that the 
present mammalian fauna of India is a complex formed by an 
admixture of western and eastern types. 

The first group is an essentially modern one, all the generic 
types contained in it, with the exception of Erinaceus (which dates 
from the Miocene), being unknown before the lower Pliocene ; 
while, if we except Erinaceus and Golunda, all occur in the 
Siwaliks. In the Carnivora, there is evidence of all the species 
except the first three being descended from Siwalik ancestors ; 
and it is quite probable that the three species of Felis may 
trace their origin to felines which lived either in the Siwaliks or 
Persia during the Pliocene, in which event the lion, and not 
the tiger, should be regarded as the characteristic large Indian 

With the probable exception of Loris, the second group is 
also a modern one ; all the forms save Loris and Golunda having 
ancestral types in either the Pliocene or Plistocene of India, and 
none of the genera being known before the former epoch. And it 
may be mentioned here that the absence of so many of the smaller 
types of Oriental mammals from the Siwaliks is no indication 
that the genera did not flourish in India during the Pliocene age, 
but is due to the strata being unsuited to the preservation of their 
remains. The remarks applicable to the second group will like- 
wise befit the third. 

On the other hand, the fourth and fifth groups appear to have 
less connection with the Siwaliks, while several of the types are 
older ones. For instance, we have no proof of the existence of 
oxen nearly allied to Bos gaurus in the Siwaliks, although such 
are found in the Indian Plistocene ; neither is there any evidence 
of a Siwalik tapir. Tupaia is a near relative of the European 
Miocene Galerix and the Oligocene Lanthanotherium ; as is 
Gymnura of the Oligocene Necrogymnura ; while Hylobates is 



represented in the European Miocene 1 . The reasons for regard- 
ing the tiger as a comparatively modern immigrant into southern 
India have already been stated. A Siwalik origin is, however, 
indicated for Simla ; but concerning the other genera the palaeon- 
tological history is unfortunately a blank. 

The affinity between the faunas of West Africa and the Malayan 
sub-region has been already alluded to ; but there are also indica- 
tions of a connection between that of the latter area on the one 
hand and that of southern India and Ceylon on the other, as 
exemplified by the occurrence of Nycticebus in the Malayan sub- 
region and of Loris in south India and Ceylon. To explain the 
latter connection, Dr Blanford 2 has discussed the possible exist- 
ence of a direct land-communication between the two areas ; but 
this connection, as he admits, scarcely seems necessary, since in 
such cases the true explanation would seem to be the survival of 
old types in the tropical forest-regions. And here it may be 
noticed that the Malayan types common to, or represented by 
allied forms in West Africa, are such as either have representatives 
in the Indian Pliocene or Plistocene, or such as we might naturally 
expect to meet with there if small forms were commonly preserved. 
For instance, Simia and Anthropopithecus, and Dorcatherium and 
Tragulus, are all represented in the Siwaliks, and Atherura 
occurs in the Madras Plistocene. This being so, what is more 
likely than that lorises, linsangs, and palm-civets, of types more or 
less intermediate between the existing Malayan and West African 
representatives of those groups, should have flourished in India 
during the Pliocene era ? Nannostiurus, again, should certainly 
be expected to occur in the Indian Pliocene. On this point I 
think Dr Wallace 3 was on the right track when, in writing of the 
Malayan sub -region, he observed that "Here alone, in the 
Oriental region, are found the most typical equatorial forms of 
life-organisms adapted to a climate characterised by uniform but 
not excessive heat, abundant moisture, and no marked departure 
from the average meteorological state throughout the year. These 

1 If my memory serves me right, I have been shown teeth from the upper 
Oligocene Phosphorites of France closely resembling those of Hylobates. 

2 Manual of Geology of India, rst Ed. pt. 1. p. Ixviii. 

3 Geographical Distribution of Animals, vol. I. p. 335. 


favourable conditions of life only occur in three widely separated 
districts of the globe the Malay Archipelago, Western Africa, and 
equatorial South America. Hence, perhaps, it is that the tapir and 
trogons of Malacca should so closely resemble those of South 
America ; and that the great anthropoid apes and crested horn- 
bills of Western Africa should find their nearest allies in Borneo 
and Sumatra." 

In addition to the resemblances between the mammalian fauna 
of the Malayan sub-region and that of West Africa, there are, 
however, equally well-marked differences between the former and 
that of Ethiopia in general. Among Burmese and Malayan types 
wanting in Africa, we have especially to note Tapirus, Gymnura, 
Tupaia, Hylobates, and sElurus. From the small size of their 
representatives it would be unfair to argue anything from the 
absence of the last four of these from the Siwaliks, but the case is 
very different with regard to Tapirus, which ought surely to have 
been found did it exist there. As this genus is equally wanting 
in the Pikermi and Persian Pliocene, while it occurs in that of 
France, Germany, England and China, it is a fair inference that it 
has reached the Malayan countries by a route lying north of India. 
And the occurrence of sElurus in the English Pliocene suggests 
that the same may be the case with that genus. If this be so, 
it is not an improbable hypothesis that the other genera men- 
tioned, all of which have representative types in the European 
Tertiaries, may have migrated eastwards by a similar route ; and 
in this connection it is especially noteworthy that such of the 
genera in question as enter India at all, occur only in the eastern 
or southern districts. 

With regard to the date of the separation of Ceylon from 
India, the numerous species of mammals common to the two 
areas show that this must have taken place at a very recent date, 
comparatively speaking ; although, as aforesaid, at a period when 
several of the mammals now inhabiting southern India had not yet 
occupied that portion of their distributional area. 

When discussing the possibility of a former land-connection 
across the Bay of Bengal between Ceylon and southern India on 
the one hand, and the Malayan countries on the other, Dr Blanford 
was careful to point out that the ocean-bed afforded no evidence 


in favour of such a line of communication. This feature, together 
with certain marked differences between the mammals of the two 
areas, appears to afford a conclusive argument that these countries 
have never been much more closely connected than they are at 
present. Had any more extensive connection existed, we should 
surely expect to find antelopes, gazelles, and perhaps asses, in the 
more open districts of upper Burma ; while the Bay of Bengal 
would scarcely have formed such a sharp line limiting the eastward 
range of wolves, foxes, hyaenas, and the other mammals mentioned 
in the list on page 289, as it actually does. That list is confined 
to existing types, but if fossil forms had been included, Hippo- 
potamus might have been added, since the extinct Oriental 
representatives of that genus do not range further east than Burma 
(whither they evidently migrated down the river-valleys from 
northern India), no species being known from the Tertiaries of 
China, Japan, or the Malayan islands. These circumstances, 
together with the depth of the sea in the Bay of Bengal, seem to 
disprove the suggestion of Dr Wallace 1 that a continuous tract of 
land formerly connected Borneo and the rest of Malaysia with 
the central peak of Ceylon, and extended eastwards to Hainan. 
Within the limits of the present volume it would be quite 
impossible to give a detailed description of the 
SuWegion mammalian faunas of the Malay Peninsula and 
Islands ; and I have accordingly selected that of 
the Bornean group, as an example of what may be called the 
typical Malayan sub-region, as distinct from Java, which differs 
markedly in its fauna from Borneo and Sumatra. The chief 
reason for selecting Borneo is that its fauna has been carefully 
worked out by Mr A. H. Everett 2 and Mr C. Hose 3 , from whose 
papers the following list of mammals (exclusive of bats), with some 
emendations and additions, has been compiled ; species, like the 
rat, mouse, and buffalo, which have obviously been introduced, 
being omitted. Mr Everett includes within the Bornean group 
the island of Palawan, and states that the group may be defined 
" by a line which starts from a point immediately to the west of 

1 Op. dt. p. 359. 2 Appendix, No. 14. 

3 Descriptive Account of the Mammals of Borneo, Diss, Norfolk, 1893. 


St Julian Island in the Tambelan Archipelago, and being drawn 
south of the Great Natuna (Bungoran Island), passes northward of 
Labuan and thence follows the loo-fathom line, so as to embrace 
Balabac, Palawan (Paragua), the Calamianes, and the Cuyo 
Islands, and, returning along the same line of soundings on the 
southern side of Palawan, is drawn immediately to the islands of 
Cagayan Sulu, and Sibutu, whence it is continued through the 
Makassar Straits south of the Paternoster, Lauriot (Laset Ketjil), 
and Solombo islets, and in a north-westerly direction through the 
Karimata Strait back to the island of St Julian." In the following 
list the genera and species peculiar to the group are printed in 
italics, those which are confined to the Palawan sub-group having 
that name placed after them in brackets. The distribution of the 
other forms has been indicated as far as practicable. 
PRIMATES. Simia satyrus. Sumatra. 

Hylobates leuciscus. Java to Philippines. 

,, muelleri. 

Semnopithecus maurus. Malay Peninsula and Java. 
,, chrysomelas. 


,, everetti. 

,, rubicundus. 

,, frontatus. 

Nasalis larvatus. 

Macacus arctoides. Burma, China, E. Tibet, etc. 
nemestrinus. Burma, Malay Peninsula, 

Sumatra, and Java. 

cynomolgus. Burma to Philippines. 
Nycticebus tardigradus. Burma to Philippines. 
Tarsius spectrum. Sumatra, Java, and Banka. 
INSECTIVORA. Chimarrogale himalayica. E. Himalaya and hills 

north of Burma. 

Crocidura fuliginosa. E. Himalaya. 
dor ice. 



Gymnura rafflesi 1 . S. Tenasserim, Malay Penin- 
sula, and Sumatra. 
Hylomys suilla. Burma, Malay Peninsula, and 


Ptilocercus lowi. Also in some neighbouring islands. 
Tupaia murina. 

,, javanica. Java and Malay Peninsula. 
,, melanura. 

,, ferruginea. Burma, Malay Peninsula, Suma- 
tra, and Java. 
,, splendidula. 

tana. Sumatra, Natuna Islands. 
,, dor satis. 
,, montana. 

Galeopithecus volans. Malay Peninsula, S. Tenas- 
serim, Siam, Sumatra, and Java. 

CARNIVORA. Felis planiceps. Malay Peninsula and Sumatra. 
,, badia. 
temmincki. E. Himalaya, Burma, Malay 

Peninsula, and (?) Sumatra. 
,, bengalensis. India to Philippines. 
,, marmorata. E. Himalaya to Sumatra. 
,, nebulosa. E. Himalaya to Formosa. 
Viverra tangalunga. Malay Peninsula, Sumatra, 

Philippines, and Celebes. 
Linsanga gracilis. Java and (?) Sumatra. 
Paradoxurus leucomystax. Malay Peninsula and 

,, hermaphroditus. India to Java and 


,, philippinensis. Philippines. 

Arctogale leucotis. Sikhim, Burma, Malay Penin- 
sula, and Sumatra. 
1 Mr Jentink regards the Bornean form as a distinct species (G. alba]. 


CARNIVORA (cent.}. 

Hemigale hardwickei. Malay Peninsula and Su- 
,, hosei. 
Arctictis binturong. E. Himalaya, Burma, Siam, 

Malay Peninsula, Sumatra, and Java. 
Helictis everetti. 

Cynogale bennetti. Malay Peninsula and Su- 
Herpestes brachyurus. Malay Peninsula. 

,, semitorquatus. Sumatra 1 . 
(?) Canis rutilans. Malay Peninsula, Sumatra, and 

Ursus malayanus. Arakan, Tenasserim, Malay 

Peninsula, Java, and Sumatra. 
Mydaus meliceps 2 . Java and Sumatra. 
Mustela flavigula. India to China. 

,, nudipes. Malay Peninsula and Sumatra. 
Lutra sumatrana. Malay Peninsula, Sumatra, and 

,, cinerea. India to Java and China. 

RODENTIA. Sciuropterus pulverulentus. Malay Peninsula. 

,, horsfieldi. Malay Peninsula and Java. 

setosus. Sumatra. 

,, genibarbis. Java. 

,, nigripts (Palawan). 

Pteromys nitidus. Malay Peninsula and islands. 

Rhithrosriums macrotis. 
Sciurus bicolor. E. Himalaya to Siam and 

(?) Celebes. 
prevosti. Malay Peninsula, Sumatra, and 


,, hippurus. Malay Peninsula and islands. 

1 Jentink, Notes Leyden Museum, vol. XVI. p. 210 (1894). 

2 The form from Calamianes has been separated as M. marc Jut. 



Sciurus brookei. 

tenuis. Malay Peninsula and islands, to 


,, lowi. 
,, notatus. Malay Peninsula, Sumatra, 

Java, etc. 
,, insignis. Malay Peninsula, Sumatra, and 


,, ho set. 
,, everetti. 
,, steerei (Palawan). 
,, laticaudatus. Malay Peninsula. 
,, soricinus, Java and Sumatra. 
Nannosciurus exilis. Malay Peninsula and Su- 

,, whiteheadi, 

,, melanotis. 

Mus infralnteus. Sumatra. 
,, muelleri. Sumatra. 
,, sabanus. 

,, neglectus. Philippines. 
,, jerdoni. E. Himalaya, Tenasserim, Java. 
,, alticola. 

,, ephippium. Sumatra, Philippines. 
,, margaretta. 
,, raja. 

,, ochraceiventer. 
,, whiteheadi, 
,, bxodon. 
,, baluensis. 
Chiropodomys major. 

,, pusillus. 

Hystrix crassispinis . 

,, pumila (Palawan sub-group). 
,, muelleri. Sumatra. 
Trichys gnentheri. 


UNGULATA. Bos sondaicus. Burma, Malay Peninsula, Java, 

and Bali. 

moellendorffi ' (Palawan sub-group). 
Cervus unicolor 2 , var. Probably introduced. 
Cervulus muntjac. 

Tragulus napu. S. Tenasserim to Java and Sumatra. 
,, nigricans (Palawan sub-group). 
,, javanicus. Malay Peninsula to Cochin 


Sus vittatus. Java, Sumatra, Amboyna, Batjian. 
,, verrucosus. Java, Ceram. 
,, barbatus*. 
longirostris. Java. 
Rhinoceros sumatrensis. Assam to Siam, Malay 

Peninsula, and Sumatra. 
Tapirus indicus. S. Tenasserim, Malay Peninsula, 

and Sumatra. 

Elephas indicus. Probably introduced. 
EFFODIENTIA. Manis javanica (Palawan sub-group). 

In this list the genera Nasalis, Trichys, and Rhithrosciurus are 
peculiar to the group, while Ptilocercus is almost so ; and, includ- 
ing the latter, there are no less than fifty-one species peculiar to 
Malaysia. Of these a very large number are common to Sumatra 
or the Malay Peninsula, or both together, while a smaller number 
occur in Java. There are but six species common to peninsular 
India, among which the Indian elephant and the sambar may have 
been introduced ; but there are ten which are found in the Eastern 
Himalaya or Assam. The most remarkable among these is the 
Himalayan water-shrew (Chimarrogale himalayica), which is only 

1 Founded on a skull from the island of Busuanga, in the Calamianes, 
which probably indicates only a race (? introduced) of the buffalo. 

2 The so-called C. equinus is regarded by Dr Blanford as not specifically 
distinct from C. tinicolor. This being so, it is probable that the Bornean 
form described by Mr C. Hose (Ann. Mag. Nat. Hist. ser. 6, vol. xn. p. -206) 
as C. brookei is also a variety. 

3 Sits ahanobarbus, Huet, from Palawan, and S. calamianensis, Heude, 
from the Calamianes, are identified by Dr Nehring (Sb. Ges. Nattirf. Berlin, 
1894, pp. 190, 191) with this species. 


found in the Eastern Himalaya, the mountains north of Burma, 
and Mount Kina Balu, in North Borneo ; a musk-shrew ( Croridura 
fuliginosa\ common to the Eastern Himalaya and Borneo, being 
likewise unknown elsewhere. These two instances alone are 
sufficient to prove that Borneo must have been in immediate 
connection with the lands to the north-west within the period 
during which the living species of mammals have come into exist- 
ence; while the restriction of the water-shrew to the mountains 
seems likewise to imply a former lowering of the temperature of 
the whole region sufficient to enable the creature to pass from the 
one area to the other, or perhaps rather to have allowed of its 
existence in the intermediate lowlands, whence it migrated to its 
present isolated haunts. That Borneo was connected with the 
mainland during the Pliocene epoch is proved by the occurrence 
in that island of the Siwalik Mastodon latidens, the tooth figured 
on page 173 being of Bornean origin. 

The large number of species common to Borneo, Sumatra, and 
the Malay Peninsula also shows that these three areas must have 
been very recently in connection ; but the excessive number of 
peculiar Bornean forms seems to indicate that the former island, 
with the adjacent islets, was the first to be isolated. Even so. 
however, the extraordinarily large percentage of distinctive types 
is most remarkable. Regarding the relationship of the Palawan 
sub-group to Borneo, Mr Everett writes that "although the general 
facies of the mammalian fauna of the sub-group is clearly Bornean, 
it is to be noted that no species appears to be peculiar to the 
group as a whole, a fact which suggests the inference that closely 
connected as Borneo has undoubtedly been with Balabac and 
Palawan, and isolated as they have been together from the main- 
land of Asia, there has also been much isolation of Borneo and 
Palawan inter se" 

From Sumatra and Borneo, which have so much in common, 
and in a somewhat less degree from the Malay Peninsula, Java 
differs very remarkably as regards its mammalian fauna ; a large 
number of typically Malayan forms being absent, while others as 
characteristically Indian are present. In the first place, the 
orangs (Simia), common to Borneo and Sumatra, are absent : 
and the elephant and tapir are likewise wanting, the former 


certainly existing in a wild state in Sumatra, although it has been 
considered that its occurrence in Borneo is due to human intro- 
duction 1 . Although the Javan rhinoceros (J?. sondaicus}, as we 
have seen, is common to eastern Bengal, Burma, and Java, its 
reputed occurrence in either Borneo or Sumatra does not appear 
to rest upon any solid basis of fact 2 ; while the Sumatran species 
(R. sumatrensis], which is common to Borneo, is wanting from 
Java. It has, indeed, been stated 3 that certain teeth from the 
Plistocene of Borneo are referable to the last-named species, but 
the molars of both kinds are so alike that it is almost, if not 
quite, impossible to distinguish between them. A noteworthy 
circumstance is that whereas there is no Siwalik species allied 
to R. sumatrensis, yet R. sondaicus is almost indistinguishable 
from the Siwalik R. sivalensis, and is thus proved to be a very 
ancient Indian type. As another instance of the distinctness of 
the mammalian fauna of Java from that of Borneo and Sumatra, 
we may take the case of the banteng (Bos banteng}, which is 
wanting 4 from both those islands, but is present in Java, the 
Malay Peninsula, and Burma. Again, the genus Hemigale, of 
which the type species is common to the Malay Peninsula, 
Sumatra, and Borneo, is quite unknown in Java. That the latter 
island was directly connected with the mainland is, of course, 
proved by such species of existing mammals as are common to 
the two areas ; but if further evidence be needed, it is to hand in 
the fossil mammals which have been obtained from Pati-Ajam, in 
Java 5 . These comprise Elephas trigonocephalus, E. bombifrons, 
E. clifti, E. namadicus, E. hysudricus, Sus hysudricus, Bos siva- 
lensis, and Cervus lydekkeri. With the exception of the first and 
last (which may prove not to be distinct likewise), all these are 
Indian forms, E. namadicus belonging to the Plistocene, while the 
others pertain to the Siwalik fauna; and it may be added that 

1 Dr Wallace states that Ursus malayanus is absent from Java, but accord- 
ing to Dr Blanford this is incorrect. 

2 See Jentink, Notes Leyden Museum, vol. xvi. p. 231 (1894). 

3 Busk, Proc. Zool. Soc. 1869, p. 409. 

4 In the Fauna of British India Mammalia, p. 490, Dr Blanford gives 
Borneo, and perhaps Sumatra, as part of the habitat of the banteng, but the 
animal is omitted from fauna of the former by Mr Everett. 

5 K. Martin, Sammlungen Geol. Reichsmusetuns in Leiden, vol. IV (1887). 


the first three belong to the group of stegodont, or intermediate 

In endeavouring to explain the relationship of the Javan fauna 
to that of the rest of the Malayan sub-region, Dr Wallace 1 was 
first of opinion that Java, which was evidently isolated before 
Sumatra and Borneo, had a brief land-connection with the Siamese 
peninsula, independently of those two islands. This view, how- 
ever, was subsequently abandoned 2 , and the following hypothesis 
proposed. From the evidence of certain Tertiary rocks in Java 
believed to be of Miocene age, it is considered probable that at 
the epoch in question that island "would have been at least three 
thousand feet lower than it is now, and such a depression would 
probably extend to considerable parts of Sumatra and Borneo, so 
as to reduce them all to a few small islands. At some later 
period a gradual elevation occurred, which ultimately united the 
whole of the islands with the continent. This may have continued 
till the glacial period of the northern hemisphere, during the 
severest part of which a few Himalayan species of birds and 
mammals may have been driven southward, and ranged over suit- 
able portions of the whole area. Java was then separated by 
subsidence, and these species became imprisoned there ; while 
those in the remaining part of the Malayan area again migrated 
northward when the cold had passed away from their former 
home, the equatorial forests of Borneo, Sumatra, and the Malay 
Peninsula being more especially adapted to the typical Malayan 
fauna which is there developed in rich profusion. A little later 
the subsidence may have extended further north, isolating Borneo 
and Sumatra, but probably leaving the Malay Peninsula as a ridge 
between them as far as the islands of Banka and Billiton. Other 
slight changes of climate followed, when a further subsidence 
separated these last-named islands from the Malay Peninsula, and 
left them with two or three species which have since become 
slightly modified. We may thus explain how it is that a species 
is sometimes common to Sumatra and Borneo, while the inter- 
vening island (Banka) possesses a distinct form 3 ." 

1 Geographical Distribution of Animals, vol. I. p. 359. 

2 Island Life, p. 360. 

3 As exemplified in the case of the birds of the genus Pitta. 


Although not taking into account the relationship between the 
fauna of Borneo and that of the Palawan sub-group, this may 
be accepted as, on the whole, a very probable explanation of 
the facts of the case. It may be added that while the Javan 
rhinoceros (. sondaicus] is, as already stated, closely allied to the 
Siwalik R. sivalensis, the nearest ally of R. sumatrensis appears to 
be the extinct R. schleiermacheri of the European Miocene, thus 
affording one more instance of affinity between the typical Ma- 
layan fauna and that of the middle Tertiaries of Europe. 

Recent investigations on the mammals from some of the small 
chain of islands lying to the south-west of Sumatra, 
such as Nias and Sipora in the Mentawi group, to- 
gether with Christmas Island 1 , lying still further to Christmas 


the south, have shown that they differ very markedly 
from those of the larger islands. From Sipora, in addition to 
bats, Mr O. Thomas 2 records the following species, of which those 
peculiar to the island are printed in italics : 

Semnopithecus potenziani. 

Macacus nemestrinus. Widely spread. 

Tupaia ferruginea, var. hypochrysa. Java. 

Paradoxurus, sp. 

Pteromys nitidus. Widely distributed. 

Sciuropterus lugens. 

,, aurantiacus. Banka. 

Sciurus melanogaster. 
,, fraterculus. 

Mus siporanus. 

raja. N. Borneo. 

From Christmas Island the same writer 3 records a peculiar 
species of fruit-bat (Pteropus natalis], a variety of a widely-spread 
musk-shrew (Croddura fuliginosa), and two peculiar rats (Mus 
macleari, and M. nativitatis], remarkable for their large size. 
Regarding the bearings of the faunas of these islands on the 
general problem of distribution, Mr Thomas writes that the collec- 

1 This must not be confounded with the island of the same name in 

2 Ann. Mus. Civ. Genova, Ser. 2, vol. xiv. pp. 660 672 (1895). 

3 Proc. Zool. Soc. 1887, pp. 511514, and 1888, pp. 532 534. 


tion from Sipora "does not show the very slightest special 
relationship to Sumatra, and therefore lends weight to the view 
that the Mentawi chain is the remnant of a long peninsula or 
island, similar in shape to, but separate from the Malay Peninsula 
or Sumatra. Further than this I cannot at present go, mainly 
because we know so little of the small terrestrial mammals of the 
other islands of the chain, those of the Nicobars being almost 
unknown, and those of Simalu, Sibiru, and Pagi entirely, while 
in Nias-and Engafio the collections consist mainly of bats. Still 
the few indications there are, such as the relations to each other of 
Pteropus nicobaricus, modiglianii, and natalis, and of Mus siporamis 
and macleari, show that the mammals, like the other animals, pre- 
sent a general similarity throughout the chain the whole way from 
the Nicobars to Christmas Island." 

Hitherto the Philippine Islands (exclusive of Palawan, Cala- 

mianes, etc., which are classed with the Bornean 
sub-re^on. 6 group ') have been regarded as forming a portion of 

the Malayan sub-region ; but the discovery of a very 
peculiar mammalian fauna in the mountains of Luzon 2 clearly 
proves their right to form a sub-region apart. This mountain 
fauna, which it is highly probable may also prove to be existent in 
Mindanao, is evidently a very ancient one, showing certain indi- 
cations of affinity with that of Australia; while the plain fauna 
is of a more modern and generally Oriental type. Curiously 
enough, the indications of affinity with the fauna of Celebes are 
by no means strongly marked. Unfortunately, there is at present 
no knowledge of the palseontological history of the mammalian 
fauna of the group. The following species of mammals, exclusive 
of bats, have been recorded from this sub-region 3 ; the names of 
such genera and species as are peculiar being printed in italic type. 
PRIMATES. Hylobates leuciscus. 

Macacus cynomolgus. A distinct race 4 . 

Tarsius philippinensis 1 '. 

Nycticebus tardigradus. 

1 Vide supra, p. 294. 2 See Thomas, Appendix, No. 31. 

3 In addition to the paper cited above, see Bourns and Dear, Appendix, 
No. n. 4 Often separated as M. philippinensis. 

5 Meyer, Abh. Mtts. Dresden, 1894, Art. i, p. i. 




INSECTIVORA. Galeopithecus volans 1 . 

Tupaia everetti. 
CARNIVORA. Viverra tangalunga. Ranges to Celebes. 

Paradoxurus philippinensis. Also Bornean. 
Felis bengalensis. 

RODENTIA. Sciurus philippinensis. 
,, samarensis. 

Nannosciurus concinnus. 
Chrotomys whiteheadi. 
Xeromys silaceus. ' j 
Phlaomys cumingi. 
,, pallidus*. 
Crateromys schatenbergi. 
Rhynchomys soricoides. 
Carpomys melanurus. 
,, phczurus. 

Mountains of Luzon. 


Batomys granti. 
Mus luzonicus. \ 

,, chrysocomus. \ 


,, ephippium. > 

Bos mindorensis. 
Cervus philippinus. 

,, alfredi. 
Sus celebensis, var. ! 

Mountains of Luzon. 

Mountains of Luzon. 

Elsewhere only in Celebes. 

With the exception of the Tarsius, which is now regarded as a 
peculiar species, the Primates are all wide-ranging forms, as is also 
the case with Galeopithecus volans, Viverra tangalunga, and Felis 
bengalensis. A relationship with Borneo is indicated by the Para- 
doxurus and two species of Mus ; while the pig is typically a 
Celebean form. The tamarao (J3os mindorensis] has its nearest ally 
in the anoa of Celebes, but, as mentioned in an earlier chapter, 


1 The Philippine race has been separated as G. 

2 Doubtfully distinct. 

3 Equal S. marchei, Huet. 



it is suggested that the animal in question is really a hybrid 
between the latter and the Indian buffalo. The two species of 
deer are small forms allied to the race of the sambar inhabiting 
Java and Borneo ; the first in the list being uniformly coloured, 
while the second is spotted with white at all ages. The so-called 
Cervus marianus of Luzon appears to be inseparable from C. 

It is remarkable that the six genera peculiar to the group all 
belong to the Muridce, and that five of these are known only from 
the mountains of Luzon. Moreover it is quite probable that the 
species from the latter locality referred to the Australian genus 
Xeromys ought really to be generically distinguished. It is among 
the Murtdce that evidences of Australian affinities are alone ex- 
hibited. As these murines have already been noticed on page 277 
it will be unnecessary to allude to them further in this place. 

Next to the peculiar rodents of the mountains, the most remark- 
able feature about the fauna of the typical Philippines is the 
absence of such a number of the most characteristic Malayan 
genera of mammals. Among the Primates the deficiency of orangs 
(Simia) is perhaps not very remarkable, but the total lack of 
langurs (Semnopitkecus) and the presence of only a single species 
of Macacus and Hylobates are most noteworthy. The macaque is, 
however, distributed over all the islands of the group, and differs 
from other forms of its species in its extremely light coloration, so 
that it is scarcely likely to have been introduced by human agency. 
Of Malayan genera which are absent, special note may be taken of 
Linsanga, Arctogale, Arctictis, Cynogale, Herpestes, the wild dogs of 
the sub-genus Cyon, Ursus, Tragulus 1 , and Elephas. Almost 
equally well-marked peculiarities are exhibited by the bird fauna. 

Apart from the tamarao, which has yet to be proved entitled 
to rank as a valid species, the lowland mammalian fauna of the 
Philippines is such as might have well reached the group by means 
of a narrow connection of limited duration with some portion of 
the Malay countries ; say, for instance, with Borneo by way of 
Palawan. That such a connection must have been comparatively 
recent is indicated by the identity of several of the Philippine 

1 Represented by T. nigricans in Palawan. 


species with Malayan forms and the absence of any peculiar 
genera save Phlceomys. The absence of such a number of 
Malayan types indicates, however, either that the connection must 
have been exceedingly brief, or that a large number of species 
formerly inhabiting the islands have been destroyed by sub- 
mergence. On the other hand, the presence in the group of a 
considerable proportion of widely spread continental genera of 
birds suggests a more free communication with China than at 
present exists ; such communication having not improbably taken 
place by way of Formosa. The mountain fauna of Luzon doubt- 
less indicates an earlier type of colonisation. 

Note. Since the foregoing was written two papers have appeared on the 
Fauna of the Natuna Islands; viz. O. Thomas, Novitates Zool. Vol. n. pp. 
2628, and Thomas and Hartert, I.e. pp. 409 429. 

2O 2 



Characteristics of the Mammalian Fauna Mammals of the Eastern Holarctic 
Region Plistocene Fauna of the Holarctic Region Geographical Changes 
since the Plistocene Western Division of the Region Arctic Sub -region 
European Sub-region Central Asian Sub-region Tibetan Sub-region 
Mantchurian Sub-region Mediterranean Sub-region Kashmir Ca- 
nadian Sub-region Transition Zone. 

BY far the most extensive of all the zoological regions of the 
globe is that which is equivalent to the whole of the Palsearctic 
and the greater portion of the Nearctic region of Messrs Sclater 
and Wallace, the one to which Dr Heilprin (after a suggestion 
of Professor A. Newton) applied the name Holarctic. In defining 
this region, Dr Heilprin separated from it a Sonoran "transitional 
region " in the Western Hemisphere, and a similar Mediterranean 
or Tyrrhenian tract in the Eastern. Of these the former is 
now accepted as a definite region ; but our knowledge of the 
distribution of species in the Eastern Hemisphere is either too 
imperfect, or the interdigitation of the two faunas is too complete 
to admit of the full definition of a Mediterranean region. Accord- 
ingly, without prejudice as to what it may be possible to ac- 
complish in this direction in the future, the Mediterranean area 
is provisionally included in the Holarctic region. It is, however, 
important to observe that the reservation by Dr Heilprin of the 
two transitional tracts already named justifies the use of the term 
Holarctic even if both such tracts be raised to the rank of separate 
regions ; and there is accordingly no necessity for the adoption of 
Dr Blanford's term Aquilonian as equivalent to the restricted 
Holarctic. Used in the sense here indicated, the Holarctic region 
will comprise all that portion of Arctogasa lying north of the 


Sonoran region in America, and of the Ethiopian and Oriental 
regions in the Old World. The whole area is extra-tropical ; and, 
as Dr Heilprin remarks, " no other region can compare with the 
Holarctic in the manifold variety of its physical characteristics. 
Every form of terrestrial configuration, or condition of soil and 
climate that may be represented in any other region, is also repre- 
sented here, and on an imposing scale. From the ice-bound fields 
of the far north to the burning desert-wastes of Turkestan on the 
south, and from the deep forest-grown lowlands to mountain- 
summits soaring thousands of feet above the level of perpetual 
snow, we pass through all those various gradations of climate 
which respectively characterise the Frigid, Temperate, and Torrid 
zones. Densely covered forest-tracts, supporting, as in the north, 
a sombre growth of pine and other coniferous trees, or, as in the 
south, a vegetation of almost tropical luxuriance, alternate with 
broadly open grass or pasture lands (tundras of Siberia, American 
prairies and plains), which in some cases support over enormous 
areas only a very scanty vegetation, and in others display a profuse 
variety of vegetable productions. It is in this region that, in 
addition to a most bountiful development of desert tracts, we meet 
with the most elevated table-land (the Central Asian), and, at the 
same time, with the greatest expanse of lowland on the surface of 
the globe, the great plain of Siberia and north-eastern Europe." 

Although the essential unity of the greater portion of the 
Nearctic and Palaearctic regions has long been fully recognised by 
the American zoologists, several attempts to bolster up the alleged 
distinctness of these artificial divisions have recently been made in 
England 1 , one proposal being to recognise an Arctic or Boreal 
circumpolar province cut off from both areas, although this is 
practically begging the whole question. 

If I rightly understand his view, Dr C. H. Merriam 2 , who is 
an ardent advocate for the zoological unity of the more northern 
portions of both hemispheres, would distinguish a Boreal circum- 
polar region common to both hemispheres ; while in America he 
recognises south of this a Transition region, followed still more to 
the south by the Sonoran. In the Old World he would have an 

1 Appendix, Nos. 28 and 35. 

2 Ibid. No. 19, pp. 24, 63. 


analogue of the Sonoran practically equivalent to the Mediter- 
ranean sub-region of European writers but says nothing about 
an analogue of the Transition; from which I infer that he 
would use the term Boreal really as practically equivalent to the 
Holarctic, if the Sonoran and Mediterranean areas were subtracted. 
In the New World the Boreal, exclusive of the Transitional, is 
divided into an Arctic and a Coniferous Forest Boreal zone, the 
latter being frequently spoken of as the Boreal " zone," in contra- 
distinction to the Boreal circumpolar "region"; the Arctic zone 
including the tract beyond the limit of trees. The distinction 
between the Boreal and Transition areas is certainly not of 
regional value ; and as the term Boreal is used in several senses, 
it had better give place to the earlier Holarctic. 

As has been partially indicated in earlier chapters, and as will 
be more fully noticed in the sequel, there is undoubtedly a marked 
distinction between the mammals of North America as a whole 
and those of Europe and northern Asia, but this has been con- 
siderably exaggerated by including the Sonoran region in the old 
Nearctic, and is overshadowed by the number of genera and 
species common to the two areas and unknown elsewhere. Could 
a Mediterranean region be satisfactorily denned, the homogeneity 
of the mammalian Holarctic fauna would be still more apparent ; 
but this, from the great mingling of northern and southern types 
which has taken place in the Old World, is, I think, impracticable, 
As has been already mentioned, it is probable that the western 
and eastern halves of the Holarctic region have never had more 
than a comparatively small area of communication by way of 
Bering Strait, and, therefore, the further south we travel in the 
two areas the more distinct do the faunas become; while only 
such forms as are capable of withstanding a certain degree of cold 
have ever been able to cross at all. It may be added that the 
evidence for the unity of the Holarctic region is by no means 
solely dependent upon the mammalian fauna, but is supported by 
many other groups of animals. To take an instance from the 
insects, I may quote from Mr W. F. Kirby l , who writes as follows : 
" Had I been dealing with Lepidoptera only, I would certainly 

1 Joitrn. Linn. Soc. Zool. 1873, p. 432. 


have united Dr Sclater's Palaearctic region and Nearctic region ; 
for although the species of North American Rhopalocera are 
seldom identical with those of northern Asia and Europe, still the 
genera are the same with scarcely an exception, except a few 
representatives of South American genera, which have no more 
right to be considered Nearctic species than the similar chance 
representatives of African 1 forms in North Africa or south-west 
Europe, or of Indian forms in south-east Europe, have to be con- 
sidered Palaearctic species." 

On the other hand, North America differs remarkably from the 
eastern half of the Holarctic region as regards its land molluscs. 
Thus the Rev. A. H. Cooke 2 writes, that " no district in the world 
of equal extent is so poor in genera, while those which occur are 
generally of small size, with scarcely anything remarkable either in 
colouring or form. The elongated land-shells (Clausilia, Bull- 
minus] so characteristic of Europe are entirely wanting, but a few 
Bulimulus, of Neotropical origin, penetrate Texas, and from the 
same sources come a few species of Glandina (as far north as 
South Carolina), Holospira (Texas), and Helicina" Probably this 
poverty is largely due to the unsuitableness of the greater part of 
North America to molluscan life ; aided by the circumstance that 
land-molluscs are just the creatures that would have been unable 
to pass over from Asia by way of Bering Strait. The batrachians, 
again, which differ most remarkably in their distribution from 
mammals, are not indicative of the unity of the Holarctic region, 
the American types being very different from those of the Eastern 

According, however, to Mr F. E. Beddard 3 , " the earth-worms 
offer the best evidence of any group in favour of the Holarctic 

Although during the Plistocene era even subsequently to the 
passing away of the glacial period elephants, rhino- 
ceroses, and hippopotami abounded over the greater tics ^he 
part of Europe, while species of the two former 
groups ranged as far north as Siberia, and macaques 

1 The author obviously means Ethiopian. 

2 Cambridge Natural History Mollusca, p. 339 (1895). 

3 Appendix, No. 5, p. 80. 


were found in France and England, yet the Holarctic region is 
now characterised by the absence of all these animals, save a few 
species of Macacus on its southern borders ; and if a Mediterranean 
region could be satisfactorily defined, even these, as well as hyaenas 
and certain other southern types, would likewise be excluded. 
The fruit-bats constituting the family Pteropodidce. are likewise 
practically wanting in this region; and Effodientia are quite 
unknown. On the other hand, carnivores, such as wolves, foxes, 
bears, martens, weasels, and the glutton, are abundant ; while the 
rodents are specially represented by such types as the marmots, 
beavers, voles, and picas ; and the ungulates comprise the bisons, 
nearly all the sheep, the true goats (absent in the western half of 
the region), and all the typical deer. 

Referring in more detail to the peculiar generic types common 
to the region as a whole, we have, first of all, among the Insecti- 
vora the typical or true shrews (Sorex] which belong to that 
section of the Soricidcz characterised by having the tips of the teeth 
stained brownish-red in the main characteristic of this region, 
although in America they extend southwards into the Sonoran. 
In the allied family of the Talpida, the mole-shrews ( Urotrichus], 
which are near relatives of the European desmans, are represented 
solely by one Japanese, and a second North American species, 
although the latter is frequently separated generically as Neuro- 

The peculiar genera of Carnivora are but few, although certain 
groups are either confined to, or very strongly represented in, the 
region. For instance, among the Felidce the true lynxes which 
although generally included in the genus Felts, are by some 
regarded as entitled to form a genus by themselves are absolutely 
confined to this region, where they range as far south as Spain. 
The bears (Ursus], also, are very strongly represented; brown- 
coloured species being peculiar to the Holarctic area, as is the 
very distinct polar species to its Arctic portions. The peculiar 
sea-otter, the sole representative of the genus Latax, has a distri- 
bution very similar to that of the mole-shrews ; this animal occur- 
ring on the coasts of Kamschatka and the Kurile Islands, as well 
as on those of the Aleutians. The wolverene (Gulo}, which is like- 
wise the only member of its genus, has a more extended range, 

IX.] RODENTS. 313 

being found throughout the forest regions of northern Europe, 
Asia, and North America, while in the Plistocene era it ranged as 
far south as England. Although marine mammals are for the 
most part omitted in this work, the walruses (Trichechidcz), which 
now have a circumpolar distinction, can scarcely be omitted, since 
these animals never wander far from land. Remains of the 
existing forms have been disinterred from the peat of the English 
fens, while tusks of fossil species have been discovered in the 
Pliocene Crag of the east of England, and also in the corre- 
sponding deposits of Belgium. 

Passing on to the rodents, we have in the Sciuridce the ground- 
squirrels or chipmunks ( Tamias] practically confined to the region. 
Although in America they also extend into the Sonoran, in Europe 
they are unknown in the Mediterranean area. The pouches in 
the cheeks for storing food and the alternate dark and light longi- 
tudinal bands down the back serve to distinguish the chipmunks 
from other squirrels. Fossil remains of the genus, probably 
belonging to existing species, occur in the Plistocene of Europe 
and North America. The family of the beavers (Castoridcz) seems 
always to have been mainly confined to the Holarctic region, one 
of the two existing species ( Castor fiber] being European, and 
dating from the English Plistocene, while the other (C. canadensis] 
is North American. Whereas, however, the latter ranges south- 
wards into the Sonoran region, the former is unknown in the 
Mediterranean sub-region. Fossil species of this genus occur in 
the upper Tertiaries of Europe and North America, where the 
extinct Chalicomys is likewise met with ; but no beavers are known 
from either the Siwaliks or the Pikermi beds. Although there are 
few generic types of Muridce peculiar to the whole region, such as 
there are are important. Foremost of these are the great tribe of 
the voles (Microtus 1 }, constituting the typical representatives of a 
sub-family nearly allied to the cricetines, but distinguished by the 
two longitudinal rows of tubercles on the crowns of the molar teeth 
being modified into alternating triangular prisms, and likewise by 
these teeth being generally of the hypsodont type. In the Old 
World they are found from the Arctic zone to Asia Minor, while 

1 Commonly known by the later name of Arvicola. 


in America they enter the Sonoran region. As fossils, they appear 
to be first known from the upper Pliocene Crag of England ; and 
were thus probably evolved within the Holarctic region from the 
more generalised cricetines at a comparatively late epoch. Nearly 
allied are the lemmings (My odes), which are, however, a more 
northern type, unknown in the Mediterranean sub-region, and 
likewise in the Sonoran. Still more northern, and indeed circum- 
polar in its range, is the banded lemming, which alone represents 
a genus (Cuniculus) distinguished from the last by the absence of 
external ears, the short and thick fur on the feet, the rudimentary 
first toe of the fore feet, and the elongation of the two middle 
claws of the same. The second of the two families peculiar 
to the region is that of the picas, or tailless hares (Lagomyidce), 
comprising small hare -like animals with short ears, of which 
all the living forms are included in the single genus Lagomys. 
While the majority of the picas are confined to the highlands 
of Central Asia, some are found on the first snowy range of the 
Himalaya, and both south-east Europe and the Rocky Mountains 
severally possess a representative. Fossil forms are common in 
the middle and upper Tertiaries of Europe, as far south as Sardinia. 
Although the hare-tribe (Leporidcz) have an almost cosmopolitan 
distribution, the majority of species of Lepus are inhabitants of the 
Holarctic region, Central Asia being especially rich in representa- 
tives of the genus. 

Among the Bovidtz, the bisons, which form a well-marked 
group of the genus Bos, may be regarded as now characteristic of 
the region, although the American Bos americanus ranges into the 
Sonoran. In addition to the European B. bison, which was for- 
merly spread over the greater part of Europe and during the 
Plistocene extended into Arctic America, there is also the some- 
what aberrant yak (B. grunniens) of Tibet, In the Plistocene the 
range of the group was somewhat more extensive, remains of an 
extinct species having been found in deposits of that age in 
Texas ; and there is likewise another from the Pliocene of northern 
India. The sheep constitute a group mainly characteristic of the 
Holarctic region ; their headquarters being the Central Asian 
plateau, where they are more numerous than anywhere else in 
the world, although one species (Ovis vignei) impinges on the 


north-western frontier of the Oriental region, while the single 
North American form also enters the Sonoran. In a fossil state 
it is possible that sheep occur in the Indian Siwaliks, but else- 
where they are first known from the Forest-bed of the Norfolk 
coast, belonging to the early part of the Plistocene epoch. More 
nearly allied to the sheep than to the goats, the musk-ox (Ovibos) 
of Arctic America is now extinct in the Old World, but as it is 
abundant in the Plistocene of Europe and Asia, where it ranged as 
far south as England, it is surely entitled to rank among the forms 
common to the western and eastern halves of the Holarctic region. 
Nearly allied extinct species occur in the Plistocene formations of 
the United States. Among the Cervidce. there are three types 
common to the entire region. The first, or Elaphine group, 
includes the typical members of the genus Cervus, as represented 
by the red deer (C. elaphus) of the Old World and the wapiti (C. 
canadensis) of North America; this group being characterised, 
among other features, by the general presence of both a brow- 
and a bez-tine to the antlers, although the latter is wanting in 
the North African variety of the red deer, and also in the Tibetan 
C. thoroldi. The alliance between the wapiti and some of the 
forms inhabiting Central Asia is so close as to render it doubtful 
whether they are really anything more than varieties of a single 
species. The single species of elk (Alces) is common to both 
halves of the region ; and the same is also the case with the 
reindeer (jRangtfer), which although now not found to the south 
of Europe, ranged during the Plistocene era into the south of 
France. The genera or groups which may be regarded as charac- 
teristic of the entire Holarctic region may be tabulated as follows, 
those which are practically peculiar being printed in italics : 


SORICID^E. Sorex. In America ranges into Sonoran. 
TALPID^E. Urotrichus. Japan and N. America. 


FELID^E. Felis, the true lynxes solely Holarctic. 

MUSTELID.E. Latax. \ One species common to both 

Gulo. J hemispheres. 
TRICHECHW&. Trichechus. 




Tamtas. Ranges into Sonoran. 



CASTORID^E. Castor. *) .. 

,, ,,-. \ Also Sonoran. 

MURID^E. Microtus. } 



LA COM YIDSE. Lagomys. 

LEPORID^:. Lepus, the greater number of species Hol- 


Bos, the bison group chiefly Holarctic, al- 

though the American species reaches 

the Sonoran. 
Ovis, just touches Oriental, and also reaches 

Ovibos, now extinct in the Old World, where 

it was common in the Plistocene. 

The arguments for the unity of the Holarctic region are, how- 
ever, by no means confined to the case of genera, for there are a 
number of species which either have a circumpolar range, or 
which are represented by closely allied forms in the opposite 
hemisphere. It is true, indeed, that many of these are now more 
or less exclusively Arctic in their distribution, but some range a 
long distance to the south ; while during the Plistocene epoch this 
was the case with the majority. The following list includes the 
more important species which are either common to the eastern 
and western halves of the region or have representative forms 
in the two hemispheres ; those which are strictly Arctic having 
the letter P appended : 

1. Common lynx (Felis lynx). Canadian lynx (F. cana- 

densis). P. 

2. Wolf (Cants lupus). 

3. Fox (Cants vulpes). 

4. Arctic fox (Cants lagopus). P. 

5. Brown bear ( Ursus arctus). 


6. Polar bear ( Ursus maritimus). P. 

7. Sea-otter (Latax lutris). 

8. Pine-marten (Mustela martes). American marten (M. 


9. Weasel (Mustela vulgaris). 

10. Wolverene (Gulo luscus). 

11. Walrus (Trichechus rosmarus). 

12. Arctic vole (Microtus rutilus). P. 

13. Common lemming (Myodes lemmus). American lemming 

(M. obensis). 

14. Banded lemming (Cuniculus torquatus). P. 

15. European beaver (Castor fiber}. American beaver (C. 


1 6. Mountain hare (Lepus timidus^}. 

17. Bison (Bos bison). American bison (B. americanus). 

1 8. Kamschatkan sheep (Ovis nivicola). Bighorn (O. cana- 


19. Musk-ox (Ovibos moschatus). 

20. Central Asian deer (Cervus eustephanus). Wapiti (C. 


2 1 . Elk (Alces machlis}. 

22. Reindeer (Rangifer tarandus}. 

If the Plistocene be taken into consideration there may be 
added the mammoth (Elephas primigenius) and the horse (Equus 
caballus), remains of both of which have been discovered in 
Eschscholtz Bay, where those of the European bison also occur. 

In this list it may be noticed that the North American lynx is 
so closely allied to the European form that it has been a question 
whether it is really more than a local variety. Although the 
American wolf has been separated as a distinct species, it is now 
generally identified with the European form ; the same being also 
the case with the so-called cross-fox of North America, which, 
together with another form from the Himalaya, and a third from 
North Africa, may be considered a mere variety of the ordinary 

1 This name is commonly applied to the English hare, although it properly 
belongs to the more northern species. The so-called Polar hare (Z. glacialis) 
of Arctic America appears to be only a variety. 


fox. Much discussion has taken place with regard to whether any 
of the bears of North America are distinct from the common brown 
bear ( Ursus arctus), with its many Asiatic varieties. According, 
however, to one of the latest memoirs on this subject 1 , all these 
forms appear mere varieties of the latter, the so-called cinnamon 
and black bears presenting more distinctly marked differences 
than does the grizzly. The American marten is so nearly related 
to the European marten and the Asiatic sable that it is almost 
impossible to point out valid characters by which the three forms 
can be specifically distinguished. In the case of the walrus, the 
Pacific form is distinguished by certain external features from the 
one inhabiting the Atlantic coasts, although these are scarcely of 
sufficient importance to be ranked as specific. In regard to the 
Arctic vole, it may be mentioned that although it is typically a 
polar form, yet it is represented in southern Europe by the bank- 
vole (Microtus glareolus) and in the United States by Gapper's 
vole (M. gapperi\ both of which may be regarded as southern 
climatic offshoots from the northern stock. Among the other 
species of rodents it will suffice to mention that the European 
and American beavers are merely distinguished from each other 
by the relative lengths of the nasal bones of the skull. And it 
may be added that the Kamschatkan wild sheep is so closely 
related to one race of the bighorn, or Rocky Mountain sheep, that 
it is very questionable whether the two are really entitled to 
specific distinction. The same is also the case with the two deer 
mentioned in the list. Reference has already been made to the 
circumstance that the musk-ox is extinct in the eastern division 
of the region. A few of the American forms, such as the bear, 
beaver, bison, and bighorn, enter the limits of the Sonoran region. 

Although, as will be shown immediately, there are a large 
number of generic types respectively confined to its eastern and 
western divisions, the lists given above, especially the one relating 
to the species, are amply sufficient to demonstrate the essential 
unity of the Holarctic region. None of the other zoological 
regions have anything like the number of common or representa- 

1 A. E. Brown, Proc. Ac. Philad. 1894, pp. 119, 129. Merriam, P. BioL 
Soc. Washington, vol. x. pp. 65 83 (1896), regards the N. American bears as 
forming several distinct species. 


tive species which characterise the two divisions of the present 
one. It must, moreover, be remembered that in the case of the 
other regions we have taken the whole of the peculiar generic 
types into consideration, although many of them are confined to 
small portions of such regions, whereas in the present instance 
only those which range over nearly the whole area have been 
mentioned. If, for instance, we were to take the genera respec- 
tively confined to the Indian and Malayan areas of the Oriental 
region, it would be found that the distinction between the two 
areas would be nearly or quite as marked as are the two great 
divisions of the Holarctic, while in the matter of species the 
differences between the two would be far greater. There would 
accordingly be stronger grounds for making an Indian and a 
Malayan region than there are for the separation of a Palsearctic 
and a Nearctic. 

Having now discussed the leading mammalian types character- 
istic of the Holarctic region as a whole, it remains 

to notice those confined to its eastern division, t f 

after which such as are restricted to its western half Holarctic 
will be taken into consideration. And here it is 
advisable to repeat that since the communication between eastern 
Asia and North America by way of Bering Strait appears always 
to have been of very limited extent, as we proceed south in the 
two great continental areas the difference between their faunas 
appears more and more strongly marked. Accordingly, if it were 
possible definitely to establish a Mediterranean region, the dis- 
tinction between the faunas of the eastern and western divisions 
of the Holarctic would be much less than is the case under the 
arrangement here followed. 

Passing by the bats entirely, no notice will be taken of groups 
which, like the hedgehogs (Erinaceus), are spread over several of 
the regions of Eastern Arctogaea, since these are not in any way 
characteristic of the eastern division of the Holarctic region 1 . 
The first mammal that presents itself for notice is accordingly the 
water-shrew, which is the sole representative of the genus Crosso- 
pus, and belongs to that division of the Soricidce in which the 

1 Such types have been discussed when considering the fauna of Eastern 
Arctogaea, supra, p. 181. 


teeth are stained red ; the especial characteristic of the genus being 
the thickly-fringed feet and tail. The water-shrew is a typical 
Holarctic mammal, ranging as far east as the Altai mountains and 
unknown in the Mediterranean sub-region. In the second division 
of the same family, or that in which the teeth are white, there is a 
peculiar shrew from the Kirghiz steppes nearly allied to the 

FIG. 66. RUSSIAN DESMAN (Myogale moschata}. 

widely-spread musk-shrews (Croridura), but constituting a genus 
(Diplomesodon) by itself. To the same sub-family belongs the 
Tibetan water-shrew (Nectogale], which is likewise the solitary 
representative of its genus, and is closely allied to Chimarrogale, 
which has, as elsewhere stated, one Oriental and one Japanese 
species. Both these types are accordingly closely connected with 
the Oriental region, although nothing is known of their past 
history. Far more characteristic of the eastern half of the region 


under consideration are the two species of desman (Myogale), 
which are aquatic insectivores, with long trunk-like snouts, some- 
what intermediate between the shrews and the moles. Of the two 
living species, the smaller is confined to the region of the Pyrenees, 
extending as far north as the Department of the Landes, while 
the other is now restricted to south-eastern Russia, although its 
fossilised remains have been discovered in the Plistocene Forest- 
bed of the east of England. Extinct species occur in the Mio- 
cene and upper Oligocene of the Continent. A slate-coloured 
insectivore, with the external form of a shrew but the skull of a 
mole, inhabiting Eastern Tibet, constitutes the genus Uropsilus ; 
and the more mole-like creature known as Scaptonyx is likewise 
from the same locality. In any case, these two animals only just 
enter the border of the region, so that they cannot be regarded as 
characteristic Holarctic types ; and, indeed, the Moupin district 
of Eastern Tibet is included by Dr Wallace in the Oriental region, 
although it is assigned by others to the Holarctic. Although two 
species occur to the south of the Himalaya, the true moles (Talpa) 
are very characteristic of the eastern Holarctic region, the common 
species ranging from England to Japan, and dating from the epoch 
of the Norfolk Forest-bed; while fossil species, some of which 
have been separated as Protalpa, range through the Tertiaries of 
Europe to the epoch of the upper Oligocene. By some writers 
Talpa moschata of Eastern Tibet is distinguished as Scaptochirns. 

Among the Carnivora the widely spread Old World genera 
Genetta, Herpestes, and Hyczna enter the Mediterranean sub-region, 
but are unknown elsewhere in the Holarctic area at the present 
day. The Ursidce include a most remarkable generic type known 
as ALluropus, represented by the parti-coloured bear of Tibet, in 
which the cheek-teeth present a decided approximation to the 
extinct Hytznarctus and also resemble those of the panda 
(jElurus). This genus is another of the border-forms between the 
Holarctic and Oriental regions. On the other hand, the badgers 
(Meles) are very characteristic of the area under consideration, 
ranging from England through the rest of Europe to Japan and 
China, where one species enters the Oriental region, being found 
as far south as Hongkong. Remains of extinct badgers occur 
in the lower Pliocene of Persia. 

L. 21 


The list of more or less peculiar generic types among the 
rodents is relatively large. Foremost among these stands a very 
large flying-squirrel {Eupetaurus}, inhabiting the regions north of 
Kashmir and Tibet, and distinguished from all other members 
of the family to which it belongs by its tall-crowned (Jiypsodonf] 
cheek-teeth. In the dormouse family (Myoxida) the squirrel- 
tailed species, which is the sole representative of the genus 
~Myoxus in its restricted sense, and the common dormouse, alone 
constituting Muscardinus, are exclusively European ; fossil forms 
occurring through the upper and middle Tertiaries. 

In the Muridce. the hamsters (Crtcetus), if regarded as generi- 
cally distinct from their allies the white-footed mice (Sitomys] of the 
New World, are absolutely characteristic of the eastern division of 
the Holarctic region, where they range over a large portion of 
Europe and northern Asia. Extinct species are abundant in the 
European Tertiaries. Two genera of the mole-like rodents, 
having the dentition of voles, but approximating in the form of 
their body and limbs to the moles, constitute a sub-family which is 
also restricted to this area. Of these mole-voles, the first genus 
(Ellobius] is represented by one species from Russia and another 
from Afghanistan, while the second (Siphneus) includes several 
forms from North and Central Asia; fossil species of the latter 
having been described from the Plistocene of the Altai and the 
Pliocene of northern China. The great mole-rat (Spalax typhlus) 
of southern Europe, Persia, Mesopotamia, Syria and Egypt, repre- 
sents the only genus among the Spalacidce. which is confined to 
the present area. The Dipodidce, on the other hand, contain 
several characteristic eastern Holarctic generic types. The most 
aberrant of these are the rat-like animals constituting the genus 
Sminthus, of which one species inhabits eastern and northern 
Europe and Central Asia, while a second is found in Kashmir, 
and a third in the Kansu district of China. Of the more typical 
forms, the true jerboas (Dipus), characterised by having only three 
toes, are exclusively Holarctic, ranging from Egypt into Central 
Asia, where they always frequent desert districts. Of the genera 
with five toes, Euchoretes is represented by a single long-snouted 
and long-eared species from the neighbourhood of Yarkund ; and 
Platycer corny s, which differs by its flattened and lancet-shaped 




tail, includes several species, extending from Siberia to Nubia, 
so that the genus just enters the Ethiopian region. It is repre- 
sented in the Plistocene of northern Asia. Lastly, there is the 
genus Alactaga, of which there are several species, mostly inhabit- 
ants of Northern and Central Asia. The typical A. jaculus 
extends, however, into Persia and southern Russia, while in the 
Plistocene it ranged as far west as Germany ; and another species 
is an inhabitant of Afghanistan. Although mainly an Ethiopian 

FIG. 67. HEAD OF SPANISH IBEX (Capra pyrenaica). 

and Neotropical family, the Octodontidce. have an Holarctic repre- 
sentative in the gundi (Ctenodactylus\ which inhabits the borders 
of the Sahara in the neighbourhood of Tripoli. The extinct 
Pellegrinia, of the Sardinian Pliocene, also belongs to the same 

21 2 


Among the artiodactyle ungulates there are six genera, either 
entirely or mainly confined to the eastern Holarctic region. Of 
these, the true goats that is to say, the members of the genus 
Capra, as distinct from the Oriental and Arabian Hemitragus are 
almost exclusively Holarctic, although C. walie inhabits the 
Abyssinian highlands, and C. siniatica, of Palestine and upper 
Egypt, may also enter the confines of the Ethiopian region. All 
the goats, it may be observed, are essentially mountain-dwelling 
animals, and the occurrence of the same species of ibex (C. 
sibiricd) in both the Altai and Himalaya is a clear proof of the 
former prevalence of colder conditions, as without these the ani- 
mal could not have passed from the one range to the other. The 
sheep also in spite of the existence of outlying North American 
species, and a variety of one Central Asian form (Ovis vignei] 
which enters the north-western confines of India are mainly cha- 
racteristic of the area under consideration, attaining their great- 
est numerical development, and also their maximum size, in the 
highlands of Central Asia. The range of the genus includes almost 
the whole of the eastern Holarctic region, the mouflon (O. musimon) 
inhabiting the Corsican islands, and the aberrant arui (O. tragela- 
phus] being found in northern Africa. It is possible that fossil 
sheep occur in the Indian Siwaliks (where remains of a goat allied 
to the Himalayan markhor are also met with), and a large species 
is definitely known from the Norfolk Forest-bed. The next on 
our list is the remarkable goat-like antelope from the hills to the 
north of the Assam known as the takin (Budorcas], which is allied 
to the Oriental Nemorhczdus, and is therefore probably an immi- 
grant into the region from the southward. Allied to this group is 
the chamois, or gemse (Rupicapra), now confined to the higher 
mountain-ranges of Europe from the Pyrenees to the Caucasus, 
but which during the Plistocene epoch ranged over many of the 
lowlands. Among the true antelopes, the addax (Addax], an ally 
of the oryx group, is an exclusively Mediterranean type, inhabiting 
North Africa and Syria, where there are also representatives of 
other genera which are typically Ethiopian. More thoroughly 
Holarctic are the two peculiar but allied genera Saiga and Panthol- 
ops, each represented by a single living species. The saiga is now 
confined to the steppes of western Asia and Eastern Europe, but 




during the Plistocene epoch extended as far westwards as Germany, 
France, and England. The chiru, as the representative of the 
second genus is called, is, on the other hand, an exclusively 
Tibetan form ; and it is believed that a fossil species occurs in the 
later Tertiary formations of the same area, where, curiously enough, 
a rhinoceros also existed. Although gazelles (Gazella) have repre- 
sentatives in both the Oriental and Ethiopian regions, they are 
mainly characteristic of the desert districts of the eastern Holarctic 
region, being especially numerous in North Africa, Syria, and parts 

FIG. 68. MUSK-DEER (Moschus moschiferus). 

of Central Asia. An inhabitant of the cooler regions of Asia, 
where it extends from the south of Siberia to Kashmir and Cochin 
China, the musk-deer (Moschus) represents a peculiar sub-family 
of the Cervidoe, confined to the region under consideration. A 
second species has been described from Kansu, in north-western 
China, and it is not improbable that the genus is also represented 
in the Indian Siwaliks. Although agreeing with the musk-deer in 
the absence of antlers and the presence of long tusks in the upper 


jaw of the males, the Chinese water-deer (Hydropotes), from the 
valley of the Yang-tsi-kiang, belongs to the more typical Cervidce. 
Another genus (Elaphodus) more nearly allied to the muntjacs is 
also Asiatic, being represented by one species from near Ningpo, 
in China, and by a second from Moupin, in eastern Tibet. Of 
the true deer (Cervus) there are two groups confined to the area 
under consideration. Firstly, there is the elaphurine group, repre- 
sented solely by the aberrant David's deer (C. davidianus\ of 
northern China; and, secondly, we have the damine group, of 
which the fallow deer (C. dama) of the Mediterranean countries 
and the Persian fallow deer (C. mesopotamicus] are the living 
forms. Allied types occur in the East Anglian Forest-bed, and 
the gigantic extinct Irish deer (C. gigantens] must likewise be 
included in the group, all the members of which have the antlers 
more or less palmated. 

As regards the camels (Camelus), there is some difficulty in 
arriving at a satisfactory conclusion, since although a feral race of 
the Asiatic C. bactrianus is met with in the deserts bordering Kash- 
gar, it is now pretty well ascertained that really wild camels exist 
nowhere in the world. Still, however, as fossil species of the 
genus are met with in the Pliocene of the Siwalik Hills (on the 
borders of the Oriental and Holarctic regions) and in the Plisto- 
cene of Algeria, it is probable that the group is of Holarctic origin 1 . 

The foregoing survey may be summarised as follows : the 
names of such genera or groups as are mainly or exclusively con- 
fined to the eastern division of the Holarctic region being printed 
in italic type. 

Insect! vora. 

SORICID^:. Crossopus. 

Diplomesodon. C. Asia. 
Nectogale. Tibet. 

TALPID/E. Uropsilus. \ _.. 

_, -V \ E. Tibet. 

Scaptonyx. ) 

Talpa. Also enters Oriental. 
URSID^E. sEluropus. Tibet. 

MUSTELID^E. Meles. Enters E. Oriental. 

1 See page 281. 










Eupetaurus. Tibetan. 





Sip /incus. 




Euchoretes. Central Asian. 

Platycer corny s. Enters Ethiopia. 


Ctenodactylus. North African. 


Capra. An outlying Ethiopian species. 
Ovis. One N. American species, and one 

from Central Asia entering Oriental. 
Budorcas. Tibetan. 
Rupicapra. European. 
Addax. Mediterranean. 
Saiga. Central Asian. 
Pantholops. Tibetan. 
Gazella. A large proportion of the species 

E. Holarctic. 
Cervus. The Elaphurine and Damine 

groups exclusively E. Holarctic; the 

former Central Asian, and the latter 


Elaphodus. E. Tibet and China. 
Hydropotes. Chinese. 
Moschus. Asiatic. 
(?) Camelus. 

With the possible exception of the Camelidce, none of the 
families in the foregoing list are peculiar to the area in question 
a feature presenting a marked contrast to the lists of the character- 


istic mammalian genera of the Ethiopian and Oriental regions 
given above. The total number of genera which can in any sense 
be considered as peculiar to the eastern half of the Holarctic 
region does not exceed thirty ; and among these Uropsilus, Scapt- 
onyx, Elaphodus, and Hydropotes can only be regarded as intruders 
from the Oriental region ; while Ctenodactylus and Addax are 
manifestly Ethiopian types. Indeed, if a Mediterranean region 
were established, the whole of these, and probably also the true 
Tibetan forms, would have to be removed from the lists. Apart 
from the absence of peculiar families, the list bears no comparison 
as regards numbers with that of the mammalian genera distinctive 
of the Ethiopian region ; while, with the aforesaid deductions, it 
is also considerably inferior to that of the Oriental region. All 
this confirms the conclusions already drawn as to the inad- 
visability of regarding the area under consideration as a separate 
zoological region. 

The Pliocene and earlier Tertiary faunas of this area having 
already been considered in connection with Eastern 
Arctogsea in general in an earlier chapter, we may 

Eastern Hoi- p ass O n to a brief review of the Plistocene mammals 

arctic Region. 

of the eastern division of the Holarctic region, pre- 
paratory to the consideration of the sub-regions into which the 
latter is divided. The Plistocene period may be taken in England 
to commence with the Forest-bed of the Norfolk coast, which 
overlies the topmost of the Pliocene Crag series, and is itself 
overlain by the glacial deposits. To a later epoch of the same 
period belong the brick-earths and gravels of our river valleys, as 
well as the cavern-deposits ; many of these being either of post- 
or inter-glacial age. 

During the Plistocene period two very remarkable differences 
from the existing state of things have to be noticed. In the first 
place, the eastern Holarctic region was at that time very much 
less distinctly differentiated from either the Ethiopian or the 
Oriental than is the case at the present day ; macaques, hyaenas, 
the lion, rhinoceroses, hippopotami, and elephants abounding in 
Europe even as far north as England. The second point is the 
curious mixture of remains of existing species of mammals respec- 
tively characteristic of hot and cold climates met with in many 


parts of England and France. For instance, the glutton, rein- 
deer, Arctic fox, and musk-ox are animals whose presence seems 
indicative of a more or less decidedly Arctic climate ; many of the 
voles, picas, jerboas, and susliks, together with the saiga antelope, 
appear to point with equal force to the prevalence of steppe-like 
conditions ; while the hippopotamus and spotted hyaena seem as 
strongly in favour of a subtropical state of things. Nevertheless, 
remains of several of these groups have been found in such close 
association as to leave no doubt that the animals lived and died 
hard by where they are now buried. Much evidence on this point 
has been collected by Sir H. H. Howorth 1 , who writes as follows : 
" Cuvier, whose prejudices were the other way, was long ago con- 
strained to write of the remains of reindeer found with those of 
the mammoth and rhinoceros in the cave at Breugue : ' II ne faut 
pas douter qu'il [the rhinoceros] n'ait dte enseveli avec lui [the 
reindeer] a Breugue. Ses os y e'taient pele-mele avec ceux de ce 
grand quadrupede, enveloppes dans la meme terre rouge, et 
revetus en partie de la meme stalactite.' In the high-level 
gravels of the Thames valley the mammoth and woolly rhinoceros 
occurred with the Elephas antiquus ^ while in the low-level gravels 
the Rhinoceros leptorhinus and the hippopotamus occurred with 
the bison and the musk-ox 2 , together with a worked flint. 
The lemming and the reindeer occurred with the lion and 
hyaena at Bleadon, the lemming with the lion and hyaena 
at Wookey-Hole. The reindeer and the grizzly bear were 
associated with the hippopotamus at Cefn, and the E. antiquus 
with the mammoth at Durdham Down. The hippopotamus and 
the E. antiquus have been found with the reindeer and bison in 
Kirk dale Cave ; the hippopotamus with the wild boar, the rein- 
deer, the mammoth, and the E. antiquus at Brentford. Lartet 
says that in France remains of the hippopotamus have been found 
in one cave, that of Arcy, in which the reindeer has also occurred, 
accompanied by a worked flint. At St Acheul and in the Somme 
valley the same two animals have occurred together, and also at 
Levallos, in the Seine valley. At Viry Noureuil, near Chauny 

1 The Mammoth and the Flood (London, 1887), pp. 114, 115. 

2 The author uses the term musk-sheep for this animal ; a few other verbal 
alterations have been made in the quotation. 


(Aisne), the mammoth and Rhinoceros antiquitatis have occurred 
with the hippopotamus, the reindeer, and the musk-ox. At 
Bicetre, close to Paris, the lion is associated with northern voles, 
a marmot, a lizard, and a snake. At Montmorency the mole and 
the hedgehog have occurred with the hamster, the suslik, and the 
pica. In Auvergne, M. Pomel has found an elephant and the 
woolly rhinoceros with a cat, a suslik, and a hamster, together with 
snakes, lizards, frogs, and with shells such as are still found in the 
district. In Germany it is the same. At Westeregeln the lion 
and the spotted hyaena, the mammoth and rhinoceros were found 
with the marmot, the suslik, the lemming, the pica, and the rein- 
deer. At Thiede, the mammoth, woolly rhinoceros, the horse, the 
ox, the reindeer, the Arctic fox, the lemming, and the pica; and 
so we might continue throughout the majority of the German 

Many attempts have been made to reconcile these apparently 
contradictory facts ; one of the older views being that while the 
tropical types of mammals lived during warm interludes, they 
migrated southwards with the incoming of colder conditions to 
give place to the more Arctic fauna. The associations mentioned 
above render it, however, perfectly certain that such an explanation 
is not the right one. On the other hand, it must be remembered 
that there is yet much to be learnt about the effects of climate on 
mammals ; and the mammalian fauna of the Tibetan plateau shews 
that many types of animals formerly regarded as more or less 
essentially tropical or sub-tropical are capable of withstanding a 
winter climate of extreme severity. Thus in parts of Tibet, as 
well as in Kashmir, langurs and macaques may be seen leaping 
among the snow-clad branches of pines. Still it is perhaps diffi- 
cult to understand how two such animals as the hippopotamus 
and the reindeer could have inhabited the same locality contem- 
poraneously 1 . 

In spite of this association of Arctic and sub-tropical forms, 
there appears, however, to be evidence of a northern and southern 

1 The present writer is not prepared to accept the view of Mr A. H. Keane 
(Mthnology, Cambridge Geographical Series, p. 65, 1896) that the reindeer has 
only recently become adapted to a northern habitat. Among other circum- 
stances, its remains are unknown from the Forest-bed. 


type of Plistocene fauna, England being apparently somewhere 
near the border-line where the two met, and, at times at least, 
overlapped each other. Probably all or nearly all of the living 
European mammals were in existence at the same time ; but most 
of these need not be referred to here, attention being concentrated 
on those which are either extinct, or are now inhabitants of other 
regions or districts. The following list includes the more im- 
portant of these forms ; those which are decidedly northern types 
being indicated by a *, and those which appear essentially southern 
with a f. The scientific names of extinct species and genera are 
printed in italic, and of those still living in ordinary type. 


t Barbary Ape. Macacus inuus. 
f ,, ,, pliocenus. England. 

f ,, suevicus. Switzerland. 

f ,, ,, tolosanus. France. 


fLion. Felis leo. 
f Kafir Cat. Felis caffra. 
t Sabre-tooth. Machcerodus latidens. 
f Spotted Hyaena. Hyaena crocuta. 
t Striped Hyaena. ,, striata. 

Cave Bear. Ursus spelceus. 

* Arctic Fox. Canis lagopus. 
European Wild Dog. ,, (Cyon) europ&us. 

Hunting Dog. Lycaon anglicus. England. 

* Wolverene. Gulo luscus. 


t Maltese Squirrel. Leithia melitensis\ 

* Giant Beaver. Trogontherium cuvieri, 

* Northern Vole. Microtus rutilus. 
Sardinian Pica. Lagomys sardus. 

1 Originally described as a gigantic dormouse, but shown by the writer in a 
communication to the Proceedings of the Zoological Society, 1895, p. 860, to be 
allied to the Sciurida. 



Aurochs. Bos taurus, var. primigenius. 
*Musk-Ox. Ovibos moschatus. 
t Barbary Sheep. Ovis tragelaphus. 
f Spanish Ibex. Capra pyrenaica. 

English Gazelle. Gazella anglica. 
t Saiga Antelope. Saiga tartarica. 

Irish Deer. Cervus giganttus. 

f Hippopotamus. Hippopotamus amphibius. 

fPentland's Hippopotamus. ,, pentlandi. 

* Woolly Rhinoceros. Rhinoceros antiquitatis. 
Megarhine ,, ,, megarhinus. 

fLeptorhine ,, ,, leptorhinus. 

t Etruscan ,, ,, etruscus. 

* Elasmothere. Elastnotherium sibiricum. 

* Mammoth. Elephas primigenius. 
t Straight-tusked Elephant. antiquus. 

t Southern Elephant. meridionalis. 

( ,, melitensis. \ Maltese 

T Dwarf Elephants. ._ . . \ , . , 

( mnaidnensts. I Islands. 

t African Elephant. ,, africanus. 

In this list the Barbary ape is now confined to North Africa 
and Gibraltar. The lion, although now restricted to Africa, India, 
Persia, and Mesopotamia, ranged during the historic period into 
Thessaly ; while the Kafir cat is solely African. The sabre- 
toothed tiger of the caves was the last survivor of a genus common 
in the Pliocene, which in the Plistocene is unknown further north 
than Cromer. The spotted hyaena, whose remains are so abundant 
in the English caves, is, as we have seen in an earlier chapter, now 
restricted to southern Africa ; while the striped species, which 
dates from the upper Pliocene, now ranges from north Africa to 
India. The cave-bear, a gigantic extinct species, was distinguished 
from the brown bear by the more complex structure of its molar 
teeth. Of the Canidce, the European wild dog has its nearest living 
ally in the Altai, the other species of the group being Oriental ; 
while the European hunting-dog, which is known only by a single 
jaw from the Glamorganshire caves, appears to have been closely 


related to the living Cape species. Among the rodents, it is 
only necessary to mention that the giant beaver (Trogontheriuni) 
represents a distinct genus ranging from the Norfolk Forest-bed to 
Siberia; and also that the Maltese squirrel (Leithia) was restricted 
to the islands from which it takes its name. In the ungulates, 
the aurochs l was the gigantic ancestor of the domestic cattle of 
the present day, but is unknown living in a wild state. The arui, 
or Barbary sheep, is now restricted to north Africa; while the 
Spanish ibex is confined to the mountains of the Iberian penin- 
sula, its fossil remains occurring in the Gibraltar caves. The Irish 
deer, distinguished by its great size and widely-spreading antlers, 
was an ally of the fallow-deer, with which it is connected by 
means of another extinct species or variety (C. ntffi); and it may 
be mentioned that there are species of extinct deer from the 
Forest-bed, which it is unnecessary to name in this place. The 
latter deposits are the source of the known remains of the English 
gazelle. The common hippopotamus, which dates from the upper 
Pliocene of Italy, is now exclusively confined to Ethiopian Africa, 
but in the Plistocene is known to have wandered as far north as 
Yorkshire. Pentland's hippopotamus is a smaller species from 
Italy and the Mediterranean islands, where there may be a second 
still smaller form. Of the rhinoceroses, R. antiqiiitatis, which is 
exclusively Plistocene, ranged from Central Europe to Siberia ; its 
remains being dredged abundantly, in common with those of the 
mammoth, from the Dogger Bank, in the North Sea. The relation- 
ship of this species to the extinct Indian Rhinoceros platyrhinus 
and the living African R. simus has been alluded to in a previous 
chapter. The other three European species of the genus, which, 
like the last, were two-horned and devoid of front teeth, date from 
the Pliocene, and form a group differing remarkably in dental 
characters from R. antiquitatis. While two of these species were 
southern types, the third accompanied the mammoth and woolly 
rhinoceros in Siberia. A near ally of the rhinoceroses, the huge 
Siberian Elasmotherium, differed remarkably in the structure of its 
cheek-teeth, which are tall-crowned, and shew some approximation 
to those of the horses. 

1 The European bison is frequently miscalled the aurochs. 


From a distributional point of view, the European Plistocene 
elephants are of especial interest. Foremost and best-known of 
these is the mammoth (Elephas primigeniiis], which, as stated in 
an earlier chapter, was a very near ally of the existing Indian 
species, although distinguished as we know from the evidence of 
specimens preserved in the frozen soil of Siberia by its coat of 
woolly red hair, among which were intermingled long bristly black 
hairs. Curiously enough, traces of this woolly coat have been 
detected in the Indian elephant, so that it is probable that this 
species originated in some part of Asia where the climate is colder 
than is that of India. Regarding the range of this species, 
Professor Boyd Dawkins 1 remarks that "the mammoth is very 
abundant in the caverns and river-deposits of Britain and of France, 
and is known to have ranged over the Pyrenees into Spain, 
from the discovery of specimens in the zinc-mines of Santander. 
It has been proved by Prof. E. Lartet and Dr Falconer to have 
lived in the neighbourhood of Rome when the volcanoes of central 
Italy were active, and poured currents of lava and clouds of ashes 
over the [site of the] imperial city. It is common in northern and 
southern Germany, but it has not been found in Europe north of 
a line passing through Hamburg, or in any part of Scandinavia or 
Finland. It occurs in the auriferous gravels of the Urals ; and in 
Siberia, as is well known, it formerly existed in countless herds, 
being buried in the morasses in large numbers, in the same manner 
as the Irish elks at the bottom of the Irish peat-bogs. The 
admirable preservation of some of the carcases is undoubtedly due 
to their having been entombed directly after death, and then 
quickly frozen up, a process which need not necessarily imply 
climatal conditions unlike those of the present time in Siberia." 
That the mammoth ranged across Bering Strait into Arctic 
America, is proved by the discovery of its remains in the frozen 
soil of Eschscholtz Bay; but in the greater part of North America 
it was replaced by the closely-allied E. columbianus. In eastern 
Europe there existed a variety or species known as E. armeniaais, 
of which the molar teeth still more closely resemble those of the 
Indian elephant than do those of the typical form. The straight- 

1 Early Man in Britain (London, 1880), p. 106. 


tusked elephant (E. antiquus) is a more southern Plistocene 
type, of which the molars are to a certain extent intermediate 
between those of the living Indian and African species. Still 
more southerly in its distribution is the gigantic southern elephant 
(E. meridionalis), of which the remains are found in the upper 
Pliocene of Italy, as well as in the Plistocene Forest-bed of 
Norfolk, and equivalent strata at Dewlish, in Dorsetshire. The 
Maltese Islands were the habitat during the Plistocene epoch of 
the two or three species of dwarf elephants, which appear to have 
been nearly allied to the existing African species, but whose size 
was diminished by the smallness of the areas where they flourished. 
Lastly, the African elephant, which is now restricted to Ethiopian 
Africa, has left evidence of its existence during the Plistocene 
epoch in Algeria, Spain, and Sardinia. 

The fauna of the Forest-bed period, among which the mam- 
moth, megarhine rhinoceros, and Irish deer are wanting, is, as 
already stated, of pre-glacial age, and, on the whole, indicative of a 
fairly warm climate, although there is some evidence that the 
musk-ox then ranged as far south as England. At the close of 
this epoch, the southern elephant, together with a small bear 
known as Ursus arvernensis, appear to have become extinct. Soon 
after, glacial conditions made their appearance, causing much dis- 
turbance and migratory movements among the original southern 
pre-glacial fauna, and bringing an incursion of northern forms like 
the reindeer, Arctic fox, wolverene, and musk-ox, as well as of 
species from the eastern steppes such as the Saiga antelope and 
the Kirghiz jerboa (Alactagd), together with mountain animals like 
the chamois, the ibex, and the marmot, into the lowlands of south- 
western Europe. Among the northern forms that then spread 
themselves southward were the mammoth and the woolly rhino- 
ceros, which at this epoch appear to have attained their maximum 

Unfortunately, there is much uncertainty as to the part played 
by the glacial epoch in the extermination of the large mammals 
characterising Plistocene Europe. By most English geologists the 
brick-earths of the Thames valley, which contain remains of rhino- 
ceroses and elephants in abundance, as well as those of monkeys 
more sparingly, are regarded as of post-glacial age ; but Prof, von 


Zittel 1 considers them pre-glacial, or more probably inter-glacial. 
If they are either inter- or post-glacial, it is clear that the cold was 
not the exterminating cause ; and it is quite possible that many 
or all were killed off by man, although this could scarcely be the 
case with the Siberian fauna. 

Be this as it may, there is good evidence that when northern 
forms, such as the reindeer, wolverene, and banded lemming, had 
once obtained an entrance into central and southern Europe, they 
remained there for a considerable time, since they were present 
during the latter portion of what is known as the palaeolithic 
epoch. With the advent of the present climatic conditions came 
in the present woodland fauna of central Europe, constituting 
what has been termed the squirrel- or bison-epoch ; and from that 
date, when animals became domesticated, man has exercised a 
large influence on the fauna. 

It is important to notice that, in spite of the mingling of 
northern and southern types in England, France, and Germany, to 
which allusion has already been made, there seems to have been 
a distinction between the northern and the Mediterranean fauna 
throughout the whole of the later Plistocene epoch, such forms as 
the Barbary sheep and the fallow-deer being essentially southern, 
although the hippopotamus, as we have seen, extended as far 
north as Yorkshire. 

Although the later Plistocene fauna was spread not only over 
Europe, but also through North and Central Asia, a number of 
the characteristic European types, such as the hippopotamus, ibex, 
chamois, fallow-deer, cave-bear, and wild cat, were wanting in 
Asia. In that area forms are met with which are still character- 
istic of the same districts. As examples may be noticed : the 
Mongolian gazelle (Gazella gutturosd], the Himalayan ibex (Capra 
sibirica), the Persian wild goat (Capra cegagrus), the argali (Ovis 
argali), the musk-deer (Moschus moschiferus], the tiger (Felts 
tigris] of which the remains have been found even within the 
Arctic Circle together with a number of smaller forms, such as 
Siphneus aspalax, Ellobius talpinus, Spalax typhlus, Sminthus 
vagans, Tamias asiaticus, and Mustela zibellina. Here, then, 

1 Appendix, No. 36, p. 189. 


are clear indications of a Central Asiatic sub-region as far back 
as the Plistocene epoch. 

The foregoing brief survey of the Plistocene mammals of the 
eastern division of the Holarctic region enables 
certain deductions to be drawn as to geographical 
changes which have taken place in the area since the Plistocene - 
that epoch. 

In the first place, the occurrence of remains of the tiger in the 
New Siberian, or Liakov Islands, lying far within the Arctic 
Circle, indicates the union of those islands with the Siberian main- 
land ; and this greater extension of the land at the north-eastern 
extremity of Asia would naturally lead to the conclusion that there 
was also a land-connection with Alaska across Bering Strait. 
That such was really the case is proved by the discovery during 
the voyage of H. M. S. " Blossom," in the years 1825-28, of 
remains of the horse, mammoth, and bison, in the frozen soil of 
Eschscholtz Bay, Kotzebue Sound, Alaska 1 ; this evidence being 
confirmed by the occurrence of the musk-ox in the European 
Plistocene, as it is also by the number of species of mammals still 
common to the more northern parts of the two hemispheres. And 
here it may be mentioned that, according to the researches of the 
Russian geologists, Siberia, instead of being covered like northern 
Europe with a continuous ice -sheet during the glacial epoch, 
had only a number of comparatively small glaciers, so that the pre- 
glacial fauna was able to exist here at a time that it could not live 
in Europe. Still the frozen condition of the subsoil, and the 
formation of ground-ice in the rivers, rendered the preservation of 
the carcases of mammoths, rhinoceroses, bison, and musk-oxen an 
easy matter. 

Passing to south-western Europe, the occurrence of remains of 
the African elephant in Sicily and Spain, together with the 
presence of small allied species in the Plistocene of Malta, and 
likewise of remains of the Barbary sheep and Barbary ape in 
southern Europe, indicates a free land-communication between 
Europe and Africa, both by way of the Straits of Gibraltar, and 
likewise between Italy, Sicily, and Tunis; Malta being then also in 

1 Beechey's Voyage to the Pacific and Behring's Straits in H.M.S. 
"Blossom," Vol. II. 

L. 22 


connection with the mainland. Probably also it was by one or 
both of these routes that the hippopotamus and spotted hyaena 
passed between Europe and Africa, as it is scarcely likely that the 
former animal, at least, travelled round by way of Turkey and 
Syria. Writing of the Maltese islands, Leith-Adams 1 observes 
that "although from their smallness the islands furnish only scant 
evidences of the complicated and extensive oscillations of level 
to which the original area has been subjected from first to last, 
nevertheless the data I have furnished are at the least suggestive, 
and, in conjunction with the fossilised remains, seem to lead to 
the belief, that in the first place there was an upheaval of a large 
tract of land in this portion of the Mediterranean at some period 
towards or after the close of the Miocene epoch. In the second 
place, that during the Quaternary [Plistocene] period, the whole, 
or at least all excepting perhaps the tops of the Benjemma heights 
and Gozo hills, were again submerged ; and, thirdly, that a re- 
elevation of the land took place, ending in the present insular 
fragments. Perhaps in the first case there was a connection or 
contemporaneity in the upheaval of the Miocene beds of Malta, 
Sicily, Italy, Candia, the Red Sea, Egypt, Arabia, Cerigo, Azores, 
Algeria, Southern France, and Spain. Thus the islands of the 
inland sea may represent portions of a land area now occupied 
more or less by water. When this area began to sink is not 
apparent, but the fact that the same elephant and hyaena now 
living in Africa existed in Sicily, shews that there was a land- 
connection between the two at a very recent epoch." 

Regarding the nature of the former connection between Italy, 
Sicily, and Malta, Dr Wallace 2 writes that a comparatively shallow 
sea or submerged bank incloses Malta and Sicily, and " that on 
the opposite coast a similar bank stretches out from the coast of 
Tripoli, leaving a narrow channel, the greatest depth of which is 
240 fathoms. Here, therefore, is a broad plateau, which an eleva- 
tion of about 1,500 feet would convert into a wide extent of land 
connecting Italy with Africa ; while the same elevation would also 
connect Morocco with Spain, leaving two extensive lakes to repre- 

1 The Nile Valley and Malta (London, 1870), p. 211. 

2 Geographical Distribution of Animals, Vol. I. p. 201. 


sent what is now the Mediterranean Sea, and affording free com- 
munication for land animals between Europe and North Africa." 

Probably the dwarf elephants of Malta were developed from a 
larger form, closely allied to or identical with the African elephant, 
after the separation of the island itself from the mainland. With 
regard to Leith-Adams' idea of the subsequent submergence of 
Malta, it is pretty certain that this could not have been complete, 
since that island is inhabited by a large species of weasel (Mustela 
africand) common to Egypt, and perhaps the south of Italy 1 ; 
this animal being doubtless a survivor from the old fauna of the 
Plistocene land connecting Italy, Sicily and Malta with northern 

In north-western Europe there are equally conclusive evidences 
of the connection of the British Islands with the Continent during 
the Plistocene epoch. On many parts of the English coasts there 
occur, for instance, submerged forests dating from a comparatively 
recent epoch, which, when exposed during exceptionally low tides, 
are seen to contain the stumps of trees in their original upright 
position,. and with their roots still implanted in the soil. Forests 
of this kind are found near Torquay and Falmouth, as well as on 
several parts of the Welsh coasts and in Holyhead harbour ; the 
submergence which has taken place in the case of the one at 
Falmouth being estimated at upwards of 70 feet. Again, the pre- 
glacial Norfolk Forest-bed, so often alluded to in the foregoing 
pages, affords evidence of an extensive submergence on the east 
coast of England ; this being supplemented by the Dogger Bank 
in the North Sea, from which, as already mentioned, such numbers 
of remains of the mammoth, as well as those of the woolly rhino- 
ceros and other mammals, have been dredged. Additional evidence 
in favour of the same subsidence is afforded by the numerous 
ancient river-channels and valleys found in many parts of Britain, 
which are situated at depths of from one to two hundred feet 
below the present level of the land, and frequently cut right across 
the existing drainage lines, so as to connect valleys now com- 
pletely distinct. These ancient channels, which are now completely 
choked with sand, mud, or gravel, have only been revealed by the 

1 See Thomas, Proc. Zool. Soc. 1895, pp. 128 131. 



aid of the borer, but their evidence is, nevertheless, unimpeach- 

From these and other lines of evidence, we learn that during 
the Plistocene epoch not only was England connected with France 
across the English Channel, but that the land extended up the 
North Sea at least as far as the Dogger Bank ; the Ouse, the 
Thames, the Rhine, and perhaps the Elbe originally uniting to 
form one mighty river, discharging far up in the North Sea. 
During a portion of this period Ireland was in connection with 
the British Islands; and it has been suggested by Leith- Adams 1 
that the connection was with Scotland, owing to the circumstance 
that, with the exception of the cave-bear, all the living and extinct 
Irish mammals have been recorded from Scotland, while a number 
of the English Plistocene mammals appear never to have reached 
the latter country. On the other hand, Dr R. F. Scharff 2 , from a 
study of the freshwater fishes and molluscs, is of opinion that 
" Ireland was in later Tertiary times connected with Wales in the 
south and Scotland in the north ; whilst a freshwater lake occupied 
the present central area of the Irish Sea. The southern connec- 
tion broke down at the beginning of the Plistocene period, the 
northern connection following soon after. There is no evidence 
of any subsequent land-connection between Great Britain and 
Ireland." Since the above was written Dr Scharff (Mem. Soc. 
ZooL France, vol. vin. pp. 436-474, 1895) has further developed 
his views on the origin of the Irish fauna. He concludes that 
all the Irish mammals reached the island in the early Plistocene 
(Forest-bed); such British forms as are unknown in Ireland being 
considered to have reached Britain later, when Ireland was 

Further reference to the former connection or connections 
between Britain and the Continent will come more conveniently 

The generic and specific mammalian types common to the 
eastern and western divisions of the Holarctic region 


Division of the having been already referred to, we may at once 
proceed to the consideration of those characteristic 

1 Proc. Roy. Irish Acad. Ser. 2, Vol. III. (1883). 

2 Appendix, No. 25. 


of the western division. And here it may be mentioned in respect 
to the two areas, that whereas many generic types of animals were 
unable to pass from the one to the other owing to the high 
latitude of the strip of connecting land, yet in other cases the 
geographical limits of the range of certain genera in the Old World 
form also an important factor in the case. As stated a few para- 
graphs back, many of the characteristic European Plistocene 
mammals, such as the hippopotamus, the fallow-deer, and the 
cave-hyaena, never extended into the Asiatic portion of the Hoi- 
arctic region, so that these and many other forms never could 
have had an opportunity of crossing Bering Strait, even had 
they been capable of existing in such a high latitude. 

Excluding bats and seals, the following genera of mammals 
will be found confined to the western half of the Holarctic region, 
although some of these range southwards into the Sonoran. There 
are also certain genera which appear to be typically Sonoran, 
whose range includes part of the western Holarctic region, but 
these are best considered in the light of intruders from the 
south l . 

Among the shrews of the western Holarctic there are two 
species, viz. Sorex palustris, of the Rocky Mountains, and*S. hydro- 
dromus, of Unalaska Island, which differ from all their allies in the 
presence of long fringes of hair to the feet, although they resemble 
ordinary species of the genus in the characters of their dentition 
and tail. In consequence of these differences these aquatic shrews 
have been referred by some writers to a separate genus, under the 
name of Neosorex ; although such distinction is considered by Dr 
Merriam unnecessary. There is, however, one genus of Insectivora 
(Condylura), represented by the star-nosed mole, absolutely charac- 
teristic of this area. Allied in structure and habits to the Old 
World moles, which are totally wanting in America, this animal 
takes its name from the presence of a star-like ring of fleshy 
appendages at the extremity of the muzzle. 

The Carnivora include no peculiar genera 2 ; but the Rodentia, 

1 It may be well to mention here that the majority of American zoologists 
regard as genera a number of groups to which the present writer would not be 
disposed to grant more than sub-generic rank. 

2 Mephitis, Taxidea, etc., appear to be of Sonoran origin. 


which, as in the eastern division, are very numerous, comprise one 
family, as well as several genera, restricted to this area. In the 
Sciuridce the marmot- like genus Cynomys ranges into the Hoi- 
arctic, but is considered by Dr Merriam as chiefly characteristic of 
the Sonoran region; and the same is the case with the white- 
footed mice (Sitomys) of which there is but a single Holarctic 
representative, while the Sonoran species are very numerous and 
also with the wood-rats (Neotoma), of which a sub-genus is 
restricted to the Holarctic. The family peculiar to the region is 
that of the Haplodontidtz, or sewellels, represented by two species 
of the genus Haplodon, from the districts west of the Rocky 
Mountains. Closely allied to the squirrels, these rodents are 
distinguished from the latter by the absence of postorbital pro- 
cesses to the frontal bones of the skull, the depressed skull, and 
the rootless, or hypsodont, cheek-teeth ; all these characters indi- 
cating a more specialised type. In the Muridce, the voles of the 
genus Phenacomys connect the more typical members of the 
group with cricetines like the wood-rats (Neotoma). Several 
species have been described. A more southern type is the single 
representative of the allied genus Synaptomys, distinguished by its 
grooved upper incisors \ its molar teeth resembling those of the 
lemmings, while its skull is of the same structure as in the true 
voles. According to Dr Merriam, this animal is restricted to the 
southern part of the Holarctic area, or what he terms the Transi- 
tion region. In the same great family the well-known aquatic 
musk-rat, or musquash (Fiber), may be considered an Holarctic 
type, since it is found in the " barren-grounds " on the borders of 
the Arctic sea, although it ranges southwards into the Sonoran. 
Closely allied to the voles, with which it agrees in the characters 
of the skull and teeth, this animal differs by the long, compressed, 
nearly naked, and reticulate tail ; the naked-soled feet being partly 
webbed, and the whole body adapted to an aquatic mode of life. 
Its fossil remains occur in the Plistocene of the United States. It 
is noteworthy, as a negative characteristic of the Holarctic area, 
that no members of the exclusively New World family Geomyidce 
are found within its limits. On the other hand, in the family 
Dipodida, the j urn ping-mice of the genus Zapus, of which several 
species are recognised by North American zoologists, are solely 




Holarctic, the typical Z. hudsonianus dating from the Plistocene 
epoch. A distinctive feature of the western Holarctic region is 
the absence of true porcupines (Hystrix), their place being taken 
by the Canadian porcupine (Erethizon], which belongs to the same 
sub-family as the South American porcupines, although distin- 
guished, among other characters, by its short and non-prehensile 
tail. It is a native of the wooded portions of Canada and the 
United States, and its remains have been discovered in a cave in 

FIG. 69. ROCKY MOUNTAIN GOAT (Haploceros montamis] 

Among the ungulates, the remarkable animal known as the 
Rocky Mountain goat, which alone represents the genus Haplo- 
ceros, and differs from all other ruminants by the extreme shortness 
of the cannon-bone in both the front and hind limbs, is exclusively 
an inhabitant of the western Holarctic region. The same is now 


the case with the musk-ox (Ovibos), but as this animal ranged over 
Europe and northern Asia during the Plistocene, it can scarcely be 
regarded as distinctive of the western area. Of other peculiar 
New World ungulates, the prong-buck (Antilocapra) and certain 
deer of the genus Cariacus are found within the Holarctic region, 
but the former seems to be essentially a Sonoran type, while the 
latter, although probably also of Sonoran origin, occurs through 
Central and South America. 

The Tertiary genera of mammals peculiar to North America 
may be best considered in the chapter devoted to the Sonoran 
region, to which they for the most part belong ; and this portion 
of the subject may be accordingly concluded by tabulating the 
existing genera or groups peculiar to the area under consideration. 
These will stand as follows, viz. : 


SORICID.E. Sorex. The sub-genus or genus Neosorex. 
TALPID^E. Condylura. 


MURID^:. Phenacomys. 

Synaptomys. Confined to the southern 
portion of the area. 

Fiber. Enters Sonoran. 
DIPODID^E. Zapus. 

HYSTRICID.E. Erethizon. 


BOVID^E. Haploceros. 

Even if we add to the above certain other sub-generic types, 
such as the spruce-squirrels (Tamiasdurus) and the bushy-tailed 
wood-rats (Teonoma), and likewise take into account the number 
of Old World types (in many cases widely-distributed ones) that 
are absent, it can scarcely be urged that such an assemblage is 
sufficient to constitute a zoological region by itself. Those of my 
readers desirous of consulting lists of the species inhabiting the 
Arctic and Boreal zones of Dr Merriam, will find them in his 
memoir 1 . 

1 Appendix, No. 19, pp. 24, 25. 


In America, probably owing to the north and south trend of 
the mountain-ranges, the glacial period has had an even more 
marked effect than in the Old World. On this subject Dr 
Merriam 1 writes that "not only are the pre-Plistocene animals and 
plants now represented imperfectly and in greatly reduced num- 
bers, but the areas at present inhabited by their descendants, 
except in the case of the Boreal forms, are insignificant in com- 
parison with their former extent. It should be remembered that 
the refrigeration of the glacial epoch has only in part disappeared. 
In earlier Pliocene times, characteristic representatives of sub- 
tropical faunas and floras existed northwards over much of the 
United States and Canada, and in still earlier times reached the 
Arctic circle. During the advance of cold in the glacial epoch 
these forms were either exterminated or driven southward into the 
narrow tropical parts of Mexico and Central America. The retreat 
of cold at the termination of this period was not complete, and 
our continent has never regained its former warmth. Hence the 
expelled species were not permitted to advance more than a short 
distance into the region formerly occupied by them, and the 
tropical species have been held back, and at the present day are 
not found except along the extreme southern confines of our 
territory [the United States]. For example, peccaries in early 
Plistocene times ranged northward over a large part of western 
North America, while at present they are restricted to parts of 
Texas and Louisiana below the Red River of the south ; and 
capivaras, tapirs and other tropical forms whose fossil remains 
have been found in many parts of the United States have not been 
able to return. The same is true of plants, for the palms, tree- 
ferns, and numerous other tropical types that formerly ranged over 
much of our country are now either altogether extinct or exist 
only in the tropics. 

"The llama and many plants now inhabiting the Andes may 
be looked upon as representing a class of cases in which Boreal 
forms were driven so far south that they actually reached the great 
mountain-system of South America and spread southward over its 
elevated plateaus and declivities to the extreme end of the conti- 
nent in Patagonia and Tierra del Fuego." 

1 Appendix, No. 19, p. 44. 




Coming to the consideration of sub-regions, we have first of 
all the Arctic sub-region, which corresponds to the 
Boreal sub-region of Dr Heilprin, and the Arctic 
zone of the Boreal region of Dr Merriam, and is of 
circumpolar extent. According to the former writer, in the Old 
World it may be defined as the tract lying to the north of a line 
starting from about the 66th parallel of latitude on the Norwegian 
coast, and passing south-eastwards to the coast of eastern Asia 
in about the 5oth parallel, thus including the greater part of 
Kamschatka, and Amurland. In America, according to Dr 
Merriam's map, after running just inside the shores of Newfound- 

FIG. 70. MUSK-OX (Ovibos moschatus). 

land and Labrador, the boundary line bends southwards after 
passing Cape Chudleigh to coincide with the southern shore of 
Hudson Bay, and then takes a north-westerly direction so as 
to include within the sub-region only a narrow strip on the north- 
eastern coast of Alaska, and a somewhat broader one on the 
north-western shore of the same. In the Old World the boundary 
line coincides approximately with the northern limit of the cultiva- 
tion of cereals, and also with that of the southern migrations of 
the reindeer; but in America certain reindeer (which are regarded 


by the American zoologists as specifically distinct from thecircum- 
polar "barren-ground" variety) extend considerably further to the 
south. For the most part of its extent, the mammals inhabiting 
this sub-region are few in number, a large proportion of them 
having a circumpolar range. Among them may be included the 
Arctic fox, polar bear, wolverene, the ermine or stoat, the eastern 
and western species of lemming (Myodes), the banded lemming 
(Cuniculus torquatus), the Arctic vole (Microtus nitilus}, Parry's 
suslik (Spermophilus empetra], the musk-ox, and the reindeer ; 
several of these being restricted to the sub-region. The sea-otter 
(Latax) frequents the shores of Alaska and Kamschatka, but also 
ranges as far south as the Kurile Islands and California, so that it 
is not confined to the sub-region. During the Plistocene epoch, 
as we have seen, such animals as the mammoth, horse, bison, and 
tiger were inhabitants of this tract ; the latter animal being still 
found in eastern Siberia. Towards Amurland and the Kams- 
chatkan peninsula, the fauna becomes somewhat less scanty ; the 
large Kamschatkan sheep (Ovis nivicola) being here met with, as 
well as a true deer, and the brown bear. 

Of other groups of animals inhabiting the more typical portions 
of this region, there may be noticed among the birds the ptarmigan 
(Lagopus], the snowy owl (Nyctea scandiaca), the Greenland falcon 
(Falco candicans), the eider-duck (Somatcria mollissima), as well as 
various species of divers (Colymbus] and guillemots (Uria and 
Lomvia}, together with the little auk (Mergulus alle\ Dr Heilprin 
writes that " Captain Markham observed the footpiints of the 
polar hare in the snow-bound ice in latitude 83 10', and the 
antlers of a reindeer were picked up by the officers under Sir 
George Nares, in latitude 82 45' (Grinnell Land). A skeleton of 
the latter animal, recently picked by wolves, was also obtained in 
latitude 80 27'. Traces of the rock-ptarmigan (Lagopus rupestris] 
have been met with as far north as latitude 83 6', and the snow- 
bunting (Pkctrophanes nivalis) in latitude 82 33'. The reptile- 
fauna is very limited, no serpent, apparently, passing beyond the 
sixty-seventh parallel of latitude, and no lizard above the seventieth. 
The fishes, which include the common perch and pike, are mainly 
salmonoids. Insects are fairly numerous, and even in the far 
north the number of species is considerable." 


During the Plistocene the region within the Arctic circle en- 
joyed a decidedly less rigorous climate than it at present pos- 
sesses. In Baron von Toll's recent expedition to the New Siberian 
Islands 1 , where, as previously stated, remains of the tiger have been 
obtained, it was discovered " that under the perpetual ice, in a 
freshwater deposit, which contained pieces of willow and bones of 
post-tertiary mammals (the mammoth-layer) were complete trees 
of Alnus fruticosa, fifteen feet long, with leaves and fruit. It was 
thus evident that during the mammoth-period tree-vegetation 
reached the seventy-fourth degree of latitude, and that its northern 
limit was at least three degrees further north than it is now." 

The next sub-region is the European, which may be taken to 
include all that part of Europe lying between the 
Arctic sub-region in the north, and the line of the 
Pyrenees and Alps, continuing eastwards along the 
northern shore of the Black Sea to the Caucasus and the Caspian 
Steppes. This area includes the typical fauna of the eastern 
Holarctic region, among its more or less characteristic mammals 
being (in the north) the elk also ranging into America , the red 
deer (unknown in America, but represented by a variety in North 
Africa), the roe, the bison, the chamois, the Alpine ibex, the 
typical variety of the brown bear, the badger, the wolverene (in the 
north), the Alpine marmot (Arctomys marmotta), the dormouse, 
hamster, mole, and hedgehog \ several of these being, however, 
common to the Arctic and Central Asian sub-regions. The des- 
mans (Myogale) are restricted to this sub-region ; and the same 
was probably the case with the aurochs (Bos taurus, var. primi- 
genius], the ancestral stock of our domestic cattle. Finally, the 
Caucasus is the home of two or three peculiar species of goats 
(Capra cylindricornis and C. caucasica] known as ture. 

It will be unnecessary, even if this could be accomplished, to 
give a complete list of the mammalian fauna of this sub-region, 
but it is essential to refer to the comparative poverty of the fauna 
of the British Islands as compared with that of the Continent. 
The following list includes all the mammals (exclusive of bats) 
known to have inhabited the British Islands within the historic 

1 See Knowledge, 1895, p. 106. 


period ; those which are now extinct having an asterisk prefixed 
to them, while such as occur in Ireland have the letter I added. 
Those that have been introduced by man have a t before them. 
The list stands as follows, viz. : 

Hedgehog. Erinaceus europaeus. I. 

Mole. Talpa europsea. 

Common Shrew. Sorex araneus. 

Lesser ,, ,, minutus. I. 

Water-Shrew. Crossopus fodiens. 

Wild Cat. Felis catus. 
*Wolf. Canis lupus. I. 

Fox. vulpes. I. 

Pine-Marten. Mustela martes. I. 

Polecat. ,, putorius. 

Stoat. erminea. I. 

Assogue. ,, hibernica. I. 

Weasel. ,, vulgaris. 

Badger. Meles taxus. I. 

Otter. Lutra vulgaris. I. 
^Brown bear. Ursus arctus. I. 

Squirrel. Sciurus vulgaris. I (? introduced), 
* Beaver. Castor fiber. 

Dormouse. Muscardinus avellanarius. 

Harvest-Mouse. Mus minutus. 

Wood-Mouse. ,, sylvaticus. I. 

Yellow-necked Mouse. ,, flavicollis. 

Common Mouse. ,, musculus. I. 

f Black Rat. rattus. I. 

t Brown Rat. ,, decumanus. I.. 

Common Field : Vole. Microtus agrestis. 

Bank-Vole. glareolus. 

Water-Vole amphibius. 

Common Hare. Lepus europaeus. 

Mountain Hare. ,, timidus. I. 
t Rabbit. ,, cuniculus. I. 

*? Wild Cattle. Bos taurus. 

Red Deer. Cervus elaphus. I. 


t Fallow Deer. Cervus dama. I. 

Roe Deer. Capreolus caprea. 
*Wild Boar. Sus scrofa. I. 

The total number in this list is only 28, out of which at least 
four are introduced. With the exception of the recently-described 
assogue 1 , which is intermediate between the stoat and the weasel, 
and is peculiar to Ireland, the whole of these mammals are com- 
mon to the Continent. As shown in an earlier portion of the 
present chapter, during the Plistocene epoch Britain possessed a 
fauna apparently identical with that of the Continent ; and there 
must accordingly be some good reason for its present poverty in 
mammalian life as compared to the latter area. The difference is 
accounted for by Dr Wallace, through the occurrence of one or 
more periods of subsidence, which took place during the close of, 
or subsequent to, the Glacial epoch ; after which England again 
became united to the Continent, when its present fauna entered, 
the period of connection being, however, of comparatively short 
duration, and thus permitting of the passage of only a moiety of 
the continental forms, or those which happened at the time in 
question to be inhabiting the districts nearest to the connecting 
line. Only a certain number of the mammals which thus crossed 
into Britain have ever succeeded in reaching Scotland ; and it is 
from this country, if we accept the views of Dr Scharff, referred to 
above, that Ireland appears to have received its still more im- 
poverished mammalian fauna. 

It will be seen that the foregoing hypothesis attributes the 
clean sweep supposed to have been made of the original British 
fauna to the effects of submergence, and not to the ice-sheet. On 
the other hand, Mr G. W. Bulman 2 , who doubts whether the sub- 
mergence has been sufficient for this, attributes such extermination 
as he believes to have taken place solely to the effects of an ice- 
sheet. And he further believes that a number of the original 
British mammals survived in the southern and south-western 
counties of England, whence they re-populated Britain on the 
disappearance of the ice-sheet, without there having been any 

1 See Thomas, Natural Science, Vol. vi. p. 377 (1895). 
- Appendix, No. 12. 


subsequent connection with the Continent. The difficulty con- 
nected with this explanation is that it apparently necessitates a 
pre-glacial or early glacial age for the mammaliferous deposits of 
the Thames valley, which are almost certainly inter-glacial or 
post-glacial. The whole subject of glaciation is, however, so 
complicated and involved, that it is almost impossible to form 
workable theories as to the exact mode of the repopulation of 
Britain after the changes which took place during the glacial 

In contrast to the British Isles, which are eminently of the 
continental type, may be cited Iceland, lying near the border-line 
between the Arctic and European sub-regions, which is as 
markedly oceanic in its character. Beyond an occasional ice- 
borne polar bear, Iceland possesses only the Arctic fox, and a 
mouse, which has been stated to be a peculiar species ; the fox 
having doubtless been originally introduced from the north on 
floating ice. 

According to the scheme of Dr Heilprin, the next sub-region 
on the list is that of Central Asia, which includes 
the countries bounded on the west by the European, sSb-regi* 1 * 11 
and on the north by the Arctic sub-region, and 
extends eastwards as far as Mantchuria and China proper, being 
bordered on the south by the Kuenlun and Nanshan mountains 1 . 
A large portion of the western districts of this tract are open 
steppes or deserts ; and in such tracts several peculiar types of 
rodents, such as the Kirghiz jerboa (Alactaga) and the Yarkand 
jerboa (Euchoretes\ are met with, while the saiga antelope (Saiga), 
and the Mongolian gazelle ( Gazella guttu rosd) are likewise charac- 
teristic types. Susliks (Spermophilus), marmots (Arctomys), and 
picas are very abundant ; and the place of the European wild cat 
is occupied by Pallas's cat (Felis manul), the tiger being also 
sparingly found in the western districts, where the ounce is like- 
wise met with. In part of this sub-region the red deer is replaced 
by a variety or species known as Cervus xanthopygus, while 
Yarkand is the home of a variety of the Kashmir stag (C. cash- 
mirianus], and the Thian-Shan possesses the very fine and wapiti- 

1 Dr Heilprin included the Tibetan plateau in this sub-region. 


like form described under the name of C. eustephanus ; all these 
deer being mostly inhabitants of forest-districts. The Tatarian 
roe (Capreolus pygargus), inhabiting suitable localities in the moun- 
tains forming the watershed between the Russian and Chinese 
empires and Turkestan, is also generally regarded as specifically 
distinct from its western ally. The sub- region is also the chief 
home of the magnificent sheep known as argali, among which 
the splendid Pamir sheep (Ovis poll) ranges from the Pamirs to 
the Altai, while the true argali (O. ammon) if the Tibetan O. hodg- 
soni be really distinct is also restricted to this sub-region, where 
it is now confined to northern Mongolia, although it formerly 
inhabited the Altai. The ibex of the Altai is, however, identical 
with the Himalayan and Tibetan Capra sibirica. 

Although included by Dr Heilprin in the preceding, the area 
typified by the Tibetan plateau is regarded by Dr 
Blanford ' as constituting a sub-region by itself. 
Typically this region is bounded on the north by 
the ranges of the Kuenlun, Altyn Tag, and Nanshan, and extends 
eastwards to China proper, while to the west it must be taken to 
include Ladak and the upper Indus valley as far as Gilgit 2 . To 
the south it extends to the main chain of the Himalaya. The 
following list of mammals is given by Dr Blanford as distinctive of 
this sub-region ; the names of such species and genera as are 
entirely or mainly confined to the area being printed in italics. 

Insect! vora. 

Crocidura aranea. 
Nectogale elegans. 


Paradoxurus laniger. 
Canis lupus, var. laniger. 

,, vulpes, var. flavescens. 

,, ferrilatus. 

,, deccanensis, var. 
Mustek foina, var. 

,, larva ta, 

1 Proc. Zool. Soc. 1893, p. 449. 

2 See Blanford, Fauna of British India, Mammalia, p. v. 


Carnivora (cont.\ 

Mustela canignla. 

alpina, var. temon. 

Meles leucura. 

sEluropus melanoleuctts. 
Ursus pruinosus. 


Eupetaurus rinereus. 
Arctomys himalayanus. 

Mus sublimis. 
Microtus blythi. 
,, strauchi. 
Siphneus fontanieri. 
Lagomys curzonice. 

,, rutilus. 

,, erythrotis. 

melanostomus . 

,, ladacensis. 
Lepus oiostolus. 
,, hypsibius. 


Equus hemionus, var. kiang. 
Bos grunniens. 
Ovis hodgsoni. 

vignei, var. 

,, nahura. 
Capra sibirica. 
Pantholops hodgsoni, 
Budorcas taxicolor. 
Gazella picticaudata. 
Cervus affinis. 

,, thoroldi. 
Moschus moschiferus. 
L. 23 


This list includes all the species inhabiting the plateau at 
elevations exceeding 12,000 feet. Dr Blanford writes that "many 
of the forms named only inhabit small portions of the area, and 
whilst Bos grunniens, Pantholops hodgsoni, and Gazella picti- 
caudata, with several rodents, appear to be peculiar to the high 
plateaus above 14,000 feet, the two species of Cervus are pro- 
bably found in brushwood at a rather lower elevation in the more 
broken region of Eastern Tibet, where the rainfall is heavier and 
the vegetation more abundant. 

"As was printed in the paper in the Geological Magazine 1 , 
there is, so far as I am aware, no equally peculiar mammalian 
fauna to be found in any continental area of equal extent, and for 
a parallel it is necessary to turn to some island like Celebes, that 
has long been isolated from all surrounding lands." 

This, however, is not all, for there occur at Hundes, on the 
Tibetan plateau, mammaliferous strata yielding, among other 
remains, bones of a rhinoceros, and of an antelope which is 
apparently generically identical with the chiru (Pantholops), now 
inhabiting the same area. The isolation and development of this 
most peculiar fauna is intimately connected with the date of 
elevation of the Himalaya. After pointing out that both the fossil 
chiru and the fossil rhinoceros appear to have inhabited the area 
when it had attained something approaching its present enormous 
elevation, Dr Blanford 2 writes as follows : " Bearing in mind that 
the isolation of the Tibetan plateau is far less perfect as regards 
mammals than that of any island, and that some of the forms 
the Carnivora especially found in Tibet are evidently very recent 
immigrants, it is a reasonable conclusion that the peculiar fauna 
of the Tibetan plateau has been distinct from that of neighbouring 
countries since middle Tertiary times. 

" But what has caused the isolation of the Tibetan fauna ? 
Why in this one continental tract is there a generic and specific 
differentiation of the mammalia, of which no other example 
exists ? There is only one character in which Tibet is different 
from other continental areas, its great height. This alone renders 
the climate of Tibet so different from that of other parts of 

1 Decade 3, Vol. ix. p. 161 (1892). 

2 Geol. Mag. op. cit. p. 165. 


Central Asia, which are equally cold and barren. It seems a 
reasonable inference that the elevation of the Tibetan plateau 
dates back to middle Tertiary times. 

" It is of course probable that the elevation was gradual ; and 
although the area may have been sufficiently high at the close of 
the Miocene period to produce a difference in climatal conditions, 
the greater part of the upward movement may have been post- 
Miocene, and a great part post-Pliocene." 

Bordering as it does upon the tropics, where it abuts against the 
Oriental region, the Mantchurian sub-region is not 
easy to define, since the intermingling of Holarctic 
and Oriental types is very strongly marked on its 
southern confines. Starting somewhere about the Amur river, it 
may, however, be taken to include the Japanese islands, Mant- 
churia, Corea, and northern China ; its southern limit being placed 
approximately in the latitude of Fuchau. Westwards it may be 
taken to include Moupin, in Eastern Tibet, although this district 
is referred by Dr Wallace to the Oriental region. 

From all the other sub-regions, with the exception of the 
Mediterranean, the Mantchurian is distinguished by the presence 
of monkeys belonging to the genera Macacus and Semnopithecus, 
some of these occurring in Japan and others in Eastern Tibet. 
Of the latter, one (Semnopithecus roxellance) is peculiar, and the 
other is identified by Mr H. O. Forbes with the widely-spread 
Oriental Macacus arctoides. Among the Carnivora, the Oriental 
genus Helictis enters this sub-region, one species occurring in the 
neighbourhood of Shanghai ; while Japan is the home of a peculiar 
long-haired dog ( Cants procyonides), which is frequently separated 
generically under the name of Nyctereutes, although it unquestion- 
ably pertains to the typical genus. Perhaps, however, the most 
characteristic mammals are the deer. Foremost among these are 
a group of small deer belonging to the genus Cenms, and distin- 
guished from the red deer group by the invariable absence of a 
bez-tine to the antlers, each of which has but four points. These 
deer are further characterised by the coat of the adult being 
spotted in summer with white, but uniformly brown in winter, and 
also by the black lateral margins to the white blaze on the hind- 
quarters. The species include the Japanese deer (C. sica), 



common to Japan and North China, the larger Mantchurian deer 
(C. mantchuricus\ and Dybowski's deer (C. dybowskii) from the 
upper Ussuri district of Mantchuria, in the neighbourhood of 
Vladivostock. Elsewhere the group is represented in Formosa, 
and also in the Caspian provinces of Persia. In addition to 
these, there are the hornless Chinese water-deer (Hydropoles), and 
the two species of tufted deer (Elaphodus)\ the latter being closely 
allied to the Oriental muntjacs. What is known of the palseonto- 
logical history of the southern portion of this area indicates that 
during the Pliocene epoch its mammalian fauna was closely allied 
to that of the Siwalik Hills, thus showing that at this time there 
was no distinction between the Oriental and Holarctic regions, 
which even now grade imperceptibly into one another in this 

The remains of fossil elephants from Japan 1 are referable to 
Elephas clifti, insignis, and namadicus, of which the two first are 
common to the Siwaliks, while the third occurs typically in the 
Plistocene Narbada beds of India. From the known distribution 
of these elephants, it is probable that Japan was connected with 
the mainland during the Pliocene epoch by way of the Corean 
peninsula, although Dr Wallace is of opinion that its latest con- 
nection was to the north. Of existing animals common to Japan 
and the mainland, allusion has already been made to Cervus sica : 
and another common type is the giant salamander Megalobatra 
chus. The latter genus is represented in a fossil state in the 
Miocene of Baden, and as it is closely allied to the North American 
CryptobranchuS) there is clear evidence of the eastern migration of 
this ancient type, of which the two survivors are respectively con- 
fined to China and Japan on the one hand, and North America 
on the other. Further evidences of affinity between the fauna of 
Japan and North America are afforded by the circumstance that 
one species of the mole-like genus Urotrichus is confined to the 
former islands, while the other is an inhabitant of the north- 
western districts of the latter continent. The sea-otter (Latax] is 
likewise common to the coasts of Japan, the Kurile Islands, and 
Kamschatka, and the Pacific shores of North America. More 
remarkable, however, is the fact that a North American scincoid 
1 See Naumann, Palceontographica, Vol. xxvm. Art. r (1881). 


lizard (Eumeces quinquelineatus] is represented in Japan by a form 
(E, marginatus] so closely allied that the two were long considered 
inseparable, although they are now regarded as distinct 1 . All 
these facts are indicative that Japan was formerly joined to both 
Corea and Kamschatka, whence land was continued across Bering 
Strait to unite the Old World with Alaska. 

Although, as already stated, the Mediterranean or Tyrrhenian 
sub-region has strong claims to be regarded as Mediterra 
representing a region by itself, it may be more con- nean Sub- 
veniently considered here than later on in the 
chapter. In addition to such parts of Africa and Arabia as lie 
to the north of the Ethiopian region, this sub-region includes 
Spain, those parts of Europe situated south of the Alps, together 
with Turkey, Asia Minor, Persia, Baluchistan, and Afghanistan. 
Whether Kashmir should be regarded as an aberrant outlier of 
this region, I am not yet satisfied. Although gerbils (Gerbillus*) 
are also found in the Oriental and Ethiopian regions, their 
distribution in the Holarctic is very nearly coincident with the 
limits of the present sub-region. 

Whereas to the north of the Mediterranean Sea a large pro- 
portion of the mammals are more or less typically Holarctic, in 
North Africa and Syria those with an Ethiopian facies are met 
with, and an Oriental element makes its appearance in the eastern 
districts of the sub-region. Even in Africa, however, some of the 
forms have an Oriental facies, the Barbary ape (Macacus inuus) 
belonging to a genus whose home is now in the Oriental region, 
and which is totally unknown in the Ethiopian. As a wanderer 
from the Ethiopian region, mention may first be made of a species 
of jumping -shrew (Macroscelides) met with in Barbary, while 
among the octodont family of the rodents, the gundi, forming 
the sole representative of the genus Ctenodactylus, has its nearest 
allies in Ethiopia, although it is confined to North Africa. The 
Barbary ape, although occurring on the rock of Gibraltar, where it 
may have been introduced, is otherwise confined to North Africa. 

1 See Boulenger, Cat. Lizards, Brit. Mtis. Vol. III. p. 369. 

2 Many writers separate certain species as Meriones, but as the two groups 
are connected by G. indicus (see Lataste, Proc. ZooL Soc. 1884, p. 88), such 
distinction seems superfluous. 


In the Carnivora, the striped hyaena, which is also an inhabitant 
of India, is widely spread in this sub-region, ranging through 
western Asia to northern Africa. The common genet (Genetta 
vulgaris\ which belongs to a genus otherwise exclusively Ethio- 
pian, is mainly confined to this region, inhabiting southern France, 
Spain, Turkey, North Africa, and Palestine. A nearly similar 
distribution characterises the common mungoose, or ichneumon 
(Herpestes ichneumon], which frequents southern Spain, Asia 
Minor, North Africa, and Palestine. The large weasel (Mustela 
africanus) common to Egypt, Malta, and perhaps the south of 
Italy has been already referred to in an earlier part of this chapter. 
In addition to Ctenodactylus, the rodents possess another and more 
widely-spread generic type confined to this sub-region in the form 
of the great mole-rat (Spalax typhlus), whose range includes south- 
eastern Europe, Persia, Mesopotamia, Syria, and Egypt. In the 
same order the common porcupine (ffystrix cristata], although 
ranging into West Africa, is found but little, if at all, to the north 
of the present sub-region, where it is common to northern Africa 
and southern Europe. 

Among the ungulates the addax antelope (Addax nasomacu- 
lata\ although allied to Ethiopian types, is solely Mediterranean, 
its home being North Africa and Syria. More closely allied to 
the Ethiopian fauna are certain hartebeests of the genus Bubalis, 
the smaller of which (B. mauritanica] is common to North Africa, 
Syria, and Arabia, while the second {B, major] inhabits Tunis. 
The same is the case with the Beatrix antelope (Oryx beatrix] 
of Western Arabia and Bushire. In gazelles, this sub-region is 
remarkably rich, doubtless from the number of sandy or desert 
tracts it contains. Algeria is the habitat of the three species 
known as Gazella loderi, G. kevella, and G. rufina, while G. 
dorcas ranges through Egypt, Algeria, Syria, Palestine, and a part 
of Asia Minor, and G. subgutturosa roams from Persia through 
Afghanistan and Turkestan. The aberrant sheep known as the 
arui (Ovis tragelaphus] is now restricted to North Africa; and the 
mouflon (O. musimon), although its fossil remains have been found 
on the Continent, appears to be now restricted to Corsica. An- 
other species peculiar to the sub-region is the Armenian sheep 
(O. gmelini) of eastern Persia and Asia Minor, represented by a 

IX.] KASHMIR. 359 

closely-allied form in Cyprus. Of the goats, the Spanish ibex 
(Capra pyrenaica) is restricted to the mountains of Spain; while 
the Sinaitic ibex (C. sinaitica) represents the group in Palestine 
and upper Egypt. Among the Cervidcz, the two species of fallow- 
deer were originally confined to this area, the common Cervus 
dama being a native of the Mediterranean countries, while the 
Persian C. mesopotamicus is found in the mountains of Luristan, in 
Mesopotamian Persia. In North Africa the ordinary red deer is 
represented by a variety distinguished by the invariable absence of 
a bez-tine to the antlers. A connection with the Tibetan sub- 
region is afforded by the wild asses inhabiting the desert-plains 
between the Red Sea and the Indus, since both these and the 
Tibetan form are but varieties of a single species (Equus hemi- 
onus). Lastly, Ethiopian affinities are exhibited by the occurrence 
of a species of hyrax (Procavid] in Syria. In the early part of the 
present century the hippopotamus still inhabited lower Egypt, 
while, as we have seen, the lion, which is now common in parts of 
Persia, was found within the historic period in Thrace. At a still 
earlier date, both these animals, as well as the spotted hyaena, 
extended as far north as England. 

On the whole, therefore, the fauna of this sub-region is a very 
mixed one ; and this fact, together with the difficulty in defining 
its boundaries, suggests the need of further deliberation before the 
area is raised to the rank of a separate region. The former con- 
nections between southern Europe and Africa having been alluded 
to in an earlier part of the chapter, require no further notice in 
this place. 

Very difficult to determine is the position which should be 
given to the valley of Kashmir, since its fauna 
exhibits such a mingling of Oriental and Holarctic 
types that it might almost be as well assigned to one region as the 
other. Holarctic affinities are, however, exhibited by the occur- 
rence of a species of the red deer group, Cervus cashmirianus, and 
likewise by one of the rodent genus Sminthus, of which the 
second species inhabits northern and eastern Europe, and the 
third Kansu, in western China. A variety of the brown bear is 
also indicative of Holarctic affinities, and this is still more 
markedly the case with the spiral-horned goat known as the 


markhor (Capra falconeri], of which one variety inhabits the Pir 
Panjal range, on the south side of the valley, while the others are 
found in the districts to the north and west of Kashmir. The 
musk-deer, again, is another essentially Holarctic type. On the 
other hand, the occurrence of a langur -(Semnopithecus) and a ma- 
caque (Macacus) points to a connection with the Oriental fauna ; 
and a Kashmir mungoose (Herpestes auropunctatus] is identical 
with one from India. There are, however, none of the exclusively 
Oriental genera in Kashmir ; and this fact, coupled with the 
absence of all deer of the sambar-group, leaves little doubt that 
the valley really belongs to the Holarctic. Whether it should 
be regarded as pertaining to the Mediterranean sub-region, or as 
forming a distinct sub-region by itself, must be reserved for future 

Passing to the western division of the Holarctic region, the 
tract lying to the southward of the circumpolar Arctic 
sub-region, designated by Dr Merriam the Boreal 
zone of his Boreal region, may be conveniently 
termed the Canadian sub-region. Its northern boundary is, of 
course, identical with the southern limits of the Arctic sub-region, 
and the area includes the greater part of the Dominion of Canada, 
although a long strip runs down the line of the Rocky Mountains, 
and another along the Pacific coast, into the United States. Indeed, 
Dr Merriam includes in this sub-region all the higher plateaus of 
Wyoming and Colorado, so that the sub-region embraces a 
number of small disconnected areas on its south-western ex- 
tremity, and it is consequently impossible to define its limits by 
description. It may be stated, however, that on the eastern side 
of the continent the sub-region extends from Hudson Bay to the 
middle of Lake Michigan, while on the western coast it stretches 
from near the extremity of Alaska to San Francisco ; a big loop 
extending northwards of Montana nearly to latitude 55. 

The mammalian fauna of the Canadian sub-region is that of 
the western division of the Holarctic region generally, and includes 
those forms mentioned on page 344. According to Dr Merriam, 
the following genera from this sub-region do not range further 
south than the undermentioned Transition zone ; namely, Condy- 
lura, Urotrichus, Gulo, Latax, Arctomys, Haplodon, Phenacomys, 


Myodes, Cuniculus, Zapus, Erethizon, Lagomys, Cervus, Alces, 
Rangifer, and Haploceros. On the other hand, the following, 
which are as clearly of northern origin, penetrate as far south 
as the Sonoran region, which some of them enter. These are 
Sorex, Mustela (only the members of the sub-genus Putorius), 
Ursus, Fiber, Microtus, Castor, Tamias, Bos, and Ovis. 

Between the Canadian sub-region of the Holarctic and the 
Sonoran region is interposed a tract whose fauna 
contains a mixture of Canadian and Sonoran forms, Transition 


and it is consequently termed by Dr Merriam the 
Transition zone. Under this somewhat indefinite title the area 
may best be left. It is described by the author just cited as 
follows 1 : "The humid division of this zone, known as the 
Alleghanian fauna, covers the greater part of New England (except 
Maine and the mountains of Vermont and New Hampshire), and 
extends westerly over the greater part of New York, southern 
Ontario, and Pennsylvania, and sends an arm south along the 
Alleghanies, all the way across the Virginias, Carolinas, and 
eastern Tennessee, to northern Georgia and Alabama. In the 
Great Lake region this zone continues westerly across southern 
Michigan and Wisconsin, and then curves northward over the 
prairie-region of Minnesota, covering the greater parts of North 
Dakota, Manitoba, and the plains of the Saskatchewan ; thence 
bending abruptly south, it crosses eastern Montana and Wyoming, 
including parts of western South Dakota, and Nebraska, and forms 
a belt along the eastern base of the Rocky Mountains in Colorado 
and northern New Mexico, here as elsewhere occupying the 
interval between the Upper Sonoran and Canadian zones. 

" In Wyoming the Transition zone passes broadly over the 
well-known low divide of the Rocky Mountains, which affords the 
route of the Union Pacific railway, and is directly continuous with 
the same zone in parts of Colorado, Uta, and Idaho, skirting the 
Canadian boundaries of the Great Basin all the way around the 
plains of the Columbia, sending an arm northward over the dry 
interior of British Columbia, descending along the eastern base of 
the Cascade Range and the High Sierra to the southern extremity 

1 Appendix, No. 19, p. 30. In this extract the word Canadian has been 
substituted for Boreal. 


of the latter, and occupying the summits of the Coast Ranges in 
California and of many of the desert ranges of the Great Basin. 

"The Transition zone, as its name indicates, is a zone of 
overlapping of Canadian and Sonoran types. Many Canadian 
genera and species here reach the southern limits of their distribu- 
tion, and many Sonoran genera and species their northern limits. 
But a single mammalian genus (Synaptomys) is restricted to the 
Transition zone, and future research may show that it inhabits the 
Canadian region also." 

FIG. 71. MUSQUASH (Fiber zibet hi cus). 

The writer adds, however, that there are a considerable number 
of species mostly rodents restricted to this zone. The follow- 
ing Canadian genera, namely, Condylura, Urotrichus, Ursus, 
Arctomys, Tamias, Fiber 1 , Zapus, Erethizon, Cervus, and Ovis, 
almost or completely disappear in this zone ; while the following 
intruders from the Sonoran, namely Scalops, Bassariscus, Spilo- 
gale, Perognathus, Thomomys, Geoitiys, Cynomys and Antilocapra 
do not range further north, several of them, indeed, only intruding 
into the zone in a small area in the west. 

1 Penetrates the Sonoran along the lines of streams where cool currents of 
air are carried down. 


Limits Characteristics of Mammalian Fauna Extinct Groups of Mammals 
characteristic of Western Arctogaea Distinctness of the Region Dual 
Origin of Groups. 

As stated in the introductory chapter, wherever one zoological 
region of the globe has no definite physical barrier by which 
it is separated from the next well-marked region, there must 
always occur an intermediate tract where the characteristic types 
of the faunas of the two regions inosculate and intermingle. That 
this is the case with that area of North America denominated the 
Sonoran region has been indicated at the close of the preceding 
chapter, and the existence of the Transition zone, which seems, 
on the whole, to pertain to the Holarctic region, unfortunately 
prevents the Sonoran from being defined with the precision which 
would be possible had this area a high mountain-barrier on its 
northern frontier. 

In a map of the small dimensions of the one accompanying 
this volume it is impossible to show with any 


attempt at accuracy the complex nature of the 
northern boundary of this region, which will, however, be found 
accurately laid down in the map illustrating Dr Merriam's memoir 1 . 
According to the latter writer, " the Sonoran region as a whole 
stretches across the continent from the Atlantic to the Pacific, 
covering nearly the whole country south of latitude 43, and 
reaching northward on the Great Plains and Great Basin to about 
latitude 48. It is invaded from the north by three principal 
intrusions of Canadian 2 forms along the three great mountain- 

1 Appendix, No. 19. 

2 Boreal in the original. 


systems already mentioned [Alleghanies, Rocky Mountains, and 
Cascade and Sierra Nevada ranges] ; while to the southward it 
occupies the great interior basin of Mexico, and extends into the 
tropics along the highlands of the interior. It covers also the 
peninsula of lower California, the southern part of which seems 
entitled to rank as an independent subdivision." 

Later it is stated that the region "may be divided by tempe- 
rature into two principal transcontinental zones, Upper Sonoran 
and Lower Sonoran ; and each of these in turn may be subdivided 
into arid and humid divisions." 

The proposal to form a separate region for such an insignifi- 
cant area as the southern extremity of California seems unnecessary, 
although its fauna may differ considerably from that of the typical 

Omitting mention of the bats, the mammalian genera charac- 

Characteris teristic of the Sonoran region may now be taken into 

tics of Mam- consideration. Commencing with the Insectivora, 

tnalian Fauna. , , .. 

the Sonadcz are represented by the peculiar genus 
Notiosorex, which is closely allied to the Oriental Soriculus, 
but has only 28 in place of 30 teeth. Of this genus the two 
species do not range north of this region, although they also 
enter Central America 1 . The short -tailed, or earless shrews 
(Blarina), with either 32 or 30 teeth, are also mainly Sonoran, 
although ranging northwards into the Holarctic, and southwards 
into Guatemala. In the Talpida the three species of the mole-like 
genus Scalops, characterised by having 36 teeth, webbed hind feet, 
and a short and nearly naked tail, are mainly Sonoran, although 
passing into the Transition zone. On the other hand, the two 
species of Scapanus, distinguished by the possession of 40 teeth, 
and the hairy tail, have a distribution very similar to Blarina, 
although they do not enter Central America. 

In the Carnivora the raccoon-family (Procyonidce) is very 
strongly represented, although none of the genera are absolutely 
peculiar to the region. The genus Bassariscus a near ally of the 
true raccoons, and possessing two species is nevertheless mainly 
Sonoran, although it ranges into the Transition zone of the 

1 Teste Dobson. 

X.] ITS FAUNA. 365 

Holarctic and also into Central America. The true raccoons, on 
the other hand, cannot be regarded as distinctive of the region, 
since they range from South America into the Canadian sub- 
region of the Holarctic ; and the coatis (Nasua) are now highly 
characteristic of the Neogaeic realm. Indeed Dr Merriam con- 
siders both genera as intruders from the latter realm, but this can 
scarcely be regarded as the correct view. The family is repre- 
sented in the two halves of the northern hemisphere (in the eastern 
by sElurus], in both of which it dates from the Pliocene, and, 
as it is unknown in South America till the Plistocene or late 
Pliocene, it is evidently one of northern origin ; the American 
forms having probably attained their maximum development in 
the Sonoran region. Much the same is the case with regard to 
the skunks among the Mustelidcz ; these being probably an original 
Sonoran type which has spread northwards into the Holarctic 
region and southwards into the Neogaeic realm. Of these, the 
single species of climbing skunk (Sptlogale) is mainly Sonoran, 
although it also enters the Transition zone of the Holarctic, and 
likewise Central America. Of the other members of the group, 
the typical skunks (Mephitis) range from Hudson Bay to Guate- 
mala ; while the allied genus Conepatus is found from Texas to 
Patagonia. In the same family the American badgers (Taxided), 
although ranging well into the Holarctic, are regarded by Dr 
Merriam as of Sonoran origin. These badgers, it may be ob- 
served, differ from the true badgers of the Old World by the form 
and characters of their cheek-teeth, the last upper molar being 
proportionately much smaller. 

Turning to the rodents, the well-known prairie -marmots 
( Cynomys], which occupy a position intermediate between the true 
marmots and susliks, are regarded by Dr Merriam as of Sonoran 
origin, although they extend into the Holarctic. In the Muridce 
the peculiar cricetine genus Rhithrodontomys which, together with 
the allied South American Rhithrodon, differs from the other 
members of the sub-family to which it belongs by its grooved 
upper incisors appears to be restricted to the region under con- 
sideration. The white-footed mice (Sitomys), although distributed 
over the whole of the New World, seem to attain their maximum 
specific development in the Sonoran, to which the two sub-genera 


Onychomys and Oryzomys are restricted. Yet their near alliance 
to the Old World hamsters indicates that the group must have 
had a northern origin, although the genus may have attained 
its present distinctive features within the Sonoran area. The 
genus Sigmodon, which differs from the last in the pattern of the 
molar teeth, and is represented solely by the rice-rat, does not 
range north of the Sonoran region, although extending into South 
America as far as Ecuador. The wood-rats (Neotoma), in which 
the molars simulate the prismatic appearance of those of the voles, 
.are also largely Sonoran, although they extend into the Canadian 





sub-region of the Holarctic, where they are represented by a 
distinct sub-genus (Teonoma). The round-tailed musk-rat of 
Florida (Neofiber) is an exclusively Sonoran type, although it is 
regarded by Dr Merriam merely as a sub-genus of Microtus. 
Highly characteristic of the region are the pouched rats, consti- 
tuting the genera GeomyidcR. Of these, the typical genus Geomys * 
extends northwards into the Transition zone and southwards into 
Central America ; while the nearly-allied Thomomys> in which the 
upper incisor teeth are smooth instead of grooved, penetrates 
into the Canadian sub-region of the Holarctic, although unknown 

1 Subdivided into eight genera by Merriam, North American Fauna, Part 
viii., Washington (1895). 

X.] ITS FAUNA. 367 

in Central America. Both these genera are represented in the 
Pliocene of the Sonoran area. In the same family the three 
genera of kangaroo-rats known as Dipodomys, Perodipus, and 
Microdipodops appear to be confined to the region ; and the same 
is the case with the allied genus Hderomys, although Perognathus 
passes northwards into the Transition zone. 


In the Ungulata, the deer belonging to the peculiar American 
genus Cariacus are very abundant in the Sonoran region (where 
those of the typical genus Cervus are entirely wanting), although 
they also range into the Canadian sub-region of the Holarctic, and 
extend right through South America. Since, however, they are 
wanting in the earlier Tertiary deposits of the latter area, as they 
are at all epochs in the Old World, there can be little hesitation in 
regarding them as essentially Sonoran types. Even more decidedly 
is this the case with the prongbuck (Antilocapra), the sole type of 
the family Antilocapridce, which is distinguished from the Bovidce. 
by the horn-sheaths of the males being branched and periodically 
shed from their bony supports. Although the prongbuck pene- 
trates a considerable distance into the Canadian sub-region of the 
Holarctic, its true home is the prairie-district of the Sonoran lying 

3 68 



to the westward of the Mississippi. Possibly a small deer-like 
animal from the Tertiaries of the same area known as Cosoryx, 
may have been the ancestral stock of the prongbuck. Lastly, the 
peccaries (Dicotyles), which are now chiefly South American, appear 
to have been originally Sonoran types which have migrated south- 
wards ; their fossil remains being common in the Tertiaries of the 
United States, whereas they are unknown in South America before 

FlG. 75. HEAD OF MALE MULE-DEER (Cariacus macrotis). 

the Plistocene. Their near affinity to the earlier Tertiary pigs of 
the Old World indicates that at a more remote date they spread 
from a more northerly starting-point. 

With regard to the armadillo (Tatusia) found in the Sonoran, 
this is clearly a very recent immigrant from the Neogseic realm ; 
and although opossums (Didelphys) were abundant in North 
America during the early portion of the Tertiary epoch, it is not 
improbable that the same explanation will hold good for their 
existing Sonoran representatives. 




The following list includes such exclusively New World genera 
of mammals (apart from bats) which are represented in the 
Sonoran area; those which may be regarded as more or less 
nearly confined to this region being printed in italics. To appre- 
ciate fully the significance of this list, reference must, however, be 
made to the series of Holarctic genera given on p. 360, which 

FIG. 76. HEAD OF MALE PRONGBUCK (Antilocapra americana). 

are more or less completely restricted to the Canadian sub-region 
of that great region, and the intervening Transition zone. The 
Sonoran list is as follows, viz. : 



Notiosorex. Also Central America. 
Blarina. Enters Canadian sub-region of Holarctic. 
L. 24 


Insectivora (cont.). 

Scalops. Enters Transition zone. 

Scapanus. Enters Canadian sub-region of Holarctic. 



Bassariscus. Enters Transition and Central America. 
Procyon. N. to S. America. 
Nasua. Also South American. 


Spilogale. Enters Transition and Central America. 

Conepatus. Texas to Patagonia. 

Mephitis. Extends into Canadian sub-region and 

Central America. 
Taxidea. Enters Holarctic. 


Cynomys. Extends into Holarctic. 



Sitomys. The whole of America. 

Sigmodon. Southwards to Ecuador. 

Neotoma. Ranges into Holarctic. 


Geomys. Extends into Transition zone and Central 

Thomomys. Ranges into Canadian sub-region. 




Perognathus. Ranges into Transition zone. 




Antilocapra. Ranges into Canadian sub-region. 


Ungulata (cont.). 

Cariacus. Greater part of America. 


Dicotyles. Also South American. 


Tatusia. South American. 



Didelphys. South American. 

Although the Transition zone undoubtedly forms an unsatis- 
factory item in regard to the distinctness of the Sonoran region, 
yet when we look at the difference of its mammalian fauna as a 
whole from that of the Canadian sub-region of the Holarctic, and 
the close similarity between the latter and the fauna of northern 
Europe and Asia, there can be but little hesitation in regard to 
the acceptance of Dr Merriam's view that the Sonoran is a valid 
zoological region of the Arctogaeic realm. 

In a previous chapter the groups of mammals, both living and 
extinct, confined to the eastern division of the 
Arctogaeic realm have been already noticed, while oraupa of 
in the present one reference has been made to such 

existing types as are restricted to the western half of Western 

, 6 /F . ., Arctogaea. 

of the same realm. It now remains to consider 
briefly some of the leading extinct groups which are found only in 
the latter area ; and the consideration of these comes most appro- 
priately here, seeing that the majority of these peculiarly American 
types are of Sonoran origin, a large number of their remains 
having been obtained from the States of New Mexico, Kansas, 
Nebraska, and Dakota, which lie within that region, or from 
Colorado, Wyoming, and Montana, which are situated within the 
Transition zone. 

Although, in common with the higher Primates, lemuroids are 
now quite unknown in North America, they were well represented 
there during the Puerco epoch of the lower Eocene by three 
families. The first of these the Chriacida includes animals 






having the same number of teeth as the allied Tertiary European 
family Adapidce, but all characterised by their more primitive 
structural features. Indeed these early lemuroids appear to 
present considerable resemblances to the creodont Carnivora, 
and differ from all the other members of the sub-order to which 
they belong by the great elongation of the bony symphysis con- 
necting the two branches of the lower jaw at the chin. Several 
other genera, in addition to the typical Chrtacis, are assigned 
to this family. The second group is that of the Anaptomorphidce, 
which is represented in the Puerco Eocene by a genus known as 
Indrodon, and in somewhat higher beds by the typical Anapto- 
morphus. Although in other respects coming closer to existing 
types than is the case with the Chriacidcz, the present family is 
broadly distinguished by the tritubercular structure of the upper 
molar teeth. The third peculiar North American family of the 
lemuroids is that of the Mixodectidcz, typically represented by 
Mixodectes of the Puerco Eocene. 

Among the extinct creodont Carnivora there are two families 
apparently restricted to the Tertiaries of North America, namely, 
the Miacidce, and the Mesonychidcz, the former of which presents 
such strongly marked affinities to the modern Carnivora that it 
is frequently assigned to that group. The second family, on the 
other hand, as represented typically by the genus Mesonyx of the 
Uinta or lowest Oligocene, is characterised by the very simple 
structure of the whole series of cheek-teeth, which are not unlike 
the pre-molars of some of the higher carnivores. One of the 
species of the typical genus attained dimensions as large as those 
of a bear. In the widely distributed family Hyanodontida, an 
exclusively North American genus is Patriofelis, which is regarded 
as a specialised offshoot from Oxyana. 

Among the ungulates there are several extinct families con- 
fined to North America. In the group forming a transition 
between the pigs and the ruminants there is first of all the family 
of the oreodonts (Cotylopida), which make their appearance in 
the middle Oligocene, and continue to the Miocene and lower 
Pliocene 1 . These ungulates, which were allied to the genus 

1 By American geologists the term Oligocene is not generally used, so that 
the whole of the Tertiary strata are classed as Plistocene, Pliocene, Miocene, 






Ancodus, common to the Tertiaries of both hemispheres 1 , and 
were represented by a large number of generic types, have 
crescentic columns to the short-crowned cheek-teeth, the upper 
molars usually carrying four such columns ; while the lower canine 
is approximated to the incisors, its usual form and function being 
assumed by the first pre-molar. The last upper pre-molar is 
simpler than the molars ; and while the feet have usually four toes 
each, in the typical genus Cotylops a rudiment of the thumb is 
retained in the front pair, as in Ancodus. In Cotylops and most 
of the other genera the molars of the upper jaw have but four 
columns, but in Protoreodon there are five ; a feature serving to 
connect the family with the Anthracotheriida, from which group 
the oreodonts are probably descended. A nearly-allied but more 
specialised family is that of the Agriochceridce, as represented by 
the genus Agriochcerus* , of the upper and middle Oligocene, in 
which the toes were developed into claws, instead of being 
incased in hoofs. Here it may be mentioned that while the 
peccaries (Dicotylida) are now exclusively New World types, and 
the pigs (Sutdce) restricted to the Old World, the Tertiaries of both 

and Eocene. Introducing the former term, the series may be approximately 
classified as follows, viz. : 

PLISTOCENE. Equus Beds. Eqtius^ Elephas primigenius. 

PLIOCENE (Upper). Blanco Series. Pliauchenia. 

(Lower). Loup-Fork. Protohippus, Hipparion. 

MIOCENE. Deep River. Anchitherium^ First Mastodons. 

OLIGOCENE (Upper). John Day. Miohippus, Ancodus. 

(Middle). White River. Agrwchcerus, Titanotherium. 

(Lower). Uinta. Amynodon, Mesonyx. 
EOCENE (Up. and Mid.). Bridger. Pachynolophus, Paltzosyops. 

(Lower). Wahsatch. Hyracotheritim, Coryphodon. 

(Lowest). Puerco. Neoplagiaulax , Poly mastodon. 

In this series the Deep River beds are identified with the European 
Miocene (supra^ p. 117) by the presence of Anchitheritim, and the first appear- 
ance of Mastodon ; while the existence of Ancodus in the John Day and Upper 
White River beds correlates them with the Upper and Middle Oligocene, 
Hyracotherium and Coryphodon serving to identify the Wahsatch beds with the 
Lower Eocene. 

1 Supra, p. 161. 

2 Equal Artionyx. 




hemispheres contain intermediate types such as Hyotherium and 
Chcerohyus which are apparently the ancestral stock of both 


Another very peculiar type of North American Tertiary ungu- 
lates is represented by Protoceras, from the upper division of the 
White River Oligocene, which forms a family (Protoceratidce} by 
itself. In these creatures the feet approximate to the ruminant 
type; but the skull, as shown in the accompanying illustration, 


has at least two pairs of large bony processes, probably covered in 
life with horns, and a pair of large upper tusks, in both of which 
respects it exhibits a curious parallelism with the perissodactyle 


ungulates. No trace of these singular artiodactyles has hitherto 
been detected in the Old World. 

The Camelidce. seem to have been primarily a North American 
family, which originated in the Sonoran region, and of which one 
branch (Lama) subsequently migrated south, while the other 
(Camelus) crossed Bering Strait into the Old World. In the 
upper Pliocene there occurs Pliauchenia, with only three lower 
pre-molars, and in the lower Pliocene Loup-Fork beds Procamelus 
with four of these teeth ; while the earliest representative of the 
family is Leptotragulus of the Uinta Oligocene. 

In the perissodactyle section of the same order the family 
Titanotheriidce is mainly North American, although, as stated on 
page 107, a representative of the typical genus Titanotherium has 
been discovered in the Tertiaries of the Balkans. Titanotherium 
includes huge rhinoceros-like animals, with low-crowned molar 


teeth of the type of those of Chalicotherium, and frequently having 
the nasal region of the skull surmounted by large bony protuber- 
ances. The genus is characteristic of the Uinta and the lower 
division of the White River Oligocene. An earlier type of the 
same family is typified by the smaller and less specialised hornless 
animals from the Bridger Eocene, known as Palaosyops, which, 
together with certain allied forms, constitute a peculiar sub-family 
confined to America. This family, like the camels, appears there- 
fore to have originated in the Sonoran region, whence a few 
representatives wandered eastwards into the Old World. 

In the generalised ungulate sub-order termed Amblypoda, 
of which the lower Eocene coryphodons were the earliest 
representatives, North America possesses an absolutely peculiar 


family in the UintatheriidcR. These were huge, somewhat ele- 
phantine ungulates, with five short toes to each foot, long tusk- 
like canine-teeth in the upper jaw, and the skull surmounted with 
three pairs of large bony protuberances ; their molar teeth being a 
specialised form of the Coryphodon type. They occur in the 
middle division of the Bridger, or the one above the zone yielding 
Coryphodon, and may accordingly have been the descendants of 
that genus. These uintatheres appear to have been restricted to 
the " Bad Lands " of the Sonoran region and adjacent districts of 
the Transition zone. 


Passing by certain other forms of less interest, attention may 
be directed to a peculiar group of aberrant mammals forming 
the Tillodontia, which appear to be mainly North American, and of 
which the serial position cannot be precisely determined. They 
are restricted to the Eocene, and seem to combine the characters 
of the modern Ungulata, Rodentia, and Carnivora. In the genus 
Anchippodus, forming the type of the family AnchippodontidcB, the 
skull approximates to that of a bear, the cheek-teeth are of an 
ungulate type, and there is a pair of large chisel-like incisors 
(preceded by a small functionless upper pair) in each jaw, very 
similar to those of the rodents and hyraces. A second family, 


Psittacotheriida, is represented by the genera Psittacotherium and 
Calamodon, in which the cheek-teeth grew permanently, instead of 
developing roots. 

The following list exhibits the chief families or minor groups 
of characteristically North American mammals, which are either 
entirely wanting or but sparingly represented in the Old World ; 
those that are extinct being indicated by a t, and the absolutely 
characteristic forms being printed in italics. 


t Anaptomorphidce. Anaptomorphus. 
t Mixodectida. Mixodectes. 
t Chriacidce. Chriads. 


Procyonidae. Represented in the Old World only by 


t Miaridce. Miacis, Didynrictis. 
t Mesonychidcz. Mesonyx. 
fHyaenodontidse. Patriofelis. 


Haplodon tidce. 


Dicotylidce. Represented by ancestral types in Tertiaries 

of E. Hemisphere. 

t Cotylopida. Cotylops, Mesoreodon, Protoreodon. 
t Agriochceridce. Agrioch&rus. 
t Protoceratida. Protoceras. 

Camelidae. t Pliauchenia, \Procamelus, M^eptotraguius. 

Antilocaprida. Antilocapra, t Cosoryx. 
t Titanotheriidae. Palceosyops, Limnohyops, Telmatotherium. 
t Uintatheriidce. Uintatherium. 


t Psittacotheriida. Psittacotherium, Calamodon. 
t A nchippodontidce. A nchippodus. 


In an earlier chapter 1 a list has been given of the leading 
Distinctness mammalian families common to the two divisions 
of the Region. o f Arctogaea, and since in the foregoing chapter it 
has been shown that many of the peculiar American families 
are more or less intimately related to some of those common to 
the two areas, it is manifest that throughout the Tertiary period 
eastern and western Arctogaea must have had a land-connection 
towards the north, so that there was an interchange of the fauna 
of the more northern districts. Those American types which 
penetrated as far south as what is now the Sonoran area would, 
however, naturally tend to become isolated, and thus develop 
into the families which may be regarded as characteristic of that 
region. So far, therefore, from this area being merely a part of a 
so-called Nearctic region, there are indications that it was differ- 
entiated from the Holarctic at a time when the existing zoological 
regions of the eastern half of the Arctogaeic realm were still unde- 
fined. Indeed from the community of the Pliocene fauna of 
southern Europe, Asia Minor, Persia, northern India, and south 
China, it seems probable that the only divisions of the Arctogaeic 
realm that could have been attempted would have been into (i) a 
Sonoran region, (2) a Holarctic region, comprising the northern 
districts of America, Asia, and Europe, (3) what may be termed a 
Mediterraneo-Oriental region, including southern Europe, north 
Africa, and the whole of southern Asia ; and (4) a Malagasy 
region, which would then, or perhaps somewhat earlier, have 
included Ethiopian Africa. 

As to the amount of interchange which took place during 
Dual Origin Tertiary times between the mammals of the eastern 
of Groups. and western divisions of Arctogaea, and as to whether 

similar generic types may have been developed independently in 
the two areas, it is almost impossible to arrive at any satisfactory 
conclusion. The suggestion that Equus has thus been independ- 
ently evolved in the two areas, has been already mentioned 2 , and 
this idea receives support from some very remarkable observations 
recently made on the invertebrates inhabiting certain European 
and North American caves. 

1 Sttpra, p. 176. 2 Supra, p. 168. 


With a quotation from Mr G. H. Carpenter's interesting 
paper 1 on this subject, the present volume may be fitly closed. 
After describing the inhabitants of the Mitchelstown Cave, in 
Ireland, the author writes that the spring-tail (Lipurd] "is hardly 
to be separated from a species found in the caves of Carniola, and 
the Sinella is almost identical with one inhabiting the caves of 
North' America ; while the spider is apparently the same as a cave- 
dweller from the Mediterranean district of southern France, which 
probably occurs in the North American caverns also. Had we to 
do with animals of the upper fauna, these results, though highly 
interesting, would not be without parallel in species already 
known.... But the occurrence of cave-dwelling species with so 
wide a range is a truly remarkable phenomenon. The caves 
cannot be of any great geological age. Any possible geographical 
connection which would permit the migration of subterranean 
animals between southern Europe and Ireland, or between Ireland 
and North America, seems altogether out of the question within 
any period during which the fauna can have been specifically 
identical with that of the present day. The only conclusion is 
that from ancestors, presumably of the same genus, which took to 
an underground life in such widely-separated localities, the similar 
conditions of the caves have evolved descendants so similar that, 
when compared, they cannot or can hardly be specifically distin- 
guished from each other. Should the identifications stand the 
test of a comparison of types, we shall have proof that the inde- 
pendent development of the same species, under similar conditions, 
but in widely-distant localities, has taken place. It must be 
granted, however, that cave-conditions are so marked and excep- 
tional, that it might not be safe to argue from them as to what 
may have occurred in the upper world." 

Although the author of this passage is perfectly correct in his 
statement that there is a vast difference between cave-life and 
open-air life, yet if animals which appear to belong to one and the 
same species can be proved to have had a dual origin in the one 
case, it can scarcely be considered impossible that similar instances 
may occur in the other. And if such dual origins exist among 

1 Irish Naturalist, Vol. IV. pp. 25 35 (1895). 


species, there is surely no reason why they should not occasionally 
occur in the case of genera. It would, therefore, seem by no 
means improbable that the species of the genus Equus which 
inhabited the eastern and western halves of the northern hemi- 
sphere during the close of the Tertiary period may have been 
evolved from the closely-allied but separate ancestral equine 

The matter does not, however, by any means end here. In 
an earlier chapter 1 it has been shown that the same species of a 
genus of fish (Galaxias) occurs in countries so remote from one 
another as New Zealand and Australia on the one hand, and 
Patagonia on the other. With the evidence of the cave-animals 
before us, is it absolutely impossible that these apparently iden- 
tical fishes can have been evolved independently of each other ? 
Should this be so, it will engender increased caution in drawing 
any inferences as to former land-connection from the evidence 
of single animals. But such instances of independent evolution, 
if they do occur, must be of extreme rarity, and will in no case 
interfere with deductions drawn from the presence of a number 
of allied species or genera of animals in distant countries. 

1 Suprh, p. 125. Recently another species has been described from South 
Africa. See Steindachner, SB. Ac. Wien, vol. ciii. p. 460 (1894). 



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L. 25 


Aard-varks, 187, 249, 256, 268 

Aard-wolf, 235 

Abderites, in 

Abderitidcz, 1 1 1 

Acaremys, 90 

Acdestis, no 

Acomys, 198 

Ac other uhim.) 192 

Acrobates, 37 

Adapis, 181, 191, 219 

Addax, 245, 324, 358 

Adelphomys, 91 

^ 157, 191, 202 
$) 321 
212, 275 
sEpyceros, 245 
sEpyornis, 222 
sEpyprymnus, 36 
Age of animal groups, 7 
Agriochceridiz, 375 
Agriochceriis, 375 
Agutis, 89, 137 
Alactaga, 323, 351 
^&w, 164, 315 
Amblotherium, 53 
Amblypoda, 173, 377 
Amblyrhiza, 87, 137 
Ameghino, on age of Santa Cruz fauna, 


Ammodorcas, 245 
AmphictiSy 191 
Amphicyon, 158, 191 
.4 ?/>// icy nod on ,193 
Amphidozotherium, 156, 191 
Amphilestes, 53 
Amphiproviverra, 109 
Amphisbcend) 132 
Amphisbcenidtz, distribution of, 132; 

fossil, 132 
AmphitheriidcB, 52 
Amphitherium, 53 

itragulus, 1 94 

^? nalcitheriu m, 105 

Anaptomorphidce, 373 

Anaptomorphus, 373 

Anchilophus, 166, 192, 193 

Anchippodontid&i 378 

AnchippodttS) 378 

Anchithei'ium, 166, 196 

Ancodus, 161, 193, 203, 375 

Ancyclotherium, 172, 200 

Anoa, 47, 164 

Anomaluridce, 237 

Anomalurus, 238 

Anomodontia, 151 

AnoplotheriidcZ) 185 

Anoplotherium, 185, 193 

Anops, 132 

Antarctica, 56, 128; extent of, 129 

Antarctogsea, 26 

Anteaters, 101, 136 

^4 ntech inom ys, 3 9 

Anthracotheriida, 161, 192 

Anthracotherium,) 163, 192, 193, 194,. 


Anthropopithecus, 180, 202, 231, 256 
Antillia, 139 

Antillean sub-region, 136, 137 
Antilocapra, 367 
Antilocapridce, 367 
Antelopes, African, 242 
Amirosorex, 272 
Africa connected with S. America, 


Africa, route of migration into, 256 
Aplin, Mr, on mammals crossing 

rivers, 14 

Aquilonian region, 26 
Arabia, South-eastern, fauna of, 262 
Arapaima, 134 
Arch&lurus, 157 
Archceoniys, 91 



Arctictis, 273 

Arctic sub-region, 360 

Arctic zone, 310 

Arctocyon, 159 

Arctogale, 273 

Arctogsea, 25, 26, 27, 144, 210 

Arctogsea, Eastern, 179 

Arctogasan region, 26 

Arctogaeic fauna, features of, 147 

Arctogseic realm, 27, 144 

Arctotherium, 72 

Arctomys, 159, 267, 351 

Arc t onyx, 276 

Area, 2 

Armadillo, giant, 136 

Armadillos, 94, 368 

Artionyx, 375 

Arui, 35 

Arvicanthis, 239 

Asses, 359 

Astrapotheria, 81 

Astrapotheriutn, 81 

Atherura, 278 

Atlantic, its recent origin, 24 

Atlantosaurus, 35, 151 

Auk, 347 

Anlacodus, 91, 240 

Australia, how it received its fauna, 
54; its connection with S. America, 
55 125; i ts relations with New 
Zealand and other islands, 58, 59 

Australian region, 25, 27, 31 

Austro-Columbian region, 25 

Austro-Malayan region, 27, 28, 45 

Avahi, 217 

Awantibo, 256 

Aye-aye, 219 

Babirusa, 47 

Bachitherium, 164, 192 

Badgers, 321 

Balanoptera, 68 

Banded anteater, 40 

Bandicoots, 38 

Banteng, 278 

Barbets, 254 

Barriers to dispersal, 10, 14 

Bassariscus, 73, 136, 364 

Bathyergus, 23 r, 239 

B atomy s, 278 

Bats, Notogseic, 42 ; Neogseic, 70 

Baur, Dr, on relations of Galapagos 

islands, 141 
Bdeogale, 235 
Bears, American, 318; their absence 

from Ethiopian Africa, 258 
Beavers, 313, 318 

Beddard, on earth-worms, 126, 311; 
on Fernando Noronha, 140 

Bering Strait indicates a line of con- 
nection between Asia and America, 

178? 337 
Bettongia, 30 
Birds, Malagasy, 254; Ethiopian, 


Bison, 314 

Black-buck, 278 

Blanford, Dr, on permanency of conti- 
nents, 24 ; on zoological regions, 27 ; 
on a former connection between the 
southern continents, 128; on con- 
nection between Africa and S. 
America, 128; on survival of old 
forms in Madagascar, 131; on the 
birds of Madagascar and Africa, 
254 ; on Oriental sub-regions, 266 ; 
on the fauna of India, 267 ; on land 
connection between Ceylon and 
Malaysia, 293 ; on the Tibetan 
fauna, 354 

Blarina, 364 

Blastomeryx, 74 

Blesbok, 244 

Boa, 132 

Boas, distribution of, 132 

Bontebok, 244 

Boreal region, 146, 309, 360; zone, 

Borhycena, 109 

Borneo, its fauna, 294 

Bos, 165, 206, 278, 314; depressicornis, 
47, 164; mindorensis, 47 

Boselaphus, 206 

Both riospondyhis, 152 

Bothremys, 131 

Bothriceps, 152 

Bourbon, 213 

Bovidce, 164, 186, 196; Holarctic, 314; 
Oriental, 279 

Brachydiastematotherium, \ 70 

Brachyurns, \ 36 

Brachyteles, 136 

Bradypodida, 100 

Brady pus, 100 

Bramatherium , 186, 204 

Brazilian sub-region, 135 

British Isles, their connection with 
the Continent, 339 

Brontornis, 60 

Bubalis, 206, 244, 358 

Budorcas, 324 

Buffalo, 279 

Bulman, Mr, on the submergence of 
Britain, 350 




Buphaga, 254 

Burmese sub-region, 267 

Buried river-channels of Britain, 339 

Cadurcotherium^ 82, 170, 192 

Ccenolestes, no 

C&notheriidce, 163, 192 

Ccenotherium, 163, 192, 193 

Calamodon, 379 

Callithrix, 136 

Caloprymnus, 36 

CamelidcE, 163, 185, 326, 377; Neo- 

gseic, 74 

Camelus, 163, 185, 204, 377 
Camels, 326 
Campos of Brazil, 65 
Canadian sub-region, 360 
Cane-rat, 91 

Canidce, Oriental, 274; Neogaeic, 72 
Cam's, 195, 202, 235, 274; antarc- 

ticus, 140; dingo, ^i;frocyonides, 


Cannabateomys, 91 

Capra, 206, 230, 280, 324, 359, 360 

Capreolus, 352 

Capivara, 88 

Capromyida, 90 

Capromys, 91, 137 

Caracal, 249 

Cardiotherium, 89 

Carcttochelyidcz, 62 

Carettochelys, 62 

Cariacus, 74, 344, 367 

Carioderma, 96 

Carnivora, Arctogseic, 157; Holarctic, 
312, 321; Neogaeic, 71; North 
American, 341 ; Sonoran, 364 

Carpenter, Mr, on cave-faunas, 381 

Carpincho, 88, 136 

Carpotnys, 278 

Carterodon, 91 

Castor, 313 

Castorida, 159, 313 

Castoroides, 87, 138 

Castoroidida, 87, 138 

Cassowaries, 48, 58 

Casuarius, 48 

Catamarca beds, 67 

Catonyx, 105 

Caves of Lagoa Santa, 66 

Ctf^a, 88 

Cavies, 88 

Caviidcz, 88 

Cave-faunas, 381 

Caxomistle, 136 

Cebidce, 69 

Cebochczrus, 192 

Celebes, its fauna, 47; an anomalous 

island, 49 
Cewas, 280 
Centetes, 220 
Centetidce, 131, 220 
Central American sub-region, 136 
Central Asian sub-region, 351 
Centres of evolution, 2 r 
Cephalogale, 191 
Cephalophus, 244 
Ceratodns, 133 
Ceratophora, 120 
Cercocebus, 231 
Cercoleptes, 72 
Cercomys, 91 
Cercopithecida, 231, 269; distribution 

of, 180 

Cercopithecus, 231 
Cernaysian Fauna, 153 
Cervicapra, 245 
Cervicaprince, 244, 245 
Cervidce, 164; absence of in Ethiopian 

region, 230, 258; Holarctic, 315 ; 

Oriental, 280 
Cervulus, 198, 281 
CVrzw, 164, 280, 326, 335'.35 8 > 359> 

367; eustephanus, 352; titnoriensis , 

46; xanthopygus, 351 
Cestracion, 63 

Cetaceans of American Tertiary, 68 
Cetotherium, 68 
Ceylon, when separated from India, 

2 93 

Ceylonese sub-region, 266 
Chcerokyus, 376 
Chceromeryx, 203 
Chtzropotamida:, 161, 192 
Cheer opotamus, 161, 192 
Cheer optis, 39 

Characiniidce, distribution of, 134 
Chcetomys, go, 136 
Chalicomys, 159, 194, 196, 313 
Chalicotheriidce, 196, 200 
Chalicotherium, 170, 192, 200 
Chameleons, distribution of, 222 
Chamois, 324 
Chelonians of Notogsea, 62 
Chevrotains, 164, 281 
Chimarrogale, 272 
Chilian sub-region, 135 
Chimpanzees, 180, 231, 256 
Chinchillas, 89, 136 
Chinchillidee, 89 
Chipmunks, 313 
Chiromyidce, 181 
Chiromys, 219 
Chiropodomys, 278 



Chiroiiectes, 107 

Chiru, 325 

Chirnromys, 41 

Chlamydophorus, 94, 96, 137 

CJilamydotherium, 96 

CholcepuS) 10 1 

Chriacidie, 371 

C/iriacis, 373 

Christmas Island, 303 

Chromididiz, 134 

Chrotomys, 277 

Chrysochloridiz, 230, 234 

Ckrysochloris, 230, 234 

Civets, 182; Oriental, 273 

Clccnodon, 159 

Claviglis, 238 

Coatis, 72, 365 

Cobus, 200, 206, -245 

Cwlogenys, 89, 136 

Colics, 254 

Collide?, 254 

Colobus, 231 

Colymbus, 347 

Comoro Islands, 213 

Conepatus, 73, 365 

Condylura, 341 

Conihtrus, 41 

Connection between Africa and S. 

America, 127 ; Australia and S. 

America, 125 
Connochcetes, 244 
Continental islands, 10 
Continents, permanency of, 22 
Cooke, Mr, on S. American molluscs, 

124; on Antillean molluscs, 139; 

on N. American land Mollusca, 

Cope, Prof, on N. American fossil 

cetaceans, 68 
Coral his, 132 
Coryphodontidce, 174 
Coryphodon, 174, 378 
C0.SWJ.*-, 368 
Cotton-rat, 88 
Cotylopidce, 373 
Cotylops, 375 
Coypu, 91, 137 
Crateromys, 277 
Cricetince, 160, 183, 239 
Cricetomysy 239 
Cricetus, 160, 192, 196, 322 
Crocidura, 195, 272 
Crocodilus porosus, 62 
Crossarchus, 235 
Crossopus, 319 
Cryptobranchus, 35 
Cryptodira, distribution of, 8 

Cryptoprocta, 220 
Ctenodactylidcz, 90 
Ctenodactylus, 91, 212, 240, 323, 357, 

35 8 

Ctenomys, 90 
Cuniculus, 314 
Cuscuses, 36, 46 
Cusimanse, 235 
Cycloturus, 102 
Cyncelurus, 202, 234, 249, 272 
Cynic tis, 235 
Cynodictis, 158, 191 
Cynodon, 191, 193 
Cynopithecus niger, 47 
C>, 235, 274 
Cynogale, 273 
Cynomys, 342 

Dacrytherium, 185, 192 

Dactylomys, 91 

Dactylopsila, 37 

Dcedicurus, 98 

Damaliscus, 244 

Damuda-Talchir flora, 153 

Dapedoglossus, 134 

Daphcenus, 158 

Dasornis, 60 

Dasymys, 238 

Dasyprocta, 98, 137 

Dasyproctida, 89 

Dasypodidce, 94 

Dasypus, 94 

Dasyuridce, 39 ; Neogaeic, 108 

Dawkins, Prof. B., on the range of 

the mammoth, 334 
Decastis, no 
Deer, 164, 267, 367 ; Oriental, 280 ; 

their absence from Ethiopian 

Africa, 230, 258 
Deep-sea deposits in islands, 23 
Degu, 90 
Dendrogsea, 26 
Dendrogale, 270 
Dendrolagus, 36 
Dendromys, 238 
Deomys, 239 
Desman, 321 
Deserts as barriers, 15 
Diadiaphoriis , 80 
Diatryma, 60 
Dichobumts, 163, 192 
Dichodon, 192 
Dichodontidce, 185, 192 
Dicroceros, 196 
Dicotyles, 73, 368 
Dicotylida, 74, 375 
Dicynodonts, 151 



Didelphyidce, Neogseic, 107 
Didelphys, 51, roy, 192, 193, 194, 


Dingo, 42 
Dinocyon, 158, 195 
Dinomyidce, 89 
DinornithidfB, 58 
Dinosauria, 151 ; wide distribution 

of, 8 

DinotheriidcB) 187 

Dinotherium, 173, 187, 196, 200,206 
Dist<echurus, 37 
Dispersal, barriers to, 10 
Distribution of groups, 9 
Distributional Areas, 19 
Distribution of families, 20 ; genera, 

19 ; orders, 21 ; species, 19 
Diplomesodon t 320 
Dipodidce, 240, 342 ; not Hystrico- 

morpha, 159 ; E. Holarctic, 322 
Dipodomys, 367 
Diprotodon, 38 
Diprotodonts, 34 
Dipus, 322 
Diver, 347 

Dobson, Dr, on Dipodidiz, 159 
Dolichotis, 88, 136 
Dolichopithecus, 180 
Dorcatherium, 164, 186, 196, 197, 

198, 204, 242, 256 
Dorcopsis, 36 
Dremotherium, 194 
Dromatherium, 54 
Dormice, 322 ; African, 238 
Dorcatragus, 245 
Dromicid) 37 
Dromiciops, 107 
Dryolestes, 54 
Dryopithecus, 180 
Dual origin of groups, 380 
Duckbill, 32 
Duikerboks, 244 

Earth-worms, 126; of Patagonia al- 
lied to those of Australasia, 126 

East Central African sub - region, 

Eastern Arctogsea, 179; its Tertiary 
mammalian fauna, 190 

Echidna, 33 

EchidnidcBi 33 

Echinomys, 91, 136 

Echinops, 220 

Echinothrix, 48 

Edentata, 92 

Edentates, doubtful fossil forms, 187 

Effodientia, 187, 192 

Elands, 247 
ElaphodiiS) 326, 356 
Elasnwtherium, 333 
Elephas, 172, 206, 247, 282 
Elephants, 172 ; Stegodont, 172 ; 

African, 247 ; dwarf Maltese, 339 ; 

European Plistocene, 334 ; Japan- 

ese fossil, 356 
Elephantida:, Oriental, 282 
Elk, 164, 315 
El lolnus, 322 

Elotherium, 161, 192, 193 
Emballonuridce, S. American, 70 
Enhydriodon, 195, 203 
Endrina, 217 
Eocardia, 91 
Eocardiidce, 91 
Eodidelphys, 107 
Epanorthidce, no 
Epanorthus, no 
Epomopfioriis, 221, 232 
Eqttida, 165 ; S. American, 75 ; Ori- 

ental, 282 
Equus, 1 68, 206, 247, 282, 359; its 

dual origin, 380 
Erethizon, 90, 343 
Ericulus, 220 
Erinaceida, 181, 271 
Erinaceus, 271, 233 
Eriodes, 136 
Eriomys, 89, 136 
Ethiopia and India, their connection, 


Ethiopian Mammals, 230 
Ethiopian Region, 25, 227 ; its past 

history, 255 
Eucardiodon, 89 
Eucholczops, 107 
Euchoretes, 322, 351 
Eumeces, 357 
Eupetaurus, 322 
En pier es, 220 
European sub-region, 348 
Etismilus, 157, 191 
Eiitat2is, 94 
Eutheria, 34 

Everett, Mr, on Malayan fauna, 294 
Extinct mammals of Western Arcto- 

ga, 371 
Extinction of large mammals, 18 


Falkland Islands, 140 
Fallow deer, 326, 359 
Faunas, community of early, 148 
Felidcc, African, 234, 249 ; Holarctic, 
312 ; Neogaeic, 71 ; Oriental, 272 



Felis, 198, 202, 249; concolor, 71; 

MO**/, 351 ; megalotiS) 46 
Fernando Noronha, 140 
/W?r, 342 
Fish-eating rats, 88 
Fishes, community between those of 

Australasia and S. America, 125 
Flora, community of Southern Palseo- 

zoic, 153 

Flying phalangers, 37 
Flying-squirrels, African, 237 
Forbes, Dr, on Antarctica, 128, 129 
Forest-bed, its fauna, 335 
Fossa, 220 
Foxes, 317 

Galago, 217, 231 

Galaginft, 217 

Galapagos Islands, 140 ; Tortoises of, 


GaleopitheciiS) 268, 270 
Galerisctts, 237 
Galerix, 197, 270 
Galictis, 73 
Galidia, 220 
Galidictis, 220 
Garzoniidce, 1 1 r 
Gastornis, 60 
Gaur, 278 
Gazella, 199, 206, 245, 325, 358; 

gutturosa, 351 
Gazelles, 245 
Ge/oats, 192, 193 
Gemsbok, 245 
Genetta, 182, 321, 358 
Genets, 182 
Geogale, 219 
Geographical changes in Europe since 

the Plistocene, 337 
Geological sequence, 7 
Gcomytd&i 366 
Geomys, 366 
Georhychus, 239 
Gerbillince, 183 
Gerbillus, 357 
Giant Salamander, 356 
Giant birds of Patagonia, 60; the 

Eocene, 60 
Gibbons, 180, 269 
Giraffa, 186, 189, 204, 242 
Giraffidce, 186, 242 
Glacial epoch, 9; period, in America, 


Glossotherium, 105 
Glyptodon, 96 
Glyptodontidce, 96 
Gnus, 244 

Goats, 324 

Golunda, 278 

Gorilla, 180, 231 

Graphiurus, 238 

Greenland Falcon, 347 

Gregory, Dr, on communication be- 
tween Atlantic and Pacific, 1 39 ; on 
modern origin of Atlantic, 23, 135 

Grison, 73 

Groove-toothed mice, 88 

Ground-sloths, 102 

Guanaco, 74, 136 

Gundi, 357 

Gymnura, 271 

Gymnobelideus, 37 

Guto, 312 

Habrocoma, 90 

Hamsters, 160, 322 

Hapalemur, 219 

Hapalidtz, 69 

Hafalomys, 278 

Hapalotts, 41 

Haploceros, 165, 343 

Haplodon, 342 

Harnessed Antelopes, 246 

Harpyia, 42 

Hawaiian region, 27, 30, 45 

Hedgehogs, 181 

Hedley, Mr, on the isolation and con- 
nections of Australia, 59 

Hegetotherium, 85 

Heilprin, Dr, on migrations of mam- 
mals, 15; on zoological regions, 26, 
29 ; on the Boreal Region, 346 ; 
on the Boreal Fauna, 347 

Helaletes, 165 

Helicophora, 200 

Helictis, 276, 355 

Helladotherium, 186, 198 

Helmsley, Mr, on Galapagos flora, 

Helogale, 235 

Hemicentetes, 220 

Hemicyon, 195 

Hemigale, 273 

Hemigalidia, 220 

Hemimeryx, 203 

Hemipsalodon, 258 

Hemitragus, 206, 209, 230, 279, 288, 

Herpestes, 182, 235, 321, 358, 360 

Heteromys, 136, 367 

Himalayan sub-region, 266 

Hipparion, 167, 200, 206, 247 

Hippidiuni, 75 

Hippohyiis, 203 



Hippopotamida, 184, 241 

Hippopotamus, 204, 241, 256; cross- 
ing from Africa to Madagascar, 223 

Hippoligris, 247 

Hippotragus, 199, 206, 245 

Holarctic region, 26, 27; physical 
features and extent, 309 ; character- 
istics of the fauna, 311; its unity, 
316; eastern division, 319; Plisto- 
cene Fauna of the eastern division, 
328 ; western division, 340 

Holochilus, 88 

Homacodon, 163 

Homalodontotheriunt) 82, 170 

Homunculus, 69 

Hornbills, 254 

Horse, 317 

Horses, S. American, 75 

Hose, Mr., on Malayan fauna, 294 

Howorth, Sir H. H., on the mixture 
of northern and southern types of 
mammals in the Plistocene, 329 

Humidity, influence of, 5 

Hundes, mammalian remains in, 354 

Hunting-dog, 236 

Hunting-leopard, 249, 272 

Hildas, 91, 137 

Huxley, Prof., on zoological regions, 
25,28; on northern origin of south- 
ern birds, 130 

Hyana, 182, 198, 235, 249, 273, 321 

HycBnarctus, 158, 195, 198, 202, 321 

HyaenictiS) 182, 198 

Hycenidce, an Old World group, 182 

Hycenodon, 158, 192, 193, 203 

Hy&nodontida, 373 

Hydaspitherium, 186, 204 

Hydraspis, 132 

Hydrochcerus, 88, 136 

Hydromyin<z, 40 

Hydromys, 40 

Hydropotes, 326, 356 

Hylobates, 180, 269 

Hylomys, 271 

Hyopotamus, 161 

ffyopsodtts, 155 

Hyotherium, 194, 196, 203, 376 

Hyperodapedon, 63 

Hypertragulus, 164 

Hypocetus, 68 

Hypselornis, 60 

Hypsipetes, 254 

Hyraces, 248, 268 

Hyrachyus, 166, 192 

Hyracodon, no, 170 

Hyracoidea, relationship to toxodonts, 
85, 248 

Hyracotherium, 161, 165 
Hystricomorpha, their abundance in 

Neogaea, 88 
Hystricidce, S. American, 90; Ori- 

ental, 278 
Hystricinte, 184 
Hystrix, 196, 198, 203 ; javanica, 46 

Ibex, 359 
Iceland, 350 
IchthyomyS) 88 

Icticyon, 72, 136 

Ictitherium, 182, 198 

Ictonyx, 236 

Idiurus, 238 

Iguanidce, distribution, 132; fossil, 

Iguanadon, 151 

Iguanas, fossil in Europe, 62; of Fiji 

and Friendly Islands, 62 
India and Ethiopia, their connection, 

209, 257 

Indian region, 25 ; sub-region, 266 
Indo-Malayan sub-region, 266 
Indo- African region, 26 
Indo-Chinese sub-region, 266, 267 
Indrisince, 217 
Indrodon, 373 
Introduced mammals, 17 
Insectivora, Neoggeic, 70; Arctogaeic, 

156; Holarctic, 312; North Amer- 

ican, 341 ; Sonoran, 364 
Ireland, its connection with Britain, 


Irish deer, 326, 333 
Irrisoridce, 254 
IsectolophuS) 165 

Islands, continental, 60; oceanic, 10 
Isomys, 239 
IssiodoromyS) 89, 92 
Ixocincla, 254 

Java, distinctness of its fauna, 300 
Jerboa-rats, 41 
Jumping-mice, 342 
Jungle-cat, 249 

Kangaroo-rats, 366 

Kangaroos, 35 

Kinkajou, 72 

Kirby, Mr, on Holarctic Lepidoptera, 


Kudus, 246 
Kurtz, Dr, on Argentine fossil flora, 

Labyrinthodontia, 152 



Lagidium, 89, 136 

Lagomyidce, 159, 314 

Lagomys, 196, 314 

Lagopus, 347 

Lagorchestes, 36 

Lagostomatidce, 89 

Lagostomus, 89, 136 

Lagos tr op hits, 36 

Lagothrix, 136 

Lama, 74, 136, 377 

Lamprotornis, 254 

Langiirs, 269, 360 

Laniarius, 254 

Lanthanotheriiun, 195, 197, 270 

Zate*, 312, 356, 347 

Le Conte, Dr, on separation of N. and 
S. America, 118 

Leberon Fauna, 197 

Leith-Adams, Dr, on the Maltese 
Islands, 337 

Lcithia, 333 

Leopard, 249 

Lepidosiren, 133 

Lepidosirenida, distribution of, 133 

Leporida, 160; Holarctic, 314; Neo- 
gaeic, 92 

Leptarctiis, 72 

Leptodon, 200 

Leptomanis, 187, 192 

Leptomeryx, 164 

Leptotragtdus, 163, 377 

Lepus, 203 

Lemmings, 314 

Lemuroids, 181, 191; Arctogaeic, 155; 
character of, 216; early entrance 
into Ethiopian and Malagasy re- 
gions, 223; extinct, 371 

Leimiridce, 181, 217, 270 

Lemur, 217 

Libytherium, 186 

Limacomys, 238 

Linsanga, 235, 256, 273 

Lion, 249, 272 

Listriodon, 196, 203 

L^thocran^^ls, 245 

Litopterna, 75 

Lomaphorus, 28 

Lomvia, 347 

Loncheres, 91, 136 

Lophiodontidce, 165, 192 

I^ophiodon, 165, 187 

Lophiomys, 239 

Lophiomeryx, 192 

Lophuromys, 239 

Z0rw, 256, 270 

Loxomylus, 87, 137 

Lutra, 202 

Lycaon, 236 
Lynxes, 312, 317 

Macacus, 180, 202, 269, 355, 357,. 

360 ; cynomolgus, 46 ; maurus, 47 
Machcerodus, 157,195,202; Neogseic r 

7 1 

Macropus, 35 

Macrauchenia, 77 

Macropodidce , 35 

Macroscelides, 233, 357 

Macroscelididce, 233, 268; fossil, 191 

Macrotherium, 172, 196, 200 

Madagascar, 213 

Madoqtia, 244 

Malabar sub-region, 266 

Malacomys, 238 

Malagasy region, 26, 213; mammals, 
214; molluscs, 223 

Malayan sub-region, 267, 294 

Malaysia and W. Africa, affinity of 
faunas, 292 

Malta, part of the mainland, 337 

Mammals, the oldest, 9; importance 
of, 9 ; classification of, 9 ; swimming 
powers of, 13; means of dispersal of, 
13; introduced by man, 17; extinc- 
tion of, 18; of Oriental region, 267 

Mammoth, 317 

Man-like apes, distribution of, 180 

Manidce, 187; Oriental, 283 

Mam's, 187, 249, 284 

Man's influence on distribution, 16 

Maraga Fauna, 197 

Marmots, 267 

Marmosets, 69 

Marsupial mole, 40 

Marsupialia, 33 

Marsupials, 33; Notogseic, 34; pa- 
Iseontological history of, 50 ; fossil, 
51; survival in Asia, 55, 57; date 
of migration into Notogaea, 60; 
Neogaeic, 107 

Mantchurian sub-region, 355 

Martens, 318 

Mastacomys, 41 

Mastodon, 196, 200, 206, 172; S. 
American, 86 

Mauritius, 213 

Medio-Columbian region, 26 

Mediterranean region, 26, 319; sub- 
region, 357 

Megaladapis, 218 

Megalobatrachus, 356 

Megalonyx, 105 

Megalotheriida, 102 

Megalotherium, 103 



Metes, 321 

Mellivora, 202, 237, 256, 276 

Mehirsus, 202, 274 

Mentawi, 303 

Mephitis, 73, 365 

Mergulns, 347 

Merriam, Dr, on effects of tempera- 
ture, 4; on W. Indies, 140; on a 
Boreal region, 309; on the effects 
of a glacial period in N. America, 
345 ; on the Transition zone, 360, 
361 ; on the Sonoran region, 363 

MerioneS) 357 

MerycopotamuS) 203 

Mesohippus, 167 

Mesomys, 91 

Mesonychidce, 371 

Mesonyx, 373 

MesoptthecW) 1 80, 197, 200 

Mesosaurus^ 133 

Metatheria, 33 

Mexican sub-region, 136 

Miacidce, 373 

MicroMotheriida, 109 

Microbiotheriu m, 109 

Microgale, 220, 234 

Microchcerus, 155, 156, 181, 191 

Micropholis, 152 

Micropus, 352 

Microtinat) 160 

Microtus, 267, 313, 318 

Minas Geraes, its caves, 66 

Mixodectes, 373 

Mixodectidtz, 373 

Moas, 58 

Moles, 321 

Mole-rat, 322, 358 

Mole-voles, 322 

Moluccas, 46 ; their isolation, 49 

Mongolian Pliocene, 201 

Monkeys, S. American, 69 ; distri- 
bution and dual origin of, 179, 180 

Monte Hermoso, its mammaliferous 
beds, 67 

Monotremata, 32 

Monotremes, 32 ; date of migration 
into Notogaea, 60 

Morenia, 91 

Morosaurus, 152 

Moschus, 204, 325 

Mouflon, 358 

Mountains as barriers, 15 

Mule-deer, 368 

Multituberculata, 33 

Muntjacs, 281 

Muridce, 160, 183; Notogaeic, 41; 
African, 238; Holarctic, 313; E. 

Holarctic, 322 ; N. American, 342 ; 
Sonoran, 365 
Murince, 160; an Old World group, 

Muscardinus, 322 

Musk-deer, 325 
Musk-kangaroo, 36 
Musk-ox, 315, 318, 344 
Musk-rat, 342, 366 
Musk-shrews, 272 
Musophagidce, 254 
Mustela, 195, 198, 202, 236, 358 
Mustelzdce, 236; Neogseic, 73; Sono- 
ran, 365 

Mus, Notogaeic species, 41 
Mus armandvillei, 46 
Mydaus, 276 
Mylodon, 103 
My odes, 314 
Myogale, 321 
Myolagus, 196 
Myopotamus, 91, 136 
Myoscalops, 239 
Myosorex, 233 

Myoxidce, 322 ; African, 238 
Myoxus, 192, 196, 322 
Myrmecobius, 40 
Myrmecophaga, 102, 136 
Mynnecophagidce, 101 
Mystromys, 238 
Myxopoda, 132 

Nandinia, 235, 256 

Nannosciurus, 238, 277 

Nanotragus, 244 

Nasalis, 270 

Nasua, 72, 365 

Nearctic region, 25 

Necrogymmirus, 182, 191, 271 

Nectarinia, 254 

Necrolestes, 71 

NecrotnaniS) 187, 192 

Nemorhtedus, 280, 324 

Neofiber, 366 

Neogaea, 25, 27, 64; its mammali- 
ferous deposits, 66 ; its early fauna 
distinct, \\.i\ distinctness of its 
modern fauna, 123; its peculiar 
birds, 123; its land-molluscs, 124; 
connected with Notogaea, 125; con- 
nected with Afi-ica, 127 

Neogseic fauna, origin of, 124 

Neogaeic realm, 27, 64 ; its sub-regions, 


Neoplagiaulax, 156 
Neoremys, 91 
Neotoma, 342, 366 



Neotragus, 244 

Neotropical region, 25, 27, 64 

Nesocerodon, 92, 89, 192 

Nesocia, 203 

Nesodon, 83 

Nesotragus, 245 

Neumayr, Dr, on a land-bridge across 
Atlantic, 133 

NeurotricJnis, 312 

New Guinea, its mammals, 31 

New Siberian Islands, 348 

New Zealand mammals, 45 

New Zealand, its relations with Aus- 
tralia, 58, 59 

Nimravus, 157 

North America, affinity of its fauna 
with that of E. Asia, 57 ; its relation 
to Asia, 310 

North American fauna, 340; Tertia- 
ries, sequence of, 375 

Nothropus, 107 

Nothrotherium, 107 

Notiosorex, 364 

Notogcea, 25, 27, 28; its isolation 
comparatively modern, 57, 58 ; its 
chelonians, 62 ; its right to form a 
realm, 62 ; northern origin of its 
fauna, 62 

Notogseic Realm, 27, 28 

Notary ctes, 40 

Notoryctida, 40 

Nototheriuni) 38 

Novo-Zelanian region, 25 

Nyctea, 347 

Nyctereutes, 355 

Nycticebus, 256, 270 

Oceanic islands, 10 

Oceans, permanency of, 22 

Octodon, 90 

Octodontidce, 137,323; Neogaeic, 90; 

African, 240 
Oligocene fauna, 190 
Onohippidium* 75 
Onychogale, 36 
Onychomys, 366 
Opossums, 368; European fossil, 51 ; 

Neogaeic, 107 ; originally a northern 

group, 1 08 
Orangs, 180, 268 
Oreodonts, 373 
Oreopithecus, 181 
Oreotragus, 245 
Or ibid, 245 
Orias, 199, 206, 247 
Oriental Region, 25, 27, 264; its past 

history, 288 

Ornithorhynchus, 32 

Ornithorhynchida, 32 

Ornithogaea, 26 

Orthomys, 91 

Orycteropodidce, 187, 249 

Orycteropus, 187, 200, 249, 268 

Oryx, 199, 245, 358 

Oryzomys, 366 

Oryzorictes, 220 

Osborn and Earle, Messrs, on Puerco 

Fauna, 154 

OsteoglossidcE, distribution of, 134 
Osteoglossum, 134 

Ostriches, 255,256; distribution of, 129 
Otocyon, 202, 236 
Otomys, 238 
Ovibos, 165, 315, 344 
Ovt's, 165, 280, 314, 324, 358 
Ovis antmon, 352 ; hodgsoni, 352 ; 

poll) 352 

Oxycena, 192, 373 
Oxyodontotherium, 79 

Pacas, 89, 136 
Pachynolophus , 166, 192 
Pachyruchida, 85 
Pachyruchus, 85 
Pachyuromys, 238, 239 
Pala, 245 

Palsearctic region, 25, 26 
Palcecerinaceus, 182 
Palaeogsea, 25 
Palceomanis , 187 
PalcEomephitis, 198 
Palceomeryx, 196, 204 
Palaeontology, importance of, 6 
Palaoprionodori) 191 
Palceorias, 199 
Palaorycteropus, 187, 192 
Palczosyops, 170, 377 
Palawan group, 294 
PalcEotapirus, 165 
Palaotheriidce, 166, 192 
Palaotherium, 166, 187, 192, 193 
Palaotragus, 186, 199 
Palhyana, 182, 198 
Palorchestes, 36 
Pampas, 66 
Pampean beds, 66 
Panda, 212, 275 
Pangolins, 187, 249, 283 
Panochthus, 98 
Pantholops, 324 
Papio, 1 80, 202, 231, 256 
Paracetus, 68 

Paradoxurus, 256, 273 ; hermaphro- 
ditus, 46 ; muschenbroecki, 47 



Parana beds, 88 

Patagonian beds, 67 ; cavy, 136 

Patriofelis, 373 

Peccaries, 74/368, 375 

Pectinator , 91, 240 

Pedetes* 240 

Pelea, 245 

Pellegrinia, 91, 212, 240, 323 

Pelomedusa, 131 

Pelomedusidce, distribution of, 131, 


Peltephilus, 96, 112 
Pelycodus, 155 

Penguins, a southern group, 131 
Peragale, 38 
Peramelidce, 38 
Perameles, 38 
Peratherium, 51 
Perimys, 90 

Perissodactyles, Arctogseic, 165 
Permanency of continents and oceans, 


Perodicticus, 231, 256, 270 

Perodipus, 367 

Perognathjts, 367 

Petauroides , 37 

Petaurus, 37, 46 

Petrogale, 36 

Peiromys, 240 

Phacochcems, 185, 203, 242, 268 

Phalanger^ 36, 46 

Phalangeridie, 36 

Phalangers, 36 

Phascologale, 39 

Phascolomyidcc, 38 

Phascolomys, 38 

Phascolonus, 38 

Phascolotherium, 52 

Phenacodus, 166, 174 

Phenacomys, 342 

Philippine sub-region, 267 ; its fauna, 


Phlczomys, 277 
Phororhachis, 60 
Phosphorites, 191 
Phyllostomatidce, 70 
Picas, 314 

Pichiciagos, 94, 96, 137 
Pikermi Fauna, 197 
PithechiruS) 278 
Pithed a, 136 
Plagiaulax, 147 
Plagiodon, 91, 137 
Plantain-eaters, 254 
Platanistid<z, 112 
Plattmys, 132 
Platycercomys, 322 

Pkctrophanes, 347 
PlesiarctotnyS) 160 

Plesictis, 191, 193 

Plesiospermophilus, 192 

Plesiorycteropus, 249 

Pleurodira, distribution of, 8 ; 131 

Plexoch&rus , 89 

Pliauchenia, 163, 377 

Pliocene Fauna of Arctogsea, 197 

Pliolagostomus, 89 

Pliopithecus, 180 

Plohophorus, 98 

Podocnemis, 131 

Poebrotheriuni) 1 63 

Pcecilogale, 236 

Poiana, 235, 256 

Poly mastodon, 149 

Polynesian region, 27, 29, 45 

Polyprotodonts, 34 

Pontoporia, 112 

Pouched mice, 39 

Pouched rats, 136, 366 

Porcupines, 184 ; S. American, 90 

Potamogale, 234 

Potamogalidcc, 220, 234 

Pot or ous, 36 

Pottos, 231, 256, 270 

Prehensile-tailed mammals in Neo- 

gsea, 124 
Priodon, 94, 136 
Proboscideans, Neogaeic, 86 ; Arcto- 

goeic, 172 

Proboscis Monkey, 270 
Procamehts, 163, 377 
Procavia, 248 
Procaviidce, 248, 268 
Procoptodon, 36 
Procyon, 72 
Procyonid(Z, 364 ; Neogseic, 72 ; 

Oriental, 275 
ProdidelphyS) 107 
Prodremotherium, 164, 192 
Proechidna, 33 
Progenetta, 195 
Prolagostomus, 89 
Protneles, 198 
Prongbuck, 367 
PropalceohoplophontS) i oo 
Protalpa, 156, 321 
Protapirus, 165, 192 
Protechinomys, 1 92 
Proteleidce, 235 
Proteles, 235 
Proterotheriidce, 80 
Proterotherium, 80 
Prothylacinus, 40, 108 
Protoceras, 376 



Protoceratida, 376 
Protogonia, 174 
Protohippus, 168 
Protolabis, 163 
Protopterus, 133 
Ptotoreodon, 375 
Prototheria, 32 
Protragoceros, 199 
Proviverra, 159, 192, 193 
Pseudalurus, 195, 198 
Pseud h apalops, \ o 7 
Pscitdochirus, 37 
Psendocyon, 195 
Pseudorhynchocyon, 191, 233 
Pseudoscittrus , 192 
Psittaci, distribution of, 129 
Psittacotheriidce, 379 
Psiltacotherium, 379 
Psittacula, distribution of, 130 
Ptarmigan, 347 
Pterodon, 158, 192, 193 
Pteromys, 268, 277 
Pteropodidce, Notogseic, 42 ; distribu- 
tion of, 221, 225, 232 
Pteropiis, 221, 232, 268 
Ptilocercus, 270 
Puerco Fauna, 153 
Puma, 71 
Pyrotheria, 85 
Pyrotherium, 85 

Quagga, 247 

Quercy Phosphorites, 191 

Raccoons, 72, 364 

Rafts, 15 

Rangifer, 164, 315 

Raphiceros, 245 

Rat-kangaroos, 36 

Ratitse, distribution of, 129 

Ratite birds, their migration, 58, 60 

Realms, 25, 27 

Regions, 25, 27 

Reindeer, 164, 315 

Reptiles, antiquity of, 8 

Ratels, 237, 256 

Rhea, 130 

Rheebok, 245 

Rhinoceroses, 169; African, 247, 256; 

European Plistocene, 333 
Rhinoceros, 169, 192, 193, 206, 247, 

256, 281 

Rhinocerotutei 169; Oriental, 281 
Rhithrodon, 88, 140, 365 
Rhithrodontomys, 365 
Rhithrosciurus, 277 
Rhizomys, 183, 203, 240 

Rhynchocyon, 233 

Rhynchomys, 278 

Rhynchosaurns, 63 

Rietbok, 245 

Rio de la Plata, its characters, 67 

River-channels, buried, W. Indian, 


Rivers as barriers, 14 
Rocky Mountain Goat, 343 
Rodents, Arctogseic, 159; Ethiopian, 

237 ; Holarctic, 313; Neogseic, 86 ; 

North American, 342 ; Notogaeic, 

41 ; Oriental, 277 ; Sonoran, 365 
Riipicapra, 324 
Range of Carnivora, 5 
Ruscinomys, 91 

S. American Tertiaries, their age, 68 
S. America connected with Australia, 

125; connected with Africa, 127 
S. American element in Ethiopian 

fauna, 128 
Sable Antelope, 245 
Sab re- tooths, Neogaeic, 71 
Sahara, its action as a barrier, 259 
Saharan sub-region, 260 
Saiga, 324 
Sam bar, 280 
Samos Fauna, 197 
Samotheriiim, 186, 198 
Sandwich Islands, 45 
Santa Cruz beds, 67 ; fauna, its 

origin, 124 
Sarcophilus, 39 
Seal ops, 364 
Scapanus, 364 
Scaptochirus, 321 
Scaptonyx, 321 
Scelidotheriuin, 105 
Scharff, Dr, on the origin of the Irish 

fauna, 340 
Schizodon, 91 
SciuridcE, Holarctic, 313 
Sciuroides, 192 
SciurofteruS) 195, 277 
Sciurus, 195, 277 ; prevosti, 48 
Sclater, Dr, on zoological regions, 25, 

26; on the antelopes of Somaliland, 


Scleromys, 91 
Scotonycteris, 221, 232 
Scott, Prof, on origin of Canis, 158 ; 

on Dipodidce, 159; on the Equidtf, 


Sea-otter, 312 ; 356 
Secondary Reptiles, wide distribution 

of, 151 



Secretary-birds, 254 

Selvas, in 

Semioptera, 48 

SemnopithecuS) 180, 202, 269, 357, 


SerpentariuS) 254, 255 
Sewellels, 342 
Seychelles, 213; indicate a line of 

connection between India and 

Africa, 224 
Sheep, 324 
Shrews, 267, 272 
Side-necked tortoises, distribution of, 

131* 13* 

Siderolites, 191 

Sifaka, 217 

Sigmodotii 88, 366 

Szmia, 180, 202, 268 

Simiidcz, 268 

Simocyon, 198 

Siphneus, 322 

Sitomys, 88, 137, 160, 322, 342, 365 

Siwalik Fauna, 201 

Sivatherium, 186, 204 

Skunks, 73, 365 

Sminthopsis, 39 

Sminthus, 322, 359 

Snow-bunting, 347 

Snowy Owl, 347 

Solenodon, 70, 138 

Solenodontida, 70 

vSomaliland, its fauna, 261 

Sonoran region, 26, 27, 363 ; its dis- 
tinctness, 380 

Sorex, 191, 198, 267, 272, 312 

Soricidce, 136, 156, 319; Ethiopian, 
2 3 2 33 > Holarctic, 312; So- 
noran, 364 

Soriculus, 272 

South American region, 26 

South America, separated from N. 
America, 117; invaded by northern 
animals in the Pliocene, 119 

Southern Pateozoic Continent, 153 

South African sub-region, 261 

Spalacidce, 183, 239, 322 

Spalacopus, 91 

Spalacotheriidce, $2 

Spalax, 183, 322, 358 

Spencer, Mr, on buried West Indian 
river-channels, 138 

Spermophilus, 159, 351 

Spheniscida, a southern group, 131 

Sphenodon, 8, 63 

Sphodromys, 90 

Spiny anteater, 33 

Spiny rats, 136 

Spilogale, 365 

Sparassodonta, 108 

Squirrels, Flying, 237 

Star-nosed mole, 341 

Station, 2 

SteatomyS) 239 

Stegodont elephants, 206 

Stenodelphis, 1 1 2 

Stenoplesictis , 182, 191 

Stereornithes, 60 

Stereo stern um , 133 

Sternothoerus* 131 

Sthenurus, 36 

StiromyS) 90 

Stonestield slate, its marsupials, 52 

Strepsiceros, 199, 206, 246 

Strut hio, 255; distribution of, 129 

Submerged forests of Britain, 339 

Sucker-footed bats, distribution of. 

13 r 

Suida:, 184, 375; Oriental, 281 
Sumatra, 300 
Siiricata, 235 
Sus, 184, 203, 242, 267,281; absence 

of, in Ethiopian region, 230 ; cele- 

bensis, 48 ; papuensis, 42 
Swamp-deer, 280 
Swayne, Capt., on the fauna of So- 

maliland, 261 
Syria, formerly connected with Africa, 

Systemodott) 165 

Takin, 324 

Talpa, 156, 191, 271, 321 

Talpidce, 156, 271, 312; absence in 

Ethiopia, 230; Sonoran, 364 
Tamandua, 102 
Tamarao, 47, 279 
Tamias, 159, 313 
Tatniascvurus ) 344 
7\ipiridce, 165; Oriental, 282 
Tapirs, 74 

Tapirus, 73, 165, 194, 282 
Tarsiidce, 181, 270 
Tarsipes, 37 

TarsiuSi 2 70 ; fuscomanus, 47 
Tasmania, 31 
Tasmanian devil, 39 
Tatusia, 94, 368 
Tayra, 73 
Tchitrea, 254 
Telephomis, 254 
Temnocyon, 158 
Temperature, influence of, 3 
Tenrec, 220 
Teonoma, 344, 366 



Tertiary mammalian fauna of Eastern 

Arctogoea, 190 
Tesiudo, distribution of, 129 
Tetraceros, 206, 244, 280 
Tetraconodon, 163, 204 
Tetracus, 193 

Thames Valley, brick-earths of, 335 
Theridomyidic, 89, 91, 240 
Theridomys, 91, 192 
TheropithectiSi 231 
Thomas, Mr O., on the fauna of S. E. 

Arabia, 262 ; on the fauna of Sipora, 


Thomomys, 366 

ThylacimiS) 39 

Thylacoleo, 37 

Thyropoda, 132 

Tibetan sub-region, 352 

Tierra del Fuego, 140 

Tiger, 272 

Tillodontia, 378 

Timor, 46 

Titanomys, 194 

Titanosaurus, 152 

Titahotheriidce, 170, 377 

Titanotheritim, in the Balkans, 57, 

I7> 377 

Tobago, continental, 137 

Tolypeutes, 94 

Tortoises, Galapagos, 141 

Tosca, 66 

Toxodon, 82 

Toxodontia, 82 

Toxodontidce, 83 

Toxodontothcrium, 83 

Tragelaphus , 199, 246 

Tragocerosy 199 

Tragulida, 164, 196, 204, 242; Ori- 
ental, 281 

TraguluS) 164, 186, 204, 281 

Transition zone, 361 

Tratratratra, 219 

Tree-porcupines, 136 

Triaulacodus , 91, 240 

Trichechida, 313 

Triconodon, 51 

Triconodontida:, 51 

Trichosums, 36 

Trichys, 278 

Triclis, 36 

Trigonia, 63 

Trinidad, continental, 137 

Trilophomys, 238 

Tntylodon, 149 

Trogonidcz, distribution of, 129 

Trogontherium , 333 

Tropical region, 140 

Trygenycteris, 22 r, 232 

Tucotuco, 90 

Tup aia, 270 

Ttipaiidce, 268, 270 

Tuatera, 8, 63 

Ture, 348 

Tylas, 254 

Ty pother iidce, 84 

Typolherium, 84 

Tyrrhenian sub-region, 357 

Uintatheriidcz, 174, 378 

Ungulates, E. Holarctic, 324; Neo- 
gaeic, 73; N. American, 343 ; num- 
ber of African, 241; Oriental, 278; 
Sonoran, 367 

Uria, 347 

Uromys, 41 

Uronycteris, 42 

Uropsihts, 321 

Urotrichtis, 156, 312, 356 

Ursidce, their absence from Ethiopian 
Africa, 258; Holarctic, 321 

Ursus, 202, 274, 318; absence of, in 
Ethiopian region, 236 ; ornatus, 

Vampire bats, 70 

Vandeleuria, 278 

Varanus priscus, 62 

Vicunas, 74, 136 

Viscachas, 136 

Vishnutherium, 186, 204 

Viverra, 182, 191, 195, 234, 273; 

tangalunga, 47 
Viverricula, 273 
Viverridcz, African, 234; an Old 

World group, 182; Oriental, 273 
Voles, 160, 267, 313, 318 

W. Africa and Malaysia, affinity of 
faunas, 292 

Wallabies, 36 

Wallace, Dr, on extinction of Plistocene 
fauna, 18; on zoological regions, 25; 
on the Moluccas, 48 ; on Celebes, 48 ; 
on origin of Australian fauna, 54 ; 
on isolation of Australia, 58 ; on S. 
American fauna, 123 ; on connection 
between S. America and Australia, 
125; on W. Indies, 138; on con- 
nection of S. and N. America, 153 ; 
on the past history of Ethiopia, 
255; on land connection between 
India and Africa, 257; on Oriental 
sub-regions, 266; on relationship 
between W. African and Malayan 



faunas, 292 ; on Javan fauna, 302 ; 
on the former union of Malta and 
Sicily with Italy, 337 

Wallace's line, 10 

Walruses, 313 

Wapiti, 315 

Wart-hogs, 256, 268 

Water-buck, 245 

Water-chevrotain, 242 

Water-deer, 356 

Water-shrew, 319 

West Indian sub-region, 137 

West Indies connecting N. and S. 
America, 139 

Western Arctogsea, its extinct mam- 
mals, 371 

\Vhite-footed mice, 88, 342 

Wildebeests, 244 

Wolverene, 312 
Wombats, 38 
Wood-hoopoes, 254 
Wood-rats, 342, 344, 366 

Xenurus, 94 
Xeromys, 40, 277 
Xerus, 195, 197 
Xiphodon, 185, 192, 193 
Xotodon, 83 

Yak, 314 

Zaglossits, 33 

Zapus, 342 

Zebras, 247 

Zoological realms and regions, 25, 27 

Zoster ops, 254 



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