Sennreeneee seeeaeeonn i [oe - GEOGRAPHY OF MAMMALS -. bh @ = Fae Wie eee ole YOouNG FEMALE GORILLA (From an animal living in the Gardens of the Zoological Society of London, 1896.) [Garden-Guide, 1896.] : 46 7 T\ are ZT iiAiis ahelf - < Cane THE GEOGRAPHY OF MAMMALS BY WILLIAM LUTLEY SCLATER, M.A., F.Z.S. DIRECTOR OF THE SOUTH AFRICAN MUSEUM, CAPE TOWN AND PHILIP LUTLEY SCLATER, M.A., Pu.D., F.RB.S. SECRETARY TO THE ZOOLOGICAL SOCIETY OF LONDON “ Benedicite omnes bestie et pecora Domino: laudate et superexaltate eum in seecula.” LONDON KEGAN PAUL, TRENCH, TRUBNER & CO. L™ PATERNOSTER HOUSE, CHARING CROSS ROAD 1899 The rights of translation and of reproduction are reserved Printed by BALLANTYNE, Hanson & Co. At the Ballantyne Press PREFACE THE first seven chapters of this volume were written by my son, Mr, W. L. Sclater, and published in The Geographical Journal (1894-97). They are now reprinted here, with some slight alterations, by the kind permission of the Council of the Royal Geographical Society. The eighth chapter, on Marine Mammals, was read by me as a paper at a meeting of the Zoological Society of London on March 16, 1897, and was subsequently pub- lished in the Society’s Proceedings. It is re- produced here, with slight alterations, by the kind permission of that Society. The remaining chapters of this volume, in which the distribution of Mam- mals is treated of systematically, have been pre- pared by me specially for the present work. Of the fifty illustrations contained in the text, which have been selected to show some of the chief Mammals typical of the different Regions, forty have been drawn by Mr. J. Smit specially for the present work. Of the remaining ten, five have been kindly lent to us by the Zoological Society of London Vili PREFACE and five by Messrs. Adam and Charles Black, the publishers of Flower and Lydekker’s standard work on Mammals. It is hoped that most of the leading facts and the conclusions to be drawn therefrom as to the dis- tribution of existing Mammals have been correctly stated in the course of this work, and that it may be of some use to students of this most attractive branch of Zoology. At the same time, it should be recollected that science in all its branches moves fast nowadays, and that statements which are perfectly correct one day may be falsified at any moment by new discoveries and by fresh investi- gations. I must not conclude these remarks without offering our best thanks to Mr. W. E. de Winton, F.Z.S., and Mr. F. E. Beddard, F.R.S., F.Z.S., for their kind assistance in the compilation of the Tables of Genera, and in the correction of the proofs of this volume. P. L. SCLATER. 3 HANOVER Square, Lonpoyn, W., March 1, 1899. CONTENTS CHAPTER I PAGE INTRODUCTION : ; ; : : : P ; : 1 LIST OF THE earner AUTHORITIES REFERRED TO IN CHAPTER I. : ; : : : 3 5 ; ee. Les CHAPTER II THE AUSTRALIAN REGION— SECTION I. BOUNDARIES OF THE AUSTRALIAN REGION LS » II. GENERAL VIEW OF THE MAMMAL-FAUNA OF THE AUSTRALIAN REGION : 5 OI, » III. SUB-DIVISION OF THE AUSTRALIAN REGION “ail » IV. THE AUSTRAL SUB-REGION . - 5 : weve F V. ANALYSIS OF THE AUSTRAL MAMMAL-FAUNA . 33 » VWI, PAST HISTORY OF THE AUSTRAL SUB-REGION. 35 » VII. THE PAPUAN SUB-REGION . 5 : - 5 BxS » VIII. ANALYSIS OF THE PAPUAN MAMMAL-FAUNA . 42 5 LX. THE MAORIAN SUB-REGION . F : : Se) a X. THE POLYNESIAN SUB-REGION . ‘ ‘ eo » A&I. THE HAWAIIAN SUB-REGION : . « 249 LIST OF THE PRINCIPAL AUTHORITIES RuratenD TO IN CHAPTER II. ; ; ; 4 : F ; F 5 ED CHAPTER III THE NEOTROPICAL REGION— SEcTION J. BOUNDARIES OF THE NEOTROPICAL REGION . 52 » II, GENERAL VIEW OF THE MAMMAL-FAUNA OF THE NEOTROPICAL REGION ‘ : 5 da! » III. SUB-DIVISION OF THE NEOTROPICAL REGION . 65 ix xX CONTENTS SECTION IV. THE ANTILLEAN SUB-REGION , V, THE CENTRAL AMERICAN SUB-REGION , VI. THE GUIANO-BRAZILIAN SUB-REGION , VII. THE PATAGONIAN SUB-REGION . ; ; ; , Vill. THe PAst HISTORY OF THE NEOTROPICAL MAMMAL-FAUNA . , ‘ A : ; A LIST OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHAPTER III. . ; c ; F ; ; CHAPTER IV THE ETHIOPIAN REGION— SECTION I. BOUNDARIES OF THE ETHIOPIAN REGION , F II. GENERAL VIEW OF THE MAMMAL-FAUNA OF THE ETHIOPIAN REGION . ; ; ; , III. SUB-DIVISION OF THE ETHIOPIAN REGION ,» LV. THE MALAGASY SUB-REGION oa 3 V. THE WEST AFRICAN SUB-REGION , VI. THE CAPE SUB-REGION » WII. THE SAHARAN SUB-REGION . é , ; , VIII. THE Past HISTORY OF THE ETHIOPIAN MAMMAL-FAUNA ‘ , 5 i - f LIST OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHAPTER IV, CHAPTER V THE ORIENTAL REGION— SECTION I. BOUNDARIES OF THE ORIENTAL REGION , 7 » II. GENERAL VIEW OF THE MAMMAL-FAUNA OF THE ORIENTAL REGION ., , : » III. SUB-DIVISION OF THE ORIENTAL REGION . » 1V. THE INDIAN SUB-REGION a VY. THE BURMO-CHINESE SUB-REGION » VI, THE MALAYAN SUB-REGION , » VII. THE CELEBESIAN SUB-REGION : » VIII THE PAST HISTORY OF THE ORIENTAL MAMMAL- FAUNA : ; , ; : ; ¢ LIST OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHAPTER V. 84 88 98 101 110 113 117 119 122 123 125 131 134 137 141 146 149 152 CONTENTS CHAPTER VI THE NEARCTIC REGION— SEcTION I. 1 ” BOUNDARIES OF THE NEARCTIC REGION GENERAL VIEW OF THE MAMMAL-FAUNA OF THE NEARCTIC REGION . SUB-DIVISION OF THE NEARCTIC REGION . . THE CANADIAN OR COLD SUB-REGION . THE WESTERN OR ARID SUB-REGION . . THE EASTERN OR HUMID SUB-REGION . THE PAST HISTORY OF THE NEARCTIC MAMMAL- FAUNA LIST OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHAPTER VI. CHAPTER VII THE PALZARCTIC REGION— SECTION I. rT: ” BOUNDARIES OF THE PALMARCTIC REGION GENERAL VIEW OF THE MAMMAL-FAUNA OF THE PALMARCTIC REGION . SUB-DIVISION OF THE PALHARCTIC RRRION . THE EUROPASIAN SUB-REGION . THE EREMIAN SUB-REGION . . THE CHINESE SUB-REGION : 5 . THE PAst HISTORY OF THE Paranonte MAMMAL-FAUNA CHAPTER VIII MARINE MAMMALS— SEcTION I. INTRODUCTORY REMARKS 9 +e ] Il. Ill. LAE DISTRIBUTION OF PINNIPEDS DISTRIBUTION OF SIRENIANS DISTRIBUTION OF CETACEANS xl PAGE 153 154 161 164 167 169 171 176 177 178 185 187 189 191 193 Xil SECTION V. DISTRIBUTION SECTION I. DISTRIBUTION SECTION I. II. . DISTRIBUTION OF THE DOG-LIKE CARNIVORA . . DISTRIBUTION OF THE BEAR-LIKE CARNIVORA . . DISTRIBUTION OF THE MARINE CARNIVORA . SUMMARY AND DEDUCTIONS iVale Il. CONTENTS DIVISION OF THE MARINE AREA OF THE GLOBE INTO SEA-REGIONS . 5 : f ; THE NorRTH ATLANTIC SEA- Rawr OR ARCTAT- LANTIS . . THE MID-ATLANTIC SEA-REGION, OR MBESAT- LANTIS. . THE INDIAN SEA-REGION, OR INDOPELAGIA . . THE NORTH PACIFIC SEA-REGION, OR ARCTI- RENIA c j ; : . THE MID-PACIFIC ul: HnGiow OR Maawes . THE SOUTHERN SEA-REGION, OR NOTOPELAGIA . CONCLUSIONS . CHAPTER IX OF MONKEYS AND LEMURS— INTRODUCTORY REMARKS a : GENERAL DISTRIBUTION OF THE Quiswemens . DISTRIBUTION OF THE OLD WORLD MONKEYS . . DISTRIBUTION OF NEW WORLD MONKEYS. . DISTRIBUTION OF THE MARMOSETS . SUMMARY AND DEDUCTIONS . DISTRIBUTION OF THE LEMURS . . SUMMARY AND DEDUCTIONS CHAPTER X OF THE ORDER CARNIVORA— INTRODUCTORY REMARKS A : DISTRIBUTION OF THE CAT-LIKE CARNIVORA PAGE 208 209 210 211 212 213 214 215 218 219 220 229 232 233 235 237 239 240 245 247 250 251 CONTENTS xii CHAPTER XI DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS— Section I. INTRODUCTORY REMARKS . : ; ; ; “356 , II. DISTRIBUTION OF INSECTIVORES . : ; . 257 , III. DISTRIBUTION OF BATS— (a) INTRODUCTORY REMARKS , ; . 263 (b) CHIEF PoINTS IN THE wee ee OF BATS . - - 4 : : : 264 (c) SUMMARY AND DEDUCTIONS AS REGARDS THE ORDER CHIROPTERA c ; 2 200 , IV. DISTRIBUTION OF RODENTS. ‘ : ; . 270 . V. SUMMARY AND DEDUCTIONS ; : ; 3 200 CHAPTER XII DISTRIBUTION OF HYRAXES, ELEPHANTS, AND UNGULATES— Section I. DISTRIBUTION OF HyRAXES. é 5 : . 288 » II. DISTRIBUTION OF ELEPHANTS . - , . 284 , III. GENERAL DISTRIBUTION OF THE UNGULATES . 286 » LY. DISTRIBUTION OF RHINOCEROSES . . : « 200 # VY. DISTRIBUTION OF TAPIRS . ; : : . 288 , VI. DISTRIBUTION OF HORSES . : 5 ; . 289 5, WII. DISTRIBUTION OF BOVINE MAMMALS , : » 9290 , VIII. DISTRIBUTION OF PRONGBUCKS . : : . 293 ,» LIX. DISTRIBUTION OF GIRAFFES ; ; : . 294 ,» & DISTRIBUTION OF DEER : : : : . 295 , Al. DISTRIBUTION OF CHEVROTAINS . 4 2 5 PASE » XII, DISTRIBUTION OF CAMELS . : ; : . 298 ,, XIII. DISTRIBUTION OF SWINE-LIKE UNGULATES 5 Phe » XIV. SUMMARY AND DEDUCTIONS ; ; ; » 9302 CHAPTER XIII DISTRIBUTION OF THE CETACEANS AND SIRENIANS , me oUn DEDUCTIONS . ; 4 : A ‘ . : ; ‘ 5 BHO) XIV CONTENTS CHAPTER XIV DISTRIBUTION OF EDENTATE MAMMALS— PAGE SECTION I. INTRODUCTORY REMARKS . i c ; eat! II. DISTRIBUTION OF THE NEW WORLD EDENTATES 312 III. DISTRIBUTION OF THE OLD WORLD EDENTATES 314 IV. SUMMARY AND DEDUCTIONS ; é j sells 39 ” ” CHAPTER XV DISTRIBUTION OF MARSUPIALS AND MONOTREMES— SECTION J. INTRODUCTORY REMARKS . ; 3 4 . 318 II. DISTRIBUTION OF THE MARSUPIALS . ; ots! III. DISTRIBUTION OF DIPROTODONT MARSUPIALS . 319 IV. DISTRIBUTION OF POLYPROTODONT MARSUPIALS 321 ” ” is V. SUMMARY AND DEDUCTIONS é cs : . 324 » VI. DISTRIBUTION OF MONOTREMES . : ; + #826 DEDUCTIONS : : ; ; : ‘ ; ; : . 827 FINAL CONCLUSIONS . : : c 5 : E : - 328 INDEX . c ; : : é : : : : : ° . dol 3” LIST OF MAPS . Outline Map of the World, showing the six Regions of the Geographical Distribution of Mammals . Map of the Australian Region, showing its Division into five Sub-regions . . Map of the Neotropical Region, showing its Division into four Sub-regions . . Map of the Ethiopian Region, showing its Division into four Sub-regions. . Map of the Oriental Region, showing its Division into four Sub-regions . Map of the Nearctic Region, showing its Division into three Sub-regions . Map of the Palearctic Region, showing its Division into three Sub-regions - Outline Map of the World, showing the six Sea-regions To face page 16 ; 50 ” 82 53 122 , 152 ” 176 i, 196 5 216 b LIST OF ILLUSTRATIONS IN THE TEXT Gorilla, Young Female . Frontispiece THE AUSTRALIAN REGION PAGE Fic. 1. The Duek-bill (Ornithorhynchus anatinus) 23 », 2 The Thylacine (Thylacinus cynocephalus) 25 », 3 The Notoryet (Notoryctes typhlops) Al », 4. The Common Wombat (Phascolomys mitchelli) 28 », 5. The Red Kangaroo (Macropus rufus) 31 », 6. The Papuan Echidna (Proechidna bruijnt) 38 THE NEOTROPICAL REGION Fic. 7. The Quica Opossum (Didelphys opossum) 56 ve 8. Hoffmann’s Sloth (Cholopus hoffmannt) 57 » 9. The Great Ant-eater (IZyrmecophaga jubata) 58 , 10. The Three-banded Armadillo (Zolypeutes tricinctus) . 59 , Il. The Lama (Lama peruana) . 60 » 12. The Barrigudo (Lagothrix humboldti) . 64 THE ETHIOPIAN REGION Vic. 13. The Cape Aard-vark (Orycteropus capensis) 89 » 14 The White-bellied Pangolin (Manis tricuspis) 90 » 15. The Hippopotamus (Hippopotamus amphibius) . 91 ,» 16. The Giraffe (Girafia camelopardalis) 92 » 17. The Aard-wolf (Proteles cristatus) : 95 18. The Chimpanzee (Anthropopithecus troglodytes) 97 XV1 24, 25. 26. 27. 28. 30. 36. 37. 38. 39. 40. LIST OF ILLUSTRATIONS IN THE TEXT THE MALAGASY SUB-REGION . The Fossa (Cryptoprocta ferox) . The Tenree (Centetes ecaudatus) . . The Indris (Jndris brevicaudatus) - The Ruffed Lemur (Lemur varius) . The Aye-aye (Chiromys madagascariensis) . THE ORIENTAL REGION The Nylghaie (Boselaphus tragocamelus) The Gayal (Bibos frontalis) . The Binturong (Arctictis binturong) The Kaguan (Galeopithecus volans) The Orang (Simia satyrus) THE NEARCTIC REGION . The Virginian Opossum (Didelphys virginiana) . . The Prong-buck (Antilocapra americana) . . The Rocky-mountain Goat (Haploceros montanus) . The Musk Ox (Ovibos moschatus) . 3. The American Bison (Bison americanus) The Star-nosed Mole (Condylura cristata) . THE PALHARCTIC REGION The Bactrian Camel (Camelus bactrianus) . The Musk-deer (Moschus moschiferus) The Chamois (Rupicapra tragus) The Barbary Sheep (Ovis tragelaphus) The Aluropus (4/uropus melanoleucus) The Water-shrew (Crossopus fodiens) . xVil PAGE 102 104 105 106 107 126 127 128 130 144 —_— Oo GM SG — ~I 158 159 160 XVill LIST OF ILLUSTRATIONS IN THE TEXT MARINE MAMMALS . The Patagonian Sea-lion (Otaria jubata) . The Atlantic Walrus (Trichechus rosmarus) . The Common Seal (Phoca vituline) . The Manatee (Manatus americanus) . The Dugong (Halicore dugong) . The Right Whale (Balena mysticetus) . The Sperm Whale (Physeter macrocephalus) . The See-see (Platanista gangetica) - The Common Dolphin (Delphinus delphis) . . The Beluga (Delphinapterus leucas) THE GEOGRAPHY OF MAMMALS CHAPTER I INTRODUCTION (Puate I., p. 16) Ir has long been evident to naturalists that the ordinary political divisions of the earth’s surface do not corre- spond with those based on the geographical distribution of animal life. Europe, for instance, the most important of all the continents politically speaking, is for zoological geographers, as well as for physical, but a small fragment of Asia. Again, the strip of Africa which borders the Mediterranean and extends to the Sahara agrees closely, as regards its animal life, with Europe, and is altogether different from the great mass of the African continent. Proceeding to America we find that physical geographers, as well as political, divide the two great masses of the New World at Panama. But those who study distribution have ascertained that Central America and Southern Mexico belong zoologically to South America, and they are con- sequently obliged to place the line of demarcation much further north. Let us, therefore, dismiss from our minds for the moment the ordinary notions of both physical and political A a THE GEOGRAPHY OF MAMMALS geography, and consider how the earth’s surface may be most naturally divided into Primary Regions, taking the amount of similarity and dissimilarity of animal life as our sole guide. In order to endeavour to solve this problem, let us select the mammals, as the most highly organised and altogether the best-known group of the animal kingdom, and examine the geographical distribu- tion of this class of animals over the world’s surface. Mammals are divided by naturalists into eleven large groups, called “Orders.” As regards their distribution, however, these Orders fall into two very different categories, according as they live on land or in the water—terrestrial and marine. For out of the eleven Orders, one of the principal divisions of the Carnivora—the Pinnipeds or seals, and two other Orders in their entirety—(the Cetaceans or whales, and the Sirenians or Manatees) are specially adapted for existence in water. Land is, therefore, a barrier to their extension, whereas, on the contrary, in the case of the ordinary terrestrial mammals, land is the means by which they extend their ranges, and seas and rivers form their restraining boundaries. We will for the present put aside the marme mammals, and address ourselves to the discussion of the distribution of the nine terrestrial Orders, namely :— 1. Primates. 6. Ungulata. 2. Chiroptera. 7. Edentata. 3. Insectivora. 8. Marsupialia. 4, Carnivora. 9. Monotremata. 5, Rodentia. Now, as is generally agreed by naturalists, one of the most certain and best ascertained points in the classifica- tion of mammals is, that these nine Orders can be grouped INTRODUCTION a primarily in three natural divisions (which may, in fact, be considered as Sub-classes) of nearly equal value. These three Sub-classes are, as named by Professor Huxley—the Prototheria, embracing only the Order Monotremata— the Metatheria, equivalent to the Order Marsupialia, and the HLutheria, which includes all the remaining Orders from the Edentata to the Primates. Let us, therefore, consider the distribution of the members of these three Sub-classes on the earth’s surface. When we come to examine the ranges of these groups on the map, we shall find that the Monotremes are wholly confined to Australia and New Guinea; that the Marsupials pre- dominate in Australia, and are only met with elsewhere in South America (one or two species of Opossum occurring in North America, but being probably only recent in- truders from the south); and that the typical mammals or Eutheria occupy the rest of the world. Again, after examining the distribution of the seven Orders of typical mammals, we remark the following significant facts :— 1. The absence of Insectivores in South America. 2. The great prevalence of Edentates in the same country; the Sloths, Armadilloes, and Ant-eaters, consti- tuting three out of the five known families of this Order, being entirely confined to South and Central America. Taking these main facts as our guide, we may divide the land-surface of the Earth as follows, into three divisions :— (1) Land where Marsupials peeved); 10 Heater Australia, New Guinea) ,, Notogea. ians except Rodents and the adjacent islands. and Bats; Mono- tremes 4 THE GEOGRAPHY OF MAMMALS (2) Land where ma ians and Marsupials occur; no Insecti-| America south of the Isth- vores ; many Eden- | mus of Tehuantepec. tates; no Mono- tremes .. Ly eogad. (3) Land where Euther-) Europe, Asia, Africa, Asia- ians only occur ; few tic Islands down to Wal- Edentates, no Mar- lace’s line, and North } Arctogea. supials? nor Mono- America down to the tremes . : Isthmus of Tehuantepec. The fault of this division is that it leaves the great mass of land in the Northern Hemisphere undivided and rather unmanageable. But this northern land is easily separable into four sections, although it should be under- stood that these four sections are not of equivalent value to the two other primary divisions. Thus we obtain a division of the land-area of the globe for mammals into six areas, which are called Regions (see Plate I., p. 16), and which may be shortly defined and named as follows :— Australia, New Guinea, and adjacent ) ee Auiberte oe islands up to Wallace’s line . : egron. “< (Central America south of the Isthmus of Tehuantepec, the West Indies, and} II. Neotropical Region. South America F : EA NovToGMa. Nroa Africa south of the Atlas, and Madagascar III. Ethiopian Region. South Asia, the Philippines and Islands of the Indian’ Archipelago down to Wale} IV. Oriental Region. lace’s line, and Celebes . ; ARCTOGA. 1 As will be shown later on, this statement is not absolutely correct as regards North America, as at least one species of marsupial occurs within its limits. INTRODUCTION 5 (ver America down to the Isthmus ) ee ee r e vegvon. of Tehuantepec Europe, Africa north of the Atlas, and ) Northern Asia . (continued. ) ARCTOGHA VI. Palearctic Region. We will now take a brief survey of the principal features of these six regions—as shown in the accompany- ing chart (Plate I, p. 16) and their most characteristic mammal-forms. 1.—AUSTRALIAN REGION Extent.—Australia, New Guinea, and Moluccas up to Wallace’s line, New Zealand, and the numerous islands of the Pacific. | Characteristics.—Absence of nearly all Eutherian Mam- mals, except a few Rodents and Bats; presence of six distinct families of Marsupials with one hundred species, and the only two known forms of Monotremes. 2.—NEOTROPICAL REGION Name.—veos, new, and tpotrixos, 7.e., tropical land of the New World. Extent—America, south of the Isthmus of Tehuantepec, and the West Indies. Characteristics.—Monkeys of the peculiar families Cebide and Hapalide# ; absence of Frugivorous Bats, and presence of Vampires (Phyllostomatide); abundance of the Porcupine family; absence of Insectivores and Civets, also of Elephants; presence of Tapirs ; no Ru- minants except Deer and Lamas; presence of Sloths, Ant-eaters, and Armadilloes; one family of Marsupials— Opossums. 6 THE GEOGRAPHY OF MAMMALS 3.—ETHIOPIAN REGION Name.— AiOiores, ancient name for negroes. Extent.—Africa, south of the Atlas; Arabia up to the Persian Gulf, and Madagascar. Characteristics. — Chimpanzee and other Monkeys; absence of Bears and Deer; presence of Lion, African Elephant, Hyrax, Rhinoceros, Hippopotamus, Wart-hog, numerous Antelopes, Giraffe, Pangolin, Ant-bear—general richness in large and highly-organised Ungulates. 4,—ORIENTAL REGION Extent.—Southern Asia, south of the Palzearctic Region, and islands of Indian Archipelago down to Wallace’s line, including Celebes. Characteristics—Orangs, Gibbons, and other peculiar Monkeys. Flying Lemur, Tiger, and other cats, Indian Elephant, Rhinoceros, Malayan Tapir, Manis. Generally, it may be said that the peculiar forms of the Oriental Region are fewer than in the Ethiopian Region, and that the Oriental Region has Bears, Deer, and Tapirs, which are wanting in the latter. 5.—NeEaARcTIC REGION Name.—veos, new, and dpx«tos, north, 2e., northern district of the New World. Extent—North America, down to the Isthmus of Tehuantepec. Characteristics.—General mammal-fauna, very like that of the Palearctic Region, but mixed up with endemic INTRODUCTION mw forms and intruders from the south. Bears, Beavers, Sheep, and Deer similar; Prong- buck, Pouched Mice, and Musquash peculiar; Raccoon and Opossum, probably derived from the south. 6.—PALHARCTIC REGION Name.—rraxatos, ancient, and a@pxtos, north, as embrac- ing the whole northern area of the Old World. Extent—Whole of the Eastern Hemisphere north of a line on the south of the Atlas, and running eastward through the south of Palestine and Persia, along the Himalayas, through Central Asia and the centre of China to the Pacific. Characteristics.—Absence of Monkeys, Lemurs, and Frugivorous Bats; abundance of Carnivores—Ounce, Lynxes, Wolves, Foxes, Bears, and Weasels; Rodents— Marmots, Beavers, Pikas; Ungulates — Sheep, Deer, Chamois, and Musk-deer; no Elephants nor Hyrax. This division of the Earth’s surface into six regions was first proposed by one of the authors of the present work in an essay on the distribution of the Class of birds read before the Linnzan Society in 1857 (9). It was further elaborated and upheld in an address given to Section D. of the British Association at the Bristol Meeting in 1875 (10), and in a Paper published in The Jbis in 1891 (11). The same system was adopted by Mr. Wallace in his standard work on “Geographical Distribution” (13), and was there shown to be applicable to the other principal groups of terrestrial animals. Moreover, the names then bestowed on the six great primary Regions are now in general use among naturalists in all countries. Mr. 8 THE GEOGRAPHY OF MAMMALS Wallace, who has devoted many pages to the discussion of this subject,! has come to the conclusion that, admitting that these six regions are not precisely equal in rank, and that some of them are more isolated than the others, they are in geographical equality, compactness of area, and facility of definition beyond all comparison better than any others which have been suggested for the purpose of facilitating the study of geographical distribution. Notwithstanding Mr. Wallace’s strong support, how- ever, it is right to say that this system has not been univer- sally accepted. Professor Huxley (6) in 1868 proposed to separate the world into two divisions—Arctogeea and Notogea, the former containing the Nearctic, Palearctic, | Ethiopian, and Oriental Regions, and the latter the Neo- tropical and Australian Regions. He adopted the Nearctic, Palearctic, Ethiopian, and Oriental Regions as subdivisions of Arctogzea, and only stipulated for the formation of a Circum-polar province independent of the Nearctic and Palearctic Regions. Notogza Professor Huxley divided into three provinces (a) the Austro-Columbian (=the Neotropical), (b) the Australian (=the Australian Region minus New Zealand), and (c) the New Zealand province. From this it will be seen that Professor Huxley’s scheme does not really diverge materially from the system here employed; the chief points of difference being (a) the uniting together of the Australian and Neotropical Regions into Notogzea; (b) the formation of independent Circum- polar and New Zealand provinces. With regard to the first point, almost the only bond of union between the Australian and Neotropical Regions, so far as mammals, at any rate, are concerned, is the presence of Marsupials 1 “Geographical Distribution of Animals,” vol. i., chap. iv. INTRODUCTION 9 in both regions. But the Marsupials of Australia seem to have but a very remote connection with those of South America, and there is at present no palzeontological evi- dence of the former occurrence of the Australian forms, or of forms allied to them, outside of Australia itself On the other hand, the presence of fossil opossums (Didel- phyidx) in the Eocene beds of France, shows that the South American forms were formerly more widely spread. Professor Huxley has also cited the Parrots (Psittaco- morphx) “as helping, together with the three-toed Ratite, to bind together the widely-separated portions of the south world.” But on referring to the account of the distribu- tion of the Parrots in Salvadori’s recently published cata- logue (8), it will be found that out of the six families into which he divides the group, five are practically confined to the Australian Region, and that the remaining one is widely spread throughout the tropical regions of both hemispheres. The most recent arrangement of this family, therefore, gives little support to Professor Huxley’s arguments. Looking, again, to the distribution of the Ratitz (wingless birds), we find the Neotropical form (the Rhea) more closely connected with the Ostrich, the Ethiopian form, and that they both differ considerably from the Emus, Cassowaries, and Kiwis, the three Australian repre- sentatives of this order. Thus, then, there seems to be 1 Recently Senor Ameghino has described from the Santa Cruz beds of Patagonia, which are probably of Eocene age, certain fossil mammals which he has referred to the Dasyuride, one of the Australian families. Again, Mr, Thomas’ Cenolestes (see P. Z. S. 1895), is also believed to be allied to the Australian Diprotodonts. If these relationships should turn out to be correct, it will indicate further evidence of some connection between South America and Australia, though at a considerably remote epoch of geological time. 10 THE GEOGRAPHY OF MAMMALS scarcely any ground for connecting the Neotropical and Australian Regions under one name. Before discussing the other differences between this scheme and that of Huxley, it will be as well to mention the diverging views of some other naturalists. Of these the chief is Professor Heilprin, of Philadelphia, who in his “Geographical and Geological Distribution of Animals,” (5) “in accordance with a suggestion by Professor Newton,” has proposed to unite the Nearctic and Palearctic Regions into a single realm—the “Holarctic”—and_ to separate the Pacific Islands from Australia as the “Poly- nesian Realm.” Again, Mr. J. A. Allen, of New York, in his recently published essays (1 and 2), has shown con- siderable independence of thought in this matter. In the introduction to the later of them, which deals chiefly with the distribution of North American mammals, Mr. Allen gives an account of the influences which, in his opinion, mainly determine the geographical distribution of life, dwelling first on the great importance of temperature and moisture, and afterwards on the inter-relation of land- areas, which, he says, is “ co-eval and perhaps more than co-ordinate with climate in its influence upon the distribu- tion of life.” Next, Mr. Allen treats of the seven primary life-regions, or “realms” as he terms them, into which he proposes to divide the Earth. These are :— 1. An Arctic Realm, occupying all the country in both hemispheres north of the isotherm 32° F., this boundary corresponding very closely to that of the northern limit of trees. 2. A North Temperate Realm, occupying the whole of the northern hemisphere between the isotherms of 32° and 70° F. INTRODUCTION 11 3. An American Tropical Realm, consisting of Tropical America. 4, An Indo-African Realm, consisting of Africa, except the northern border, and Tropical Asia and its islands. 5. A South American Temperate Realm, embracing extra-tropical South America. 6. An Australian Realm, equivalent to our Australian Region. 7. A Lemurian Realm, contaming Madagascar and its islands. Mr. Allen’s views on Distribution have been criticised and answered by another American naturalist, Mr. Gill (4), who has proposed a division of the Earth into nine “realms.” These, as will be seen, although not differing in many cases from regions adopted by former authorities, are distinguished by an entirely new set of names, as follows :— (1) The Anglo-gzean (= Nearctic Region). (2) The Eury-gzean (= Palearctic Region). (3) The Indo-gzean (= Oriental Region). (4) The Afro-gzean (= Ethiopian Region). (5) The Dendro-gzean (=the tropical half of the Neo- tropical Region). (6) The Amphi-geean (=the temperate half of the Neotropical Region). (7) The Austro-gean (= Australia, New Guinea, and the adjacent islands). (8) The Ornitho-gzean (= New Zealand). (9) The Neso-gzean (= Polynesia). Dr. Bowdler Sharpe (12) has also recently published his views on the zoo-geographical areas, as worked out from the distribution of birds. Dealing here only with the division of the Earth into Regions, we notice that 12 THE GEOGRAPHY OF MAMMALS although he makes many complimentary allusions. to Mr. Allen and his views, he employs in nearly all its entirety the system adopted in this work, with the exception that he recognises an Arctic Sub-region to include the more northerly parts of both the Old and New worlds. Finally, Professor Newton, who has given us his views on this subject as regards birds (7), adopts the method of divisions followed here with the two following exceptions. In conformity with the suggestion already made to Professor Heilprin, he unites the Palearctic and Nearctic Regions under the title “ Holarctic,” and he also separates New Zealand from Australia as an independent region. The chief questions in dispute, therefore, seem to be as follows :— (1) Whether the Palearctic and Nearctic Regions are to be recognised as separate ? (2) Whether Madagascar and New Zealand are to be separated as independent regions from the Ethiopian and Australian Regions respectively ? (3) Whether the Ethiopian and Oriental Regions should be joined to form one region ? (4) Whether there are any good grounds for dividing the Neotropical into two separate regions. The only way in which questions of this sort could be settled would be by constructing accurate lists of the families and genera of the various classes of the terres- trial faunas of the regions in dispute, and then carefully comparing them, in order to determine the percentage of peculiar species and of absentees. The difficulty of doing this satisfactorily is twofold. (1) The absence of any definite boundaries to most of INTRODUCTION ales the regions, and hence the difficulty in determining how many of the border-forms, which have obviously intruded from the neighbouring regions, should be counted. (2) The uncertainty as to the limits of the genera. This uncertainty has been greatly increased of late years by the action of some zoologists in proposing a multitude of unnecessary generic terms. When these two factors have been settled and the lists constructed, a further difficulty is met with, and this is one which depends very much on the individual fancy of the author, namely, as to the percentage of peculiarity which should be required to constitute a Region. Taking the first question in dispute, we find that Mr. Allen, in his paper already quoted (2), gives a tabulated list of the genera of his North Temperate realm, dividing them into North American and Eur-asiatic (= Palearctic) forms, and putting the individual genera into three cate- gories, namely, those circum-polar, or common to the Nearec- tic and Palearctic Regions (numbering thirty-two); those peculiar to each Region (7.e. twenty-nine to the Nearctic and forty-one to the Palearctic); and, finally, those which range further south into the Neotropical Region on the one hand, and into the Oriental and Ethiopian Regions on the other. Working from these tables we find that 38 per cent. of the Nearctic genera and 42 per cent. of the Palearctic genera are confined to their respective Regions, while 42 per cent. in the case of the Nearctic and 34 per cent. in the case of the Palearctic are common to the two regions. These last percentages include, however, several quite wide- spread genera which can hardly be called cireum-polar—such as Sevurus, Sciwropterus, Lepus, Lutra, Canis, and Felis. These figures show that there is, as has indeed never 14 THE GEOGRAPHY OF MAMMALS been disputed, a great amount of similarity between the Nearctic and Palearctic faunas, but not enough to justify the junction of these two great land-masses into one “Region” or “ Realm.” As for the so-called “ Arctic realm,” which consists of the land bordering the Polar Ocean and Hudson’s Bay and the great peninsula of Greenland, “beyond the limit of arboreal vegetation,” together with the similar Arctic por- tion of the old world, Mr. Allen states, no doubt correctly, that it contains a “homogeneous hyperborean fauna of circum-polar distribution.” But looking to the extreme poverty of life in these inclement latitudes, as Mr. Allen well puts it, it seems to be quite unnecessary to elevate this wretched fraction of the Earth’s surface into one of its principal constituent life-regions. The plan adopted by Mr. Wallace, of regarding it as a borderland between the Nearctic and Palearctic Regions is far preferable. The question of the recognition of Madagascar and New Zealand as independent regions will be further discussed in the articles on the Ethiopian and Australian Regions respec- tively, as will also the propriety of dividing the Neotropical into two separate regions. A few words, however, may be said here with regard to Mr. Allen’s proposal to join together the Oriental and Ethiopian Regions into one “ Realm.” According to the estimate given below (Table L, p. 16), the total number of genera found in the Oriental Region is 113, and of these thirty-nine are not found elsewhere. Of the balance—seventy-four—eight only are common to the Oriental and Ethiopian Regions, and are not found in any other Region,! whereas twenty-eight more, also known 1 These are Manis, Rhinoceros, Elephas, Golunda, Atherura, Viverra, Mellivora, and Nycteris. INTRODUCTION 15 in the Oriental and Ethiopian Regions, are likewise found in the Palzearctic Region. Furthermore, out of the eight genera above mentioned, although they are not now found in the Palearctic Region, four of them are known to have existed there during the Pliocene period. This shows, we think, very conclusively that what small resemblance there is between the mammals of the Oriental and Ethio- pian Regions is due rather to a similarity in their origin, than to any sort of direct connection between the two regions. To sum up the subject we add a table of the numbers of orders, families, and genera of mammals found in the six different Regions, together with the number of genera confined to them (endemic), the number of those slightly transgressing the Regional borders (quasi-endemic), and the number of those of wide-spread distribution in each Region. In the second table these numbers have been reduced to percentages, which give in some respects a better idea of the relative specialisation of the mammal- fauna of each Region. On examining these tables, it will be seen that the Ethiopian Region stands second in point of the number of genera confined to it, coming next to the Neotropical Region. The lists of genera from which this table has been worked out are based on those adopted in Flower and Lydekker’s (3) standard work on mammals, with a certain number of additions and corrections. In consequence of this the figures in the case of the Nearctic Region will not be found to exactly correspond with those quoted from Mr. Allen above (2). It must also, of course, be under- stood that the figures are merely approximate, and liable to continual alterations as new discoveries are made. 16 THE GEOGRAPHY OF MAMMALS TABLE I. Approximate Numbers of Families, Genera, and Species of Mammals in the six Regions. T Families. Genera. | Species. | a — | =! | Regions. < | ee a 3 =e oe) | vel) eai\ee| 2 o |#8\ee| iL 2 \62)F5| 8 | 2 |Selbe| 8 jo" a o rs ca ® | | — = | —$—> | 1. Australian .| 7| 4 | 11 | 22 |) 51 |) 6 | 27) 84) 264 2. Neotropical | 10 | 5 | 13 | 28 || 103 | 15 | 12 | 180 |) 501 3. Ethiopian .| 10] 2 | 30 | 42 || 99] 3 | 48 | 150] 577 4, Oriental 2/1 | 33 | 36 || 39 \ 1064 | MiSaieeng 5. Nearctic SWE) DO D2 eo 2/43) 66 || 192 6. Palearctic .| 0/ O | 32 | 32 | 25| 4/| 74 | 103 |) 448 TasE II. , | Numbers given in Table I. reduced to Percentages of Total | Numbers of Families and Genera. Families. Genera. Regions. lau: an ’ ate Endemic. endomier| pnread. | Endemic.) ane Berea | nas | | E | | | ipeAustralian’) = |) 3 TE.) 250 v 60> a 32 2. Neotropical . 35 Ie A a a= atl ae) 3. Ethiopian. . ZO) iin APA SB Biel Si Ges 2 32 4. Oriental =: by | Pi A yh = Pa! 8 56 DewNCAECHIC. 2%. ee tee (6) 90 || 30 3 65 6. Palearctic. . 0 0 100 24 3 71 The groups entirely confined to each region are classed as “endemic” ; those that cross the frontiers slightly as ‘‘ quasi-endemic ”’ ; all others are considered as “‘ wide-spread.” The percentages, it will be observed, on account of the omission of fractions, do not exactly make up one hundred in every case. ae | vey ee iy / / f » > SHOWING THE 6 REGIONS OF THE GEOGRAPHICAL DISTRIBUTION OF MAMMALS INTRODUCTION ily List OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHAPTER I. (1) Auten, J. A.—“The Geographical Distribution of Mammals.” Bull. U.S, Geol. Surv. IV., p. 376 (1878). (2) “The Geographical Distribution of North American Mammals.” Bull. Amer. Mus. Nat. Hist. IV., p. 199 (1892). (3) FLrowgr, W. H., and LyprKKrr, R.—“ An Introduction to the Study of Mammals, Living and Extinct.” London (1891). (4) Gitt, T.—“The Principles of Zoo-Geography.” Proc. Brol. Soc. Washington, II., p. 1 (1885). (5) Hermpriy, A.—* The Geographical and Geological Distribution of Animals.” London (1887). (6) Huxiey, T. H.—“ On the Classification and Distribution of the Alectoromorphe and Heteromorphe.” Proc. Zool. Soc., 1868, p. 294. (7) Newton, A.—Article on Geographical Distribution in the “ Dictionary of Birds,” p. 311. London (1893). (8) Satvapori, T.— Catalogue of the Psittaci or Parrots in the Collection of the British Museum.” London (1891). (9) Sctater, P. L.—“ On the General Geographical Distribution of the Members of the Class Aves.” Journ. Proc. Linn. Soc. (Zool.), vol. 1i., p. 130 (1858). (10) Address to Section D. (Biology). ‘‘ Report of the Forty- Fifth Meeting of the British Association at Bristol,” p. 85 (1876). (11) “On the recent Advances in our Knowledge of the Geographical Distribution of Birds.” bis (6), vol. ii1., p. 514 (1891). (12) SHarrr, R. B.—‘On the Zoo-Geographical Areas of the World.” Nat. Science IIL, p. 100 (1893). (18) Waxtacg, A. R.—* The Geographical Distribution of Animals.” 2 vols. London (1876). CHAPTER II THE AUSTRALIAN REGION (Puate II., p. 50) SEcTION I.—BouNDARIES OF THE AUSTRALIAN REGION Tue Australian Region, as will be seen by the map (Plate II.), includes Australia, New Guinea, and the Moluccas, together with all the Pacific Islands and New Zealand. It is divided from the Oriental, the next adjacent region, by a line drawn between the two small islands of Bali and Lombok (called Wallace’s Line), and passing thence through the Flores and Molucca Seas, between the islands of Celebes on the one side, and Sumbawa, Flores, Bouru, Sula, and Gilolo on the other. All the islands westwards of this line (z.e. Bali, Java, Sumatra, Borneo, Celebes, and the Philip- pines) are included in the Oriental Region; while all the islands to the eastwards, from Lombok to Timor, the Moluccas, and New Guinea, are referred to the Australian Region. Besides this, the Australian Region includes all the islands of the Pacific, from the Pelews and Ladrones in the north-west, to the Sandwich Islands in the north- east, the Marquesas in the south-east, and New Zealand and its neighbouring islands in the south-west. / The boundaries, as above given, correspond with those laid down by Wallace in his work on Geographical Dis- tribution, with the exception that the island of Celebes has been transferred to the oe Region. 1 THE AUSTRALIAN REGION 19 Celebes, as Mr. Wallace has fully shown in his “ Island Life” (14), is an anomalous island presenting a very difficult problem. It has doubtless relations to both the Oriental and the Australian Regions, but has besides many peculiar forms inhabiting it, which do not seem to connect it with either region. On the whole, however, the evidence of the mammals, at any rate, serves to connect it more closely with the Oriental Region, as will be seen by the discussion of the subject in the chapter dealing with that Region. Section I]—GENERAL VIEW OF THE MAMMAL-FAUNA OF THE AUSTRALIAN REGION The peculiarities of the Australian Region are very striking. Within its boundaries are found the only repre- sentatives of the lowest and most reptilian of the Orders of Mammals; these are the egg-laying forms, Ornithorhynchus, Echidna, and Proechidna, which constitute the order Monotremata (12). Of the eight generally recognised families of Marsupials, or “ Pouched Animals,” no less than six are entirely con- fined to the Australian Region, with the exception of two species of phalanger (Phalanger ursinus and P. celebensis), which have crossed the boundary into Celebes. The seventh family, the Didelphyidz, or true Opossums, are found only in the Neotropical Region, whence they have intruded into the southern part of the Nearctic Region. Of the other mammals, the Rodents and the Bats are the only orders at all adequately represented in the Austra- lian Region. To the former belong six genera of Muridx 20 THE GEOGRAPHY OF MAMMALS (Mice), all of which, except the wide-ranging Mus, are confined to this Region; the only other Rodent that occurs in it is the Javan Porcupine, which has just crossed the dividing-line between the Oriental and Australian Regions into the islands of Flores and Sumbawa. The Bats of the Australian Region (3) are very numerous, more especially in the islands to the north of Australia, where the tropical conditions are specially favourable to the development of this group. No less than twenty-eight genera, containing nearly one hundred species of Bats, are met with within the Region, and of these genera eight are not found elsewhere. This is a high percentage, only excelled in the Neotropical Region, which is extraordinarily rich in Bats, having no less than forty-two out of forty- seven genera confined to it. The five other Orders of terrestrial mammals—the Ungulata, the Carnivora, the Insectivora, the Edentata, and the Quadrumana—may be considered as practically absent in the Australian Region, though members of several wide-ranging genera have just crossed the line of separation, and are represented in some of the islands on the north of Australia; but, with the exception of the Dingo, none of these reach the actual continent of Australia. The question of the origin of the Dingo has not yet been settled. Although fossil remains of this animal have been found in the recent Tertiary deposits, it is difficult to say whether the Dingo was introduced into Australia by the aborigines or is indigenous. At the present time it appears to be found both in a wild state and in a semi-domesticated condition among the native Australians, THE AUSTRALIAN REGION Zl Section III.—SuBpDIvIsIoN OF THE AUSTRALIAN REGION The Australian Region may be most conveniently divided into five Sub-regions (see Plate IL, p. 50); these are— (1) The Austral Sub-region, containing—The island- continent of Australia, with the dependent island Tasmania. (2) The Papuan Sub-region, containing—The islands lying to the north of Australia, including (a) the Timor group from Lombok to Timor-Laut; (b) the Moluccas, of which the chief islands are Morty, Batchian, Gilolo, Bouru, and Ceram; (c) the large island of New Guinea; (d) the New Britain group, containing New Britain, New Ireland, the Admiralty Islands, and the Solomon group. (3) The Maorian Sub-region, containing—The two islands of New Zealand, together with their outliers the Norfolk, Kermadec, and Chatham Islands, as also the Auckland, Campbell, and Macquarie Islands. (4) The Polynesian Sub-region,containing—The various islands in the Pacific, from the Ladrones in the north-west to the Society and Marquesas in the south-east, of which the principal are New Caledonia, the New Hebrides, and the Fiji and Samoan groups. (5) The Hawaiian Sub-region, containing—The Sand- wich-Island group. Of these, only the first two, the Austral and Papuan Sub-regions, need be considered in detail, as there are, practically speaking, no mammalian inhabitants, except Bats, in the other three Sub-regions. 22 THE GEOGRAPHY OF MAMMALS Section I[V.—THE AUSTRAL SUB-REGION The “island-continent” of Australia, as Mr. Wallace terms it, has, as has now been positively ascertained, a large portion of its interior so parched up and barren as to be almost destitute of animal life. But all along the east and south-east coasts, where there is land of sufficient elevation to condense the vapours from the adjoining ocean, more fertile districts are found. Besides the more widely diffused Australian types, some peculiar forms are met with only on this side of the continent. Tasmania, which is, in fact, but a recently separated piece of this portion of Australia, has also a moister and less extreme climate, and contains representatives of many of the special Australian forms, besides some indications of an autochthonous fauna. The most peculiar mammals of Australia, and those which first claim our attention, are its representatives of the two forms which together constitute the Sub-class of Monotremes. These are the extraordinary genera Orni- thorhynchus and Echidna, which in their toothless jaws, in the conformation of their sternum and shoulder girdle, and especially in the structure of their reproductive organs, exhibit unmistakable signs of divergence towards the Classes of Reptiles and Amphibians. The first of these, usually known to the colonists as the “ Duck-bill” or “ Water-mole,” (see Fig. 1, p. 23) is entirely aquatic in its habits, and is met with only in the streams and waterholes of New South Wales and Tasmania, where it burrows in the banks, and swims and dives with great facility. The Echidna, or “Spiny Ant-eater,’ is more widely distributed, and, as we THE AUSTRALIAN REGION 23 shall presently see, has also representatives in the Papuan Sub-region. It inhabits the scrubs throughout the eastern districts of Australia, and is likewise occasionally found as far along the southern coast as Western Australia. In Tasmania a slightly different species (Hchidna setosa) with Fic. 1.—THE DUCK- BILL. (Ornithorhynchus anatinus). longer fur, almost concealing the spines (by some authors considered to be only a geographical race or sub-species of the typical form), is met with. After the Monotremes, the most important group of mammals, and that which constitutes by far the most prevalent feature of Australian mammal-life, is the great 24 THE GEOGRAPHY OF MAMMALS order of Marsupials. Although not absolutely restricted to Australia, since one of its component families is purely American, the Marsupials, from their prevalence, con- stitute a still more noticeable feature of the Australian fauna than the Monotremes. Of the seven families into which this order is usually divided by naturalists, six are entirely confined to the Australian Region. Moreover, the Marsupials are so abundant in Australia proper, as to quite overshadow the few representa- tives of the higher orders of mammals found within its limits. The carnivorous Dasyures (Dasywridx) forming the first of these families take the place in Australia of the true carnivora of other parts of the world. They are semi- nocturnal in habits, and prowl about at dusk in search of the smaller mammals and birds which constitute their food. In Tasmania two peculiar forms of Dasyures occur, which are not met with in the Australian continent. These are the Thylacine, a dog-like animal with a long tapering tail, noticeable as being the largest of living car- nivorous Marsupials (Fig. 2, p. 25), and in general external appearance so much resembling our familiar domestic friend that the uninitiated can hardly be persuaded that its proper place is in a different order of mammals ; and the Sarcophilus wrsinus. or Tasmanian Devil, as it is popularly called—a somewhat aberrant dasyure of a prevailing black colour, about the size and somewhat of the shape of an English badger, and remarkable for its savage and voracious disposition. Both these animals, now confined to the island of Tasmania, must have formerly extended into Australia, as their remains have been found fossil in the caves of the Wellington Valley of New South —__—_ THE AUSTRALIAN REGION 25 Wales. Altogether the Dasyures number some twenty species in Australia and Tasmania. Besides the Dasyures, a second type of the same family, but in many respects divergent in structure, is found on the mainland of Australia. This is the Myrmecobius, or Banded Ant-eater, a little animal of the size and somewhat Fig. 2.—THE THYLACINE. (Thylacinus cynocephalus. ) | pf the likeness of the common squirrel, with a long bushy tail and elegant transverse stripes across the back. The _Myrmecobius is terrestrial in its habits, and feeds princi- pally on ants. It is apparently confined to the western and central parts of the sub-region. i The Peramelidx, or Bandicoots, the second family of 26 THE GEOGRAPHY OF MAMMALS | Australian marsupials, have also teeth adapted to an insectivorous diet, although we know, from the records of trustworthy observers, that some of the species feed more or less upon vegetable substances. Such is certainly the |, case with the Rabbit-eared Bandicoot (Perogale lagotis) of | Western Australia, which is abundant over the grassy plains of that colony, and, from its burrowing habits and large hare-like ears, is commonly known as the “Native Rabbit.” Of the typical bandicoots (Peromeles), five or six species are known, distributed over various portions of Australia, each colony having its peculiar forms. They are } all purely terrestrial animals, some inhabiting the densest scrubs, and others the hot stony ridges of the upland plains. } The only remaining member of the family Peromelidz is the anomalous Pigfoot (Cheropus castanotis), a small rat- like animal with slender feet, which is confined to the hard } stony grounds of the interior of the southern Australian} colonies. A most interesting discovery has recently been made in the deserts of Central Australia of a new burrowing mar- f supial, of mole-like habits, for which it is necessary to con- ' stitute anew family. This little animal has been described | | by Dr. E. C. Stirling (9) under the name of Notoryctes| typhlops, and is apparently very rare (Fig. 3, p. 27). Asits name implies, it is quite blind, its eyes being represented. merely by pigment spots buried beneath the skin and muscles ; furthermore, its whole structure is admirably adapted for its burrowing life. Full descriptions of both its habits and anatomy will be found in the Transactionsy of the Royal Society of South Australia for 1891. We now come to the herbivorous marsupials (Diproto- dontes), of which there are three families all confined to the a THE AUSTRALIAN REGION 27 Australian Region (unless the newly-discovered Cxnolestes of South America shall be ascertained to belong to the Diprotodonts) namely, the Wombats (Phascolomyidz), the Fig. 3.—THE NOTORYCT. (Notoryctes typhlops. ) halangers (Phalangeridx), and the Kangaroos (Macro- odid#x), The Wombats are numerically of the least importance f the three families above mentioned, although the form nd general appearance of the animals of the single known enus, Phascolomys, are hardly less remarkable than those f the Kangaroos. 28 THE GEOGRAPHY OF MAMMALS There are three species of Wombat (Fig. 4) generally recognised, one of which is confined to Tasmania ; they are clumsy-looking animals, resembling in their form and actions small bears. They never climb trees, but live entirely on the ground or in burrows and holes, feeding on grass, roots, and other vegetable substances. In general structure the Wombats are closely allied to the a | { ) Fic. 4.—THE COMMON WOMBAT. (Phascolomys mitchelli. ) family, the Phalangers, although their exclusively force habits naturally involve minor points of difference, which added to the characters presented by their dentition, hav) induced naturalists to recognise them as a_separat family. | The Phalangers (Phalangeridz), which next follow, are, a much more extensively developed group of animals. presenting us with several well-marked generic forms} W. AK Johnston. Edinburg a & Lends: MAP OF THE NEOTROPICAL REGION VYING ITS DIVISION INTO 4 SUB-REGIONS. - ee oe, MAP OF THE NEOTROPICAL REGION SHOWING ITS DIVISION INTO 4 SUB-REGIONS. ee es nt Fe a ct a O68 . i on ” 7. : it velahodeh et ha i: ts tit | ott oa cs : ; ; vi: . } , A WATTS . | ‘NY nulaan- WO - i - : nate : +) Seed? j ; ' } ( : WOlWsA JADIIC 2NOLDSA-BUS b GTI . THE AUSTRALIAN REGION 29 and with about twenty-one readily distinguishable species in the Australian mammal-fauna. While the Kangaroos are mostly grazing animals, and the Wombats burrowers and grubbers, the Phalangers are essentially arboreal in their habits, and are much more strictly nocturnal than the two former groups. In the daytime the Phalangers lie concealed in the hollows of trees, issuing forth at night to feed amongst the branches upon leaves, buds, and fruits. The Koala, or “native bear” (Phascolarctos), of which form a single isolated species only is known, serves to connect the Phalangers with the Wom- bats, being allied to the latter by many characters, and amongst others by the absence of a tail, which distin- guishes it from the rest of its family. In Pseudochirus, Trichoswrus, and Dromicia, the more typical forms of the Phalangeride, which next follow, the tail is not only well developed, but is of vital importance to the animal, being used as a prehensile organ. The flying Phalangers of the genera Petawrus, Gymnobelideus, and Acrobates, do not em- ploy their caudal appendages in the same way. But this organ, which is much elongated in all these groups, and densely clothed with hair, serves, along with the membrane extended between the fore and hind legs, in the manner of the flying squirrels (Pteromys), to support the animal in the air when descending from the top of one tree to the base of another. One more very singular little animal must be enume- rated before we leave the Phalanger family—the Tarsipes (Tarsipes rostratus), small in size, but great in interest, even among the many abnormal forms of this wonderful land. The Tarsipes is of the size and general form of an ordinary mouse, but with a long slender-pointed muzzle, 30 THE GEOGRAPHY OF MAMMALS and with the nails of the toes for the most part embedded in the upper surface of the expanded fleshy pads, with which they are terminated, thus affording some resemblance to the abnormal lemuroid mammal Tarsius spectrum of the Indian Archipelago, whence its name is derived. Another peculiarity of the Tarsipes is that its food appears to be exclusively honey, no other substance having been found in the stomachs of the specimens examined, and its long and slender tongue being obviously adapted, like the bill of the humming-bird and the brush-tongue of the lories, for collecting such food. The Kangaroos (Macropodidx) must be considered as par excellence the most important group of the Australian mammal-fauna. They are at once the most numerous in species, and in the former condition of Australia, before the influx of Europeans took place, were probably likewise the most prevalent form of mammalian life as regards individuals. In his great work on the mammals of Australia, Mr. Gould has devoted the whole of the second volume to the illustration of members of this family, figuring no less than forty-four species. Mr. Thomas, in his catalogue of the Marsupialia (12), has recognised forty- five species. These are divided into eleven genera, of which the best known and largest are Macropus, Dendrolagus, and Bettongia. The first of these genera contains the larger kangaroos, in which the upper incisors are of equal length, the canine teeth are deciduous when present, and all the toes of the fore feet are of nearly equal length. One of the finest and most brightly coloured of this larger group is the Red Kangaroo (Fig. 5, p. 31), which has been introduced into Europe, and breeds freely in our zoological gardens. THE AUSTRALIAN REGION 31 Derdrolagus includes four species of Tree-kangaroos, of which, however, only one is found in Australia, Dendrolagus lumholtzi of Northern Queensland, the others being con- fined to New Guinea. They are remarkable for the fact Fig. 5.—THE RED KANGAROO. (Macropus rufus. ) that the proportionate lengths of the fore and hind limbs, unlike those of the true kangaroos, resemble those of ordinary mammals; they further differ from all other kangaroos in being arboreal in their habits, climbing trees with great facility, and living on bark, fruits, and leaves. 32 THE GEOGRAPHY OF MAMMALS Bettongia and its allied genus Potorous embrace the Kangaroo-rats, as they are often termed. These are all small, never exceeding a rabbit in size; they possess well- developed canine teeth, and have the central toes of the fore feet elongated to accommodate their digging habits. Having completed our survey of the Monotremes and Marsupials of the Australian Sub-region, we must now con- sider the Eutherian series, which, as has been already shown, plays a very subordinate part in this extraordinary fauna. Putting aside the marine mammals—the Seals, Cetaceans, and Sirenians—and confining our attention to the terrestrial groups, we find only three of the usually recognised orders, namely, the Rodents, Bats, and Carnivores, with any representatives in this strange country. And the Carnivores would be perhaps better considered as quite external to the fauna of Australia proper, since the solitary member of this group found within its limits is the semi- domesticated Dingo, which, as already suggested, has not improbably been introduced by the primitive native inhabitants. Monkeys, Insectivores, and the most useful Order of Ungulates, to which its grassy plains would appear to be, and, as we know by actual experience are, excellently adapted, are alike unknown, except as introductions, in Australia, and their functions in a state of nature seem to be performed by the various groups of Marsupials. Of the other orders, the Rodents are represented by six genera, all belonging to the Mice (Muridz). Of these four are confined to Australia proper, while two others also occur in the Papuan Sub-region; the remaining one, the THE AUSTRALIAN REGION 33 cosmopolitian Mus, or true Mouse, numbers no less than eighteen species in Australia. The habits of the Water- voles of Europe are assumed by the species of the genus Hydromys, which are modified for aquatic life, while the species of Hapalotis are found chiefly in the dry sandy scrubs, The Bats, with one exception, all belong to genera of considerably wide distribution, and the number of species known to inhabit Australia is not very great. Thirty only are described in Dr. Dobson’s catalogue of the Bats as Australian. Pteropus, the great genus of Fruit-eating Bats, is represented by a few species; so too are the widespread insectivorous genera Vesperugo and Miniopterus. One species of the former genus, the Southern Pipistrelle (V. abramus), is found almost throughout the Old World, from Sweden to North Australia. SEecTION V.—ANALYSIS OF THE AUSTRAL MAMMAL-FAUNA The full list of the Mammalian Fauna of the Austral Sub-region contains the names of about 169 species, refer- able to fifty-nine genera. Of these genera twenty-nine, just half, are not found outside the Austral Sub-region, the greater number of them (twenty-five) being Mar- supials; they include, however, three genera of Mice (Xeromys, Hapalotis, and Mastacomys) and one peculiar Bat (Rhinonycteris). Thirteen genera are confined to the Australian Region, that is, are found in the Papuan Sub- region as well as in the Austral Sub-region; of these, again, the majority are Marsupials, besides which there C 34 THE GEOGRAPHY OF MAMMALS are two genera of Rodents (Hydromys and Uronys), and one Bat (Nyctophilus). Seventeen genera which are found in Australia extend beyond the limits of the Region, to the Oriental, and in some cases range even into the Ethiopian and Palearctic Regions. Of these the greater number (fourteen) are Bats; one is a Marsupial (Phalanger), of which two species occur in Celebes; another is the widely spread genus Mus, which is found throughout the Old World; and the last is a Carnivore (Canis), of which the Australian representative is the Dingo. Canis is spread over the whole of the rest of the world, both Old and New. Turning now to the distribution of the genera within the continent of Australia itself, it will be found that out of forty-three genera (of Monotremes, Marsupials, and Rodents), twenty are fairly represented throughout the whole area, thirteen are confined to the east (several of these ranging north into New Guinea), seven are re- stricted to the western and central parts of the continent (these latter are chiefly desert forms such as Perogale, Cheropus, and Notoryctes), and the three remaining genera are found only in Tasmania. These are Thylacinus, Dasywrus, and Mastacomys. If now we reconsider the list, counting only Mono- tremes, Marsupials, and Rodents, it will be found that out of 130 species, thirty-five range from west to east to greater or less extent; forty-nine are confined to the eastern part of Australia, in many cases extending to Tasmania; and thirty-six are peculiar to Western Australia, while ten species out of the whole Mammal-fauna are peculiar to Tasmania. © Or THE AUSTRALIAN REGION SEcTION VI.—Past History oF THE MAMMAL- FAUNA OF THE AUSTRAL SUB-REGION The past history of the Australian mammals is still very obscure; the only remains of extinct species yet discovered have been found in certain bone-caves and in surface-deposits generally attributed to the Pleistocene Epoch. Putting aside the Dingo, all the Pleistocene mammals of Australia belong to the Monotremes or to the Mar- supials, and, with two exceptions, can be accommodated in still existing families. These exceptions are Noto- theriwm and Diprotodon, the latter of which was a very large animal, equalling a Rhinoceros in bulk, but both of them were probably allied to the existing Kangaroos. Recently a deposit containing bones of these and other large Marsupials has been found in a dry salt lagoon, called Lake Mulligan, in South Australia, and when these remains have been worked out, a great deal more informa- tion as regards this pliocene or pleistocene fauna may be expected. As in South America, these extinct animals appear in many cases to have attained a size far surpassing that of their degenerate descendants. None of the remains hitherto found in Australia throw much light on the origin of its remarkable fauna. But quite recently evidence of an extensive mammalian fauna has been discovered in certain beds, of probably Upper Eocene age, in Santa Cruz, Patagonia. In addition to a number of other forms, this series contains the re- mains of many Marsupials, and though the most promi- nent of them belong to the Didelphyide—the Marsupial 36 THE GEOGRAPHY OF MAMMALS family now confined to America, and apparently dis- tributed over the whole northern hemisphere during Tertiary times, but not found in Australia—a certain number of them show an Australian stamp. Some of them have even been relegated by Ameghino (1), to whose researches our knowledge of the Santa Cruzian fauna is mainly due, to the existing Australian family Dasyuride. If, on further investigation, these references shall be found to be correct, the inference would seem to be that in very remote times—probably in the early Tertiary or the late Secondary Period—there has been some sort of land- connection between South America and Australia. In such case there would be no necessity to suppose that Australia was ever directly connected with the rest of the Old World at all, none of the peculiar forms of Australian Marsupials having yet been detected in any other part of the globe. Besides the common possession of Marsupials, many other resemblances between the faunas of Australia and South America have been pointed out to occur among the Birds and Amphibians, and especially among the Fishes. Two families of freshwater fishes (Galaxiide and Haplochi- tonid#) are found exclusively in these two Regions, and are not known to occur elsewhere. Apart from speculation, however, there is no question that Australia has been isolated from all the other continents since the end of the Secondary, or at least since the beginning of the Tertiary Period of geological time. THE AUSTRALIAN REGION 37 Section VIJ.—TuHeEe Papuan SUB-REGION In contrast to Australia, the great island of New Guinea, or Papua, is traversed throughout by mountains of high altitude. The rivers rising in these ranges, aided by the suns of the tropics, produce a luxuriant vegetation, and such a country as we should suppose would be espe- cially favourable to mammal-life. Yet mammals are by no means abundant in New Guinea and in the adjacent islands which constitute the Papuan Sub-region. As is the case in Australia, the greater number of the in- digenous animals of New Guinea and the neighbouring islands consist of Monotremes, Marsupials, and Rodents, together with a certain number of the cosmopolitan order of Bats (8, 10, 11). Of the Monotremes, two species have been met with in New Guinea, both of them belonging to the family of Echidnas above referred to, Of these one species, only at present known from the south of New Guinea, is but a slightly modified form of the small Australian Echidna. But in the mountains, in various parts of New Guinea, has been lately discovered a larger representative of the same family (Proechidna), which, moreover, differs from the typical form in having only three toes on its fore limbs, and in other particulars (Fig. 6, p. 38). The Papuan Marsupials, as yet discovered, are about thirty-three in number, and embrace representatives of the Dasyures, Bandicoots, Phalangers, and Kangaroos, which are also characteristic families of the Australian mammal- fauna, There are only two genera of Marsupials peculiar to the 38 THE GEOGRAPHY OF MAMMALS Papuan Sub-region. One of these is Distechwrus, a small mouse-like animal belonging to the family of Phalangers, and remarkable for its long tail, which bears at its tip a double row of hairs on either side, and thus resembles a feather. The other genus is Dorcopsis, containing three species of animals somewhat nearly allied to the true OO Ly! Fig. 6.—THE PAPUAN ECHIDNA. (Proechidna bruijni.) Kangaroos. A third genus (Dendrolagus), containing Kangaroos specially modified for arboreal life, was formerly supposed to be peculiar to New Guinea. But a species of the same form, as already mentioned, has been ascertained of late years to exist in Northern Queensland also, thus giving further proof of the close alliance of the Papuan and Austral mammal-faunas. THE AUSTRALIAN REGION 39 Nearly all the Marsupials found in the Papuan Sub- region are confined to the island of New Guinea; a few, however—for example, the Phalangers—also inhabit the other islands of this Sub-region. The Grey Cuscus (Phal- anger orientalis) extends from Amboyna in the Moluccas and from Timor all across New Guinea, and as far east as New Britain and the island of San Christoval in the Solomons. As regards the remaining orders of mammals, a few scattered representatives of the higher forms (Ungulates, Insectivores, and Carnivores) are found in the islands of the Papuan Sub-region. Some of these have, undoubtedly, been introduced by the Malays from the neighbouring islands of the Oriental Region; but others have been described by naturalists as peculiar species. In the latter case, even if we assume that the specific dis- tinctions have been satisfactorily established, it is not probable that such species have been brought into the Papuan Region by the hand of man. It is more likely that they have migrated into it at an earlier period, since a considerable lapse of time is necessary before the effects of isolation can produce new races of sufficient distinctness from the original form to be entitled to specific separation. As regards the Papuan Ungulates, several kinds of Swine (Sus) are stated to be met with in the Papuan Region. How far these differ from one another, and whether they are really distinct from the allied wild pigs of the Oriental Region, seems a little uncertain. It is probable, however, that many of these so-called “species” of wild pig may be descendants of the domestic animal, which has run wild, as is well known, in many of the Pacific Islands. The other representatives of the Order Ungulata found in the Papuan Sub-region are three species of Deer belonging to what is 40 THE GEOGRAPHY OF MAMMALS termed the Rusine group of the genus Cervus. Two of these deer are confined to Timor and the Moluccan group of islands respectively. All three are closely allied to the Javan Deer (Cervus hippelaphus), and it is quite possible that they are merely dwarfed forms of this species. Passing over, for the present, the Rodents and Bats, the Carnivora are represented in the Papuan Region by three species. These are a Cat (Felis megalotis), of which very little is known, but which is stated on competent authority to be quite a distinct species, and to be confined to the islands of Timor and Rotti; a Palm-cat (Paradoxwrus hermaphroditus), and a Civet (Viverra tangalunga). Of these two the Palm-cat is found in both the Moluccas and the Timor group, the Civet only in the Moluccas. These animals have in all probability been introduced by the Malays, since they are both frequently kept by them as pets in captivity. The Order Insectivora is represented in the Papuan Sub-region by about five species of small Shrews (Crocidura), but, so far as is yet known, these occur only in the Moluccas and in Timor, and do not reach further east. Lastly, two species of Monkeys just enter the margin of the Sub-region, namely, the Common Macaque (Macacus cynomolgus), very widely distributed throughout the Oriental Region, which has crossed “Wallace’s Line” into Flores and Timor; and the Black Ape of Celebes (Cynopithecus niger), which has passed the narrow straits between that island and Batchian, one of the Molucca group. The Rodents are represented in the Papuan Sub-region by a considerable number of rats and mice of the family THE AUSTRALIAN REGION 41 Muride, all of them restricted to the Sub-region. One of these, remarkable for its peculiar and probably prehensile tail, is found only in the mountains of New Guinea, and has been placed in a new genus (Chiwromys). Among the Rodents also there is one very obvious intruder from the west; this is the Javan porcupine (Hystria javanica), which in this Sub-region is found only in Timor, but is very widely distributed throughout the Oriental Malayan Islands. Finally, the Papuan Sub-region, with its luxuriant vegetation and tropical forests, seems to be extremely favourable to the presence of Bats, of which there are more than sixty species known to occur within its limits. More especially is this the case with the large fruit-eating bats of the genus Pteropus, since about twenty out of the forty known species of this genus are found within this Sub- region. This genus (Pteropus) has a remarkable area of distribution, which it is difficult to account for satis- factorily. Its range extends from Madagascar and the neighbouring Mascarene Islands through the Seychelles to India, Ceylon, Burma, and the Malayan Archipelago, and includes even Southern Japan ; thence it is continued over all the Papuan Sub-region into Australia and most of the Polynesian Islands. This genus, it may be noted, seems to have a special propensity to peculiar development in insular areas. Out of about forty species of Pteropus as yet known, only two (Pteropus medius from India, and Pteropus edulis of the Malay Peninsula) are found on the main continental mass; all the other species are confined to islands, and in many cases to very small limits. For instance, one species (P. livingstoniz) is restricted to the Comoro Islands in the Indian Ocean, three others to single 42 THE GEOGRAPHY OF MAMMALS islands in the Solomon group, and a fourth is only known from the island of Lombok. Section VIII.—ANALyYsIS OF THE PAPUAN MAMMAL- FAUNA “The number of genera of mammals represented in the Papuan Sub-region is fifty; of these eight are restricted to the Sub-region (namely, Proechidna, Distechwrus, Dorcopsis, Chiuwromys, Pteralopex, Nesonycteris, Melonyc- teris, and Anthops). There are also twelve genera, nearly all marsupials, common to the Papuan Sub-region and Australia; and sixteen genera common to the Papuan Sub-region and the Oriental Region. Of these, however, only six (Sus and five genera of bats) penetrate so far eastwards as New Guinea; the other ten are stragglers over “ Wallace’s Line” as far as the Timor group and the Moluccas only. ‘Thirteen genera (Phalanger, Mus, and eleven genera of Bats) are found in both the Oriental Region and in the Papuan and Austral Sub-regions. 1 The most recent list of Papuan mammals, published by Dr. K. M. Heller (Abh. Mus. Dresd. vi., No. 8, 1896-97) gives the number of species now known as follows— Genera. Species. 1. Monotremes 3 : ; : 2 3 2. Marsupials 14 36 3. Sirenia 1 1 4, Ungulata 1 2 5. Rodentia 5 18 6. Carnivora . 2 2 7. Insectivora 1 il 8. Chiroptera . 16 39 [ls] 3| THE AUSTRALIAN REGION 43 SecTION [X.—THE MAoRIAN SUB-REGION The Maorian Sub-region includes, besides New Zealand proper, many smaller groups of islands in the sea around, such as Norfolk Island, the Kermadec group, Chatham Island, Stewart Island, Auckland Island, Campbell Island, and Macquarie Island, and probably Lord Howe’s Island, though in some respects this appears to belong rather to the Australian mainland than to New Zealand. As in the Polynesian Sub-region, there are no indi- genous terrestrial mammals found in this Sub-region, the only exception being a species of rat (Mus maoriwm). But it is quite possible, nay, probable, that this Rat, as its name seems to imply, was brought by the invading Maoris into New Zealand from Tonga, or from wherever the Maoris originated; indeed, Mr. Thomas believes it to be identical with a Polynesian species, Mus exulans (ef. Buller, Trans. N. Z. Inst., xxv. p. 49). The only two bats recorded from New Zealand are Mystacina tuberculata, the genus as well as the species being restricted to New Zealand; and Chalinobus morio, which is also found in Australia. The islands of New Zealand are indeed remarkable as being the only insular area on the globe, of any consider- able size, which are entirely destitute of mammal-life. ‘All the other large islands of the world possess a Mammal- fauna of greater or less extent related to the continent to which they are nearest, and have consequently been termed by Mr. Wallace “continental islands.” All such “continental islands” are separated by narrow seas, of no great depth, from their respective continents. New 44 THE GEOGRAPHY OF MAMMALS Zealand alone, of all the larger islands of the globe, is disconnected by a considerable breadth of ocean (about 1400 miles) and also by a deep sea (more than 2000 fathoms) from the nearest point of mainland. This fact and the absence of an indigenous Mammal- fauna show that New Zealand has not been joined directly by land with Australia recently, even in a geological sense of that term; possibly it has never been so connected at all. To determine, therefore, the geographical affinities of this Sub-region, we must turn to the birds and to the other lower groups, and so endeavour to gain an idea of the affinities of these interesting islands. In New Zealand the want of mammals has been apparently supplied in former epochs by the great development of two families of flight- less birds. One of these groups, the Kiwis (Apterygidz), is still represented by five or six species, although these birds are being rapidly exterminated by the British colonists. The other group, the Moas. (Dinornithidz), is now quite extinct, but as remains of their skin and feathers have been found in some of the caves of the Southern Islands, and as the ancient legends and songs of the Maoris contain unmistakable references to them, it is probable that they have ceased to exist only within the last few hundred years. In addition to the flightless birds, recent and extinct, New Zealand still possesses two very singular forms of Parrots (Nestor and Stringops), for the reception of which special families have been formed; and eighteen other peculiar genera of land-birds, most of which are related more or less remotely with Australian forms. Altogether there are in New Zealand fifty-seven land-birds, belonging to thirty- THE AUSTRALIAN REGION 45 four genera, of which sixteen, or nearly half, are not known elsewhere. There are also five peculiar genera of waders and aquatic birds in New Zealand,making twenty-one indigenous genera in all. Among its few reptiles, also, New Zealand numbers the very remarkable Tuatera (Sphenodon punc- tatus), which, though externally resembling a lizard, differs from all other lacertians in so many points of its skeleton and internal structure that it is usually considered to belong to a separate and distinct order of reptiles. The nearest allies of this form are found among three extinct families which make up the order Rhynchocephalia. Re- mains of these families occur in beds of Permian age in Germany, in the Keuper of Elgin (Scotland), and in the (probably contemporaneous) Gondwana beds of India, as likewise in the lower Eocenes of North America and Northern Europe. All these facts indicate a great amount of individualism in the Maorian Sub-region. But on the whole they betray an affinity to the tropical parts of Australia and to the Papuan Sub-region rather than to the temperate portion of Australia, to which New Zealand is now nearest in point of actual distance. This connection is further confirmed by the soundings of the seas round the islands, which show that, although on the west, south, and east deep water extends all round, a long submerged bank, with a depth of less than 1000 fathoms, stretches along to the north-west, and connects the shallow waters round Australia with those round New Zealand. It is probable that the land connection between the two areas, if it ever actually existed, took place somewhere along this line. As already mentioned in the first chapter of this volume, 46 THE GEOGRAPHY OF MAMMALS some writers on Geographical Distribution, especially Professor Huxley and Professor Newton, are inclined to give to New Zealand and its islands the rank of an inde- pendent Region among the primary divisions of the globe, There is, no doubt, as has just been shown, a good deal to be said for this proposal; but, on the other hand, there are even stronger reasons for retaining New Zealand as a Sub-region of the Australian Region. In the first place, we are here dealing with Mammals alone, and it seems rather absurd to assign the value of a primary Region to a group of small islands characterised by the almost entire absence of that class of animals with which we are most concerned, In the second place, looking at Regions from a more general point of view, there is a great practical convenience (as Mr. Wallace has pointed out) in keeping the more or less equal divisions of the globe as primary divisions. It seems, therefore, to be quite unnecessary to elevate so small a portion of the world into a Primary Region. Other small insular areas might, with some justice, put forward nearly similar claims. In the third place, although New Zealand possesses no indigenous terrestrial Mammals, yet the fauna, such as it is, shows an unmistakable affinity of various degrees to that of Australia, and more especially to the tropical part of that continent. It is, indeed, probable that the whole of the fauna of New Zealand has been originally derived from that source, although in the greater number of cases it has undergone considerable modification. Dr, H. O. Forbes (4) has lately published a specula- tive article on the former existence of a (now mostly submerged) southern continent, the remains of which are represented by the land round the South Pole, while THE AUSTRALIAN REGION AT former arms stretched upwards and embraced New Zealand, Eastern Australia, Tasmania, Madagascar, the Mascarene Islands, and part of South America. The grounds for this bold assumption, which, although by no means new, has not been previously developed to so great an extent, rest chiefly on the finding of the remains of a large ocydromine rail in the Chatham Islands, allied to the now extinct Aphanapteryx of Mauritius, and the fossil bones of a large coot (Fulica) allied to F. newtoni of the same island. Other evidence adduced is that of the occurrence of the Ratit#, or Struthious birds, in New Zealand, Australia, Madagascar, and Patagonia. But the distribution of Struthious birds is probably to be explained much in the same way as the distribution of other archaic forms, such as the lemurs and tapirs. They are remnants of what were formerly widely spread groups. That this is likely to be the case is shown by the recent discovery, in other parts of the world (such as the Sewaliks of India, and the Eocenes of England and France) of the remains of other extinct Ratite birds. Another piece of evidence brought forward by Dr. Forbes is the occurrence of Didunculus in the Samoan group, and of the Dodo (Didus), to which Didunculus was once supposed to be nearly related, in Mauritius. But it is now allowed that Didunculus has little near aftinity to the Dodo, and that it is in fact a mere strongly modified member of the family Columbide. Mauritius, as a matter of fact, is in every way a typical oceanic island, and there seems to be little evidence, either physical or zoological, of its having been ever connected with any other land. 48 THE GEOGRAPHY OF MAMMALS SECTION X.—THE POLYNESIAN SUB-REGION The Polynesian Sub-region includes all the numerous and scattered island groups of the Pacific, from the Ladrones and Carolines in the west to the Marquesas in the east, with the exception of the Sandwich Islands, which, owing to their many peculiarities, must be kept apart as a separate Sub-region. There is very little to be said concerning the Polynesian Sub-region so far as mammals are concerned. As is always the case with oceanic islands—that is, islands that do not seem to have ever been directly connected with any of the great land-masses of the globe—the Mammal-fauna of Polynesia is practically non-existent, the only exception being a certain number of Bats, which are creatures able to traverse the intermediate sea-areas, and so more resembling birds than ordinary mammals in their dis- tribution. There are, however, besides the Bats, three or four species of the cosmopolitan genus Mus (Mice and Rats), recorded to occur in Polynesia, whether truly indigenous or the modified descendants of introduced species it is impossible to say. Of the eleven species of Bats which have been regis- tered as Polynesian, eight are peculiar to the Sub-region, two extend into Papua, and one ranges even as far as the Oriental Region. But, looking to the extreme poverty of the Mammal- fauna, it is evident that, to ascertain the general character of the Sub-region, we must turn to the Birds. These, as shown by the excellent summary of Polynesian Ornithology THE AUSTRALIAN REGION 49 recently compiled by Mr. Wiglesworth (15), are, considering the number of islands, not numerous, but on the whole show distinctly Australian aftinities. SECTION XI.—TuHer Hawattan SuB-REGION The Hawaiian Sub-region includes only the Sandwich Islands. This group of islands is situated in the northern part of the Pacific Ocean, and is very isolated, not only from the great land-masses of Asia and America, from which it is separated by a very deep ocean more than 2000 miles across, but also from the other larger groups of the Polynesian islands such as Samoa and the Marquesas, from which it is parted by nearly the same distance. The larger islands composing the group are seven in number, all of purely volcanic origin. As would be naturally expected, there are no indigenous land-mammals in the Hawaiian Sub-region, but a single species of bat (Atalapha semota) occurs there. This bat belongs to a genus found in America, and has, no doubt, reached the Sandwich Islands from that continent. The birds, however, to which we must turn for a moment in order to gain some idea as to the composition of the Hawaiian fauna, show extreme specialisation. The greater number, not only of the species but even of the genera of this Sub-region, 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 distribution is generally wider, the Hawaiian birds are, in many cases, local. We shall, however, be able to form a better general D 50 THE GEOGRAPHY OF MAMMALS idea upon this subject when Mr. Scott Wilson’s new work on the Hawaiian Avifauna (16), now in process of issue, has been brought to a conclusion. As is the case >with the Birds, so with the , Land- shells, which have been carefully studied by the Rev. J. T. Gulick, a wonderful specialisation is found in the Hawaiian fauna. There are, therefore, ample grounds for making a separate Sub-region for this remote island group, not- withstanding its small size and the complete absence of mammalian life. List OF THE PRINCIPAL AUTHORITIES REFERRED TO IN {CHAPTER II. (1) AmEGHINO, F.—“ Revista Argentina de Historia Natural.’ Tomo 1 (1891). (2) Cotuetrt, R.—‘ On some apparently New Marsupials from Queensland.” /.Z. S., 1884, p. 381, pls. xxix.-xxxil. (3) Dosson, G. E.—“ Catalogue of the Chiroptera in the Collection of the British Museum ” (1878). (4) Forsres, H. O.—“ The Chatham Islands: their Relation to a Former Continent.” R. Geogr. Soc. Suppl. Papers, ITI., pt. 4 (1893). (5) Goutp, J—“*The Mammals of Australia.” 3 vols. London (1845-63). (6) LumHotrz, C.—‘‘ Notes upon some Mammals recently discovered in Queensland.” P. Z. S., 1884, p. 406 (7) Oainpy, J. D.— Catalogue of Australian Mammals.” Sydney (1892). (8) Peters, W., and Doria, G.—“ Enumerazione dei Mammiferi raccolti da O. Beccari, L. M. d’Albertis ed A. A. Bruijn nella Nuova Guinea propriamenta detta.” Ann. Mus. Civ. Stor. Nat. Genov., XVI., p. 665, pls. v.—xvili. (1881). (9) Srrriine, E. C.—“ Description of a New Genus and Species of Marsupialia, Notoryctes typhlops.” Trans, R. Soc. S. Austr., XIV., p. 154, pls. ii.-ix. (1891). See also op. cit., p. 283, pl. xii. (10) THomas, O.—“ On the Bats collected by Mr. C. M. Woodford in the Solomon Islands.” P. Z. S., 1887, p. 320, pls. xxv. and xxvi. (11) “ The Mammals of the Solomon Islands, based on the ACUES The RAGE SUB-REGION MAP OF THE AUSTRALIA PLATE Il. Y 20 SUB-IREGION ™ Marquesas RE°G 1 ON 5 “ ni : aie y A New SONS 2 s \ Hebrides x iste + Socetyl® \: warsty d 120 1£0 180 MAP OF THE AUSTRALIAN REGION SHOWING ITS DIVISION INTO 5 SUB-REGIONS. 160 140 eon “Athen : < uh sg THE AUSTRALIAN REGION 51 Collections of Mr. C. M. Woodford during his Second Expedition to the Archipelago.” P. Z. S., 1888, p. 470, pls. xx.-xxii. (12) THomas, O.—“ Catalogue of the Marsupialia and Monotremata in the Collection of the British Museum.” London (1888). (13) “Description of a New Genus of Muride@ allied to Hydromys.” P. Z. 8., 1889, p. 247, pl. xxix. (14) Watuaceg, A. R.—“ Island Life.” London (1880). (15) WicLEswortH, L, W.—“ Aves Polynesiz : a Catalogue of the Birds of the Polynesian Sub-region (not including the Sandwich Islands).” Abth. K. Zool. Mus., Dresden, No. 6 (1891). (16) Witson, S. B., and Evans, A. H.— Aves Hawaiienses: The Birds of the Sandwich Islands.” Parts I.-IV., 4to. London (1890-93). CHAPTER III THE NEOTROPICAL REGION (PuatE III., p. 82) Section I—BouNDARIES OF THE NEOTROPICAL REGION THE Neotropical Region is, no doubt, after the Australian, the most distinct of all-the regions. It includes not only the continent of South America, but the West Indies, Central America, and a considerable portion of Southern Mexico. As regards its northern termination, on account of the great admixture of Nearctic and Neotropical forms which takes place where the two Regions join, it is impos- sible to lay down anything but an approximate boundary. Mr. Wallace (11) draws the line from the mouth of the Rio Grande on the Atlantic side to the neighbourhood of Mazatlan, in about the same latitude, on the Pacific side, but bends it down between these two points so as to include in the Nearctic Region the whole of the high tableland down to the city of Mexico. Some American naturalists, among others Merriam and Allen (1), include in the Neotropical Region the southern extremities of the peninsulas of Lower California and of Florida. This, however, appears to be unnecessary, at least so far as the mammals are concerned, though there are certainly a considerable number of Neotropical birds and insects found in both these districts. Besides the mainland of Central and South America THE NEOTROPICAL REGION 53 and the West Indies, the Falkland Islands and the Galapagos must likewise be included in the Neotropical Region. The West Indies form an important sub-region, con- taining some forms of great interest, and will be treated of in detail below. The Falkland Islands are situated in the south Atlantic, about 250 miles east of the nearest point of Patagonia. They are, however, known to be connected with the main- land by a shallow sea of less than 100 feet in depth, and therefore present all the characteristic features of a “continental” group of islands. The only indigenous mammals are a Wild Dog (Canis antarcticus) and a Vesper-mouse (Hesperomys), which seem to be distinct, though closely allied to the mainland forms. The birds also are mostly identical with those of the mainland, though there are some just recognisable representative forms. The Galapagos, a group of five larger and ten smaller islands, are situated in the Pacific, exactly under the Equator, at a distance of from 500 to 600 miles west of Ecuador. They rise up from very deep water, and are entirely of volcanic origin. They are therefore typical “Oceanic” islands. With the exception of two Vesper- mice, slightly differing from those of the mainland, and a peculiar Bat (Atalapha brachyotis), there are no indigenous mammals (2) in the Galapagos. There are, however, a considerable number of birds in these islands, most of them not found elsewhere, and many of them restricted to individual islands in which they represent each other (7). There can be no doubt that the Galapagos have never, ~ 54 THE GEOGRAPHY OF MAMMALS at any period of their history, been joined to the mainland, and that, owing to the fact that they are situated in the region of equatorial calms, immigration from the mainland is very occasional. In this way has been gradually evolved the peculiar fauna, which, although highly specialised, shows abundant evidence of its having been derived from the nearest mainland. SEcTION I].—GENERAL VIEW OF THE MAMMAL-FAUNA OF THE NEOTROPICAL REGION The Neotropical Region is essentially one of luxuriant tropical vegetation, the great mass of the land from Central America to Uruguay being occupied by vast forests. South of the tropic of Capricorn the woods soon disappear, and the country, over a large part of its surface, becomes a flat treeless expanse covered with more or less abundant pasture known as the Pampas, while all along the western coast extends the giant range of the Andes, the eastern flanks of which are, as a rule, well watered and wooded, while the western slopes from the Gulf of Guyaquil to the island of Chiloe facing the Pacific are utterly dry and arid. There are, however, also in the higher parts of Venezuela, the Guianas, and Brazil, open grassy plains called “campos,” which rise as islands from a surrounding sea of evergreen tropical forest. The mammalian fauna of the Neotropical Region is naturally a rich one, but in the number of genera and species falls considerably short of that of the Ethiopian Region. This may perhaps be accounted for by the physical features of the country, which are certainly not THE NEOTROPICAL REGION 55) so favourable for mammalian development as the more open and varied country of Africa. This deficiency in mammalian life is, however, more than counterbalanced by the abundance of other groups of animals, more especially of birds and insects, to the development of which the luxuriant tropical vegetation seems to be especially conducive. Again, the mammalian fauna of the Neotropical Region is quite as remarkable for what it does not possess (lipotypes) as for what it has. Everything points to the conclusion that during a long geological age, probably throughout the greater part of the Tertiary epoch, South America was entirely isolated from the rest of the world. Thus the present fauna has arisen from two quite different sources —first, from the original fauna of early Tertiary times; and, secondly, from immigrants from the north, some of these being of rather long standing, and others of later arrival. Of the nine Orders of Terrestrial Mammals, repre- sentatives of eight occur in the Neotropical Region, the only Order entirely absent being the Monotremes, which are absolutely confined to Australia. The Marsupials are represented in the Neotropical Region by a single family only,’ out of seven into which this order is usually divided. This is the Didelphyide, or Opossums, of the twenty-four generally recognised species of which one (Didelphys marsupialis) ranges north with some modification into the Nearctic Region. The Quica Opossum (D. opossum) (Fig. 7, p. 56) is another well-known 1 Since this was written Mr. Thomas has described his wonderful new South-American genus Cenolestes, which seems to belong to the Australian Diprotodonts. See P. Z. S., 1895, p. 870. 56 THE GEOGRAPHY OF MAMMALS species of the group, which is found all through the Region, from Southern Mexico to La Plata. The third Order of mammals—the Edentata—is highly characteristic of the Neotropical Region. Of the five Fic. 7.—THE QUICA OPOSSUM. (Didelphys opossum. ) generally recognised families two belong entirely to the Old World; the other three—the Sloths, the Ant-eaters, and the Armadilloes (which are more nearly allied to one another than to the two Old World families)—are, with the exception of one species of Armadillo (Tatusia THE NEOTROPICAL REGION 57 novemcincta), which extends into Texas, absolutely con- fined to the Neotropical Region, and are eminently characteristic of its mammal-fauna. The Sloths (Brady- podide) of the present epoch at least, are entirely arboreal Fic. 8.—HOFFMANN’S SLOTH. (Cholopus hofimanni.) in their habits, and pass their lives suspended by their limbs on the underside of the branches of trees (Fig. 8). The Ant-eaters (Myrmecophagidz) are also mainly in- habitants of forests, and one of the three existing forms (Cyclothwrus) is exclusively arboreal. A second (Taman- dua) may be said to be semi-arboreal, but the largest—the 58 ' THE GEOGRAPHY OF MAMMALS Great Ant-eater as it is usually called (Fig. 9}—does not climb trees, though mostly found in forest-districts. These three animals are all widely distributed in the woodlands of tropical America, but never met with elsewhere. The Armadilloes (Dasypodidz) are mostly inhabitants of more open districts (see Fig. 10, p. 59). Besides the three living Fic. 9.—THE GREAT ANT-EATER. (Myrmecophaga jubata. ) [List of Vertebrates, 1896, p. 197.] families of Kdentates, there are two (the Megatheriidz and Glyptodontidz) now extinct, which are chiefly char- acteristic of the Neotropical Region, though remains of them have also been found in certain formations in North America (6). THE NEOTROPICAL REGION 59 The fourth order of mammals, the Ungulates, is very poorly represented in the Neotropical Region, four only out of the fourteen usually recognised families being found within its limits. The Peccaries (Dicotylidx) consist of only two species, of which one (D. tajyacw) ranges as far Fig. 10.—THE THREE-BANDED ARMADILLO. (Tolypeutes tricinctus. ) north as the Southern United States, and the other is confined to the Neotropical Region. A second family, the Camelide, is shared by the Neotropical Region with the Old World. The representatives of this family in the New World are the Lamas, belonging to the genus Lama (see 60 THE GEOGRAPHY OF MAMMALS Fig. 11). They are entirely confined to the higher ranges of the Andes and to the desolate plains of Patagonia. The Deer (Cervide) of the Neotropical Region all belong to two peculiar genera (Cariacus and Pudua), of which the former extends northwards throughout the United States Fie. 11.—THE LAMA. (Lama peruana.) to British Columbia, while the latter is found only in Western South America. Finally, the Tapirs (Tapiridx) are represented by four species, all of which are peculiar to this region, the only other existing Tapir known being the Indian Tapir of the THE NEOTROPICAL REGION 61 Malay Peninsula. The explanation of this curious case of discontinuous distribution is afforded by the past history of the group. During Miocene and Pliocene times, members of this genus and its allied forms were found both in North America and also throughout the Old World from France to China. This gives us direct evidence of the former much wider extension of the family of Tapirs, and bridges over the present great gap in its distribution. As already stated, the fauna of the Neotropical Region is almost as remarkable for the absence of certain families as it is for the presence of peculiar forms. This is specially noticeable in the Ungulates. There is no existing re- presentative of the four great families of the Oxen, Rhinoceroses, Horses, and Elephants in this region, though remains of the latter two groups have been found in most recent deposits of Argentina. Here they were probably immigrants from the north, which survived but a short time in this locality. Turning now to the Rodents, we find that out of the four chief divisions into which this order is separated—the Squirrels (Sciwromorpha), the Mice (Myomorpha), the Porcupines (Hystricomorpha), and the Hares (Duplici- dentata)—the first two and the last contain very few peculiar genera and no peculiar families in this Region. But, on the other hand, out of the six families of Hystrico- morpha four are restricted to this region, while of the remaining two, one (the Octodontidx) is found elsewhere only in Africa, and the other, the Porcupines (Hystricidz) is of wide distribution. Moreover, all the Neotropical genera of the Hystricomorphine division are, without exception, confined to this region. 62 THE GEOGRAPHY OF MAMMALS The Carnivores, which follow next, are well represented in the Neotropical Region, but belong generally to families of wide distribution. But one family, the Raccoons (Procyonide), with the exception of a single genus (4ilwrus), which is perhaps doubtfully referred to it, is entirely confined to the New World. On the other hand, the Viverridx, so widely spread in the Old World, are entirely absent in America. With the exception of some four or five species of Shrews, which have obviously spread southwards from the Nearctic Region, wherein they are found in considerable numbers, the Insectivores are represented in the Neotropical Region only by a single remarkable family. This is the Solenodont—a characteristic form of the Greater Antilles —absolutely unknown elsewhere. The Insectivores are usually considered to be the most generalised of all the mammalian orders, and to be the least changed descendants of the ancestral group from which most of the other orders of mammals have originated. If this be the case, it seems strange that we should find no traces of them on the continent of South America, which was, doubtless, long isolated from the rest of the world, and which still contains many representatives of primitive and declining types. The paleontological history of the Insectivores is, however, as yet very incomplete, as very few fossil forms of this order have been described. It is, therefore, possible that when future discoveries have increased our knowledge on this subject, this seeming anomaly may be explained. The Neotropical Bats (Chiroptera) are of much interest ; they are included in three families, of which two (Vesper- tilionide and EHmballonuride), although containing several peculiar genera, are found in other parts of the THE NEOTROPICAL REGION 63 world, but the third (Phyllostomatidz) is entirely confined to this region.! This family, which numbers among its members the true Vampires or blood-sucking bats (Desmodus and Diphylla), is a very considerable one, numbering at least sixty species, distributed among thirty-three genera, which are doubtless still to be supplemented by future discoveries. Finally, the Neotropical Region possesses two families of Monkeys, the Marmosets (Hapalidx) and the Capuchins (Cebidex), neither of which is found elsewhere. More- over, both these groups are distinguished from their Old World allies by very important anatomical characters, which render them absolutely distinct from the Old World monkeys and apes. As a representative of this latter family we give a figure of the Barrigudo Monkey (Lagothrix hwmboldtr) of Upper Amazonia (Fig. 12, p. 64) of which Mr. Bates has written us an excellent account in his well - known “Naturalist on the Amazons.” Summarising these statements, we find that the Neo- tropical Region is characterised by the exclusive possession of no less than ten families of mammals, namely :— Bradypodide (Sloths) ; Caviide (Guinea-pigs) ; Myrmecophagide (Ant-eaters); | Solenodontide (Solenodonts) ; Chinchillide (Chinchillas) ; Phyllostomatidz (Vampire bats) ; Dasyproctide (Agoutis) ; Hapalide (Marmosets) ; Dinomyidz (Dinomys) ; Cebidee (Capuchin monkeys) ; and by the presence of about 130 genera, of which about 103 are restricted to its boundaries. On the other hand, when we compare the fauna of the 1 One species, Macrotus californicus, has wandered as far north as California. 64 THE GEOGRAPHY OF MAMMALS Neotropical with that of other regions, the deficiencies or “lipotypes” are manifestly considerable. For example, the following ten families of mammals, all fairly well Hie. 12.—THE BARRIGUDO. (Lagothriz humboldti.) spread over the rest of the world except Australia, are entirely absent from this region :— Bovidee (Oxen). Equidee (Horses). Elephantide (Elephants). Lagomyide (Pikas). Viverride (Civets). Talpidze (Moles). Pteropidee (Fruit-bats). Lemuridee (Lemurs). Cercopithecide (Old World monkeys). Simiidee (Anthropoid apes). THE NEOTROPICAL REGION 65 Section II]—Svspivision oF THE NEOTROPICAL REGION The divisions of the Neotropical Region, as based on a consideration of the class of birds (8), are six in number, namely :— 1. The Antillean Sub-region, containing the Greater and Lesser Antilles, exclusive of Tobago and Trinidad. 2. The Central-American Sub-region, containing all that part of the whole region that is north of Panama. 3. The Colombian Sub-region, containing Trinidad and the slopes of the Andes, through Venezuela, Colombia, Ecuador, and Peru, into Bolivia. 4, The Amazonian Sub-region, embracing the whole watershed of the Orinoco and Amazons up to the hills, and including the highlands of Guiana. 5. The South Brazilian Sub-region, containing the wood region of South-East Brazil, Paraguay, and the adjoining districts. 6. The Patagonian Sub-region, containing Patagonia, Southern Argentina, and Chili, and running up the west coast of the continent to Guyaquil. This division, although perfectly good when the dis- tribution of Birds is mainly relied upon, presents consider- able difficulties in the case of Mammals, owing chiefly to our ignorance of the limits of the distribution of the greater number of the South American mammals, especially of the smaller forms. There is, however, no doubt that the Antilles or West Indies (excluding Trinidad and the other islands off the coast of Venezuela, which are connected with the mainland by quite shallow water) form E 66 THE GEOGRAPHY OF MAMMALS a very well-marked Sub-region, in which the terrestrial mammals, though not very numerous, nearly all belong to peculiar genera. The higher ranges of the Andes from Ecuador south- wards, together with the pampas of Southern Argentina and Patagonia, form another well-marked Sub-region char- acterised by a number of peculiar genera and species. But the whole remainder of the Neotropical Region from Mexico to Southern Brazil contains, so far as we under- stand it at present, a more or less homogeneous mammal- fauna, of which, however, the northern half possesses a considerable admixture of Nearctic forms, while the southern preserves a more purely indigenous facies. It will, therefore, be quite in accordance with the facts of nature, as well as convenient, to separate the northern portion of this extensive area as the Central American (or Transpanamanic) Sub-region. But as regards the southern portion, until our knowledge of the distribution of South American mammals has made greater progress, it seems best to unite the Colombian, Amazonian, and Brazilian Sub- regions of the Ornithologists into one combined Sub-region, which may be called the Guiano-Brazilian Sub-region. We shall thus have, as regards Mammals, four Sub- regions of the Neotropical Region, as follows (see Map, Plate IIL., p. 82) :— 1. The Antillean Sub-region, comprising the whole of the West India Islands except Curagao, Trinidad, and Tobago. 2. The Central-American Sub-region, comprising the low-lying and southern parts of Mexico and Central America as far as the Isthmus of Panama. 3. The Guiano-Brazilian Sub-region, comprising the THE NEOTROPICAL REGION 67 greater part of South America from the Isthmus of Panama to the southern limits of the great forest in about lat. 30° S., and from the forest of the eastern slopes of the Andes to the Atlantic, including Trinidad and the other islands off the coast of Venezuela. 4. The Patagonian Sub-region, comprising the higher ranges and western slopes of the Andes from Guyaquil to Tierra del Fuego and the pampas of Argentina and Patagonia. SEecTION [V.—THE ANTILLEAN SUB-REGION The Mammal-fauna of the Antillean Sub-region is exceedingly poor, so poor, indeed, that it seems almost doubtful whether the islands of which it is composed have ever been directly connected with the mainland of America as at present constituted. To begin with the Rodents, four genera of this order are represented within its limits, and three of these are restricted to the Sub-region. Mega- lomys (a large rat, over twelve inches in length without the tail) is allied to the Vesper-mice of the American continent, and has been obtained only in the islands of Martinique and St. Lucia, where it is now becoming very rare (10). A more important factor in the Antillean mammal-fauna is Capromys, a genus allied, according to Flower and Lydekker, to the Coypu rat of South America, but also showing some affinities to the Porcupines. There are five or six species of this genus usually recognised, of which two or three are restricted to Cuba, one is peculiar to Jamaica, and one to the Bahamas, while another species has been recently discovered in Swan Island, situated 68 THE GEOGRAPHY OF MAMMALS in the middle of the Gulf of Mexico. An allied genus (Plagiodon) with one species, differing from Capromys only in a slight modification of the teeth, is found in Hayti. It is obviously derived from the same stock. A peculiar species of agouti (Dasyprocta cristata) is found in two of the Lesser Antilles—St. Vincent and St. Thomas; the other members of this genus belong to the mainland of Central and South America. The only other Order of terrestrial Mammals represented in this Sub-region is the Insectivora, and this is the more remarkable because, as already shown, one of the special characters of the Neo- tropical Region is the almost complete absence of this group within its limits. The West Indian representatives of this group are two in number, and belong to a peculiar genus (Solenodon), which is of rather doubtful affinities but of family rank. It is allied in some respects to the moles (Talpidx), and in others to a peculiar Malagasian family, the Tenrecs (Centetidx). The Solenodonts are small creatures with a somewhat shrew-like aspect, a long snout, and a long naked tail. There are two representative species of this genus confined to the islands of Hayti and Cuba respectively. The Bats of the Antillean islands, as would naturally be expected, are somewhat more abundant than the terres- trial mammals. There have been recorded by naturalists about thirty species belonging to some twenty genera as found in the different islands, the greater number being from Cuba and Jamaica. On examining the list, it will be found that of these twenty genera three only seem to be con- fined to the West Indian Region, while thirteen are spread over the greater part of the Neotropical Region, one belongs to the Nearctic Region, and the others are of wider distribution. THE NEOTROPICAL REGION 69 The evidence of the Bats, therefore, points unmistak- ably to the inference that the West India Islands have been peopled with Mammalian life from South and not trom North America. In this connection it may be noted that Mr. Frank Chapman in an instructive article (4) on the origin of the West Indian fauna, recently published, has stated, as regards the birds, that the total number as yet recorded as met with within the limits of this Sub-region amounts to 550. Of these 303 are endemic, while the remaining 247 may be allotted to the countries from which they have been apparently derived as follows :— Common. to North and South America . : : a TG Of general distribution in the tropics. 56 South American (ten in the Windward Teavides ame ee wile in the Greater Antilles). ; : : ‘ a nls Central American . 5 : : : : : : 3 North American (all annual migrants from the north, through Florida, and the larger proportion found in Cuba) . 5 : ; : : : : P = 60 This summary gives us a clue to the origin of the more recent additions to the West Indies fauna, which is obviously by migration from the north. If, however, the relationships of the 303 endemic species of Birds are examined, it will be at once evident that they are all more closely allied to South American than to North American forms, and, like the Bats, show that the islands have been stocked with life from the south. Moreover, Mr. Chapman, as well as Mr. Wallace, has pointed out that nearly all the more distinct and most characteristic West Indian Birds are found in the Greater Antilles (i.e. Cuba, Hayti, Porto Rico, and Jamaica), and that the Lesser Antilles form a distinct group, the line of separation between the two provinces coinciding nearly with the deep channel between 70 THE GEOGRAPHY OF MAMMALS the Virgin Islands on the west, and the Anguilla group on the east. On the whole the Lesser Antilles appear to have derived their fauna direct from South America, but probably in times considerably remote, and without the | assistance of a land-connection. The almost complete _ absence of terrestrial mammals and of representatives of sedentary and non-migratory birds in the Lesser Antilles seems to show this. For instance, Grenada possesses only fifteen land-birds also found in Trinidad, and of these none belong to the sedentary families, although the two islands are only separated by an interval of seventy-five miles of sea. On the whole, therefore, it seems probable that, although the Lesser Antilles have derived the bulk of their fauna from South America, they have never been directly connected with that continent. Of the Greater Antilles, Jamaica and Cuba have by far the richest endemic faunas, whereas in Hayti and Porto Rico the total number, as well as the number of endemic species, is considerably smaller. Jamaica, therefore, with its small area (one-tenth of that of Cuba, one-eighth of that of Hayti, and a little more than that of Porto Rico), and in spite of its more isolated position, contains on the whole, so far as our present knowledge goes, the richest fauna. This may, perhaps, be explained by the fact that between the north-east corner of Honduras and Jamaica there stretches a series of more or less shallow banks, so that a comparatively slight elevation of the intervening sea- bottom would very nearly connect Jamaica with the mainland. Whether such a complete land-connection (or only an THE NEOTROPICAL REGION Fal approximation of land areas) by this route ever existed, cannot at present be detinitely settled. On the one hand, as pointed out by Mr. Chapman, the disproportionately rich fauna of Jamaica, the shallow sea, and the West Indian affinities of Swan Island (as shown by the presence of a species of Capromys) lead to such a direct connection. On the other hand, the scarcity of land-mammals in Jamaica and Cuba, and the absence of many families of Birds found on the mainland, rather point the other way. The help in these questions to be derived from paleontology is up to this time very scanty. Almost the only remains of fossil mammals that have been yet obtained from the West Indies are certain detached teeth and some fragmentary bones, found in some caves in the island of Anguilla, which is situated just to the east of the deep channel separating the Greater from the Lesser Antilles, and must, therefore, be included in the latter province. These remains have been described by Professor Cope (5), who considers them to be related to the Chinchillas, a family of rodents con- fined to South America. The nature of the Mammalian genera of the Antillean Region is summarised in the subjoined table :— Insecti- | Rodents. aa Bats. | Total. | Endemic 3 i 4 8 Nearctic . 0 0 1 1 Neotropical] : : : 1 0 boos i ts American : : : : P| geek 0 1 ag | Cosmopolitan . A 10 0 Die Win et Total . : ee 1 20 25 ~I bo THE GEOGRAPHY OF MAMMALS SECTION V.—THE CENTRAL AMERICAN SUB-REGION This Sub-region, as has been already shown, contains the coast-lands of Mexico lying along the Pacific and Atlantic shores from Mazatlan on the north on one side, and from the Rio Grande on the other, together with the whole of Central America from the Isthmus of Tehuantepec to that of Panama. As regards the fauna of this part of America, we are fortunate in being able to refer to the excellent account of it contained in the “ Biologia Centrali-Americana ” of Messrs. Salvin and Godman. The volume relating to the mammals in this work was undertaken by Mr. Alston, and finally completed after his death by one of the authors of this work in 1882 (3). Out of a total of sixty-nine genera of mammals represented in this Sub-region, only two seem to be absolutely restricted to it, and these, moreover, are genera of bats, which may possibly be found at some future time to extend into the main South American continent. Of the sixty-seven non-peculiar Central - American genera of mammals, forty-one are Neotropical types— that is, found also in one or more of the other Sub-regions of this Region—five are Nearctic, nine are found both in the Nearctic and in other Sub-regions of the Neotropical Region, and twelve are cosmopolitan, or, at any rate, found in some part of the Old World as well as in the New. These facts show conclusively the thoroughly Neo- tropical character of the Central-American Sub-region, which, although the admixture of northern forms has really made very little progress, may be defined as that part of the Neotropical Region which has been subjected to an incursion of Nearctic types. THE NEOTROPICAL REGION 73 The Marsupials in the Central American Sub-region are represented by two genera containing about seven species, most of which are also found further south. One of these —the common opossum (Didelphys marswpialis)—extends northwards into the Nearctic Region as well as far south- wards into Brazil, where, however, it has a slightly modified form. The Edentates are well represented in the Central- American Sub-region by two Sloths, three Ant-eaters, and an Armadillo, although the greater number of these are met with only in the most southern portion of the Sub- region. The Armadillo (Tatusia novemcincta) is a widely spread species, ranging from Texas throughout the Sub- region, and extending southwards to Paraguay. Central America is also remarkable for possessing two out of the four American species of Tapir exclusively confined to it; these are Baird’s Tapir (Tapirus bairdt), extending from Mexico to Panama, and Dow’s Tapir (7. dow), found only in Guatemala and Costa Rica. Very few of the typical South American Hystricomor- phine Rodents extend as far north as Central America. The greater number of the animals of this order found within Central American limits belong to the Sciurine and Murine groups, which have spread southwards from their homes in the Nearctic Region. Passing on to the Carnivora, we find nearly all the genera of this order met with in the South American continent (amongst which are the Cats, Dogs, Racoons, and Weasels) also represented in this Sub-region. The only exceptions are Jcticyon, a peculiar form of Wild Dog re- stricted to South-East Brazil, and the Bear (Ursus), a species of which is found in the Andes of Peru. 74 THE GEOGRAPHY OF MAMMALS The Bats of Central America are fairly numerous, and nearly all belong to South American genera. A large pro- portion of them are referable to the Phyllostomatidez, one of the characteristic Mammal-families of the Neotropical Region. Finally, as regards the Monkeys, the Marmosets (Hapa- lidx) appear to be represented only by a single species, which is an intruder into the extreme southern end of the Sub-region. Of the other family of American Monkeys (Cebidx), about eight species, against a total of at least sixty found in the Guiano-Brazilian Sub-region, occur in the Central-American Sub-region. Of these five are peculiar, or not yet ascertained to occur elsewhere. The following table gives the statistics of the origin and distribution of the Central-American genera of Mammals. The “Endemic” genera are those confined to this Sub- region; the “ Nearctic” genera are those common to this Sub-region and the Nearctic Region; the “ Neotropical ” genera are those common to this Sub-region and one or more of the other Sub-regions of the Neotropical Region ; “ American” designates those found in both the Neotro- pical and Nearctic Regions, and “Cosmopolitan” those met with also in the Old World. [eats 5 SU Sy soul etsesal teh Ie |= | Sale en 5 e Endemic . OOP 0.71) OF OPO iliac maa et Nearctic Rae, OO} ©} Oi} 2 i Si lt | @ | © 5 Neotropical . . . 1 4 i 6 2 Om) 22 Oy Al American . aero) (as ay MMO meee OM iarmde ih Gihe | 10) 9 Cosmopolitan. ; | 70 1°00) 0. \°4)| 4 "Os aq no sae otal 2. 2a A elle | 1 L 2 | 28 5 | 69 THE NEOTROPICAL REGION (6) The inspection of this table will show at a glance that the Central-American Sub-region is predominantly “Neo- tropical” as regards its mammals, but has received a small immigration of Nearctic forms, and possesses only few endemic types. SEcTION VI.—THE GUIANO-BRAZILIAN SUB-REGION This extensive area, in which is combined three of the Sub-regions usually allotted to Birds, is the largest and by far the richest of the four divisions of the Neotropical Region adopted in the present article. It extends from the Isthmus of Panama in the north to about 30° south latitude. But the southern frontier between this and the Patagonian Sub-region is very undecided, the Fauna of Uruguay and the northern part of the Argentine Republic containing forms characteristic of both Sub-regions. The western boundary is formed by the well-watered and forest-clad western slopes of the Andes, the waterless eastern slopes, together with the western slopes of Peru and Bolivia from the neighbourhood of the Equator downwards, being referable to the fourth or Patagonian Sub-region. The greater part of the Guiano-Brazilian Sub-region consists of forest, and the Mammalian fauna, though tolerably abundant, is not nearly so profuse as that of the Birds and Insects, which are here both developed in far greater luxuriance than in any other part of the world. This Sub-region is more especially the home of the peculiar Platyrrhine monkeys, the arboreal Sloths, and other tree-loving Mammals. Beginning at the bottom of the list, we find that nearly 76 THE GEOGRAPHY OF MAMMALS all the twenty-four species of Opossums known from the New World are found within its limits. The peculiarly modified Water-opossum (Chironectes) occurs all over its area, but also extends into the Central-American Sub- region. Among the Edentates the Sloths are the most characteristic inhabitants of its forests. But two peculiar genera of Armadilloes (Xenuwrus and Priodon) are confined to this Sub-region, and several other species of this group occur there. All three genera of Ant-eaters are also here met with. Rodents are very abundant in this Sub-region, but, with the exception of Squirrels (Sciwrus), Vesper-mice (Hesperomys), Pouched mice (Heteromys), and a single Hare (Lepus), they all belong to the Hystricomorphine group, which is so highly developed in the Neotropical Region. The Guiano-Brazilian Sub-region is also the special home of the Phyllostomine bats. Out of a total of about sixty- five species of this family forty-four are found in this Sub-region, and the greater number of them are confined to it. Such, too, is the case with the Platyrrhine monkeys. The Marmosets (Hapalid#) would be also unknown out- side the limits of the Sub-region had not a single species, as already mentioned, overstepped the northern boundary at Panama. The Capuchins (Cebidx), numbering more than sixty species and belonging to ten genera, are likewise abundant, and are found elsewhere only in the Central-American Sub-region. The following table shows at a glance the numbers of (1) the “ Endemic” genera of this Sub-region, 7.e. those not found beyond its limits ; (2) the “ Neotropical ” genera, 2.¢. THE NEOTROPICAL REGION 77 those confined within the limits of the whole Region; (3) the “ American” genera, 7.e. those occurring in other parts of the New World, but not beyond; and (4) the “Cosmo- politan,” ze. those of general distribution :— | a S| = Sali Pan a Endemic . Oa 2a Or iOrn I On elulealieZ aiezOL | Neotropical PO On shoe LnO! eh | vo eek American a SO ae | Oat 10} Cosmopolitan LOR Can ant A Ue | ON WEES att | Total FeO Se 200 10) 5 |) ete s8h) 12.105 SEcTION VII.—THE PaTAaGoONIAN SUB-REGION The Patagonian Sub-region may be most conveniently taken to begin on the south side of the bay of Guyaquil, and to extend thence southwards, embracing the whole western slope of the Andes of Ecuador and Peru. In Bolivia it widens out and includes the high plateau of Titicaca, extending thence over the whole of the Argentine Republic, Chili, and Patagonia. The most characteristic form of the mammals of this Sub-region is the Lama, which, with its allies, consti- tutes the genus Zama. Four forms, usually regarded as distinct species, are recognised by naturalists. Of these two, the Lama (LZ. peruana) and the Alpaca (L. pacos) are only met with in a domestic state, and are very variable in size and colour. Of the other two, which are met with wild, the Guanaco (L. huanacos) has the most 78 THE GEOGRAPHY OF MAMMALS extensive distribution, ranging from the highlands of Ecuador and Peru, along the Andes, to the open plains of Patagonia; while the Vicugna (LZ. vicugna), which is a somewhat smaller animal, is found only in Ecuador, Peru, and Bolivia. In addition to the Lamas, this Sub-region possesses a species of thickly haired mountain Tapir, differing from the lowland forms, and two or three peculiar Deer, of the sub-genus Furcifer, which are likewise densely furred. A third diminutive deer found in Chili is distinguished from Cariacus, the ordinary American form of deer, by ana- tomical characters, and belongs to a special genus, Pudua. A second species of this form (P. mephistopheles) from the highlands of Ecuador has lately been described by Mr. De Winton. The Rodents of the Patagonian Sub-region almost all belong to the Hystricomorphine section of the Order. Amongst them are the Chinchillas (Chinchilla and Lagi- dium), noted for their delicate fur, the Viscacha (Lago- stomus), and the Patagonian Cavy (Dolichotis). Out of the eighteen genera of this division known to occur in the Patagonian Sub-region, ten are restricted to it. Of the carnivorous Mammals of this Sub-region, one of the most interesting is the Spectacled Bear of the Andes (Ursus oratus), which affords an instance of that rare phenomenon in nature “discontinuous distribution,” the nearest allied species of bear (the black bear of North America) only coming as far south as Mexico. The presence of a bear in the Andes can only be explained by the supposition that the ancestral form migrated south- wards along the line of the Cordilleras, but has died out in the intermediate district. The Bats of the Patagonian THE NEOTROPICAL REGION 79 Sub-region present but few features of interest; they are, with one exception, all of genera found also in the Guiano- Brazilian Sub-region. | As might have been expected from the dearth of forests and the generally severe climate, the American monkeys and marmosets are entirely unrepresented in the Patagonian Sub-region, On the other hand, at least two small species of Opossum (Didelphys and Dromiciops) occur in Chili, and a very remarkable form of Armadillo (Chlamy- dophorus) is peculiar to Argentina and the high plateau of Bolivia. The following list of the mammal genera of the Pata- gonian Sub-region is constructed on the same plan as that of the preceding tables :— ae [ae 8 g : Ee io Sy a agit Persea | a ; = S Hd emic? .¥ex 2 Sy ec ol 2 De LOR od 1 i Neotropical iO 2 0 ae eae 5 | 16 American, <2 5 |) 1 1 1 0 1 2 6 Cosmopolitan |) Oa Ue 4 2 8 Potaly 7”. 2S eo 4 18 8 10 47 Section VIII.—TuHe Past History or THE NEOTROPICAL MAMMAL-FAUNA During the last few years our knowledge of the extinct mammals of the Neotropical Region has been enormously increased by the discoveries of the palzeontologists of the Argentine Republic, more particularly by the labours of 80 THE GEOGRAPHY OF MAMMALS Burmeister, Moreno, and Ameghino. A few words about this branch of the subject may be added. The oldest formation containing well-preserved remains of mammals yet investigated is in the neighbourhood of Santa Cruz, in Southern Patagonia, where the deposits are about 200 feet in thickness. The exact age of the Santa Cruz beds it is very difficult to determine, but the best authorities consider that they cannot be of earlier date than the Upper Eocene or Oligocene of Europe. Follow- ing these in point of time, are the so-called “ Patagonian ” beds of Patagonia and Uruguay, which are probably of Miocene age. The “ Araucanian” formation of Ameghino, which is well developed at Monte Hermoso, near Bahia Blanca, in Southern Argentina, seems to correspond approximately with the older European Pliocene. Finally, the later Pliocene is apparently represented by the “Pampas” formation of Argentina and Uruguay. Our knowledge of the extinct mammal-faunas of these beds is mainly due to the efforts of the Argentine palzeontologists just mentioned, but a clear réswmé of the work done will be found in a recent number of the Geological Magazine (12). In the “Santa Cruz” beds have been found remains of about 120 genera of mammals referable to the following groups :— Marsupials. Perissodactyles. Rodents. Edentates. Toxodonts. Monkeys. Among the Marsupials the most prominent forms in this formation are the Opossums (Didelphyidz), which are still found all over America, and in Tertiary times appear to have been distributed nearly all over the northern hemi- sphere. But accompanying these are other forms of the THE NEOTROPICAL REGION 81 same Order, which seem to be more nearly related to the Australian marsupials of other families; in fact, Ame- ghino has placed some of them in the (hitherto supposed to be) exclusively Australian family Dasywride. The fossil Edentates of this formation are all of the American section of the group. The Perissodactyle Ungulates are represented by two families (Protero- therudex and Macraucheniidx)not very definitely connected with other Perissodactyles, and unknown elsewhere; and the Toxodonts (another group forming a distinct sub-order of the Ungulata) are also quite unrecognised outside South America, where, indeed, they appear to have existed for a short period only. The Rodents of this formation all belong to the Hystri- comorphine section of the order, and the Monkeys are all Platyrrhine, so that, except in the case of the Marsupials where the results are to a certain extent doubtful, no clue to the origin of the Neotropical fauna is shown by this extinct fauna. When these results are compared with the Eocene Mammal-fauna of the northern hemisphere, the absence of Artiodactyles, Insectivores, Bats, Carnivores, and Lemurs, is very striking. The “ Patagonian” beds contain a Mammal-fauna only to be distinguished from the previous “Santa Cruz” series by a further differentiation of genera belonging to the same orders without any traces of foreign admixture. When, however, we reach the more recent Araucanian formation, we find, in addition to the Edentates, Toxodonts, and other typical South American forms, a number of foreign intruders, such as Tapirs, Lamas, Elephants (Mastodon), and Wild Dogs (Canis), of an entirely different aspect. There can be little doubt that these animals had migrated F 82 THE GEOGRAPHY OF MAMMALS here southwards from North America, where their remains (or those of closely allied species) have been likewise found in the nearly contemporaneous formations of the United States. Moreover, it would seem that in these bygone days, not only did the northern forms move southwards, but that also some of the southern forms emigrated north- wards. This is evidenced by the fauna of the so-called « Hquus-beds” and “ Megalonyx-beds” of a slightly later date in the United States, which contain a composite Mammal-fauna of northern forms mixed with forms usually considered to be exclusively South American— such as the gigantic armadillo-like Glyptodon, the Capy- bara (Hydrocherus), Toxodon, and others. Finally, in the age of the “ Pampas” beds, the peculiar South American Mammal-fauna seems to have reached its culminating-point, and to have far exceeded that of the present day both in number of species and in the size of the individuals. This great increase in size, which is, as a rule, accompanied by an extreme specialisation of individual organs, seems to have had a fatal effect on its possessors, as none of the larger Edentates or Toxo- donts appear to have outlived the end of the pampas formation. Along with most of the larger arrivals from the north, such as Mastodon and Equus, they became extinct. All the conclusions to be derived from this much-abbreviated account of the extinct Mammals of South America, confirm in a remarkable way the evidence of the present fauna as to this history of the Neotropical Region. Up to the last period of the Tertiary epoch, South America was certainly isolated from the rest of the world, and the connections with Australia and with Africa, if they ever did exist, must have been previous to this a THE NEOTROPICAL REGION 83 period. At the beginning of Pliocene time, during the deposition of the Araucanian formation in Argentina and the “ Hquwus-beds” in the United States, a wide bridge between North and South America, affording an easy road to migrating animals, must have existed, and this again seems to have become considerably narrowed to form the present Isthmus of Panama. List OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHaprer III. (1) Auten, J. A.—“The Geographical Distribution of North American Mammals.” Bull. Amer. Mus. N.H., iv., p. 199, 1892. (2) AuLEN, J. A.—“On a Small Collection of Mammals from the Galapagos Islands, collected by Dr. G. Baur.” Bull. Amer. Mus. N.H., iv., p. 47, 1892. (3) Auston, E. R.—‘ Biologia Centrali-Americana, Mammalia. London, 1879-82. (4) Caapman, F. M.—‘ Notes on Birds and Mammals observed near Trinidad, Cuba, with remarks on the Origin of West Indian Bird-life.” Bull. Amer. Mus. N.H., iv., p. 279, 1892. (5) Copr, E. D.—“ Description of two large extinct Rodents from Anguilla, West Indies, with remains of human art associated.” Proc Amer. Philos. Soc. Philad., xi., 1871, p. 183. (6) Letpy, J.—“ Remarks on Mylodon.” Proc. Acad. Nat. Sct. Philad., 1885, p. 49. (7) Satvin, O.—“ On the Avifauna of the Galapagos Archipelago.’ Trans. Zool. Soc., 1x., p. 447 (1876). (8) Sctater, P. L.—‘‘ Address to Section D. (Biology).” “Report | of Forty-fifth Meeting of the British Association, at Bristol,” p. 85 (1876). (9) THomas, O.—‘‘ On Some Mammals from Central Peru.” P.Z.S., 1893, p. 333. (10) Trovrssart, E. L.—‘ Note sur le rat musqué (Mus pilorides) des Antilles.” Ann. Sct. Nat. Zool. (6), xix., No. 5 (1885). (11) Wattracg, A. R.—‘ The Geographical Distribution of Animals.” 2 vols. London, 1876. (12) Zrrret, K. A. von.—‘ The Geological Development, Descent, and Distribution of the Mammalia.” S. B. &. bayer. Akad. Wiss,, xxiii., p- 137, 1893. Translation in Geol. Mag. (3), x., p. 401 (1893). CHAPTER IV THE ETHIOPIAN REGION (Puate IV., p. 122) Section I.—BouNDARIES OF THE ETHIOPIAN REGION THE Ethiopian Region (see Map, Plate IV.) contains the whole of Africa south of the Sahara, together with Southern Arabia and the island of Madagascar. As in all other cases where there is a long land-frontier between two neighbour- ing Regions, so here it is impossible to lay down anything but an approximate line of demarcation between the - Ethiopian and Palearctic Regions. The boundary usually adopted is the line of the Tropic of Cancer, which strikes Africa between Morocco and Sene- gambia, runs through the middle of the Sahara, crosses the Nile between the first and second cataracts, and passes through Arabia to the neighbourhood of Oman, on the Persian Gulf. Most of the country through which this line passes is desert, and its mammalian fauna is con- sequently meagre. Mr. O. Thomas (6) has recently published an account of a collection of mammals received at the British Museum from Oman, which shows, as would naturally be expected, that “the geographical relationships of this district are about equal with Africa and India; three of the species being distinctly African in affinities, three Indian, and the remainder either peculiar or widely spread and of no special significance.” On the whole, 84 THE ETHIOPIAN REGION 85 therefore, the line of the tropic of Cancer, adopted by Wallace, seems to be a fairly suitable boundary. Besides the island of Madagascar and its appendages, which contain one of the richest and most interesting of all known Insular Faunas, and form a very important Sub-region, which will be considered in greater detail below, the Ethiopian Region possesses other islands, These, however, are mostly Oceanic, and not of any great importance. The Azores, Madeira, Canaries, and Cape Verde groups, although geographically African, seem to have derived their animals chiefly from Europe. They therefore belong to the Palearctic Region, and will be dealt with under that head. The islands of St. Helena and Ascension, situated in the South Atlantic, are both of them of volcanic origin, and separated from the mainland of Africa by more than 800 miles of deep water. Neither of these islands possesses any Vertebrates. The only land-groups well represented in them are the Beetles and the Land-shells—a study of which shows that the affinities of these islands are to Southern Europe and Southern Africa, but that the Fauna is in all probability an exceedingly ancient one; since its pecu- liarities are very great, and opportunities of migration of new forms to these islands have been few and far between. The other islands in the Atlantic connected with Africa are Fernando Po, Prince’s Island, St. Thomas, and Anno Bon, all situated in the Gulf of Guinea at various distances from its head. Fernando Po, an island of some 40 miles in length, but separated from the mainland by a some- what shallow sea about 20 miles across is said to be 86 THE GEOGRAPHY OF MAMMALS inhabited by a considerable number of Mammals which, however, in all probability, also occur on the adjacent mainland. The other islands above mentioned, Prince’s Island, St. Thomas, and Anno Bon, are all considerably more distant from the coast of Africa, and, so far as we know, do not possess any native Mammals. A certain number of Land- birds have been obtained from these islands, and a few of these are peculiar, but they are all closely allied to forms that occur in the neighbouring West African mainland. The most important islands in the Indian Ocean are, apart from Madagascar, Socotra, the Seychelles, the Amir- ante Islands, the Comoros, and finally Réunion, Mauritius, and Rodriguez. Socotra is situated about 150 miles due east of Cape Guardafui, the extreme eastern point of Africa. Its fauna and flora were carefully investigated some years ago by Professor Bayley Balfour (1). He found that the only Mammals inhabiting the island are a Bat, which he was unable to obtain, and a Civet (Viverricula), met with also in South Asia, and probably introduced by human agency. The Seychelles are distant about 700 miles from the northern point of Madagascar in a north-easterly direction, a more or less continuous chain of islands (Amirante, Provi- dence, and Farquhar) forming a series of stepping-stones between them. But two narrow deep channels of over 1000 fathoms run between the Seychelles and Amirante on the one hand, and between Amirante and Providence on the other, thus cutting the Seychelles off from Madagascar by their deep water. THE ETHIOPIAN REGION 87 There do not appear to be any indigenous terrestrial Mammals in the Seychelles, except two Bats, which have been recorded as occurring there. The Land-birds are most of them peculiar, though belonging to genera found in Madagascar or Africa; the reptiles and amphibians are fairly numerous for islands such as these, and several of the species are not found elsewhere. It is difficult to say whether these islands have ever had a land-connection with Madagascar, but probably, if such were ever the case, it was at a remote time, geologically speaking. The Comoros are a group of several islands lying about midway between the most northerly point of Madagascar and the mainland of Africa. They are separated from Africa by the Mozambique channel, more than 1000 fathoms in depth, while the depression between them and Madagascar is considerably less. Inhabiting these islands there are three Bats. Two of these are of the genus Pieropus, which is entirely absent from the mainland of Africa, but is found in Madagascar and many islands of the Indian Ocean. There is also a peculiar species of Lemur (Lemur mayottensis) and the Indian Civet (Viver- ricula), which last is also found in Madagascar and Socotra, and has in all probability been introduced by human agency. A fair proportion of the Land-birds of the Comoros, though in some instances peculiar, belong to Madagascar forms, and everything goes to show tnat the relation of this group of islands is with Madagascar rather than with the mainland. The islands of Réunion, Mauritius, and Rodriguez are distant from Madagascar in an easterly direction 450, 550, and 800 miles respectively. They are all of volcanic origin, and separated from Madagascar by water of more 88 THE GEOGRAPHY OF MAMMALS than 2000 fathoms in depth. As would be expected, these islands are thoroughly Oceanic, and possess no indigenous Mammals or Amphibians ; while the Land-birds are few in number, and belong mostly to genera found in Madagascar. The most remarkable feature, however, of the Fauna of these islands is the former existence of a group of flightless Ground-birds now quite extinct, but some of which were found in great numbers when the islands were first dis- covered. These are the Dodos of Mauritius and Réunion respectively, and the Solitaire of Rodriguez. These birds form a distinct family—the Dididz, probably allied to the Pigeons, but of somewhat obscure affinities. It seems that the ancestors of these birds must have reached the islands in very early times,and that most of the striking pecu- liarities exhibited by them were gradually acquired after their arrival in the group. | We may, at any rate, conclude that these three islands are truly Oceanic, and that they have never had a land- connection with Madagascar or elsewhere. Section II].—GENERAL VIEW OF THE MAMMAL-FAUNA OF THE ETHIOPIAN REGION The Ethiopian Region, as will be seen by looking at the Tables of the numbers of families, genera, and species given at the end of Chapter I. (p. 16), is the richest of the six Regions as regards the total numbers of its families, genera, and species of mammals, although the percentage of peculiar forms not found in other Regions is hardly so high as in the Neotropical and Australian Regions. This may, however, be accounted for by the consideration that there is a long THE ETHIOPIAN REGION 89 land-frontier between the Ethiopian and the Palearctic Regions, though this is chiefly occupied by desert. Out of the nine orders of Terrestrial Mammals the Ethiopian Region contains representatives of seven, the Fig. 13.—THE CAPE AARD-VARK. (Orycteropus capensis. ) Marsupials and Monotremes being alone absent. The Edentates of this Region are represented by two families. One of these, of which the sole genus is the Aard-vark (Orycteropus), is quite restricted to the Region (see Fig. 13). 90 THE GEOGRAPHY OF MAMMALS The other, containing the scaly ant-eaters (Manidz) is found also in the Oriental Region (see Fig. 14). These two forms are in most respects more closely allied to one another than to any of the Edentates of the New World. It is, however, the animals belonging to the Order ™ WD am Fig. 14.—THE WHITE-BELLIED PANGOLIN. (Manis tricuspis. ) Ungulata which form so conspicuous a factor in the Ethiopian fauna. These are distributed among thirty-nine genera, of which no less than twenty-four are not found anywhere outside this Region. The antelopes, of the family Bovidy, which are most THE ETHIOPIAN REGION 91 of them large animals living in vast herds in the more open country of eastern and southern Africa, take up the greater number of these genera. These are perhaps, at the present epoch, the most notable feature of the African Fauna, but will doubtless, owing to the unceasing persecu- tion of hunters and sportsmen, become rapidly less so. Already the larger Antelopes are nearly extinct in the Cape Fig. 15.—THE HIPPOPOTAMUS. (Hippopotamus amphibius. ) Colony south of the Orange River, and there can be little doubt that, unless special precautions are taken, the large Mammals of Africa will very soon disappear, like those of the United States, before the express and repeater. Beside the numerous genera of Antelopes, the Ethiopian Region has exclusive possession of two other conspicuous forms of the Ungulates—the Hippopotamus (Fig. 15) and 92 THE GEOGRAPHY OF MAMMALS the Giraffe (Fig. 16), each of which forms a special family eget Soe se 0, min °F ea he ee ta ‘ Fig. 16.—THE GIRAFFE, (Girafia camelopardalis. ) while it shares the Rhinoceroses and the Chevrotains with the Oriental, and the Horses (Zquide) with the Palearctic THE ETHIOPIAN REGION 93 Region. The only important family of Ungulates not found in the Ethiopian Region is the Deer-family (Cervide). The total absence of this otherwise widespread family it is difficult to explain. Paleontology does not help us much, since the members of the Deer-tribe appear to have been already well established and abundant in Europe during early Pliocene times, and were contemporaneous with Ante- lopes and other Ungulates, which have availed themselves of the opportunity of spreading southwards to Africa, while the Cervidzx have apparently remained obstinately attached to the Palzarctic Region. The Sub-order Proboscidea containing the Elephants is in these days represented only in the Ethiopian and Oriental Regions. The Hyraxes (Hyracidz) are absolutely confined to the Ethiopian Region, and constitute one of its most signifi- cant forms of mammal life, as not being elsewhere met with. Turning now to the Rodents, we find that the Ethiopian Region possesses representatives of a considerable number of genera, twenty-seven of which, out of a total of thirty- nine, are confined to the Region. Two of these genera are sufficiently distinct to be entitled to family rank. These are, first, Anomalwrus, a form resembling the flying squirrels in having a flap of skin available for imperfect flight extended between the fore and hind limbs. But the Anomalures differ from the true Flying Squirrels (Pteromys) in having a long cartilaginous process ex- tending from the elbow-joint to support the parachute, and also in being provided with strong imbricated scales attached to the lower surface of the tail, which are pro- bably of assistance in climbing! 1 A new genus of Anomaluride (Zenkerella), lately described by Herr Matschie, has the imbricated scales on the tail, but no parachute. See P. Z. 8. 1898, p. 450. 94 THE GEOGRAPHY OF MAMMALS The second peculiar Rodent of family rank is Lophiomys, a curious arboreal animal allied to the rats, but differing from them in several anatomical features, and in having a long crest of hair upon the back. Only one species is known, which is restricted to North-East Africa. The distribution of a third Ethiopian family of Rodents (Octodontidz) is particularly interesting, as it is represented in South America by a considerable number of genera, and is not found elsewhere except in the Ethiopian and on the borders between the Ethiopian and Palearctic Regions. There are four African genera of this family, each with a single species, so that in the Old World the group appears to be in a state of decay. The only light thrown on this curious case of discontinuous distribution is the occurrence of a fossil genus (Pellegrinia) allied to the African forms in the Pleistocene beds of Sicily. This indicates that members of this family once had a wider distribution northwards than what they now retain. The Carnivora are well represented in the Ethiopian Region. This is especially the case with the family of Civets (Viverride), only found elsewhere in the Pale- arctic and Oriental Regions. Out of a total number of seventeen genera, three only are found outside this region, and out of the remaining fourteen, six are confined to the island of Madagascar. The Aard-wolf (Proteles), which is so distinct from its fellow Carnivores as to be allowed to constitute a family of itself (Fig. 17, p. 95), is restricted to the Ethiopian Region, and of the Hyenas (Hyznidx), only one of the three species strays outside its limits, into the western borders of the Oriental Region. Another remarkable fact about the Ethiopian Carni- THE ETHIOPIAN REGION 95 vora is the entire absence of the otherwise widespread family of Bears (Ursidx). This group has existed in Europe since the time when the Upper Pliocene beds were laid down, and remains of Bears have been also found in the Siwalik deposits of India, which are possibly Fig. 17.—THE AARD-WOLF. (Proteles cristatus. ) of even greater age. It seems, therefore, very strange that the Bears should never have reached Africa, when so many of the other genera found in the same deposits have managed to do so. The Insectivora of the Ethiopian Region are fairly numerous, and show considerable specialisation. Out of 96 THE GEOGRAPHY OF MAMMALS the six Ethiopian families three are not found elsewhere, and one of the others only extends into the Palearctic Region as far as Northern Africa. This is the family of elephant-shrews (Macroscelidz). The Bats of this Region present few particular features of interest, belonging nearly all to widely spread genera of widely spread families. Among those that are peculiar the most important is Hpomophorus, containing eight or ten species of large fruit-eating bats, with long rather horse-like heads, and expansible and peculiarly folded lips. The Zpomophori seem to take the place, in Africa, of the genus Pteropus, which, although represented by several species in Madagascar, and even in the Comoros (which are separated from the mainland by only a com- paratively narrow strait), is entirely absent from the African mainland itself. Among the Lemurs we have two families, one con- taining only a single species (Chiromyidx) confined to Madagascar, the other (Lemuwridx) containing a large number of genera, of which ten are confined to Mada- gascar, two are found on the mainland of Africa, and the other two in the Oriental Region. The Quadrumana of the Ethiopian Region, which are entirely absent from Madagascar, belong to two families, - both of them shared with the Oriental. The genera, however, six in number, are all without exception con- fined to this Region. In the first place this Region is the only home of the Gorilla and Chimpanzee (Fig. 18, p. 97), two of the Apes most nearly allied to Man in structure, and usually placed at the apex of the mammalian series. The Colobs, or Thumbless Monkeys (Colobus), represent the Langurs (Semnopithecus) of the Oriental Region in THE ETHIOPIAN REGION 97 _ Africa, and number some ten or twelve species. The long- Fig 18.—THE CHIMPANZEE. (Anthropopithecus troglodytes.) tailed Cercopitheci, or Guenons, are essentially arboreal, and are spread all over tropical Africa where trees are G 98 THE GEOGRAPHY OF MAMMALS found. They are very numerous, and from thirty to forty species have been described, some of them, such as the Diana Monkey (Cercopithecus diana) and Brazza’s Monkey (C. brazzx) remarkable for their beautifully coloured fur. Intermediate between the Guenons and the Macaques of the Oriental Region are the Mangabeys (Cercocebus), of which some six species are recognized. Finally, the ter- restrial Baboons (Theropithecus and Cynocephalus) close the Ethiopian series with some eight or ten species. Summarizing these results, therefore, we find that the Ethiopian Region is characterized by the exclusive pos- session of eleven families of mammals, namely— Orycteropodide (Aardvark), Protelidee (Aard-wolf), Hippopotamide (Hippopotamus), § Potamogalidz (River-shrew), Giraffide (Giraffe), Centetidz (Tenrec), Hyracide (Rock-coney), Chrysochloridz (Golden-mole), Anomaluride (Anomalure), Chiromyide (Aye-aye), Lophiomyide (Crested Tree-rat), and by the presence of about ninety-nine endemic genera. On the other hand, among a considerable number of families not represented in this Region, the following five are all widely spread elsewhere, and may therefore be considered as typical “lipotypes” of the Ethiopian Region :— Cervidee (Deer). Ursidee (Bears). Tapiride (Tapirs). Talpidze (Moles), Castoridz (Beavers). Section IIJ.—SupspIvisIon OF THE ETHIOPIAN REGION In dividing the Ethiopian Region into Sub-regions, it must be always remembered that one of the Sub-regions far predominates over the others in speciality and distinct- THE ETHIOPIAN REGION 99 ness, and that the remaining Sub-regions all resemble one another more or less closely, and are difficult of separation. This Sub-region, which is so distinct from the others, consists of the large island of Madagascar, together with the island-groups in its immediate vicinity, viz. the Comoros, the Seychelles, and Amirantes to the north, and the Mascarene islands (Réunion, Mauritius, and Rodriguez) to the east. Whatever other conclusions may be arrived at regarding the best mode of dividing the Ethiopian Region, every authority is, we believe, agreed on this matter, the only doubtful point being whether the Malagasy Sub-region is not well entitled to the full rank of a Region. On the African continent itself a fairly distinct Sub- region can be recognized, extending all over the forest country of Western Africa from the Senegal River over the whole Congo basin, or perhaps rather further south. The best inland boundary of this Sub-region would probably be the water-parting between the West African rivers on the one side, and the Nile on the other. There can, at any rate, be now little doubt that the West African fauna extends nearly as far eastwards as the western bank of Lake Tanganyika, Even on the shores of Victoria Nyanza, according to Herr Neumann’s (5) recent researches, some typical West African forms are met with; but for the present it will be safer to restrict the West African Sub- region to the western watershed. The Southern or Cape Sub-region, as defined by Wallace, includes only the country south of a line drawn from Walfisch Bay, just to the north of the tropic of Capricorn, to Mozambique. Since the publication of Mr. Wallace’s book, however, much addi- tional information has been obtained regarding the distri- 100 THE GEOGRAPHY OF MAMMALS bution of the mammals of Eastern Africa. Many of the animals formerly supposed to be confined to the southern end of the continent, have been shown to extend all through Nyasaland, at least as far north as British Hast Africa. It will, therefore, be advisable to extend the boundaries of this Sub-region further north. The boundary adopted in this paper, as will be seen by consulting the map (Plate IV., p. 122), runs from Angola in the west, along the southern water-parting of the Congo as far as Lake Tanganyika, passing thence to Kilimanjaro, and so on to the Indian Ocean along the Tana River. The rest of Africa, including the Sahara, the southern portion of Arabia, and North-East Africa, will form a fourth Sub-region, which, however, does not contain nearly so high a percentage of endemic genera as the other three. The Ethiopian Region may therefore be divided into four Sub-regions as follows :— 1. The Malagasy Sub-region, including Madagascar and its adjacent islands. 2. The West African Sub-region, including the great equatorial forest of Central Africa contained in the basins of the western rivers, from the Senegal to the Congo inclusive. 3. The Cape Sub-region, including all Africa south of the watershed of the Congo on the West and of the Tana on the East coast. 4. The Saharan Sub-region, consisting (if we exclude the Abyssinian plateau) chiefly of desert, or at any rate of a comparatively dry country, including the Sahara, Eastern Africa as far south as the Tana River, and Southern Arabia. THE ETHIOPIAN REGION 101 SEecTION 1V.—THE MALAGASY SUB-REGION The island of Madagascar is separated from the main- land of Africa by the Mozambique Channel, which, though only about 250 miles across at its narrowest point, is more than 100 fathoms deep throughout its extent. For our recent knowledge of the fauna of Madagascar we are chiefly indebted to the great work of Grandidier! (3). Unfortunately, up to the present time only the plates and a small portion of the letterpress of that part which deals with the mammals have been published. Out of a total number of forty-seven genera of mammals found in this Sub-region, no less than thirty-three are exclusively confined to it. Of the others, two or three occur also on the mainland of Africa, and the remaining twelve—most of which are bats—are cosmopolitan, or at any rate extend beyond the limits of the Ethiopian Region. The Ungulates are represented in Madagascar by only one form—a peculiar River-hog (Potamocherus) closely allied to the South African species, but still sufficiently distinct to have earned a title to a separate appellation. With this exception, the Ungulates, so numerous and so conspicuous a feature on the African continent, are entirely absent from Madagascar. The Rodents are represented in this Sub-region by seven genera of Mice, containing thirteen species, all confined to it. The Malagasy Sub-region possesses six peculiar forms of 1 Dr, Forsyth Major’s recent explorations in Madagascar have added considerably to our knowledge of its smaller mammals, particularly of Rodents and Insectivores, 102 THE GEOGRAPHY OF MAMMALS Fig. 19.—THE Fossa. (Oryptoprocta ferox. ) THE ETHIOPIAN REGION 103 Carnivores, of which the most remarkable is the Fossa or Cryptoprocta—a large cat-like animal allied to the Civets, but sufficiently distinct to form a separate family (see Fig. 19, p. 102) according to some authorities. Of the Civets, besides a Viverricula closely allied to the Indian Civet, there are six genera with eight species of Mongooses, all confined to the island. Of these the most remarkable is Ewupleres—a form the jaws and teeth of which are so weak and small, that it was at one time thought to belong to the Insectivora. On the whole, out of the seven genera of Carnivora found in the island, six are absolutely peculiar. Among the Insectivora of this Sub-region even greater specialisation prevails. Besides two species of ubiquitous shrews (Sorex) said to occur there, there are eight genera of this Order found in Madagascar, all of which are con- fined to the island. Geogale, a small mouse-like animal about which little is known, is said to be allied to Potamogale, an aquatic otter-like animal found only in Western Africa, The remaining seven genera make up the family Centetidx, the affinities of which are somewhat doubtful, but seem on the whole to approach the Solenodonts, a group confined to the Greater Antilles. The best known of the Centetidz is the Tenrec—the largest member of the order, between twelve and sixteen inches long, and devoid of a tail (see Fig. 20, p. 104). The young of this creature are covered with spines like a hedgehog, but these are lost in the adult state. The Bats of Madagascar, as would naturally be expected, do not exhibit the great peculiarities found among the other orders of mammals. Out of about twelve genera only one is confined to the island. The distribution of Pteropus, the genus of large tropical fruit-eating bats, often 104 . THE GEOGRAPHY OF MAMMALS called “ Flying-foxes,” is interesting. Five members of this genus are found in the Malagasy Sub-region, including two species in the Comoro islands, and although these islands are almost within sight of the mainland of Africa, not a single representative of the true Pteropus has yet been obtained on the continent. Fig. 20.—THE TENREC. (Centetes ecaudatus. ) Finally, among the Lemurs we reach the culminating point of the Fauna of this strange land. No less than eleven genera of this Sub-order are entirely confined to the Sub-region, while outside of it there are only five genera now in existence, two of them belonging to Africa proper, and three to the Oriental Region. The lemurs are none of them very large; they are all i i is THE ETHIOPIAN REGION 105 arboreal animals, spending their lives retired in the forest, and, as a rule, strictly nocturnal. Though allied to the monkeys, they have none of their vivacity and intelligence ; Fig. 21.—THE INDRIS. (Indris brevicaudatus. ) they move but slowly, and have usually very large eyes, which are no doubt necessitated by their nocturnal habits. The lemurs inhabitmg Madagascar are divided into two families: (a) the Lemuridex, with ten genera and about 106 THE GEOGRAPHY OF MAMMALS thirty-five species, to which family, also, the African and two of the Oriental genera are generally assigned: and (b) Fic. 22.—_THE RUFFED LEMUR. (Lemur varius. ) the Chiromyide. The Indrises (ndrisine) form a distinct sub-family of Lemwrid# with three genera, all re- THE ETHIOPIAN REGION 107 stricted to Madagascar (see Fig. 21, p. 105). The typical Lemurs (Lemwridx) are also only found in this island (see Fig. 22, p. 106). The second family of the Lemurine Order (Chiromyidz) contains only a single genus and species, the extremely Fig. 23.—THE AYE-AYE. (Chiromys madagascariensis. ) anomalous Aye-aye, discovered by the traveller Sonnerat in 1780 (see Fig. 23). There are generally examples of this curious animal in the Zoological Society's gardens in London, but, unless especially aroused, they are seldom seen by daylight. Their chief peculiarity is the long, thin, 108 THE GEOGRAPHY OF MAMMALS ghost-like middle finger, with which they have been supposed to extract wood-boring insects from their burrows, although their chief food in captivity certainly consists of succulent juices. Of the extinct mammal-fauna of Madagascar we know as yet too little. Dr. Forsyth-Major (4) has lately described a large lemur (Megaladapis) differing considerably from those now inhabiting the island; while remains of two small Lemurs and of two small species of Hippopotamus have been also met with. These remains, together with the bones of a large flightless bird (4/pyornis), apparently allied to other Ratite birds, are of a comparatively recent period. On the whole, however, we cannot but presume that Madagascar originally obtained its animal life from the mainland of Africa. The striking differences between the present faunas of Africa proper and Madagascar are doubt- less due to the fact that the great bulk of the existing African fauna is of comparatively modern origin, and came from the Northern continent at the end of the Miocene or the beginning of the Pliocene times, whereas Madagascar was cut off from Africa before this eruption of Northern forms took place. Madagascar, therefore, appears to contain a sample of the ancient Ethiopian fauna, which has been almost exterminated on the mainland, but has survived here under the protection afforded by its separation from the adjacent continent. The fauna of the Malagasy Sub-region may be summarized by the exclusive possession of— (1) Seven genera of the family Muridz, among the Rodents. (2) Six genera of the family Viverridz, among the Carnivores. THE ETHIOPIAN REGION 109 (3) One genus (Geogale) of the family Potamogalide (shared with the West African Sub-region) and six genera making the whole of the family Centetidy, among the Insectivores. (4) One genus (Chiromys), alone forming the family Chiromyide, and ten genera of the family Lemuride, out of a total of fifteen generally recognized. Almost equally important is the absence of the follow- ing groups :— (1) The orders Edentata and Ungulata (except Pota- mocherus). (2) The families Sciwridx (Squirrels), Spalacide (Blind Moles), Octodontidx, Hystricide (Porcupines), and Lepo- ridx (Hares), among the rodents. (3) The families Felidx (Cats), Canide (Dogs), and Mustelidx (Weasels), among the Carnivora. (4) The Primates, other than the lemurs. The following table shows the number of genera of each order represented in the Malagasy Sub-region and their distribution :— Ungu- | Rod- | Carni- | Insecti-|Chirop-| Pri- iaiat | antiay|| vorae i vormell eka anbees | ol | Endemic ... tse 7 6 | 8 1 iil 33 ANAC HIOS Gy 1 AEE oa ee 1 Be Pye | Cosmopolitan. .| ... ais iL 1 10 sie 12 | Tigtalae || n 7 7 OU pon lett: (ray | [V.B.—The “Endemic” genera are those confined to the Malagasy Sub-region; the “African” genera are those common to Madagascar and the mainland of Africa; and the “Cosmopolitan” genera are those which range beyond the limits of the Ethiopian Region.] 110 THE GEOGRAPHY OF MAMMALS Section V.—THE West AFRICAN SUB-REGION The West African Sub-region, as defined above, does not contain nearly so large a proportion of peculiar genera as the Malagasy Sub-region. At the same time, twelve out of a total of eighty genera of mammals that are found within its limits are not met with elsewhere. Moreover, we notice that, as a general rule, the genera inhabiting other parts of Africa are here replaced by distinct species. The Edentata are represented by one genus, Manis, the Scaly Ant-eater. Of the four known African species three are confined to this Sub-region. Of the Ungulates only one genus is endemic. This is Hyomoschus, a very curious aquatic form, which, together with an allied genus found only in the Oriental Region, forms a peculiar Family (Tragulidz) distantly allied to the deer (Cervidz). Many of the genera of Ungulates, however, are here represented by peculiar species. Instances of this are the Liberian Hippopotamus (Hippopotamus liberi- ensis), the Red River-hog (Potamocherus penicillatus), the West African Eland (Oreas derbianus), a peculiar species of wild ox (Bos pumilus), and several species of Rock- coney (Hyraz). The Rodents do not present any features of special interest, the only peculiar genera being Deomys (allied to Mus), recently obtained from the Congo district ; and the Brush-tailed Porcupine (Atherwra), of which one species is found only in this Sub-region, and the other two are confined to South-East Asia. Four out of the five known species of the peculiar Rodent-genus Anomalurus, THE ETHIOPIAN REGION spelt already referred to, are also confined to the West African Sub-region. Turning now to the Carnivores, there are found in the West African Sub-region only, two remarkable genera, Poiana and Nandinia. Of these the former is closely allied to the genus Prionodon, a beautifully marked civet- like little animal of the Oriental Region, and the latter is akin to the Palm-civets (Paradoxurus), also found in the Oriental Region, but not in Africa. There is only one genus of the Insectivora confined to this Region (Potamogale), already alluded to as being probably allied to the Madagascan Geogale. This much modified form is one of the few members of the Insectivora that has adopted aquatic habits. It is, for a member of the order, of considerable size. The Bats of West Africa, as is usually the case in every land, belong mostly to widespread forms. Out of sixteen genera only two, each containing a single species, are confined to this Sub-region, while a third (Hpomophorus) has not been found outside Africa. These three genera all belong to the family Pteropodidz, which contains the large fruit-eating bats. A few species of Lemurs still survive in the forests of the West African Sub-region. They belong to two genera, neither of which is represented in Mada- gascar. Of these, one (Galago) is also found in the other parts of Africa; the other (Perodicticus), containing two species, is met with only in the West African Sub- region. The forests of West Africa are plentifully supplied with Monkeys. Most of these belong to the genus Cercopithecus, of which, out of about forty species, thirty are met with in West Africa. Another enus, Lt? THE GEOGRAPHY OF MAMMALS Cercocebus, contains four species, all confined to this Sub- region. , Finally, it is only in these pathless and luxuriant jungles that two man-like apes, the Chimpanzee and the Gorilla, are to be met with. The Gorilla seems to be confined to the Gaboon district, but the Chimpanzee extends all over the Congo basin nearly up to the shores of Tanganyika. These two, together with the Orangs and the Gibbons of the Oriental Region, make up the family Simudz, which in structure is the most closely allied to Man of all the Monkeys. On comparing the West African Fauna with that of the rest of Africa, it will be seen that it is characterized by the exclusive presence of the following forms :— 1. Hyomoschus (the Water-chevrotain), which, together with Tragulus of the Oriental Region, forms the family of Tragulide of the Ungulata. 2. Two genera (Malacomys and Deomys) of the family Muridez, and Atherwra (Brush-tailed porcupine), found elsewhere only in the Oriental Region, among the rodents. 3. Two genera (Poiana and Nandinia) of the family Viverridx, among the Carnivora. 4, Potamogale, among the Insectivora. 5. Two genera of fruit-eating bats (Liponyx and Trygonycteris). 6. Perodicticus, a genus of Lemurs, and Cercocebus, and Anthropopithecus, among the higher Monkeys. The West African Region is further characterized by the absence of the following families, well represented in other parts of Africa: Orycteropodide (Aard-vark), Girafide (Giraffes), Equide (Zebras and Wild Asses), Rhinocerotidxe (Rhinoceroses), and Leporidx (Hares). THE ETHIOPIAN REGION 113 The following table shows the approximate number of genera of the West African Sub-region in each order and their distribution :— Eden-|Ungu-| Ro- | Carni-| Insecti- Chirop-| Pri- | motal tata. | lata. | dentia. | vora. | vora. tera. |mates.| ~OU™ | | - ere | Endemic. i 0 I 2 Se 2 AL | by [pxtrican ee Os | 11.1010 |) 2-0 i), Ad sary _ Cosmopolitan .| 1 6 8 9 3 13 0 40 | el | Poel mA Iain 20-2) 18) -4 | 16: |). & |80: | Section VI.—THE CAPE SUB-REGION This Sub-region, when extended so as to include the whole country as far north as Angola on the west, and up to the Tana river on the east, is on the whole, after the Malagasy, the most distinct of the four Sub-regions, since it possesses nineteen endemic genera out of a total of ninety- one. Furthermore, its area exhibits a greater range of temperature and humidity than the other Sub-regions, for, while in Natal and Mozambique tropical forest-conditions prevail, giving the fauna a certain resemblance to that of West Africa, in Cape Colony itself a temperate and fairly dry climate is found. Again, in Namaqua-land, to the north-west of Cape Colony, we meet with an open, dry, hot, desert country, the conditions of which closely approximate to those of the Saharan Sub-region. The distinctness of this Sub-region, apart from the mammals, is strongly marked by a very remarkable Flora, as well as by the exclusive possession of many forms among the other orders and classes of the animal kingdom. Among the Edentata, H 114 THE GEOGRAPHY OF MAMMALS besides the Scaly Ant-eater (Manis), of which one species extends as far south-east as the Transvaal, there is also found the Aardvark (Orycteropus), an animal characteristic of South Africa, though also occurring in East Africa as far north as the upper Nile basin. The relationships of this creature are very obscure, but the recent discovery of remains of a closely allied fossil form in beds of lower Pliocene age in the island of Samos, in the Aigean Sea, shows that the Aardvark, like so many other African animals, has migrated southwards from the Palearctic Region. The greater number of the genera to which the numerous Antelopes of the Cape Sub-region belong, extend northwards into the Saharan and even into the West African Sub-region, although the species of the Cape Sub-region are in many cases distinct. Among the distinct species may be mentioned the Hartebeest (Bubalis caama), the Spring-bok (Antidorcas euchore), and the Sable Antelope (Hippotragus niger). The two Gnus (Connochextes) and the Pallah (dpyceros) are confined to this Sub-region. To the portion of this Sub-region south of the Zambesi are restricted two of the African horses, the Quagga (Hquus quagga), now said to be extinct, and the Mountain Zebra (Hquwus zebra), which is fast approaching the same condition. But Burchell’s Zebra (Equus burchellt), under various slight modifications, occurs in suitable localities throughout the whole Sub-region. The White or Square-lipped Rhinoceros (Rhinoceros simus), which now only exists in two small districts of Mashonaland and Zululand, was formerly abundant in the Cape Colony and the Transvaal, but has never been met with north of the Zambesi. THE ETHIOPIAN REGION 115 Five out of the eighteen genera of Rodents found in this Sub-region are endemic. Two of these belong to the Mouse-family (Muridx), and two more to a burrowing family (Spalacidx) ; the fifth (Petromys) belongs to the family Octodontidx, which the Ethiopian shares with the Neotropical Region. The South African Sub-region, however, is especially remarkable for the large number of peculiar genera of Carnivora. Five of these genera have been formed for the reception of a number of small animals allied to the Indian Mongoose. Another, for which a separate family has been created, contains only the Aard-wolf (Proteles), an animal nearly allied to the hyenas, but with very small and rudimentary teeth, and a heavy mane of long hair. It is nocturnal, and feeds principally on carrion, being too weak to attack other living animals. The Otocyon, or Cape Hunting-dog, which constitutes another peculiar genus, is remarkable for having a greater number of molar teeth than any other mammal. Owing to this, it has been regarded by Professor Huxley as the most primitive of all existing members of its Family. Out of the six genera of Insectivora found in this Sub-region three are endemic. Of these the most interesting is the genus Chrysochloris, which forms a distinct Family, and comprises no less than eight species. These animals are sometimes known as “Golden Moles,” owing to the brilliant metallic lustre of their fur. They are in some respects allied to the Cen- tetidz, a family mentioned above as confined to Madagascar; but they are modified for a burrowing life, and externally resemble the Moles. Neither of the two remaining Orders, the Bats and the Monkeys, is represented by peculiar genera in the Cape Sub-region. This, however, is quite 116 THE GEOGRAPHY OF MAMMALS what one would expect, as members of these two Orders are essentially forest-loving in their habits, and this Sub- region consists chiefly of sparsely wooded and arid districts. On the whole, therefore, the Cape Sub-region will be found to be characterized, as compared with other parts of Africa, by the exclusive possession of the following forms :— 1. Connochetes (the Gnu), Pelea (the Rhébok) and Afpyceros (the Pallah) among the Antelopes. 2. Pachywromys and Mystromys, genera of Muridz ; Bathyergus and Myoscalops, genera of the burrowing Spalacide ; Petromys of the Octodontidy, among the Rodents. 3. Five genera of Mongooses, belonging to the family of Viverride ; Proteles, the only representative of the family Protelidx ; Otocyon (the Cape Hunting-dog); and Pecilo- gale (a small weasel), among the Carnivora. 4. Rhynchocyon, of the Macroscelide ; Myosorex (a Shrew); and the Chrysochlorid# (Golden Moles), among the Insectivora. The following table, showing the distribution of the genera, is constructed on the same plan as that of the other Sub-regions. | | Eden-|Ungu-| Ro- | Carni-| Insecti-| Chirop-| Pri- Total tata. | lata. | dentia. | vora. | vora. tera. | mates. : Endemic. . .| 0 Sra eo 8 3 Cee a0) 19 Aimicane eae a0) ol 1p "|. 6 3 0 Za col Cosmopolitan . | 1 8 a 9 3 TS |) Oy ek Motales 002) ul 426 18 20 6 bb. AP gl 1 THE ETHIOPIAN REGION 117 Section VII.—THE SAHARAN SUB-REGION The Saharan Sub-region, as its name implies, is essen- tially a dry and desert district. As here regarded, in addition to the Sahara proper, it includes the southern part of Arabia and Somaliland, both of which are distinctly arid countries. The only part of the Sub-region where there is much rainfall is in the Abyssinian highlands and the country surrounding such isolated mountains as Kenia. In consequence of these facts, and also in accordance with the fact that the Sub-region is conterminous for so long a distance with the Palearctic Region, the number of endemic genera of the Saharan Sub-region is extremely small, being only seven out of a total of eighty-five—that is, 8 per cent. as compared with 21 per cent. in the Cape, and 15 per cent. in the West African Sub-region. Of recent years a considerable number of new forms, especially of Antelopes, have been obtained from Somali- land, a country which, up to a few years ago, was quite unknown, and it is quite possible that eventually the “unknown Horn of Africa” will be found to possess a Fauna sufficiently distinct from the rest of the continent to be separated as another Sub-region. But for the present, until our knowledge is widened a little more, it will be best to keep Somaliland in the Saharan Sub-region. The most characteristic feature of the Sub-region is, perhaps, the large numbers of Antelopes.. These, as a rule, are specifically distinct from those of the Cape Sub- region. Among them may be mentioned Swayne’s Harte- beeste (Bubalis swaynei), Hunter’s Bontebok (Damaliscus hunteri), three species of Oryx (O. beisa from Abyssinia 118 THE GEOGRAPHY OF MAMMALS and Somaliland, 0. beatria from Arabia, and O. leucoryx extending from Senegal to Nubia), and a distinct species of Kudu (Strepsiceros imberbis), which inhabits the Sub- region from Somaliland as far south as the Tana. This Sub-region is also the headquarters of the Gazelles (Gazella), at least twelve species out of a total of twenty- five being found there. Three genera of antelopes are endemic; these are the Dibatag (Ammodorcas), and the Gerénuk (Lithocranius), both found in Somaliland, and remarkable for their very long necks, by means of which they are enabled to reach down branches of trees from a considerable height. The third is the Addax (Addaz), which is found throughout the Sahara, and extends into Arabia. The Rodents are well represented in this Sub-region, but there are only three peculiar genera—Lophiomys, Heterocephalus, and Pectinator. Lophiomys, a curious crested Rodent, forms a distinct family. The remarkable little animal Heterocephalus is almost entirely hairless, and apparently lives underneath the ground, burrowing in the soft sandy soil of the desert. It was originally obtained by the celebrated traveller Riippell in Abyssinia, and has since been brought from the interior of Somaliland by Mr. Lort Phillips. The third endemic genus, Pectinutor, is also from the coast of Somaliland and Abyssinia. It was first obtained by Captain Speke, and named by Blyth (P. spekev) after its discoverer. : The Saharan representatives of the Carnivora, In- sectivora, and Bats are almost all widespread forms, and present no features of special interest. Among the Monkeys, however, we find one peculiar genus—this is THE ETHIOPIAN REGION 119 Theropithecus, containing two species of terrestrial baboons, which inhabit Abyssinia and the Galla country. The other genera of African monkeys are fairly repre- sented here, though by no means so abundantly as in the West African Sub-region. The Saharan Sub-region, therefore, is characterized by the exclusive possession of the following genera :— (1) Ammodorcus, Lithocranius, and Addax among the antelopes. (2) Lophiomys (forming a distinct family), Heteroce- phalus, and Pectinator among the rodents. (3) Theropithecus among the Monkeys. The following families, found in other parts of Africa, are absent from this Sub-region: Tragulid# (chevrotains), Protelidz (aard-wolf), Potamogalidz (river-shrews), Chryso- chloride (golden moles), and Simiide (the anthropoid apes). The following is the summary of the mammals inhabit- ing the Saharan Sub-region :— | Eden-|Ungu-|} Ro- | Carni-|Insecti-| Chirop-| Pri- Total tata. | lata. | dentia. | vora. | vora. tera. | mates. = Endemic. . .| 0 3 3 0 0) 0 1 @ PTAC, 5 Gc 1 15 8 3 0 1 4 32 Cosmopolitan .| 1 9 12 if 3 14 0 46 Total . 2 27 23 | 10 3 15 5 85 Section VIII.—Tue Past History or THE ETHIOPIAN MAMMAL-FAUNA Up to the present time paleontology has afforded us little assistance in tracing out the past history of the African Mammal-fauna. With the exception of a few still 120 THE GEOGRAPHY OF MAMMALS surviving species found in a semi-fossil condition in caves, the only fossil mammal hitherto obtained from the Ethiopian Region is a form named by Owen Tritylodon, the remains of which were discovered in the Karoo beds of Basutoland, in South Africa. This formation is of Mesozoic age, and seems to correspond to the Trias of Europe. Near Stutt- gart, also, in beds of the Trias age, another specimen closely resembling that of South Africa has been obtained. Tritylodon belongs to a group which was apparently allied to the modern Marsupials, and many somewhat similar forms have been found in other Mesozoic beds in Europe and North America. The only conclusion to be drawn from this is, that before the commencement of the Tertiary epoch the whole world was, so far as is at present known, inhabited by small insignificant mammals distantly allied to the Mar- supials, which are at present restricted to Australia and South America. Besides Tritylodon, the only fossil mammals hitherto known, from the Ethiopian Region, are those which have been described by Grandidier (3) and Forsyth-Major (4) from Madagascar, as mentioned above. The resemblances between the Faunas of the Oriental and Ethiopian Regions have been noted and commented on by many writers; Mr. Allen is even disposed to join the two Regions into one realm. The number of prevailing genera, however, common to these two Regions, and not found in the Palzarctic Region, is very small, the total being, as was pointed out in the first article of this series, only eight, and of these one (Mellivora) has since then been recorded by Biichner (2) as occurring in the Trans- Caspian district, and is therefore also Palearctic. Of the other seven, four have been found, in a fossil state, in THE ETHIOPIAN REGION 120 various parts of the Palearctic Region. These are—(a) Manis (the Scaly Ant-eater), from the Pliocene of Samos; (b) Rhinoceros, which existed in various parts of Europe from the Miocene up to the Pleistocene times; (c) Elephas, which first appears in Pliocene times, and extends to late Pleistocene in Europe; and (d) Viverra, which commences earlier than the others, and also survived until Pliocene times in Europe. The three remaining genera common to India and Africa, but not hitherto found in fossil state in the Palzarctic Region, are Golunda (a Rat), Atherwra (a Porcupine), and Vycteris (a small insectivorous Bat). It is quite possible that these animals may eventually be discovered in the European Tertiaries. Besides this, the remains of a considerable number of the now endemic African genera have been found fossil in Europe. The list of these is instructive, and points almost unquestionably to the conclusion that Africa has been gradually peopled by successive inroads of animals from the North. In the Eocene beds of Europe the still existing genera are few in number; but the Lemurs, and many of the more primitive forms of the Carnivora, such as form the present fauna of Madagascar, abound. It is, therefore, probable that the separation of Madagascar from the mainland of Africa took place at about the close of the Eocene period. During the Miocene and lower Pliocene times in Europe, a large number of new genera appear for the first time, the bulk of which still survive in Africa and India, though extinct or almost driven out of the Palearctic Region. Examples of such genera are—Orycteropus, the Aardvark; Manis, the Scaly Ant-eater; Rhinoceros ; Hyomoschus, the Water-Chevrotain (probably identical 122 THE GEOGRAPHY OF MAMMALS with Dorcotheriwm, a fossil form); Giraffa; several genera of Antelopes; the Porcupine; the Squirrel; Felis; Hyena; Viverra ; Herpestes, and even the higher Monkeys. It must have been during this period that broad land- connections existed between Europe and Africa, by means of which the African continent became peopled by its present fauna. In the succeeding Pliocene times in Europe, although a number of the more distinctly African forms still sur- vive, there begin to appear certain genera, such as those of the Deer-family (Cervidx), Ursus (the Bear), and others which have never reached Africa at all. This seems to show that Africa was, at the commencement of this period, cut off from the Palearctic Region by an inter- mediate sea. List oF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHAPTER IV. (1) Batrour, I. B.—“ Botany of Socotra” (Mammals, p. xxx.) Trans. Roy. Soc. Edin., xxxi., 1888. (2) Bicuner, E.—‘‘ Ueber das Vorkommen der Mellivora indica, Kerr, im Transcaspi-Gebiet.” Notes Leyd. Mus., xv., p. 99. (3) GranprpieR, A.—“ Histoire Physique, Naturel et Politique de Madagascar,” vol. x., Mammiféres, by Milne Edwards and Grandidier, 1890. (4) Masor, C. I. Forsyra—“ On Megaladapis madagascariensis, an extinct gigantic lemuroid.” Proc. Roy. Soc., 1893, p. 176. (5) Neumann, B.—“ Bericht iiber seine Reisen in Ost und Central Afrika.” Verhandl. Gesellsch. Erdkunde Berlin, 1895 (p. 24 in separate copy). (6) THomas, O.—“ On Some Specimens of Mammals from Oman, South-East Arabia.” /P. Z. 8., 1894, p. 448. PLATE IVY. HIOPIAN REGION WAR Whavon Sundiagu Kent INTO 4 SUB-REGIONS. 10 20 MAP OF THE|ETHIOPIAN REGION CHAPTER V THE ORIENTAL REGION (PLaTE V., p. 152) SECTION I.—BOUNDARIES OF THE ORIENTAL REGION THE Oriental is the smallest of the six Regions into which the Earth has been divided for the study of zoological distribution. On the west it includes the great peninsula of India and its attendant island of Ceylon. Its boundary on this side is probably the Suliman range of hills, though the fauna of Western Sind and the Punjab, which lie between that range and the Indus, is intermediate in character between those of the Oriental and Palearctic Regions. Beyond this range the boundary runs eastwards along the slopes of the Himalayas, at an elevation of from 9000 to 10,000 feet above the sea-level. Above this height Palearctic forms are chiefly met with, below it Oriental forms mostly prevail. Hastwards of Sikkim the boundary between the Palearctic and Oriental Regions cannot be laid down with certainty, owing to our little acquaintance with the eastern part of Tibet and the adjacent portion of China. What knowledge we have of the fauna of this Region is due almost entirely to the celebrated French missionary, Pére David, who made considerable researches in Moupin, a small mountainous territory, situated at the extreme western edge of the Tibetan plateau. Pére. David's collections have been mostly described by M. 123 124 THE GEOGRAPHY OF MAMMALS Milne-Edwards (1). An examination of the list of the mammals obtained by him in this district shows that the fauna has a character intermediate between those of the Oriental and Palzarctic Regions, besides containing a con- siderable proportion of peculiar forms. As, however, most of the Oriental genera extend even further north into the Chinese province of Kansu, and some even cross into Japan, countries which are otherwise well within the Palearctic Region, it will be most convenient to draw the boundary of the Oriental Region to the south of Moupin. Beyond this point again our knowledge of the distribution of the mammals is very scanty, and though the northern part of China appears to be distinctly Palearctic, and the southern Oriental in its affinity, there is, so far as we know, a considerable admixture of forms all over this part of Asia. Probably the most convenient boundary will be found to be that adopted by Wallace—the northern edge of the basin of the Yang-tze-Kiang. This is, no doubt, to a great extent an artificial boundary, but such a fault is unavoidable in the present instance, as there is here no natural frontier to separate the two regions. In addition to the south-eastern part of Asia, the Oriental Region includes within its boundaries all the large and important islands lying between that continent and the Australian Region. The principal of these are the Chinese islands of Formosa and Hainan, the large group of the Philippines, together with Sumatra, Java, Borneo, and the adjacent islands up to Wallace’s Line. With the exception of Celebes, all these islands are truly continental in character—that is to say, are separated from the mainland by seas of less than 100 fathoms of depth. But Celebes is in some respects anoma- lous, and will be considered in greater detail below. THE ORIENTAL REGION 125 The boundary between the Australian and the Oriental Regions called Wallace’s Line, as having been first pointed out by that distinguished naturalist, runs between the two small islands of Bali and Lombok. Bali is connected by shallow water, and also by its zoological relationships, with Java; while Lombok agrees in character with Timor and the other Australian islands further east. From Bali the boundary of the Oriental Region runs in a north-eastward direction, between Celebes on the one side and the Sula islands and Gilolo on the other. Section I].—GENERAL VIEW OF THE MAMMAL-FAUNA OF THE ORIENTAL REGION The Oriental Region lies almost wholly within the tropics. The greater part of the country within its borders enjoys a bountiful rainfall, and is covered with luxuriant forests; the only portion which is less favoured being the north-western part of India and the strip of country along the northern shores of the Persian Gulf. In these districts there is very little rain, and desert conditions and a desert Fauna, somewhat resembling those of the African Sahara, prevail. The Fauna of the Oriental Region presents, on the whole, a striking contrast to that of the Australian Region. The characteristic features of the latter are doubtless due to the long isolation to which it has obviously been sub- jected, whereas the Oriental Region as regards its charac- teristic forms is more nearly allied to the neighbouring Palearctic Region, from which probably most of its in- habitants have been derived. 126 THE GEOGRAPHY OF MAMMALS The Oriental Region contains representatives of eight out of the nine Orders of terrestrial mammals, the Monotremes alone being wholly absent, while the Mar- Fig. 24.—THE NYLGHAIE, (Boselaphus tragocamelus. ) supials are barely represented by two species of Cuscus (Phalanger) found only in the island of Celebes, which have been obviously derived from the neighbouring Aus- tralian Region. THE ORIENTAL REGION 1g The Edentates, like the marsupials, are also represented by one genus only, Manis (the Pangolin), which the Oriental shares with the Ethiopian Region. The Oriental further resembles the Ethiopian Region in the variety of its forms of Ungulates, although they are not Fig. 25.—THE GAYAL, (Bibos frontalis. ) [P. Z. S. 1890, p. 595.) nearly so abundant. Three genera of Antelopes, the Four- horned Antelope (Tetraceros), the Black-buck (Antilope), the Nylghaie (Boselaphus), (see Fig. 24, p. 126), are peculiar to it, as is also a group of wild Oxen (Bibos), with three species—the Gayal (Fig. 25), the Gaur, and the Banting. 128 THE GEOGRAPHY OF MAMMALS But while the Rhinoceroses, the Wild Asses, the Elephants, and the Antelopes are common to both the Oriental and Ethiopian Region, the Oriental possesses in addition Deer, Fig. 26.—THE BINTURONG. (Arctictis binturong.) Wild Sheep, and Wild Goats. Thése three last-named groups have never established themselves in the Ethiopian Region, though a single goat (Capra walie) has penetrated THE ORIENTAL REGION 129 as far as the highlands of Abyssinia. But representatives of all of them are found in the Palearctic Region. Among the Rodents of the Oriental Region the Squirrels are especially numerous, there being upwards of fifty species found within its limits, nearly all of which are arboreal in their habits. Although there are no families of Carnivores peculiar to the Region, there is a considerable number of genera of Civets (Viverridx) not found elsewhere, such as the Para- doxures (Paradoxurus) the Binturong (Arctictis) (Fig. 26, p. 128), and the genera Prionodon, Arctogale, and Hemi- gale. The Bears (Ursidz), too, which are quite unknown in Africa, are characteristic members of the Oriental Mammal-fauna. Among the Insectivores we find two peculiar families. One of these has been formed for the reception of Galeo- pithecus, the so-called Kaguan, an animal of about the size of a small cat, with thin flaps of skin between the fore and hind limbs and tail, which enable it to make flying leaps from tree to tree (see Fig. 27, p. 130). The other family (Tuparidx) contains two genera. One of these, Tupaia, with at least twelve species, is an abnormal Shrew with a curious external resemblance to the Squirrels, with which, however, it has no real connection. The other, Ptilocercus, is distinguished from Tupaia by its peculiar tail, which is provided at the end with a bilateral fringe of long hairs. Both these families are confined to the Malayan portion of the Oriental Region. Bats are numerous in the Oriental Region, and many of the genera extend eastwards into the Austro-Malayan islands. Only four genera, each with a single species, are peculiar. I 130 THE GEOGRAPHY OF MAMMALS Finally, among the Primates there are three genera of Lemurs. Two of these (Vycticebus and Loris) are peculiar, HIG. 27.—THE KAGUAN. (Galeopithecus volans. ) but belong to the family Lemuride, and have their nearest allies in Africa. The third (Tarsius), which forms a family THE ORIENTAL REGION Lon of itself, is practically confined to the Region, although it has slightly overstepped its boundaries, being said to occur in one of the smaller Austro-Malayan islands between Sumba and Timor. Besides the Lemurs, six genera of true Monkeys are found in the Oriental Region. Three of these, the Proboscis Monkey of Borneo (Nasalis), the Gibbons (Hylo- bates), and the Orangs of Sumatra and Borneo (Simia), are strictly endemic; while the other three, although highly characteristic of this Region, have extended their range slightly across its frontiers. Summarizing these results, we shall find that the Oriental Region contains only two truly endemic and one quasi-endemic families out of a total of thirty-six which occur within its limits. These are the Galeopithecide (Flying Lemurs), Twpaiide (Tree shrews), and Tarsiide (Tarsiers). The total number of genera found in the Region is 113, out of which 38 are peculiar; 11 extend their ranges slightly beyond the limits of the Region, and 64 are widely spread. On reducing these figures to an average, it will be found that the Oriental Region contains about 38 per cent. of peculiar genera, or, if the quasi-endemic genera be added, about 45 per cent. In either case, this shows a much lower percentage of peculiarities than has been shown to exist in the three Regions previously considered. Section JIJ.—Swtspivision oF THE ORIENTAL REGION The Oriental Region, as regards its mammals, may be most conveniently divided into four Sub-regions (see Map, Plate V., p. 152). These are :— 1. The Indian Sub-region.—This comprises the whole 132 THE GEOGRAPHY OF MAMMALS of India proper from the Suliman range and the lower slopes of the Himalayas to Cape Comorin. A line drawn northwards from Calcutta to the Himalayas, forms the approximate eastern boundary between this Sub-region and the next. There should also be included in this Sub-region the island of Ceylon, and probably the narrow, low-lying strip of desert country between the Persian Gulf and the central plateau of Persia. 2. The Burmo-Chinese Sub-region.—This Sub-region includes the portion of Sikkim below 10,000 feet, Assam, China south of the northern water-parting of the Yang-tze- Kiang, the islands of Formosa and Hainan, and all the countries of the Indo-Chinese peninsula (Cochin China, Siam, and Burma), its southern land-boundary being approximately a line running to the north of the Malay peninsula from Tavoy on the west, to Bangkok on the east, at about 15° N. lat. 3. The Malayan Sub-region—The Malay peninsula, together with the great islands of the East Indian archi- pelago, Sumatra, Java, Borneo, and the Philippines, forms a third division, which may be called the Malayan Sub-region. 4. The Celebesian Sub-region, containing only the island of Celebes. This subdivision of the Oriental Region differs from that adopted by Wallace in two important points :— (a) In the combination of Wallace’s Indian and Ceylonese Sub-regions into one—the Indian Sub-region. (b) In the transference of Celebes from the Australian to the Oriental Region, and the formation of a new Sub- region for its reception. The reasons for these changes may be here briefly considered. First, as regards Ceylon, there are nineteen THE ORIENTAL REGION ILS genera of mammals found in the Indian Sub-region, which do not extend their range further eastwards into the Burmo- Chinese Sub-region, Of these, thirteen are to be met with also in the Palearctic and Ethiopian Regions, leaving only six confined to the Indian Sub-region, These six are— (1) Tetraceros (Four-horned Antelope). (2) Antilope (Black Buck). (3) Boselaphus (Nylghaie). (4) Platacanthomys (Spiny Rat). (5) Melursus (Indian Bear). (6) Loris (Slender Lemur). Of these, Loris alone is characteristic of Southern India and Ceylon (the Ceylonese Sub-region of Wallace). Melursus is found in Ceylon, but occurs also all over the peninsula of India from the Himalayas southwards. Platacanthomys inhabits the western Ghats and the Animali hills of Southern India alone, and not Ceylon; the remaining three genera are distributed over the whole of the Indian peninsula, but do not reach Ceylon. There is, therefore, only one genus of mammals con- fined to the Ceylonese Sub-region of Wallace, and this hardly seems to afford an adequate reason for separating it from the Indian Sub-region proper. The chief ground g, according to Mr. Wallace, is the existence there of a peculiar family of snakes—the Uropeltide, or Rough-tails, which are entirely confined to Wallace’s Ceylonese Sub-region. Examples of these reptiles, how- for so doing ever, have been recorded in India as far north as Ganjam, in 20° N. lat, and it seems probable that they may eventually be found all over the peninsula south of the great plains of the Indus and the Ganges. Secondly, as regards Celebes, this island certainly 134 THE GEOGRAPHY OF MAMMALS presents a difficult problem to the student of geographical distribution. But so far as the mammals are concerned, the only Australian element in Celebes consists of two species of Phalanger and a few Bats; the remaining forms, although many of them are very peculiar, have been, doubtless, originally derived from the Oriental rather than from the Australian Region. It seems more logical, there- fore, on the whole, to make the island of Celebes a separate Sub-region of the Oriental rather than of the Australian Region. This position, however, will be con- sidered in greater detail in the account of the Sub-regions. Section 1[V.—THE INDIAN SUB-REGION The Indian Sub-region has a close resemblance to the Palearctic Region. This is more especially the case in the north-western districts, where the country is practically rainless, and the fauna, owing to similarity of condition, is in many respects closely allied to that of the neighbouring desert regions of Central Asia. The desert district of the Indian Sub-region includes the narrow strip of coast land to the north of the Persian Gulf, the Punjab, Rajputana, and the northern part of the Bombay Presidency. The greater part of the peninsula of India south of the great plains is occupied by the high, and rather dry, plateau of the Deccan and of Central India, which is covered with a thin and scanty jungle. The southern slopes of the western Ghats and the greater part of Ceylon enjoy an abundant rainfall, and are clothed with a tropical forest, in consequence of which their faunas present many points of resemblance both to each other and also to that of the Malayan Sub- region, which has similar physical conditions. THE ORIENTAL REGION lie Our knowledge of the mammals of this and the next Sub-region is very fairly complete, owing to the excellent handbook recently published by Mr. W. T. Blanford (2). The Pangolin (Manis), which, with slight specific modi- fications, is also found in all the other Sub-regions as well as in the Ethiopian Region, is here the sole repre- sentative of the Edentates. The Sub-region is well pro- vided with members of the various families of Ungulates. Three peculiar genera of Antelopes, which are not found beyond the limits of the Sub-region, have been already mentioned; other genera, such as the Gazelles (Gazella), the Goats (Capra), and the Sheep (Ovis), are found in other parts of the world as well as in this Sub-region, but are absent from the remaining Sub-regions. All the families of Ungulates are common to this and the Ethiopian Region, except the Deer-family (Cervidz), the entire absence of which from Africa south of the Atlas has already been commented upon. The Rodents do not present any marked features of interest in this Sub-region. One genus, Platacanthomys, a small dormouse-like member of the family Muride, is found only in the hills of Southern India, otherwise the genera are mostly wide-spread forms. Among the Carnivores the Cats are numerous and large. This Sub-region is the proper home of the Tiger, which, however, has extended itself throughout the whole Region, and even across its boundaries westwards into Persia and Trans-Caspia, and eastwards far into China and Manchuria. The Lion, too, which is essentially an animal haunting dry and comparatively barren countries, is a member of this division of the Indian fauna. It was formerly much more abundant in the peninsula, but 136 THE GEOGRAPHY OF MAMMALS is now, apparently, restricted to a small area in Western India. As is the case with the Ungulates, so here, with the exception of the Bears (Ursidz), all the families of Indian Carnivores also range into Ethiopia. The Insectivores of this Sub-region need not detain us long. A Tree-shrew (Tupaia), an outlying member of a genus very abundantly represented in the Malay countries, is found in Southern India; the other genera, the Hedge- hogs (Hrinaceus) and the Shrews (Crocidura), are widely spread throughout the Old World. Among the Bats of this Sub-region we find that not only there are no peculiar genera, but that even the species in nearly all cases have an extended range beyond its limits. Out of about forty species, six alone are confined to the Sub-region. The Slender Loris is found only in Southern India and Ceylon, and is the single representative of the Lemurs in this Sub-region. It is a strange-looking creature, with long spidery arms and no tail. Like most of its race, it is arboreal and nocturnal in its habits. Indian monkeys all belong to the two large genera, Macacus and Semnopithecus, both of which are charac- teristic of the Oriental Region, although two or three species of them have strayed over the borders into the Palearctic Region. The following table shows, in a succinct manner, the origin and distribution of the mammals of this Sub-region. The species in the first line, reckoned as “ Endemic,” are confined to the Sub-region; those called “Oriental” do not occur beyond the boundaries of that Region; those catalogued as “ Paleearctic”” are common to that Region THE ORIENTAL REGION 137 and to the Indian Sub-region; the “Ethiopian,” in the same way, are found alike in the Ethiopian Region and the Indian Sub-region. The “Paleogean” genera are those which are found in the Indian Sub-region and in more than one of the other three Regions of the Old World. Finally, the “Cosmopolitan” genera are those found in the New World as well as in the Old. Jenks Wealee| 2 es) Bo)” > — ~~ Endemic . OMe Soleil wie Ory Opler) 16 Oriental Oy WneIealerul see Malay econ" On he Palearctic 0 Me ey ale ste OF 2a 6 Ethiopian . 1 Dela eeor On) On Onsen Paleogean 0 2a ole alee) Le Osea Cosmopolitan OF aio. F 27) 79: | 80 3208 4) Total . ie ae We ae 3) Sai" Se | Gan From this table it will be seen that the relations of the Indian Sub-region are about equally divided between the Palearctic and Ethiopian Regions; the largest number of genera are registered as “ Paleogean,” and most of these are common to the three Regions of the Old World. The relations of this Sub-region to the Australian Region are very slight; with the exception of Canis, it is only among the Bats that we find any common genera. Section V.—THE BuRMO-CHINESE SUB-REGION Owing to our imperfect knowledge of the fauna of the central part of China and of Tibet, it is impossible at present to draw up a complete list of the mammalian 138 THE GEOGRAPHY OF MAMMALS genera inhabiting this Sub-region, and it is consequently out of the question to lay down anything but a very uncertain boundary between this Sub-region and the neigh- bouring Palearctic Region. It is probable, however, that even when Western China and Tibet have been thoroughly explored, it will still be difficult to trace an absolute frontier between the Palzearctic and Oriental Regions. As we already know, Northern China and Japan contain a con- siderable number of purely Oriental species. Even the Tiger, usually associated with tropical jungles, ranges through China into the valley of the Amoor and the island of Saghalien, where a most severe Arctic winter is met with. In the same way, two species of a typically Oriental genus of Monkeys are found in North-East Asia—one (Macacus speciosus) in Nipon, the largest of the Japanese group of islands; the other (Macacus tcheliensis) in the mountains north of Pekin. On the other hand, a good many purely Palearctic forms extend into Southern China. This is more espe- cially the case among the birds, which have hitherto received a preponderating share of the attention of the naturalists and collectors in the Chinese Empire. Passing over the Edentates, represented, as in the Indian Sub-region, by two species of Pangolin (Manis), we come to the Ungulates of the Burmo-Chinese Sub-region. Here we remark the disappearance of the Antelopes, and the great development of the Deer-family (Cervidz), of which no less than fifteen species are found in this Sub- | region, One of these (Zlaphodus) is a curious little Deer with very small simple antlers and large canine teeth; it was first described by Milne-Edwards from Western Tibet, and subsequently a second species of the same genus was THE ORIENTAL REGION 139 discovered in Southern China. Another small Deer, for which a separate genus (Hy¢dropotes) has been rightly formed, has no trace of antlers at all, and in other respects differs much from the remaining members of the family. This form is entirely confined to Southern China. A third peculiar genus, belonging to the Bovidz, is the Takin (Budorcas). This ox-like Antelope is also found in Western Tibet, but extends its range southwards to the Mishmi country in the north of Assam. The Takin is one of the very few of the larger ruminants that has never been met with or shot by European sportsmen, and our knowledge of it is entirely derived from the natives. The Burmo-Chinese, like the Malayan Sub-region, is the most frequented haunt of the Squirrel family (Sciwride). No less than thirty-two species, referable to the genera Sciwrus (the true Squirrel) and to Pteromys and Sciwrop- terus, the Flying Squirrels, are found here alone. The only Rodent supposed to be truly endemic is Hapalomys, a long-tailed Rat found in Burma. The Burmo-Chinese Carnivores do not call for any special remark; one genus alone (Helictis) is strictly . endemic. It contains three or four species of small badger-like animals with arboreal habits. Among the insectivores of this Sub-region only one genus is endemic. This is Soriculus, containing some small shrew-like mammals found only in Sikkun and Assam. Several species of Mole (Yalpa), as also Anuro- sorex, and Chimarrhogale, belonging to the Shrew family (Soricidx), extend from the Palearctic Region into this Sub-region, but no farther. The Bats of Burmo-China need not detain us long; most of the genera are widely spread, and a very large 140 THE GEOGRAPHY OF MAMMALS number of them extend across Wallace’s line into the Austro-Malayan islands—a distribution shared by hardly any other of the Oriental genera of mammals. One of the Slow Lemurs, Vycticebus, is common to this and the Malayan Sub-region; it bears a certain resem- blance to the Indian genus Loris, but is distinguished by its somewhat stouter aspect and its still more sluggish habits. Among the Monkeys of this Sub-region, in addition to the two genera Macacus and Semnopithecus, inhabiting also the Indian Sub-region, a genus of the anthropoid Apes occurs. This is Hylobates, members of which are commonly known as Gibbons; they are slender animals, with very long limbs and no tail, and are entirely re- stricted to the forest districts, being exclusively arboreal in their mode of life. The following summary of the Burmo-Chinese genera of mammals has been drawn up exactly in the same way as the previous list, except that under an additional head- ing, “ Australian,” are placed two genera common to the Oriental and Australian Sub-regions :— ie aren eran ee dy | ;} a |b |] & | Sw cele se = Endemic . ee Vege ce AO. 1| 1 5)41. 3) oa et) | 0 by} Orientaly a) ee ee | CO O il 4b) 3} Ht) Dy) I? Palearctic 4 ie eRe yea il) 94 | Tet Ethiopian . eer ey tal 2 1 2! 0 TL) Oy 7 Meebo ye gg ay OF Os One® Da 10) 2) Palzeogean Re ee Lt, PSs a Gals et Oe Cosmopolitan... . .| 0 3 3 G75 Os) «oa ON aloe Goel 6 oP ae alii | IO lee | Sy AO ab | 7 | | THE ORIENTAL REGION 14] SEcTION VI.—THE Maayan SUB-REGION The Malayan Sub-region lies entirely within the tropics, and almost the whole of it is covered with a luxuriant tropical jungle. It is here, consequently, that we find the Oriental fauna in its highest state of development, and with the least admixture of forms belonging to other Regions. With the exception of the Malay Peninsula, the whole of this Sub-region consists of islands, which, however, are separated from the main continental mass by comparatively shallow water, so that an elevation of 100 fathoms would obliterate the whole of the sea between the various islands, leaving them connected with one another and with the Asiatic continent. There can be no doubt that these islands, all of which have very rich faunas, have been stocked from the mainland, and that a careful study and comparison of their component parts would go far to enable us to trace out the past history of the Region, and to find out what changes have taken place from time to time in the distri- bution of land and sea. If a careful analysis of the mammalian genera of the Sub-region be made, it will be found that the greater number of the genera found on the mainland extend to all the three larger islands, Sumatra, Java, and Borneo, and that of the remaining genera the larger proportion are common to the Malay Peninsula, Sumatra, and Borneo, and are not found in Java. This would seem to indicate that Java was separated from the mainland before Sumatra and Borneo, and this view is further borne out by the fact that individual species of a genus are frequently common to the Malay Peninsula, Sumatra, and Borneo, 142 THE GEOGRAPHY OF MAMMALS whereas in Java they are replaced by slightly different forms. ‘The Philippines, though connected with Borneo by two chains of islands, so that the straits separating the group from Borneo are nowhere very wide, contain a poor mammal-fauna as compared with Borneo, Only about fourteen genera, exclusive of bats, have reached these islands, and, with one exception, these are all widespread. But it must be recollected that the Mammal-fauna of the Philippines is still very imperfectly known, The following table gives the figures relating to the distribution of the genera within the Sub-region (excluding bats) :— Number of genera that occur in— (1) The Malay Peninsula, Sumatra, Borneo, and Java . 36 (2) The Malay Peninsula, Sumatra, and Borneo — . ; 6 (3) The Malay Peninsula and Borneo alone. ’ : 3 (4) The Malay Peninsula, Sumatra, and Java, ; } 2 (5) The Malay Peninsula and Java alone : : ‘ | As regards the Philippines, the total number of mammal genera (exclusive of bats) is fourteen, thirteen of which oceur also in the Malay Peninsula and islands; one is con- fined to the Philippines alone; in addition five Malayan genera reach Palawan, a large island lying between Borneo and the Philippines.' Although the Malayan Sub-region does not contain any endemic genera of Ungulates, there is one form the dis- tribution of which is so remarkable that special attention must be drawn to it. This is the Tapir, one species of ''This calculation was made before Mr. John Whitehead’s great dis- coveries in the highlands of North Luzon (see Ann, NV. //., ser. 6, vol. xvi., p. 160, and T'rans, Zool. Soe, vol, xiv,, pt. 6) were announced by Mr. Thomas, ‘These embrace five new generic forms of rodents, and there are probably more to follow, THE ORIENTAL REGION 143 which is found in the Malay Peninsula, Sumatra, and Borneo, the only other Tapirs now existing in the world being met with in Central and Southern America, This is one of the most interesting cases known of what is termed “discontinuous ” distribution, but the explanation of it is not very difficult. If we turn to the records of paliweon- tology, we find undoubted remains of the members of the genus Tapirus recorded in the Miocene formation of France, in North America, and also in the Pliocene of China. There can be little doubt, therefore, that the Tapir, which is a harmless beast, destitute of all means of offence and defence, has been driven out of these northern countries into the tropical forests of South America and Malaya, where the absence of competition has enabled it to survive. Among the Malayan Rodents we find the squirrels (Sciwride) even more abundant than in the last Sub- region. Two of the species belong to a separate genus (Rhithroscwwrus) which does not occur elsewhere. There are also two endemic genera of rats (Mwrid«x)— one (Phlwxomys) from the Philippines, the other (Pithechirus) from Sumatra and Java. A genus of Poreupines (7richys), which differs from Atherwrw in several important cranial characters, is con- fined to Borneo, Three genera of Malayan Carnivores are worthy of special mention, One of these is Hemigale, not very far removed from the Palm-cats, with two species, a second one having been recently discovered by Mr. Hose in the mountains of Borneo, The second is Cynogale, also be- longing to the same family. The latter, which is semi- aquatic in its habits, and bears a superficial resemblance 144 THE GEOGRAPHY OF MAMMALS to an otter, is found in the Malay Peninsula, Borneo, and Sumatra. A third endemic Carnivore, Mydaus, which, Fic. 28.—THE ORANG. (Simia satyrus. ) like the American Skunk, is remarkable for the very powerful odour emitted from its anal glands, was originally THE ORIENTAL REGION 145 described from the mountains of Java, but has since been obtained from Sumatra and Borneo. Of the Insectivores by far the most important genus in the Malayan Sub-region is the Tree-shrew (Tupaia), of which at least a dozen species are here found. - The Tree- shrews are small animals, of the general appearance of squirrels, that live chiefly among the branches of trees, and, like the squirrels, sit on their haunches and use their fore limbs for holding their food. An allied genus, with an elegant double fringe of long hair to its tail (Ptilocercus), is confined to Sumatra and Borneo. Tarsius, belonging to a distinct family of Lemurs, inhabits the forests of most of the islands of the Sub-region, as well as Celebes. It is a small animal, about the size of a squirrel, deriving its name from the fact that the tarsal bones of its foot are greatly elongated. Among the Monkeys, in addition to the three genera found also in the Burmese Sub-region, we have the Pro- boscis Monkey (Nasalis) of Borneo, very remarkable for its large and projecting nasal organ. Finally, in Sumatra and Borneo we find the Orang (Sima), of which there are possibly two species, although this is by no means certain. These large man-like apes (see Fig. 28, p. 144), which form, along with the Gibbons, and the African Chimpanzee and Gorilla, the family Simiidz, inhabit mostly the low swampy districts near the coast; they may be distinguished at once from their African cousins by the reddish-brown colour of the long hair with which they are clothed. In some respects they are the most closely allied to Man in structure of the anthropoid Apes. The following is a summary of the Malayan genera of mammals, constructed on the same plan as in the case of K 146 THE GEOGRAPHY OF MAMMALS the other Sub-regions. It will be seen that, while the total number of genera has not increased very much, the number of endemic genera is nearly doubled, as compared with those of the other two Sub-regions. : 73 2 7 ; 3 £ ae See.) et | ale ae ae =} 5 8 a || SS 12) a <3) Pp a= 3} = S i) Endemic . O10 | Cs)" cel reese Oriental 0 ik. jal 4} 3 Sys alls: Palearctic Oi eee ok elt dl oO} 2 8 Ethiopian. as ee 1) 2h OU aa 7 Australian 0 OF) Oh Oe Oh Sea ae. 2 Paleogean . Oy BY Si] Ba DL rea Gries Cosmopolitan Cis) 3 6204) Sane Total Peto" ie 17 16.) aoe wean Section VII—THE CELEBESIAN SUB-REGION As already mentioned, the island of Celebes presents a problem of considerable interest to the student of geographical distribution. Celebes is separated from the other islands, both to the eastward and to the westward, by seas of considerable depth. Compared with the other Malayan islands, its fauna is scanty. This fact, and the very peculiar shape of the island, suggest a possibility of its having been formerly of greater extent, and of having been subsequently reduced by subsidence. We will first review the mammal-fauna, and then try and deduce, from a study of it, our conclusions as to its . past history. In Celebes alone of the Oriental Region we find repre- Ceo : : 4 : THE ORIENTAL REGION 147 sentatives of the marsupials characteristic of the Australian Region. These consist of two species of Phalanger, which differ from those of the Australian islands only in slight particulars. The next interesting animal of this fauna is the Babirussa, a wild pig remarkable for the enormous size of its upper and lower canine teeth, which form, as it were, two pairs of horns on the upper side of the head. Another peculiar Ungulate, now generally referred to the widespread genus Bos, is the Anoa, which shows many primitive characters, and is entirely confined to the island. The Mice and Squirrels of Celebes are fairly numerous, and most of the species are peculiar to the island; one rat forms a special genus. Carnivores are very scarce in Celebes; Insectivores have not been recorded at all. The Bats, which are numerous, comprise a considerable number of Australian forms, and one peculiar genus. Among the Primates, Tarsivs of the other Malayan islands is also found in this Sub-region, and one Monkey, Macacus mawrus, seems to be restricted to it. Finally, one of the most remarkable of the animals of the island is the Black Ape of Celebes, belonging to a genus (Cyno- pithecus) intermediate between the Macaques and the Baboons. Cynopithecus appears to have found its way from Celebes into the adjoining island of Batchian, which belongs to the Australian Region. The following table shows the mammals of this Sub- region arranged in a form like those of the other Sub-regions :— 148 THE GEOGRAPHY OF MAMMALS a = . : . a s a Sia). | ee see Endemic . OH 0s Peak Fs 10 ieee He eae Oriental Ow Ome) sasO: Tg esi 4 Australian TS Oa ga Mada he ela) 3 Palzeogean COTE he) eee la We [ Oo) el aie Cosmopolitan COP HITT) Se hE fe (0) | PV Ui) Total «20/2 ie eed From this summary it will be seen that the total number of Mammal-genera that occur in Celebes is thirty- one, the greater number of which (twenty in all) are placed under the headings of Paleogean and Cosmopolitan. These are all widespread genera, which do not afford us any particular clue to the origin of the Celebesian fauna. Nine out of the twenty are genera of Bats, which, as has before been remarked, are by nature much less restricted in their range than the true quadrupedal mammals. Of the remaining eleven only two (Mus and Sus) have any extensive distribution in the Australian Region; the others, although they have, in one or two cases, managed to struggle into adjoining islands belonging to the Australian Region, can in no sense be viewed as Australian genera. Of the genera registered in the table as “ Australian,” two are Bats, which have apparently reached Celebes from the more easterly islands of the Australian Region, where they have a wide distribution; the other is the genus Phalanger, which has been already alluded to as being the only member of the Marsupial Order found in the Oriental Region. THE ORIENTAL REGION 149 The endemic genera of Celebes are four in number, and judging from their affinities, it is impossible to believe that they have any relation to the animals now living in the Australian Region. Everything points to their being remains of a very ancient fauna, which must have been originally derived from the Asiatic continent. The presence of the three Australian genera in Celebes does not in any way require the supposition of an ancient land-connection with that Region. This is obviously so in the case of the Bats, and the Phalanger is a strictly arboreal animal, and might easily have been drifted across a narrow strait on floating timber. On the other hand, to account for the greater proportion of Oriental forms found in the island, we are driven to the conclusion that at some time or other there has been some sort of land-connection between Celebes and the mainland of Asia. These are the prin- cipal reasons for transferring the island of Celebes from the Australian to the Oriental Region.! Section VIII.—Tue Past History or THE ORIENTAL MAMMAL-FAUNA Considerable controversy has arisen from time to time with regard to the similarities that undoubtedly exist between the faunas of the Oriental and Ethiopian Regions. Some writers have urged that, in order to account for this, some form of direct land-connection must have existed at one time or another across the Indian Ocean between 1 For the most recent information on the mammals of Celebes see Meyer, ‘‘Saugethiere von Celebes und Philippinen-Archipelago,” Abh. Mus. Dresden, vi., No. 6. 150 THE GEOGRAPHY OF MAMMALS Southern India and South Africa. Others have maintained that the points of similarity between the two faunas have been exaggerated, and that no such land-connection is required to account for the facts, which can easily be explained on the supposition of a southward emigration of northern forms due to glacial cold. If we go back to the early part of the secondary epoch of geological time, we find, very well developed in India, a geological system known as the Gondwana, composed of sandstones and shales, which appear to be of fluviatile origin. These beds have long afforded a problem to geologists, as they cannot be at all satisfactorily correlated with any formations in Europe. In South Africa, however, we find a series of beds, also doubtless of fresh-water origin, known as the Karroo formation, which contains a nearly similar set of fossil remains, and in New South Wales, again, there are formations also agreeing in the characters of their fossils with the Gondwana beds. These facts, according to Mr. Oldham (3), our latest authority on this subject, are “inexplicable, unless there has been a continuous land-communication along which plants could freely migrate, and the conclusion is vastly strengthened when we remember that throughout the great part, if not the whole, of this period, a very different type of flora was flourishing in Europe and North America.” This land-connection may be of use in explaining the distribution of some of the lower vertebrates, but is of no assistance so far as the Mammals are concerned; because in those early times it is probable that none of the families or even orders of our present Mammals had arisen. The best-known and richest of the mammal-bearing formations of India are certain beds in Sind, and the THE ORIENTAL REGION 151 Siwalik deposits lying along the foot of the Himalayas. These beds, especially the latter, contain the remains of an extensive and exceedingly interesting Mammalian fauna, which has hitherto been very inadequately ex- plored, and will probably afford abundant opportunities of discovery to the paleontologist of the future. The number of genera hitherto discovered in these formations amounts in all to about sixty, of which thirty- nine are still in existence, while twenty-five are extinct. Among the recent genera are a considerable number which, though still occurring in Africa, have become extinct in the Oriental Region; such are Bubalis, Cobus, Oreas, and Strepsiceros—all genera of antelopes, Giraffa (the Giraffe), Hippopotamus, Loxodon (the African Elephant), Cynocephalus (the African Baboon), and A nthropopithecus (the Chimpanzee), while others still survive in India. The most remarkable feature, however, of the Siwalik © fauna is the fact that, while certain of the genera are only found in Miocene beds in Europe, and not in more recent deposits, the greater number are only known, out of India, from the Pliocene and Pleistocene, so that it is very difficult to fix the age of the Siwaliks as compared with the formations of Europe. Beds containing a somewhat similar fauna, in most cases not so rich, have been discovered in Greece, near Athens, in Samos, and in one or two other localities, at least, in South-Western Europe; while north of the Alps nothing of the sort has been found of a corresponding age. The most plausible explanation of the whole matter, therefore, so far as we can say at present, is that the increasing cold at the end of the Miocene and the beginning of the Pliocene times gradually drove the ae THE GEOGRAPHY OF MAMMALS northern inhabitants southwards. It thus came to pass that, at that period of the world’s history, the Mammalian faunas of Southern Europe, South-Eastern Asia, and of India were so nearly uniform as to constitute these countries, as regards their mammals, one widely extended Region. List OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHAPTER V. (1) Minnz-Epwarps, H. nr A.—“ Recherches pour servir a histoire naturelle des Mammiféres.” Paris, 1868. (2) Buanrorp, W. T.—“ The Fauna of British India : Mammalia.” London, 1888-91. (3) OLpHam, R. D.—“ Manual of the Geology of India.” Caleutta, 1893. Scale of Miles i199 200 300 400 500 oO MAP OF THE ORIENTA ‘ ‘ ‘ ' ' ‘ ' ‘ ' ' ' ' ' 1 ' ' ' ' ' ¥ i ' ‘ ' ‘ ' sal Scale of Miles o 190 200 300 400 500 Ley 60 MAP OF THE ORIENT L REGION SHOWING ITS DIVISION INTO 4 SUB-REGIONS. CHAPTER: VI fe NEARCTIC REGION (PLATE VI, p. 176) SECTION J.—BOUNDARIES OF THE NEARCTIC REGION Wir the Nearctic Region we enter upon a subject on which there has been a great deal of controversy among the students of geographical distribution. As was pointed out in the introductory article of this volume, a certain number of writers maintain that this Region does not contain a sufficient quantity of distinctive and indigenous forms to entitle it to separation from the Palearctic Region. What should constitute a sufficient number of distinctive forms depends, of course, largely on the individual opinions of the writers, but if allowance be made for the undoubted similarities of the extreme northern parts of the Old and New Worlds, which together may be considered as form- ing a kind of intermediate district, the facts and figures given below will, we think, convince every one that the land-surfaces of the Palearctic and Nearctic Regions have now, and have had in the past, quite sufficiently dis- tinct faunas to warrant their division into two primary Regions. The boundaries of the Nearctic Region are compara- tively simple. They embrace the whole of North America as far as the southern limit of the tableland of Mexico, with which Greenland may be included. On either side of 153 154 THE GEOGRAPHY OF MAMMALS the tableland of Mexico, the Neotropical Region extends as a narrow strip along the Gulf of Mexico and the Pacific, as far north as the Rio Grande on the former coast, and to about Guaymas on the latter. There are no islands of any importance belonging to this Region that need be mentioned here. The Pacific islands on the west coast of Mexico have few, if any, mammals. The large islands of Newfoundland and Van- couver are of the true continental type, being separated from the mainland only by quite shallow water. The Antilles, or West India islands, belong entirely to the Neotropical Region, and have been already considered in a previous chapter. SEcTION I].—GENERAL VIEW OF THE MAMMAL-FAUNA OF THE NEARCTIC REGION On referring to the table given at the end of the first chapter, it will be seen that the total number of genera, and also of species contained in the Nearctic Region, is considerably less than in any of the other Regions hitherto treated of. This may be explained partly by the fact that the whole of this Region is practically outside the tropics, whereas the other Regions previously described lie to a great extent within the tropical zone, which is very favour- able to the development of a rich and varied fauna. Out of the nine Orders into which the terrestrial Mammals have been divided, two only are not represented in this Region. These are the Primates, at the head of the list, and the Monotremes, at the extreme end, the latter being confined to the Australian Region. The Marsupials THE NEARCTIC REGION 155 are represented by one species only, the well-known Virginian opossum (Fig. 29), which is found with slight modifications from the Southern States of North America Fic. 29.—THE VIRGINIAN OPOSSUM. (Didelphys virginiana.) southwards over the greater part of South America. This animal would, perhaps, judging merely by the present dis- tribution of life, be considered to have intruded into the Nearctic Region from South America, where alone members of this family still survive; but, on examining its past 156 THE GEOGRAPHY OF MAMMALS history, we ascertain that the genus Didelphys was formerly found both in Europe and in North America during Eocene and Miocene times, so that it is possible that the Virginian Fig. 30.—THE PRONG-BUCK. (Antilocapra americana.) opossum may be a survivor rather than an intruder in North America. The next order, the Edentates, is represented in this THE NEARCTIC REGION 157 Region only by a single species of armadillo (Zatusia novem-cincta), which almost certainly came up from the south, and is only met with just inside the southern borders, Fig. 31.—THE ROCKY-MOUNTAIN GOAT. (Haploceros montanus.) Among the Ungulates, the most remarkable form is the prong-buck (Antilocapra), which forms a distinct family of that order, and is entirely confined to this Region (Fig. 30, p. 156). It is allied in some respects to the Antelopes of the 158 THE GEOGRAPHY OF MAMMALS Old World, but it is unique among all the hollow-horned ruminants from the fact that it sheds its horns every year. Two other genera, belonging to the family Bovidee, are confined to the Nearctic Region; these are the Rocky Mountain goat (Haploceros), found only in the Rocky Fig. 32.—THE Musk Ox. (Ovibos moschatus. ) Mountains (Fig. 31, p. 157); and the Musk Ox (Ovibos), which ranges over the barren grounds at the extreme north of the continent, and spreads into Greenland (see Fig. 32). The latter, however, was also found in the northern parts of the Old World until a comparatively recent epoch, geologically speaking. THE NEARCTIC REGION 159 The Bisons (Fig. 33) are still common to the Nearctic and Palzarctic Region, though now nearly extinct in both hemispheres. Rodents are very numerous in the Nearctic Region. According to the tables here used, which have been compiled from Flower and Lydekker’s text-book of Fic. 33.—THE AMERICAN BISON. (Bison americanus. ) mammals, out of a total number of twenty-eight genera, thirteen are endemic. One of these, Haplodon, a small animal resembling the Prairie-dog in its habits, and found only west of the Rocky Mountains, forms a distinct family. The Carnivora are also well represented, especially the genera of Cats, Dogs, Bears, and Weasels, all of which, 160 THE GEOGRAPHY OF MAMMALS however, are widely spread. The only endemic genus is that formed for the reception of the American Badger (Taxidea), which differs from its European ally in certain anatomical features. In contradistinction to the Neotropical Region, the Insectivora are abundant in the Nearctic; there are no less <) sie Fig. 34.—THE STAR-NOSED MOLE. (Condylura cristata. ) than four genera of Moles met with, three of which are peculiar. Amongst them is the remarkable Star-nosed Mole (Condylura cristata) which carries a ring of riband-like appendages at the end of its snout. These serve, no doubt, as a sensitive organ (see Fig. 34). Finally, the Bats are neither very numerous nor of THE NEARCTIC REGION 161 great importance; only one genus (Antrozous), containing one species, being peculiar out of a total of nine. Summarizing, therefore, we find the Nearctic Region to be characterized by the exclusive possession of only two families of Mammals—namely, Antilocapridee (the Prong- bucks) and Haplodontide (the Haplodonts), and by the presence of sixty-six genera, of which twenty-one are restricted within its boundaries. On the other hand, in addition to the two orders already mentioned, Monotremes and Primates, the following important families are absent in the Nearctic Region, although fairly well spread over the Old World :— Suidee (Swine). Hyenide (Hyenas). Equidee (Horses). Erinaceidze (Hedgehogs). Myoxidee (Dormice), Pteropodidee (Fruit-eating Bats). Viverridee (Civets). Rhinolophidz (Leaf-nosed Bats). That some of these families did, however, at one time exist on the North American continent has been shown by recent palzontological discoveries. Section II].—SvuB-DIVISION OF THE NEARCTIC REGION The recent work of American naturalists, more espe- cially that of Merriam (2) and of Allen (1), has greatly increased our knowledge of the mammals of North America and of their distribution. These naturalists have further shown that the Sub-regions adopted by Wallace in his well-known text-book on geographical distribution are not altogether supported by the facts now known to us. Mr. Allen, in his paper on the distribution of North L 162 THE GEOGRAPHY OF MAMMALS American mammals, first of all excludes from what he terms the North American Region the extreme northern parts of that continent. He considers that the Arctic portion of that continent, namely, that beyond the limit of arboreal vegetation, forms, together with the similar part of the Old World, a separate Region, or, as he terms it, “the Arctic Realm.” Furthermore, the southern part of North America south of the Mexican tableland, together with the low- lying country of Mexico on either flank, he assigns to the American Tropical Realm. The remainder of the con- tinent, combined with the great mass of Europe and Asia, forms, according to this author, the North Temperate Realm. This scheme of division does not differ essentially from that of Mr, Allen. The Arctic portion of North America forms, no doubt, as Mr. Allen puts it, “part of a homogeneous hyperborean faunal area of circumpolar distribution.” Mr. Allen’s American Tropical Realm has already been treated of in the chapter of this volume dealing with the Neotropical Region (see p.52). There remains, therefore, Mr. Allen’s North American Region, which nearly corresponds to the Nearctic Region of our scheme of classification. Mr. Allen divides his North American Region into two Sub- regions—the Cold Temperate and the Warm Temperate, the two latter falling into two provinces, a Humid or Eastern and an Arid or Western. Proceeding further, he divides the Humid province into two sub-provinces, namely, an Apalachian or Northern and an Austro-riparian or Southern. The Arid or Western province is also separated into two sub-provinces—the Campestrian or Northern and the Sonoran or Southern, and, besides this, the sub- THE NEARCTIC REGION 163 provinces are divided into various minor divisions termed districts and faunas. It will be sufficient for our present purpose to divide the North American or Nearctic Region into three Sub- regions; these may be termed, (1) the Canadian or Cold Sub-region, (2) the Western or Humid Sub-region, and (3) the Eastern or Arid Sub-region. Mr. Wallace has recognized four Sub-regions in the Nearctic Region. His Canadian Sub-region corresponds fairly well to the Canadian or North Temperate of Mr. Allen, except for the fact that it has not been made by Mr. Wallace to extend southward down the mountain ranges. The Alleghany Sub-region of Mr. Wallace, again, practically corresponds to the “ Humid” of Mr. Allen. The two others, the Rocky Mountain and Californian, correspond to Mr. Allen’s “ Arid,” the Californian Sub-region being composed of a narrow strip of coast country between the Sierra Nevada and the sea, and extending from Queen Charlotte’s Sound in the north to the south-western corner of Cali- fornia. The differences, therefore, between Mr. Wallace’s and Mr, Allen’s views are not so fundamental as one would gather from the critical remarks of the latter author. The boundaries of the Sub-regions here adopted will be best understood by reference to the accompanying map (Plate VI., p. 176). The Canadian or Cold Sub-region embraces the whole of the northern portion of North America, including Greenland. The southern limit of this Sub-region com- mences, on the Atlantic side, on the coast of Maine, in the neighbourhood of Augusta, and thence runs to Quebec and through the Great Lakes. Further west it is bounded 164 THE GEOGRAPHY OF MAMMALS by the northern branch of the Saskatchewan, and ends on the Pacific coast in the neighbourhood of Queen Charlotte’s Sound. But long prolongations of this Sub-region extend down the Alleghany mountains; in the east as far as Georgia, along the Rocky Mountains as far as the Rio Grande, and along the Cascade and Sierra Nevada moun- tains as far as the Colorado river. Besides these, there are several smaller detached portions of other mountain ranges, which should be attributed to the Canadian Sub- region. The Eastern or “Humid” Sub-region is separated from the Western or “Arid” by a line running roughly nearly along the meridian of 100° west of Greenwich, and ex- tending from Manitoba to the mouth of the Rio Grande del Norte. Section 1V.—THE CANADIAN OR COLD SUB-REGION The Canadian Sub-region is especially remarkable for a number of genera which are common to it and the northern part of the Old World, but which do not extend southwards into the other Sub-regions to be presently treated of. It is further characterized by the small number of genera which are essentially New World forms, and have no connection with the Old World. Reviewing the Mammals in detail, we find that the Sub-region contains no representative of either Marsupials or Edentates. On the other hand, there are six genera of the Ungulates within its limits—a far larger proportion than that found in the other Sub-regions. Of these the only one endemic is Haploceros (the Rocky Mountain goat). This somewhat isolated Ruminant has its nearest allies in the genus THE NEARCTIC REGION 165 Nemorhzdus, of the mountains of Asia, which occurs in Japan (NV. crispus), but of which the best-known form is commonly designated the “Serow” by the sportsmen of the Himalayas. There are also no less than four genera found in the Old World, and also in the Canadian Sub- region, which do not extend further south. These are Cervus, containing the Wapiti (C. canadensis), closely allied to the Red Deer of the Old World; the Cariboo (Rangifer), which cannot be distinguished from the Reindeer of the northern part of the Palearctic Region: and the Moose (Alces machlis), which has the same distribution as the Reindeer, but is known in Europe under the name of “ Elk.” Besides these, there are two Canadian genera of this order which are found in other Sub-regions as well as in the Old World, namely, the Bison (Bison) and the Sheep (Ovis). The Bison, of which the American representative is the so- called buftalo (now, alas! nearly extinct), is closely allied to the European Bison, still found in certain parts of the Old World; while the Big-horn Sheep (Ovis canadensis) is a representative of the Wild Sheep, which are extensively distributed in the Palearctic Region. The number of genera of Rodents of the Canadian Sub-region amounts in all to eighteen, of which three are peculiar. One of these is Haplodon, to which allusion has already been made; the others are Phenacomys, a small genus of rats, and Erethizon, which contains only the Tree-porcupine of the Canadian forests. Among the members of this order, too, we find three genera common to this Sub-region and the Old World, which do not extend further south. These are Myodes (M. obesus), which is represented in Europe by an allied species, the well-known Lemming of Scandinavia ; Cuniculus, a form nearly allied to the Lemming; and 166 THE GEOGRAPHY OF MAMMALS Lagomys, the “ Pika,” or Tailless Hare, which is found in the higher mountain ranges of both the Old and the New Worlds. The Carnivora do not present many features of special interest. Two genera—Mustela, containing the Weasels, and Gulo, the Glutton, have much the same circumpolar distribution as has been already remarked on in the case of the Deer and the Lemming. The number of genera of Insectivora and Chiroptera in the Canadian Sub-region is insignificant, and they are of no interest from a distributional point of view. Viewing the fauna of the Canadian Sub-region as a whole, it will thus be seen that its greatest point of interest is its resemblance to that of the more northerly parts of the Old World. This, of course, may be easily accounted for when we recollect that the sea of Behring Siraits is quite shallow, and in places not more than about 20 miles in breadth. There can be no doubt that there was a land- connection between Siberia and Alaska in comparatively recent geological times, and that this has resulted in the commingling of the faunas of the northern parts of these two Regions, to a considerable extent. This land-bridge must have existed so recently that there has not yet been even time for, in some cases, the animals to become differentiated into appreciable species, as in the cases for example, of the reindeer and elk. . Below will be found a surmmary of the genera of the Canadian Sub-region, forty in number, which are divided into five categories much in the same fashion as has been done in the previous articles, namely :— 1. Endemic—those found only in the Canadian Sub- region. THE NEARCTIC REGION 167 2. Nearctic—those not found beyond the limits of the Nearctic Region. 3. Neogean—those found in the New, but not in the Old World. 4, Arctic and Palxogean—those which are found in the Old World, and only in the Canadian Sub-region of the New; and, finally— 5. Neogean and Palxogean — containing the most widely distributed forms. 2 . | | [Pesce s| PRE Sethe Pa TN es all Sah | = | S| = 2 | s | fsija|a| e/a) 1. Endemic 1 SOL. OF 0 | 4 2. Nearctic O 3 i) @ | & 3. Neogeam- = 2 . « « OM One Oe OLS D ics 4, Arctic and Paleeogean . BENS Ql 4Oua 0? | 79 5. Neogean and Paleeogean wy Oy Bal al 2 || Is) Total (3 |] alts} yada! 2 33 || lo) SECTION V.—THE WESTERN OR ARID SUB-REGION This Sub-region is, on the whoie, the richest of the three, both as regards the total number of genera found within its limits and also as regards the number of genera peculiar to it, which amounts to seven out of fifty-three, as com- pared with four out of forty in the Canadian, and one out of forty in the Eastern. The Western Sub-region shares with the Eastern the only representative of the Marsupials found in North America, the Virginian Opossum. Just extending, too, to within its limits occurs the only member of the Order 168 THE GEOGRAPHY OF MAMMALS Edentata, the Nine-banded Armadillo (Tatusia novem- cincta). Among the Ungulates, the Prong-buck (Antzilo- capra) is restricted to this Sub-region, and the genus Cervus of the cold Temperate Sub-region is replaced by the purely American genus Cariacus, of which the Black-tailed Deer is the representative. This Sub-region is also more particularly the home of the American Bison, which, however, ranged even in historic times eastward nearly to the Atlantic seaboard. Among the Rodents there are no less than five endemic genera, of which, perhaps, the best known is Cynomys, the Prairie- dog. The other endemic genera all belong to the family Geomyide, which contains a number of small Rodents known as Pocket-gophers. Among the Carnivora there are no genera in the Western Sub-region which are not more or less widely dis- tributed, the greater number of them, both in the case of this order and of the last, being also found in the Old World. Three genera of Moles belonging to the next order, Insectivora, though confined to North America, extend into the Eastern Sub-region. The Bats of the Arid Sub-region include among their members two genera (each with one species) which occur only in California, while four other genera are found only in the New World. One of these, Macrotus, is of special interest, since it con- tains the only member of a very large and well-marked family (Phyllostomatidx), which extends into the Nearctic Region from South America. This Sub-region, as compared with the Canadian, con- tains a far larger proportion of Neotropical genera, and, in addition, is characterized by the absence of anumber of the Palearctic forms found only in the Canadian Sub-region. THE NEARCTIC REGION 169 Such, for instance, are Rangifer (the Reindeer), Alces (the Elk), Haploceros (the mountain goat), Gulo (the glutton), and many others. The following table, which gives a summary of the genera of this Sub-region, differs only from the summary of the Canadian Sub-region in the omission of the heading “ Arctic and Palearctic,” since practically all the genera common to this Sub-region and the Palearctic Region are also found in the Canadian Sub-region. ep ees | tse [eee ne oneal let alt cas Bo) | & be ee | Ble 8 Scenes | 2s.) aah e Endemic . OPO) at| ml Ah OF 5G 62 7 Nearctic . OR Om iO By Wy By ow 9 INCOREAT mao eens tol A Uae S:\) 5) | OF aS vel Neogean and Paleogean. OF Omi 1 OF | Gy 2A see Total 1 1 AG S29 Zao 953 SEcTION VI.—THE EASTERN OR HUMID SUB-REGION So far as peculiar forms go, the Eastern Sub-region is quite the least peculiar of the three. It contains only one genus strictly confined within its limits; this is Neofiber, with a single species commonly known as the “ Round-tailed Musk-rat.” This Rodent is found only in Florida, and is much less completely aquatic in its habits than the true Musk-rat (Fiber), which is spread over the rest of North America. Taking the orders seriatim, the Marsupials are repre- sented by the widely spread Virginian Opossum ; but the Edentates do not reach the Region at all. Of the Ungulates 170 THE GEOGRAPHY OF MAMMALS only the genus Cariacus (the Virginian Deer) occurs, the Bison (though formerly inhabiting this Sub-region) not having been seen east of the Mississippi for the last forty or fifty years. The Rodents, as in the other Sub-regions, make up the great mass of the mammalian genera, numbering seventeen in all, including Neofiber. The Carnivores, Insectivores, and Bats do not differ very markedly from those of the Western Sub-region. On the whole the Eastern Sub-region is not a very well-marked division; it differs from the Canadian chiefly in the non-existence of the numerous nortuern Paleogean types found there, and from the Western Sub-region in the absence of a good many characteristic desert-haunt- ing forms, and also of several of the South American genera, which have spread up northwards from the Nearctic Region into the Western Sub-region, but which have not reached the more distant Eastern. The following table gives a summary of the genera of this Sub-region, from which it will be seen that the total number (forty) of genera is markedly less than the corresponding number in the Arid Sub-region. Pieri = (R= ear PeSetl|| po | 455" ee |e) seer eS rar S ro) & o =| oS Endemic Pear ec Os ee) Or O80 1 Neanetio: Wire mal tee ve fee pons Oa nO 5) Dilan so 9 Nepean wear a wigs ie 2 DAW Se ele ee Neogean and Paleogean . . .| 0 | 0] 9/ 5] 1 | 4 | 19 Total’ -;) ) 2/31/30 age) Oia a eceleea as THE NEARCTIC REGION lejall Section VII.—Tuer Past History oF THE NEARCTIC MAMMAL-FAUNA During the last twenty years the wonderful discoveries of American paleontologists have thrown a flood of light, not only on the past history of the Nearctic Region, but also on the evolution of many of the mammalian groups themselves. It is, therefore, very necessary, when review- ing the geographical distribution of the present mammalian fauna, to shortly recapitulate the more important results and conclusions arrived at from their writings. A very useful and comprehensive summary of this work will be found in a paper by Professor Zittel (3), lately published in the Geological Magazine. The beds which contain the remarkably perfect remains above alluded to are found only in the western part of North America. Here, apparently, there existed throughout the Tertiary Epoch a series of great fresh-water lakes, on the sides and the bottoms of which were formed an almost continuous series of deposits with the remains of the animals of the surrounding districts embedded in them. The great interest of these discoveries lies in the fact that we can here trace the gradual formation and evolution of several of the mammalian orders as they at present exist. In the oldest beds the mammals resemble one another so closely that it is often impossible to assign them very definitely to any of the existing orders, although the germs of the commencing distinctive characters can even here be traced. In the later beds the various groups gradually differentiate themselves, until in the most modern of $72 THE GEOGRAPHY OF MAMMALS the deposits the genera can all be definitely assigned to existing orders. The earliest mammals that have been found in North America come from the Trias of North Carolina, but neither these nor any of the other mammalian remains of the Secondary Period tend to assist the geographical problems involved, or are of importance in the present juncture. With the oldest Tertiary beds an entirely new fauna appears, and furnishes us with remains of forms belonging to various orders of which no traces can be found in the earlier Secondary deposits. The following is a short list of these deposits, together with their European equivalents, so far as they can be ascertained :— Puerco beds of New Mexico. Wasatch beds of Wyoming, Utah, and New Mexico. Mip-EocenE . . Bridger beds of Wyoming. Uprer Eocenr. . Uintah beds of Utah and Wyoming. Lower Miocene . White-river beds of Nebraska, Dakota, Colo- rado, and Wyoming. Lower EocENE 4 Mip-Miocene . . John-Day beds of Oregon, Nevada, and Washington. PLIOCENE . . . Loup-Fork beds of Nebraska, Colorado, Wyo- ming, Kansas, and New Mexico. LareE PiioceNE . Equus beds of Western and South-Western States, and Megalonyx beds of Eastern States. In the lowest Eocene beds, not only the Genera, but even the Orders of mammals are in almost every case different from those at present existing. The greater number of these belong to two orders—Creodontia and Condylarthra; the former the progenitors of the modern Carnivora, the latter of the existing Ungulata. These THE NEARCTIC REGION 17h) two, and indeed the other orders to which the mammals of this fauna have been assigned, all show considerable points of resemblance to one another, first in the posses- sion of five toes on both limbs, which are provided with neither claws nor hoofs, but with a structure somewhat intermediate between the two, and, secondly, in their ex- tremely small cerebral cavity. A similar, though much more incomplete Fauna has been found in certain beds of a corresponding age in Europe, the Genera of their fossil mammals being in most cases identical with those of the Nearctic Region. In the next stage, the Wasatch beds, which correspond in age nearly to the London clay of England, a further development of the same fauna is found, with, however, the commencement of certain of the modern Orders; such, for instance, as the Perissodactyla (or Odd-toed Ungulates), the Rodents, the Insectivores, and the Lemurs. Here, too, so far as the scanty remains found in Europe allow us to form a comparison, there is a close similarity between the faunas of the two Regions. In the succeeding “ Bridger beds” of Mid-Eocene age is found the earliest evidence of the still surviving genus Didelphys (the Opossum). Here also marine Mammals and Bats appear for the first time. But comparison of these remains with European forms is even more difficult than in the last case, owing to the scarcity of such fossils in beds of the same age in Europe. In the Uintah beds of the Upper Eocene we first begin to find very distinct traces of differentiation between the European and the North American faunas, although a good many of the Genera met with are still common to the two Regions. 174 THE GEOGRAPHY OF MAMMALS A great advance is found in the Fauna of the White- river beds of Miocene age. In this case the mammals can almost be referred to existing orders, but comparatively few of the genera are common to the Old and New Worlds; and it appears that, whilst during the older Eocene there was a considerable emigration of New World forms into Europe, in Miocene times the stream was re- versed, and North America received the greater number of its immigrants from this side of the Atlantic. This immigration continued during the Middle Miocene epoch, the Fauna of which has been well preserved in the John- Day beds of the extreme west. At the same time many endemic Families and forms are also met with, especially as regards the early predecessors of the Camel Family, which apparently had its origin and early development in the Nearctic Region, though now entirely absent from it. In the John-Day beds, we also find, for the first time, remains of the modern genera, Rhinoceros, Sciurus, Hesperomys, and Lepus. The succeeding “ Loup-Fork beds” contain additional recent genera, some of which, such as Hquus and Camelus, are now no longer found in the Nearctic Region, while others, such as Canis, Mustela, and Lutra, still remain there. On the whole, however, the Fauna of this epoch is still further removed from that of the corresponding period of the Old World than that of the preceding. A little later, in the so-called “ Equus-beds” of the Western States, and in the contemporaneous “ Megalonyx- beds” of the Eastern, we first find a number of Neotropical forms, such as Mylodon (a gigantic Sloth), Glyptodon (a gigantic Armadillo), Hydrocherus (the Capybara), and THE NEARCTIC REGION 175 Towodon, » member of a peculiar extinct family of Ungulates. The occurrence of all these animals indubitably proves that now for the first time a connection had been formed between the continents of North and South America. Before this epoch, no trace of a Neotropical admixture can be anywhere detected in the Nearctic mammal-fauna. Thus the evidence of palzeontology in every way sup- ports the deductions drawn from a study of the distribu- tion of recent forms, namely, that the bulk of the present Nearctic fauna has been mainly derived from the Old World, although at times the Region has been sufficiently isolated and sufficiently extensive for the independent evolution of its own characteristic forms. In accordance with these deductions, the present remaining inhabitants of the Nearctic Region may be divided into three cate- gories, as follows: (1) The Endemic Fauna, the bulk of which has had, at. some considerably remote geological period, a common origin with that of the Palearctic Region, although it has enjoyed ample time to develop and differentiate itself on its own lines. (2) A Neotropical constituent, which first appeared in the Nearctic Region in Pliocene times. (3) A comparatively modern Palearctic fragment, in which not only the genera, but frequently the species, are identical in both Regions. This portion of the fauna has probably reached the Nearctic Region by the passage which must have existed in comparatively modern times across Behring Straits. Consequently, while the Neotropical element is the stronger in the south, this last, the Palearctic element, is far more prevalent in the extreme north. 176 THE GEOGRAPHY OF MAMMALS List OF THE PRINCIPAL AUTHORITIES REFERRED TO IN CHaprer VI. (1) Auten, J. A.—“The Geographical Distribution of North American Mammals,” Bull. Amer. Mus. Nat. Hist., iv., p. 199. 1892. (2) Merriam, C. H.—“ The Geographical Distribution of Life in North America, with Special Reference to the Mammalia.” Proc. Biol. Soc. Washington, vii., p.1. 1892. (8) Zrvre, Kart von.—“ The Geographical Development, Descent, and Distribution of the Mammalia.” Geol. Mag., 3rd ser., x., p. 401. 1893. (Translated from the S. B. k. Bayer. Akad. Wiss., xxiii., p. 137. 1893). PLATE VI. = 130 wo nO 100 90 go 7 Arctic | cirdle HE NEARCTIC REGION SN Stel A nas VISION INTO 2 @TIp mremrrmnaure 150 160 4130 12 Oo nO 100 90 80 ao 20 Scale of Miles 100 9 200 400 foo 300 ieo0 ——S— ee ae ee ee | Oo Loo MAP OF THE NEARCTIC REGION SHOWING ITS DIVISION INTO 8 SUB-REGIONS. Ye 2 | 2 _— ~ ie —_ | Scale ot Miles 200 400 00 MAP SHOWING I CHAPTER VII THE PALMARCTIC REGION (PuiaTE VII, p. 196) SEcTION I.—BOUNDARIES OF THE PALHARCTIC REGION Tuis, the last of the six great Regions, consists, as its name implies, of the whole northern part of the Old World. Its boundaries have already been defined in previous articles dealing with the Ethiopian and Oriental Regions, these being the only Regions with which it marches. Speaking generally, it may be said to consist of the whole of Europe, the northern border of Africa, and Asia north of the Himalayas. Its southern boundary in Africa was taken, in the article on Ethiopia, as the Tropic of Cancer, but this, of course, is a purely arbitrary line, and runs through the centre of the Sahara Desert. It would, perhaps, be more accurate to put in its place the northern edge of the Sahara as the limit, and to include all the desert country both of Africa and Arabia in the Ethiopian Region. The question of Egypt is a difficult one, as its fauna undoubtedly contains a mixture of forms characteristic of both the Palearctic and Ethiopian Regions; on the whole, however, Egypt, up to the First Cataract, is best included in the Palearctic Region. In regard to the boundary line between the Palearctic and Oriental Regions, there can be no doubt that at the higher elevations of the Himalayas a true Palearctic fauna 177 M 178 THE GEOGRAPHY OF MAMMALS is met with. Eastward of Sikkim, however, as has already been shown, it is very difficult to draw a definite line, chiefly in consequence of our defective knowledge; but the boundary already adopted in the case of the Oriental Region seems, on the whole, to be a suitable one—namely, that of the northern water-parting of the Yang-tse- Kiang —thus leaving Moupin, and the district of Eastern Tibet explored by the French missionary, Pere David, within the confines of the Palzearctic Region. There are only two important groups of islands con- nected with this Region; these are the British Islands in the West, and the Japanese Islands in the East. The faunas of both these insular groups are of the true “continental- island” type, and differ very slightly from that of the neighbouring mainland. This is more especially the case with the British Isles, where we find among the Mammals no peculiarities worthy of mention, with the exception, perhaps, of a recently discriminated stoat (Mustela hiber- nica), said to occur only in Ireland. SEcTION I].—GENERAL VIEW OF THE MAMMAL-FAUNA OF THE PALZARCTIC REGION The Palzarctic Region, although covering a larger area than any of the other Regions, comes out only fourth as re- gards the number, both of species and of genera, of mammals represented in it, being surpassed in this respect by the Neotropical, Ethiopian, and Oriental Regions. The total number of such genera is one hundred and three, out of which twenty-five are absolutely confined within its limits, while four others are highly characteristic of it, though they just pass over its frontiers. The remainder, seventy-four THE PALAARCTIC REGION © 179 in number, are mostly widely spread. When these figures have been reduced to percentages, it will be found that only 24 per cent. of the genera are endemic, which is considerably less than in any other of the Regions hitherto treated of. Reviewing the Fauna in detail, we find that of the nine Fig. 35.—THE BACTRIAN CAMEL. (Camelus bactrianus.) terrestrial Orders, six only are represented in the Palearctic Region, the Edentates, Marsupials, and Monotremes being completely absent. Among the Ungulates, of which a considerable number of forms are found within the Palearctic sphere, there is a fair percentage of peculiarities. The Bactrian, or Two-humped camel (Fig. 35), is known to 180 THE GEOGRAPHY OF MAMMALS exist still in a wild state only in certain districts of Central Asia, while the Arabian, or One-humped camel, has never yet been met with in a truly wild condition, so that the genus Camelus may be considered as truly Palearctic. Fig. 36.—THE MUSK-DEER. (Moschus moschiferus.) [List of Vert. An. 1896, p. 171.] Two other endemic genera belonging to the Deer-family (Cervide) are Moschus, the Musk-deer, a small hornless deer found only in the higher mountain ranges of Central Asia (Fig. 36); and Capreolus, the Roe-deer, fairly well spread over the whole extent of the Region. THE PALZARCTIC REGION 181 Among the Bovidx, Saiga, an Antelope found on the steppes of Russia and Western Asia, Pantholops, another Fig. 37.—THE CHAMOIS. (Rupicapra tragus.) Antelope of the Central-Asiatic mountains, and Rupicapra, the Chamois of the European mountain ranges (Fig. 37) are confined to this Region. The important genera of 182 THE GEOGRAPHY OF MAMMALS Sheep (Ovis) and Goat (Capra) must be considered as especially characteristic of this Region, although stragglers of these forms are found in the confines of the Nearctic and Oriental Regions. They are all mountain animals, one of the best marked forms being the Barbary Sheep Fig. 38.—THE BARBARY SHEEP. (Ovis tragelaphus.) (Ovis tragelaphus) of Northern Africa (Fig. 38). Of the Rodents, the two most characteristic Palearctic families are the Dormice (Myoxidz) and the Jerboas (Dipodidz). Of the former family two genera, and of the latter four, are all confined to this Region. There are also two very THE PALZARCTIC REGION 183 peculiar genera of Carnivores met with in this Region. One of these (Zluropus) is a curious bear-like creature of a white colour, with the ears, shoulders, limbs, and rings round the eyes black, which has only hitherto been obtained in the high mountains of Eastern Tibet (see a Fig. 39.—THE MLUROPUS. (dluropus melanoleucus. ) Fig. 39). The other genus, Zlwrus, sometimes called the Panda, is also found in the same district, but extends southwards into Yunnan. Remains of a closely allied species of this genus have lately been found in the Pliocene deposits of England. This animal is usually 184 THE GEOGRAPHY OF MAMMALS placed in the otherwise strictly New World family Procyonide, which embraces the Raccoons and their allies. The only other carnivorous genus not represented beyond the limits of the Palearctic Region is Meles, containing the familiar European Badger and other species. Fic. 40.—THE WATER-SHREW. (Crossopus fodiens.) The Insectivora are found in considerable numbers in this Region, three genera of Shrews and three of Moles being restricted to it. Among these are the little Water- shrew (Crossopus) found in England, and thence through- out the Palearctic Region as far as the Altai mountains. Crossopus (Fig. 40) is distinguished by having fringes of THE PALAARCTIC REGION 185 stiff hairs along the sides of its feet and tail, which are doubtless of great assistance to it in swimming. Another shrew, Nectogale, found only in Tibet, is still better pro- vided for an aquatic existence, as it has webs between the toes of both fore and hind limbs. The most remarkable endemic representative of the family of Moles in the Palearctic Region is the Desman, Myogale, of which there are two species, one found in the Pyrenees, and the other in the streams and lakes of South-Eastern Russia. The external appearance of these animals, however, resembles much more that of a Shrew than that of a Mole. Considering that the whole of this Region hes within the temperate zone, the number of its Bats is considerable, although they mostly belong to widespread genera. The Monkeys are represented in the Palearctic Region by out- lying species of two genera, Macacus and Semnopithecus, which are both abundant in the Oriental Region. To the former of these belongs the well-known Barbary ape (Macacus inwus), which inhabits the Rock of Gibraltar and the Barbary States of Northern Africa, as well as several species of Eastern Asia. Another Macaque (J. tcheliensis) is enabled by its thickened fur to endure the extremely severe climate of the mountains north of Pekin, and is probably the most northern monkey now living. Section II].—Svup-pIvIsION OF THE PALAARCTIC REGION The sub-divisions of the Palearctic Region recognized by Wallace are four in number; these are—First, the European Sub-region, which includes Europe north of the 186 THE GEOGRAPHY OF MAMMALS Alps and the continuing mountain ranges that form the backbone of the continent; secondly, the Mediterranean Sub-region, which consists of the remainder of Europe, Northern Africa, and Western Asia as far as the borders of the Oriental Region; thirdly, the Siberian Sub-region, which includes not only the country from which it takes its name, but also the whole of the desert region of Central Asia, and reaches as far south as the Himalayas; fourth and last, the Manchurian Sub-region, containing the greater part of China proper and Manchuria together with Japan. These Sub-regions, however, do not appear to represent the true faunal divisions of the Palsarctic Sub-region quite adequately. In the first place, there seems to be a fairly continuous and unchanging fauna extending from the west of Europe all across Siberia and embracing the northern island (at any rate) of Japan. This wide area is still, to a great extent, covered with forest, and was, no doubt, mainly so beset until within comparatively recent times. Again, Wallace’s arrangement divides between two Sub- regions the vast extent of desert country that reaches from the Sahara through Egypt, Arabia, Persia, and Turkestan to Mongolia, which also appears to contain a fairly homogeneous fauna. Wallace’s Manchurian Sub- region, on the other hand, seems to be well established, and to be the most distinctive of all his Sub-regions, but as it only embraces a part of Manchuria, we prefer to call it the Chinese Sub-region. We may, therefore, distinguish three Sub-regions in the Palearctic Region as follows :— 1. The Huropasian Sub-region, containing Europe, the THE PALEZARCTIC REGION 187 whole of Siberia north of the great mountain ranges together with the island of Saghalien, and perhaps, too, the Japanese island of Yezo. In this Sub-region must also be included Asia Minor, the Caucasus, and the Elburz mountains. 2. The Eremian Sub-region, including the north of Africa, Northern Arabia, the greater part of Persia and Afghanistan, and the great desert of Central Asia, ex- tending from the steppes of Southern Russia as far as Manchuria. 3. The Chinese Sub-region, embracing the greater part of China proper, Southern Manchuria, and Japan, and extending westward to Western Tibet and the top of the southern slopes of the Himalayas. ? The boundaries of these Sub-regions will be best under- stood by referring to the accompanying map (Plate VIL, p- 196), in which they are approximately delineated ; but it must be always understood that it is in most cases quite impossible to draw a hard and fast line as the boundary between two adjacent Regions on land. Section IV.—THE EUROPASIAN SUB-REGION The ELuropasian Sub-region contains the great tem- perate forest-area of the Northern Hemisphere. In its western part, at any rate, this has been considerably modified by the hand of man, but in primeval times the forests probably extended almost without break from the Bay of Biscay to Kamtchatka. The Europasian fauna is not very rich; it comprises fifty-seven genera of Mammals, of which four only are 188 THE GEOGRAPHY OF MAMMALS restricted to its boundaries. The endemic forms among the Ungulata are Capreolus, containing the Roe-deer, a single species of which is found throughout the whole extent of the Region; and Rupicapra, the Chamois, found only in the Pyrenees, Alps, Carpathians, and Caucasus. The single endemic rodent is the familiar Dormouse (Muscar- dinus), which is apparently confined to Europe. On the other hand, the little Water-shrew (Crossopus) extends from England to the Altai mountains. There is a considerable number of genera common to the Europasian Sub-region and the Nearctic Region. With the exception, however, of two, the Elk and the Reindeer, these have mostly spread also into the other Palearctic Sub-regions. On the whole the fauna of this Sub-region has little individuality, and calls for very few remarks as to its distinctness. Appended is a list of the genera, drawn up in the same manner as in the previous tables. EUROPASIAN SUB-REGION 3 a a = g | a | A eel Wt) et eae s=ed/et= IE B/2|e|s) Bi save Blwee ee veer igem lage Endemic 2 1 @\) 1 OM Oe et Palearctic Dine Biles pl eats es Oil ew) Ff Old World . bss 4d Sel Salat aes len Holarctic 2 6 AW al 1 Ome Widespread Sl D ly sOa ON taal Os aala, Motall’. LOA AOE oe a 7 i iy) THE PALZARCTIC REGION 189 SECTION V.—THE EREMIAN SUB-REGION The Eremian or Desert Sub-region of the Palarctic Region contains representatives of a larger number of genera than the Kuropasian, and a higher percentage of endemic forms, although even here we do not find so much individual character as in some of the Sub-regions pre- viously treated of. A considerable number of the genera are common to this and the Ethiopian Region, which is, perhaps, not to be wondered at, considering the long land boundary which runs between them. Among the Ungulates only one genus is confined to this Region—the Camel (Camelus), which is now found wild only in certain desert tracts of Central Asia, being else- where known only in a domesticated condition. But remains of animals of this genus have been found in comparatively recent beds both in India and Algeria. Except for this, other indications seem to point to the fact that the Camels must have had their origin in the New World, where they are now represented only by the Lamas. But remains of several allied genera of Camelidx have been met with in the Tertiary beds of North America, where, however, they have been long extinct. Bubalis, which contains the Antelopes usually known as “ Hartebeests,” and Hyrax (the Tree-conies) are common to this Sub-region and the Ethiopian Region. Among the Rodents no less than five genera are confined to the Sub-region, the most remarkable of these being, perhaps, the Jerboas, or Kangaroo-rats, as they are called, from having their hind legs elongated for jumping purposes. The four known genera of Jerboas, which contain a large number of species, are not found 190 THE GEOGRAPHY OF MAMMALS outside the limits of this Sub-region. A fifth endemic genus of rodents is Ellobius, which is thoroughly adapted to a subterranean life, having very short limbs and tail, and rudimentary external ears. The only two known species of this genus are restricted to the Eremian Sub-region. The Eremian Carnivora, as is usually the case with this group, are mostly widespread, and this is also the case with the Bats. The Insectivora are not very numerous, but one genus is peculiar to the Sub-region ; this is Diplomesodon, a little shrew-like animal of terrestrial habits found in the Kirghiz steppes. Another genus, Macroscelides (the Elephant-shrew), though typically Ethiopian, has one species that extends into this Sub-region in Algeria and Tunis. The Monkeys are represented in the Eremian Sub- region by one species only, the well-known Barbary Ape, (Macacus inwus), which is found in Algeria and Morocco. The genera that occur in the Eremian Sub-region are shown in the following table, which may be compared with the corresponding table appended to the Europasian Sub- region :— ) EREMIAN SUB-REGION | | : . S ¢ fS = on Sei ls | Sots Endemic . iM Waovae Glyn eal! On 0 fl Palearctic et ee: OHA sode|t ellis ell 0; 0 if Palearctic and Ethiopian . Persea” liom FO: ae Old World . cy ae eT ea en MF | AOw v2 Holarctic. On. Be) 2000 1/0 4 Widespread . DEN eA SAO ee ees | eta ee Total ay) es: | LI 6 Ss el Gs THE PALZARCTIC REGION 191 SECTION VI.—THE CHINESE SUB-REGION The Chinese Sub-region bears the same relation to the Oriental Region as the Desert Sub-region does to the Ethiopian Region, and the number of genera of Mammals that are common to it and the Oriental Region is considerable. On the whole, too, this is the most specialized of all the Palearctic Sub-regions; six genera out of a total number of sixty being endemic, and several others only just crossing its borders. This Sub-region contains within its limits the highest tableland on the face of the globe, that of Tibet, the zoology of which is not so well known as it should be, owing to the persistent exclusion of European travellers from its limits. What knowledge we have of the Tibetan fauna relates chiefly to the larger animals, and among these are several very in- teresting and peculiar forms. It is, therefore, probable that when more about this Region is known, many novelties will be discovered among the smaller animals as well. The Ungulates of the Chinese Sub-region show no genera which are absolutely confined to it, but a curious little deer with short straight antlers (Zlaphodus) is highly characteristic of it, although it extends into the outskirts of the Oriental Region. Another peculiar genus is the Tibetan antelope (Pantholops), well known to Indian sportsmen, which is found only at great heights on the Tibetan plateau. A third genus, also of great interest, is the Takin (Budorcas), a large bovine animal with horns somewhat resembling those of the 192 THE GEOGRAPHY OF MAMMALS South African Gnu; this is also found only among the higher mountain ranges, and is probably confined to the Sub-region. Among the Rodents of this Sub-region there are two genera peculiar. One of these is Hupetawrus, a flying squirrel, which, unlike all others of the same group, lives in a part of the world practically devoid of forests. It is as yet known only from the extreme north-western part of the Indian Empire, but doubtless has a wider range. The second endemic genus, Typhlomys, has been formed for the reception of a curious, almost blind Mouse of Northern China. Among the Carnivora of the Chinese Sub-region the only genus of very special interest 1s dlwropus, which has already been mentioned in the general account of the Region. The Insectivora of this Sub-region are numerous, comprising as they do ten genera, of which three are endemic. These are all of them Shrews, two of which, Chimarrhogale and Nectogale, are aquatic forms with webbed toes, while the third, Anwrosorex, is probably fossorial. All the Bats belong to fairly wide-spread forms. Finally, there are two genera of Monkeys represented in this Sub-region, Macacus and Semnopithecus, but these forms belong strictly to the Oriental Region. Below is a table of the genera of the Sub-region, arranged as in the foregoing Sub-regions, which shows what a con- siderable number of forms are common to this and the Oriental Region. THE PALZARCTIC REGION 193 CHINESE SUB-REGION lee ee de chad Endemic. O} -Bap wy Bek Oe 6 Palearctic ees Pepe ocd ecbatee Ons Onn Opa ped Palearctic and Oriental et: ij) S| 8 A ea pa ey Old World . ese Ae On D2) Sool Oh i T4 | Holarctic Oh ae OM, ee eno a | Widespread . lieatyle onl: 8Galh) On rl Ontits | Total . . L137) 1 10 10") 2 “163 Section VII.—TuHe Past History oF THE PALZARCTIC MAMMAL-FAUNA Although the paleontological history of Europe, so far as it has been worked out, has been very thoroughly in- vestigated, our knowledge of its extinct mammals, at any rate, is not to be compared with that which has been acquired in the Nearctic Region. This is probably due, to a great extent, to the comparative rarity on this side of the Atlantic of fresh-water lake deposits, the examination of which, in North America, has produced such astonishing results. Passing over the Mesozoic Mammals, which throw very little light on any of the problems involved in the present case, we find in the earliest Hocene beds scanty remains of a fauna containing hardly any members of the existing orders of Mammals. In their place is a series of forms closely resembling one another in possessing five-toed and N 194 THE GEOGRAPHY OF MAMMALS plantigrade extremities, furnished with neither hoofs nor claws, but with structures somewhat intermediate between the two. Nevertheless, even among these primitive mam- mals, it 1s possible to recognize the germs of the marked characters which at the present day separate the various Orders. In North America, in beds of the corresponding age, a much more ample stock of remains of a similar fauna is met with. Later on, in the Upper Eocene beds a much larger number of Mammals appears, this fauna containing at least a hundred genera, most of them of large size, whereas to-day the European Mammal-fauna consists only of fifty-four genera, and of these more than half are of small size. At this epoch slight distinctions between the European and American forms. begin to appear, showing that even at these early times there was a commencing separation between the two great continents. In the earlier part of the Miocene age, so far as we know, no very great changes take place, but at the end of Miocene time we find in several localities wonderful as- semblages of fossil Mammals in great abundance and in an excellent state of preservation, which enable us to make a better comparison. Such localities have been discovered at Pikermi in Greece, in the island of Samos in the Aigean Sea, at Maragha in Persia, and, perhaps the most important of all of them, in the Sivalik Hills at the southern base of the Himalayas. This fauna bears a close resemblance to that of the Ethiopian Region in the present state, especially as regards the presence of Giraffes, Gazelles, and other Ungulates. North of the Alps this fauna, although represented, is not nearly so rich, many of the Antelopes and Giraffes being absent and being replaced by various forms of Deer THE PALZARCTIC REGION 195 (Cervidx), which now commence to be very much more prominent. In the true Sivalik fauna of India there are a good many types which have never yet been found in Europe; such, for instance, as the Camels, which are specially characteristic of the American Tertiary strata. Furthermore, there are found, in the American formations of this age, a large number of forms, such as Bos, Equus, Hippopotamus, and Ursus, which do not appear at all in Europe until the later Pliocene times. When the Pliocene times arrive, we begin to find a preponderating number of still existing genera present in the fossil beds, although the greater number of them have, at the present epoch, retreated southwards into the Oriental and Ethiopian Regions. This southward migra- tion seems to have gone on throughout the Pliocene period, and was probably occasioned by the increasing cold caused by the gradual advent of the great Ice-age, which now began to make itself felt over the whole of the northern part of the globe. Finally, during the Glacial period the fauna assumed nearly its present form, containing large numbers of species that still survive. At this epoch, too, a connection appears to have been formed between the Old and New Worlds in the neighbourhood of Behring Strait, by means of which an interchange of animals took place, and resulted in occasioning the similarity which forms so marked a feature on a comparison of the Nearctic and Palearctic faunas. It is this similarity that has caused certain writers on geographical distribution to unite the Palearctic and Nearctic Regions into one, whereas, as a matter of fact, palzontological evidence seems to show that, out of all the 196 THE GEOGRAPHY OF MAMMALS four Regions embraced under the term “ Arctogza,” the North American or Nearctic Region was the first to be separated from the main mass, and that the similarity is a comparatively modern element in the character of the two faunas. Te MAP OF THE PALAEARCTIC REGION SHOWING ITS DIVISION INTO 3 SUB-REGIONS. CHAPTER VIII MARINE MAMMALS (Puate VIII., p. 216) SEecTIon I.—IntTRopDUcTORY REMARKS Most of the recent writers on geographical distribution have confined their attention to terrestrial Mammals, or at any rate have but casually alluded to the marine groups of that Class. The seven previous chapters having been devoted to the terrestrial Mammals, it is proposed now to examine the principal facts connected with the distribution over the world’s surface of the Marine or aquatic members of the Class. Aquatic Mammals which pass their lives entirely, or for the greater part, in the water are, of course, subject to very different laws of distribution from the terrestrial forms. As regards aquatic Mammals, land is of course an impassable barrier to their extension, and, subject to restrictions in certain cases, water offers them a free passage. Just the opposite is the case with the terrestrial Mammals, to which in most cases land offers a free passage, while seas and rivers restrain the extension of their ranges. The groups of aquatic Mammals that are represented on the earth’s surface at the present time are three in number, viz.: (1) the Sub-order of the Carnivora, contain- ing the Seals and their _ generally called the Pinni- 198 THE GEOGRAPHY OF MAMMALS peds, which are semi-aquatic ; (2) the Sirenia, or Manatees, which are mainly aquatic; and (3) the Cetacea, or Whales, which never leave the water, and are wholly aquatic. We will first consider briefly the principal representatives of these three groups, following nearly the arrangement of them employed in Flower and Lydekker’s “Mammals Living and Extinct.” Section IJ.—DIsTRIBUTION OF PINNIPEDS The Pinnipeds, which we will take first, comprise three distinct families—the Otariidx, the Trichechidx, and the Phocide. Beginning with the Otariidz, or Eared Seals, commonly known as Sea-lions and Sea-bears, we find the greater number of the species confined to the South Polar Ocean, where they pass most of their time at sea, but, as is well known, resort to the land at certain seasons for breeding purposes. In the Atlantic Ocean, so far as is known, the Eared Seals have never been ascertained to occur much further north than the estuary of the La Plata on the American coast, where the Patagonian Sea-lion (Fig. 41, p. 199) is met with, and the vicinity of the Cape on the African coast, where Otaria pusilla is found. But in the Pacifig, on the contrary, three distinct species of Otaria are distributed all over the northern portion of that ocean. Two species of Sea-lions are also met with in the Galapagos, and they likewise occur on the coasts of Peru and Chili. I think therefore we may assume that Otaria was origin- ally an Antarctic form, but has travelled northwards along the West-American coast and is now firmly established in the North Pacific. In a parallel way in the Class of Birds, MARINE MAMMALS 199 the Albatrosses (Diomedea), which is essentially a group of the Antarctic Seas, are represented by three distinct species in the North Pacific. The second family of the marine Carnivora, on the other hand, the Walruses (T'richechidz), are entirely Arctic in their distribution—one species (Trichechus rosmarus) Fig, 41.—TH& PATAGONIAN SBA-LION. (Otaria jubata. ) [P. Z. S. 1866, p. 80.] (see Fig. 42, p. 200) being peculiar to the North Atlantic, a second nearly allied species (7. obesus) takes its place in the Northern Pacific. The third family of Pinnipeds is more numerous and varied, both in genera and species, than the two preceding, and has a more extended range. The Seals, Phocide, 200 THE GEOGRAPHY OF MAMMALS embracing about nine different generic forms, are most numerous in the Arctic and Antarctic Seas, but are also feebly represented in some intermediate localities. Be- ginning with the North Atlantic, we find several species of Phoca (Fig. 43, p. 201), inhabiting various parts of this area, and the Grey Seal (Halichwrus) and the Bladder-Seal Fig. 42.—_THE ATLANTIC WALRUS. (Trichechus rosmarus.) (Cystophora) exclusively confined to it. In the North Pacific all the four true Seals belong to the genus Phoca, and three of them are identical with the North Atlantic species; but when we descend as far south as the Gulf of California on the American coast we meet with a species of Sea-elephant (Macrorhinus) which, like Otaria, has no BY Ald ad MARINE MAMMALS 201 doubt penetrated thus far from its ancestral abode in the Antarctic Ocean. Returning to the central Atlantic we find two species of Seals inhabiting these waters, both belonging to the same genus, Monachus. One of these (M. albiventer) inhabits the Mediterranean and the adjoining coasts of the Atlantic, Fig. 43.—THE COMMON SEAL. (Phoca vitulina. ) while the other (/. tropicalis) is in these days restricted to some of the smaller and less known islands of the West Indies. The Phocide of the Antarctic Ocean all belong to genera distinct from the Arctic forms and more nearly allied to Monachus, the Seal of the Mid-Atlantic. They are of four species, belonging to as many genera: Ogmorhinus, 202 THE GEOGRAPHY OF MAMMALS Lobodon, Leptonychotes, and Ommatophoca. Besides these the Sea-elephant of the whalers (Macrorhinus) is essentially an Antarctic form, though now nearly extinct there, after Fic. 44.—THE MANATEE, (Manatus americanus.) long persecution by man, But, as already noted, it extends. or has in former days extended, far up the west coast of America, and is still occasionally found on the coast of California. j MARINE MAMMALS 203 —S = Section [III.—DIstripuTion oF SiRENIANS Only two forms of Sirenians are at the present time existing on the earth’s surface—the Manatee (Manatus) Fic. 45.—THE DUGONG. (Halicore dugony. ) and the Dugong (Halicore)—each representing a distinct family of the Order. The Manatee (Fig. 44, p. 202) is an inhabitant of the coasts and estuaries of both sides of the middle Atlantic Ocean—one species (Manatus senegalensis) occurring on the African shores, and another (M. ameri- 204 THE GEOGRAPHY OF MAMMALS canus) on the South American coast and in the Antilles. A third species (I. inwnguis), so far as we know at present, is found only in fresh water high up the Amazon. The Dugong (Halicore) (Fig. 45, p. 203) is distributed from East Africa, along the shores of the Indian Ocean and its islands, to North Australia. Three species of this genus have been established—Halicore tabernaculi from the Red Sea, H. dugong from the Indian Ocean, and 4H. australis from Australia; but it is doubtful how far these forms are actually distinguishable. Fig. 46.—THE RIGHT WHALE. (Balena mysticetus. ) (Flower and Lyd. Mamm., p. 236.] Besides Manatus and Halicore, a third quite distinct form of Sirenian was formerly an inhabitant of the North Pacific. This was Steller’s Sea-cow (Rhytina stellerv), by far the largest animal of the group, which was extermi- nated by human agency about 1768. Fortunately recent researches in Behring’s Island have been successful in supplying specimens of its skeleton for our principal museums, and Steller, its discoverer, left to posterity a good account of its habits and anatomy. MARINE MAMMALS 205 DM Section IV.—DIsTRIBUTION OF CETACEAN Adopting the recognized division of the Cetaceans into two sub-orders, Mystacoceti and Odontoceti, according as to whether their mouths are furnished with baleen (“ whale- bone”) or teeth, we will first consider the True or Whale- bone Whales, which consist of a single family Balenidz, usually divided into five genera: Balzna, Neobalena, Rhachianectes, Megaptera, and Balznoptera. Of these, Fic. 47.—THE SPERM WHALE. (Physeter macrocephalus. [Flower and Lyd. Mamm., p. 249.] Balzna (Fig. 46, p. 204), Megaptera, and Balznoptera are almost cosmopolitan—species of them, whether distinct or not is at present more or less uncertain, being met with in nearly every part of the ocean. But Rhachianectes has as yet been ascertained to occur only in the Northern Pacific, and Neobalzna in the South Polar Ocean, so that we have in these cases two well-marked local types to deal with. The Toothed Whales (Odontoceti) are more diversified than the preceding group, and are usually held to em- brace at least four existing families besides extinct forms. 206 THE GEOGRAPHY OF MAMMALS The first family, containing the Physeteride or Sperm- Whales, consists of at least six genera, Physeter (Fig. 47, p. 205), Cogia, Hyperoodon, Ziphius, Mesoplodon, and Berardius). Physeter and Cogia are inhabitants of the whole oceanic area between the tropics, extending in certain localities some way beyond them. Hyperoodon is confined to the North Atlantic. Ziphius has an extensive range, and has been found in nearly every part of the ocean. Mesoplodon is also widely distributed, but is apparently more abundant in the Southern Hemisphere. Berardius, Fic. 48.—THE SEE-SEE. (Platanista gangetica.) [Flower and Lyd. Mamm., p. 255. ] however, so far as we know at present, is restricted to the Pacific Ocean. The second (existing) family of Toothed Whales con- tains only the Platanistidx, or Freshwater Dolphins, which although, in some cases, at the present day entirely fluvia- tile, must probably have descended from oceanic forms. The three known genera are Platanista of the Ganges and Indus (Fig. 48), Znia of the river Amazon, and Pontoporia of the river La Plata; the last form making 1 Sir William Flower (‘‘ Whales, Past and Present,’ Proc. Roy. Inst., x., p. 360, 1883) rather favours the idea of a freshwater origin of the Cetaceans. MARINE MAMMALS 207 a connecting link between the two preceding genera and the marine Dolphins. The third family of Toothed Whales, containing the Dolphins, Delphinidz, is very numerous in species and embraces at least fifteen or sixteen genera, of which the Common Dolphin (Fig. 49) is a good example. But in spite of the efforts of Mr. True, who has recently given us an excellent summary of our present knowledge of them,! both the genera and species of Delphinide are still so imperfectly understood that not much can be said about their geographical distribution. Most of the forms Fig. 49.—THE COMMON DOLPHIN. (Delphinus delphis.) [Flower and Lyd. Mamm., p. 271.] appear to be very widely distributed, but it may be said generally that Dolphins are most abundant in the inter- tropical seas and less plentiful both to the north and south of them. There are, however, two forms that are exclusively inhabitants of the Northern Oceans. These are the very remarkable Narwhal (Monodon), in which the male is furnished with a single enormous horn-like tusk, and the Beluga, or White Whale (Delphinapterus), closely allied 1 See ‘A Review of the Family Delphinide,” by Frederick W. True. Bull. U.S. Nat. Mus., No. 36 (Washington, U.S., 1889). 208 THE GEOGRAPHY OF MAMMALS to the Narwhal in many points of its general structure (Fig. 50). These may be looked upon as quite isolated Fia. 50.—THE BELUGA. (Delphinapterus leucas.) [Flower and Lyd. Mamm., p. 262.] forms characteristic of the Arctic portion of the Atlantic and Pacific. Section V.—DIvISION OF THE MARINE AREA OF THE GLOBE INTO SEA-REGIONS From what has been already said, it will be evident that although many of the marine mammals have a wide distribution, others are very definitely localized; and a study of the latter will enable us to divide the oceanic portion of the globe into six Sea-regions, corresponding to a certain extent with the six Land-regions already discussed. It is proposed to name these Sea-regions, which are shown in the map (Plate VIII, p. 216), as follows :— (1) The North Atlantic Sea-region, or Arctatlantis (apxtos and ’Ar)davtis = the daughter of Atlas), consisting of the northern portion of the Atlantic down to about 40° N. lat. (2) The Mid-Atlantic Sea-reqion, or Mesatlantis (wéoos and ’Ardavtis), consisting of the middle portion of the Atlantic down to about the Tropic of Capricorn. MARINE MAMMALS 209 (3) The Indian Sea-region, or Indopelagia ("Ivésos and 7réXayos), containing the Indian Ocean down to about the same degree of S. lat., and extending from the coast of Africa on the west to Australia and the great Oriental islands on the east. (4) The Worth Pacific Sea-region, or Arctirenia (dpxros and eipyvn = pax), containing the northern portion of the Pacific Ocean down to about the Tropic of Cancer. (5) The Mid-Pacific Sea-region, or Mesirenia (uécos and edpyvn), containing the inter-tropical portion of the Pacific Ocean ; and finally, (6) The Southern Sea-region, or Notopelagia (vortos and zéAayos), containing the whole of the South Polar Ocean all round the globe south of the above-mentioned limits. We will now proceed to consider shortly the char- acteristic mammals of these six Sea-regions. SEcTION VI.—THE NortH ATLANTIC SEA-REGION, OR ARCTATLANTIS Amongst the Pinnipeds two well-marked generic forms, the Grey Seal (Halicherus) and the Bladder-Seal (Cysto- phora), are exclusively confined to Arctatlantis. The True Seals (Phoca) and the Walrus (Trichechus) are found in this region and in Arctirenia; and of the former genus three species (P. vitulina, P. grenlandica, and P. barbata) are actually common to both these Sea-regions, while the Walruses (Trichechus rosmarus and T. obesus) of the two Sea-regions are perhaps somewhat doubtfully distinguish- able. It may be easily understood how this has come to pass, O 210 THE GEOGRAPHY OF MAMMALS because the Seals and Walrus in the course of time, during unusually mild summers, may have extended themselves along the north coast of the American continent into the Northern Pacific. But Arctirenia, as we shall presently show, is markedly distinguishable from Arctatlantis by the presence of Hared Seals (Otaria), which are utterly unknown in the whole of the Atlantic area. Otaria is in fact, as regards Arctatlantis, a lipomorph. | The Sirenians are entirely absent from the North Atlantic and constitute another lipomorph of that area. Coming to the Whales, we find the Mystacoceti well represented in the North Atlantic by Balena, Megaptera, and Balxnoptera; but of these the two latter are almost universally distributed over the ocean, and Balxna recurs again in the North Pacific as well as in more southern latitudes, so that there is no genus of Whalebone Whales peculiar to Arctatlantis, Proceeding to the Odontoceti, the case is different. Amongst the Physeteridey, Hyperoodon is confined to Arctatlantis. Arctatlantis therefore may be said to be well characterized by the possession of at least three genera of marine mammals not found elsewhere, viz., Halichwrus, Cystophora, and Hyperoodon, and by the complete absence of the Eared Seals (Otariide). Section VII—Tae Mip-ATLANTIC SEA-REGION, oR MESATLANTIS Mesatlantis has certainly not so many forms of marine mammals confined to its area as Arctatlantis, but there seem to be good grounds for its separation. As we descend MARINE MAMMALS AMAL towards the tropics the true Seals (Phocinx), which are constituted to live in colder water, gradually fall off in number, and in Mesatlantis are no longer met with. But in their place we find the genus Monachus, or Monk-Seal, restricted to Mesatlantis, one species (M. albiventer) occur- ring in the Mediterranean and on the North African coast, and a second (M. tropicalis) being found in the West Indies. Mesatlantis is likewise the true home of the well- marked Sirenian genus Manatus, one species of which (MV. americanus) frequents the coast of America and another (M. senegalensis) that of Africa. As regards the Cetaceans, we are not at present able to say that Mesatlantis, although well furnished with many generic types of this Order, has any one peculiar to it. We must therefore rest content with assigning two genera of marine mammals, Monachus and Manatus, as charac- teristic forms or topomorphs of the Sea-mammal-life of Mesatlantis. Section VIJI.—TuHer InpIAn SEA-REGION, OR INDOPELAGIA The marine Carnivora, so far as we know, are entirely foreign to Indopelagia, but the Sirenians are well repre- sented by the Dugong (Halicore), which pervades all its northern coasts from North Australia to India and the Red Sea and down the African coast to the confines of British East Africa. Whether the species of Halicore found at different points within this area are the same or different is still a matter of discussion, but there can be no doubt that Halicore is an exclusive inhabitant of 2r2 THE GEOGRAPHY OF MAMMALS Indopelagia. As regards the Whales of Indopelagia, we know that Physeter, Cogia, and Ziphius, and numerous forms of Delphinide occur there, but are not aware of any Cetacean that is entirely restricted to this Sea-region. SecTion [X.—THEr Nort Paciric SEA-REGION, OR ARCTIRENIA As was pointed out when speaking of Arctatlantis, Arctirenia has one genus of Phocide (Phoca) in common with the North Atlantic, and three of the species of this genus appear to be actually identical in these two Sea- regions, whilst a fourth Phoca (P. fasciata) is only found in the North Pacific. The Walrus (T7richechus) is again a form of marine mammals common to both the great northern Sea-regions. But the feature of Pinnipedian life that absolutely distinguishes Arctirenia from Arctatlantis is the presence in the former of three (if not four) well- marked species of the Eared Seals (Otariidz), which are absolutely unknown in the vast extent of the Atlantic down at least to 30° S. lat. Arctirenia has unfortunately lost its Sirenian, Steller’s Sea-cow (Rhytina stelleri), the largest modern represen- tative of this formerly prevalent group, which since the days of the Pleistocene has greatly diminished in numbers, but I think we may still treat Rhytina as one of the characteristic forms of the Arctirenian Sea-region. The North Pacific is also, even at the present day, the sole possessor of a remarkable genus of Whalebone Whales which combines the long head and elongate form of Balz- onptera with the absence of the dorsal fin of Balena. MARINE MAMMALS 2d This is the Grey Whale, Rhachianectes glaucus of Cope, which, in these days, is confined to the North Pacific, and does not range farther south than the 20th parallel in that ocean. At the same time it should be stated that indications have been discovered that a nearly allied form existed in the Atlantic in previous geological ages, though this is by no means certain. Besides Rhachianectes, Balena, Megaptera, and Balxnoptera are all represented in the North Pacific, and also many species of Delphinide of which little is at present known. But Rhytina and Rhachianectes are the only genera of Marine Mammals absolutely confined to Arctirenia. SECTION X.—TuHE Mip-Paciric SEA-REGION, oR MESIRENIA The Eared Seals, Otaria, must have necessarily passed through Mesirenia in their passage from south to north, though the only record of their actual presence in the central part of the Pacific is, so far as we know, the recent discovery of them in the Galapagos. It should be stated, however, that Tschudi records the occurrence of two species of Otaria on the islands of the coast of Peru, and that in 1802 Humboldt met with an Eared Seal on the Island of San Lorenzo, in the Bay of Callao, which is only some 12° south of the Equator. Like Otaria, the Sea-elephant (Macrorhinus) has appa- rently in former ages travelled up the South American shores and established itself as far north on the coast of California as about 34° N. lat. The Californian Sea- elephant has been discriminated by Gill as a distinct 214 THE GEOGRAPHY OF MAMMALS species (Macrorhinus angustirostris), but its differences from the southern form (M. leoninus) seem to be but trifling. As regards the Cetaceans of Mesirenia, our information is at present very imperfect, and there is little to say except that species of Megaptera, Balenoptera, Physeter, Cogia, and Ziphius certainly occur there, besides many repre- sentatives of the widely spread Delphinide. SECTION X].—THE SOUTHERN SEA-REGION, OR NOTOPELAGIA The wide ocean which surrounds the Southern Pole on every side, and extends up to 40° S. lat., seems to present, as regards its marine mammals, a nearly homogeneous fauna, which we will now briefly consider. In the first place it contains representatives of four genera of true Phocide — Ogmorhinus, Lobodon, Leptonychotes, and Ommatophoca, which are peculiar to the southern seas, and are quite distinct from all their northern representa- tives in the Arctic Ocean. The Sea-elephant, Macrorhinus, is also a denizen of Notopelagia, though, as we have already seen, it has wandered north along the South American coast far into Mesirenia. Like Macrorhinus, Otaria also, containing the group of Eared Seals, appears to have been originally an Antarctic group, and the greater number of its species are still found in the Southern Ocean. But the Otariz have travelled still further north than Macrorhinus, and three, if not four, species are, as already stated, in these days well- established inhabitants of Arctirenia. MARINE MAMMALS 215 The Sirenians are absent from WNotopelagia, but Cetaceans of every kind are abundant. Besides one or more representatives of the true Whalebone Whale (Balzna), Notopelagia has a smaller representative of the group (Neobalena) entirely restricted to its area. It has also representatives of Megaptera and Balxnoptera, though it is doubtful how far they are even specifically distinct from some of their northern representatives. Among the Toothed Whales (Odontoceti) we find a large Ziphioid form, Berardius, restricted to the Pacific area, while Ziphius and Mesoplodon also occur there. The Dolphins (Delphinidz) are likewise numerous, and present some distinct species, but not, so far as our present knowledge extends, any generic forms that do not occur elsewhere. But Notopelagia is sufficiently distinguished from all the five more northern Sea-regions by possessing four genera of Seals and two of Cetaceans entirely restricted to its area. SEcTION XII.—ConcLUSIONS It has therefore been shown that, for the geography of marine mammals, the ocean may be conveniently divided into six Sea-regions (Plate VIII. p. 216), which are as follows :— I. Regio Arctatlantica, characterized by its Seals (Phocinx), of which two genera, Halichewrus and Cysto- phora, are peculiar, whilst Phoca is common to it and Arctirenia; by the absence of Sirenians; and by the possession of a peculiar genus of Cetaceans (Hyperoodon). II. Regio Mesatlantica, sole possessor of the Monk- 216 THE GEOGRAPHY OF MAMMALS Seals, Monachus, amongst the Pinnipeds, and of the Sirenian genus Manatus. Ill. Regio Indopelagica, characterized by the presence of the Sirenian Halicore and by the absence of Pinnipeds. IV. Regio Arctirenica, with Phoca like the Regio Arctatlantica, but having Otaria also; the home of the (now extinct) Sirenian Rhytina and of the endemic Cetacean Rhachianectes. V. Regio Mesirenica, without true Seals (Phocinx), but having Oteria and Macrorhinus from the south; no Sirenian being known there. VI. Regio Notopelagica, characterized by four endemic genera of Phocidx, and by the presence of many Otarie ; without Sirenians, but with two endemic forms of Cetaceans (Neobalena and Berardius). In conclusion, attention may be called to some of the more remarkable points in the general distribution of the marine mammals and to their apparent significance. In the first place it is evident that the Pacific has much more in common with the Notopelagian Region than the Atlantic. Otaria and Macrorhinus, quite unknown in the Atlantic, extend themselves to the northern ex- tremity of the Pacific, the former pervading that ocean up to Behring’s Strait, and the latter reaching to the Californian coast. It follows that in former ages there must have been some barrier in the Atlantic which did not exist in the Pacific to stop their progress northwards. The only barrier one can imagine that would have effected this must have been a land uniting South America and Africa, across which they could not travel. Adopting this hypothesis, we have at the same time an explanation of the presence of the Manatee on both the American SNOIQSYH-VaAS 9 SHL ONIMOHS € ‘MIA ALY 1d tophora peroodon iT] ree NTIS - Monae us _ <5 fe chi a | : Sa Eaaeee8 i | BO 10 cenit | Serene 80 60 £0 ee Otaria Macrorhi OUTLINE MAP OF THE WORL|2 SHOWING THE 6 SEA-REGIONS. PLATE VIII. cy i) ey Sf. Lei << Cy . —_- ah Mm gle. . fie? ee ees M 3 a eo — a oie ss —— ne = ‘ ta ‘ - = 4 Fewirruiie | 1 } jase Le camden feet tpg irs i ‘ \ : cS 7 ‘ « ov oe) whe fe oH veciviry “ Be ceyreerean se sapiens Ss ns ere ‘ ; a" 4 aot 258 30 ene wy _ a aay rl ot Fle &u 4 * x sid Yas \ . / me MARINE MAMMALS 7 and African coasts. The Manatee could hardly live to cross the Atlantic. It is only found close to the coast, in estuaries and rivers, where it browses on sea-weeds and other vegetable food in shallow water. How did it travel from America to Africa (or vice versd), unless there were a continuous shore-line between them? The same may be said of the Monk-Seal (Monachus), of which one species lives in the Mediterranean and on the African coast and islands, and another in the West Indies. We can hardly believe that these creatures could easily traverse the whole Atlantic. The hypothesis of a former barrier of land between Africa and America, which we know is supported by other facts of distribution,’ would alone ex- plain the difticulty. On the other hand, in the Pacific we find no such break between the north and south. The aquatic mammals of Notopelagia have evidently had free access to the whole of the Pacific for a long period, and have well availed them- selves of this facility. Again, while the great Southern Ocean exhibits a considerable uniformity of marine mammalian life, we see the Northern waters divided into two distinctly recogniz- able Regions by the interposed masses of land. All these facts, with the one exception of the supposed Atlantic barrier, would tend in favour of the now generally ac- cepted doctrine that the principal masses of land and water are not of modern origin, but have existed mainly in their present shapes throughout all ages. 1 Cf. Wallace, Geogr. Distrib., vol. i., p. 156. CHAPTER IX DISTRIBUTION OF MONKEYS AND LEMURS Section I.—IntTRopucToRY REMARKS HaviInGc now completed the discussion of the great Zoological Regions of terrestrial and marine mammals and their leading peculiarities, it is proposed to take the subject in another way, and considering the Orders, or great primary groups of mammals, one after the other, to sketch out the mode in which their leading forms are distributed over the surface of the world. We shall then see whether the conclusions thus arrived at appear to lead to similar results to those attained by making geographical divisions our primary subject of study. In doing this it will be more convenient to take the most highly organized groups of mammals first, and to descend gradually to the lowest, revers- ing the arrangement used in the previous chapters of this work. The divisions and names now employed are taken from the last edition of the “List of Ver- tebrated Animals in the Zoological Society’s Gardens,” (1896), but do not differ materially from those used in Flower and Lydekker’s “Mammals Living and Ex- tinct,” which we have hitherto mainly followed. These DISTRIBUTION OF MONKEYS AND LEMURS 219 divisions, commonly called “Orders,” are fourteen * in number, as follows :— I. QUADRUMANA. VIII. PRoposcipEa. Il, LemMuREs. IX. UNGULATA. III. Carnivora. X. CETACEA. IV. INSECTIVORA. XI. SIRENIA. V. CHIROPTERA. XII. EpEnNTATA. VI. RopENTIA. XIII. MARsupiatia. VII. Hyracess. XIV. MonotrEemMata. SEcTION IJ.—GENERAL DISTRIBUTION OF THE QUADRUMANA The Order of Quadrumana, or Monkeys, of which about 200 species are now recognized, is generally divided by zoologists into four families: (1) the Simiidw or Apes, (2) the Cercopithecide or Old-World Monkeys, (3) the Cebidx or New-World Monkeys, and (4) the Hapalide or Marmosets. Of these families the two first are exclusively inhabitants of the tropical or sub-tropical districts of the Old World, while the two last are as severely restricted to the hotter portion of the New World, and form, in fact, two of the most characteristic groups of the Neotropical Region. We thus see that the division of the Quadrumana into families according to their structure is in complete accordance with the distinctness of the geographical areas in which they are found. The two first and highest families of Monkeys, it should be remarked, are much more nearly allied to one another than they are to the two families which inhabit the New World. They are sometimes called “Catarrhines,’ on account of the narrowness of the nasal septum and the consequent downward direction of the nostrils. They all 220 THE GEOGRAPHY OF MAMMALS agree together in possessing the same number of teeth and the same general structure of these organs as Man, who, if considered only from a material point of view, must certainly be referred to the same group of animals. On the other hand, the two families of Quadrumana belonging to the New World have each of them a dentition peculiar to themselves and different from that of the Catarrhine Monkeys, to which they are inferior in every point of organization. The first of these two families, the Cebidx, have been designated “ Platyrrhines,” in reference to the broad septum which separates the nostrils from each other, and thus distinguishes them from the Catarrhine. Monkeys of the Old World. The second family of New World Monkeys, the Marmosets, are unquestionably the lowest of the Quadrumana, and have some superficial resemblance to the Squirrels and other Rodents. Secrion III.—DuistripuTIon oF THE OLD WorLD MoNnkKEYS The Catarrhine Monkeys, as already stated, are re- stricted to the Old World, and in present days, as we shall see, mainly to the tropics, though a few species are found farther north, and other forms, now extinct, have left their remains in the tertiary and _ post-tertiary formations of more temperate countries. The Catarrhine Monkeys form two families, the Simiid# or Man-like Apes, and the Cercopithecidxe or Monkeys proper. The Man-like Apes, in the present con- dition of the world’s surface, are confined entirely to the equatorial Regions of Africa and Asia. They are usually rp DISTRIBUTION OF MONKEYS AND LEMURS 22% referred to three genera: (1) Anthropopithecus, which em- braces the Black Apes of tropical Africa, (2) Simia, which contains the Brown Apes of tropical Asia, and (3) Hylobates, which contains the Gibbons, likewise restricted to tropical Asia. It has been a subject of much discussion among zoologists as to which of these three forms of Simiine life comes nearest to man in structure. Some have advo- cated the claims of the Orang to this distinction, but it is now generally held that his black cousins of the Ethiopian Region ought to be placed at the head of the series. It has likewise been maintained by some authorities that in certain respects the Gibbons (Hylobates) come nearer to man than either of the above-mentioned forms, but this opinion has not met with general support. We will therefore commence our survey of the distribution of the Anthropoid Apes with their African representatives, the Chimpanzee and the Gorilla. The Chimpanzee (Anthropopithecus troglodytes) is certainly widely spread over tropical Africa, but we have not as yet acquired sufficient details as to the exact limits of its distribution. Nor are naturalists at all agreed as to whether one or more species are comprehended under the name of Chimpanzee, though it is generally allowed that the Bald-headed Chimpanzee (A. calvus), a well-known specimen of which lived in the Zoological Society’s Gardens for many years, must be considered distinct from the ordinary A. troglodytes. But in examining the distribu- tion of the Chimpanzee it is hardly necessary to seek to determine this vexed question more nearly. Commencing on the western coast of Africa, the range of the Chimpanzee appears to begin with the wooded 222 THE GEOGRAPHY OF MAMMALS districts somewhat to the south of the river Gambia, and to continue all down the coast-region into Angola, south of the Congo. Here the dry and open country of south-west Africa commences, and, as the Chimpanzee is essentially a forest animal, it could not be expected to be met with farther in this direction. The Chimpanzee in its inland range appears to extend over the whole wood- district of the great Congo valley, and in the Niam-Niam country, perhaps, passes over into the basin of the Nile. It was heard of by Livingstone in the forests on the western side of Lake Tanganyika where it is said to be called “Soko,” and the late Emin Pasha sent to England the skull of a Chimpanzee stated to have been shot by himself near Lake Albert Nyanza, which seems hardly to differ from the West African form. Whether Emin Pasha’s assertion that the Chimpanzee occurs in Uganda and Unyoro up to 32° E. lat. is correct is very doubtful. It seems to have been made on native authority, and has not, so far as we are aware, been corroborated by more recent travellers. But as regards its existence in the Niam-Niam country we have Emin Pasha’s express statement that he re- ceived a living specimen of the Chimpanzee as a present from one of the chiefs of that nation. The Gorilla (A. gorilla), which by some authors is referred to a different genus from the Chimpanzee although its general structure is not materially different, is confined to a very much more limited area than that of its smaller brother (A. troglodytes). So far as we know at present, it seems to be entirely restricted to Gaboon and the ad- jacent districts of French Congo. Whether the extra- ordinary accounts formerly given of the strength and 1 “min Pacha in Central Africa,” p. 355. DISTRIBUTION OF MONKEYS AND LEMURS 223 ferocity of the Gorilla are based on fact seems to be still quite doubtful. The few specimens of this animal that have reached Europe alive have not given much support to the stories told about it. The genus Anthropopithecus, therefore, with its two, or perhaps three, species, may be regarded as one of the most characteristic forms of the mammal-life of the Ethiopian Region to which it is absolutely confined. Passing on to the next genus of Anthropoid Apes, we find that the Orangs (Simia) are in like manner one of the most characteristic mammal-forms of the Indian Region, to which they exclusively belong. As in the case of the Chimpanzee, there has been the same difference of opinion among naturalists as to whether there is only one or several species of Orangs. The Orang is found only in the dense forests of Borneo and Sumatra. The forms met with in these islands respectively have been treated as specifically distinct, and the Bornean form has also been divided into several species. But, on the whole, the prevalent opinion of modern writers is that there is only one variable species of Orang, which we call by Lin- neus name Simia satyrus. In Sumatra this Ape is mainly found in the lowlands of the eastern coasts of the island, in the districts of Palembang and Jambe. In Borneo it is more numerous, and has a wider range, extending all over the low forest-covered swamps between the coasts and the mountains of the interior, but in some places ascending to a considerable height in the lower hills. The third and only remaining genus of Anthropoid Apes contains the Gibbons, or Long-armed Apes (Hylobates), which are likewise restricted to the Oriental Region, but have a much wider distribution than the Orang. The 224 THE GEOGRAPHY OF MAMMALS Gibbons vary much in the colour of their fur, and numerous: doubtful species have been based upon such differences. The most recent authorities are not inclined to allow more than seven or eight well-marked species of Gibbons. Of them the most distinct is the Siamang (Hylobates syn- dactylus) of the Malay Peninsula and Sumatra, which has the second and third toes of the feet joined together by a thin web of skin, and has other slight peculiarities, which have induced some naturalists to place it in another genus. But this seems to be quite unnecessary. The remaining typical Gibbons are thinly distributed over all the three great islands of the Malay Archipelago, Sumatra, Java, and Borneo, extending into the Sulu group between Borneo and the Philippines. On the continent they range high up the Malay Peninsula, even so far north as Assam and Bhootan, where they are represented by the Hoolock (H. hoolock). They are also found in Siam, Cambodia, and Annam, and one species (H. havnanus), has been discovered in the Chinese island of Hainan. Thus of the three generally recognized genera of Simiidx, or Anthropoid Apes, one is of the Ethiopian, the two others of the Oriental Region. We now come to the second family of Catarrhine Monkeys, the Cercopithecidx, or what are usually called the Old World Monkeys. Of these, about one hundred and twenty species are known, divisible into about nine well- marked genera. Most of these monkeys, as we shall show, belong to the Oriental and Ethiopian Regions, but the genera which inhabit these Regions respectively are quite distinct. Two genera alone, Macacus and Senvnopithecus, have repre- sentatives within the limits of the Palzearctic Region. The Langurs (Semnopithecus) of the Oriental Region, DISTRIBUTION OF MONKEYS AND LEMURS 225 which we will take first of the group, remarkable for their slender body and the excessive length of their tails, are distributed over Southern Asia and the adjoining islands of the Malay Archipelago, and are numerous in species, as many as twenty-nine or thirty being of probable validity. Dr. Blanford describes as many as fourteen of them in his “ Fauna of British India,” as met with within the limits of India, Ceylon, and Burma. Of these, one, S. schistuceus, has mounted high into the Himalayas, where it ascends to an elevation of 10,000 feet and is never met with below 5000 feet. But this species is undoubtedly derived from the plains of India, being a very close ally of S. entellus, which has a wide distribution over the northern provinces. Another well-marked species of this genus is found high up in Eastern Tibet and North-western China, where it inhabits the mountain forests and has been named S. roxellane from its conspicuous turned-up nose. Borneo, Sumatra, and Java, have each of them distinct species of this wide-spread genus. Closely allied to the Langurs (Semnopithecus) is the very curious Long-nosed or Proboscis-monkey of Borneo, which seems to be confined to the lowlands near the mouths of the rivers in Sarawak and the adjoining dis- tricts of that island. It is the sole representative of the genus Nasalis. Passing back to the Ethiopian Region, we find in the place of Semnopithecus and Nasalis the genus Colobus, remarkable for the absence of a thumb on its hand, but in other respects closely allied to its Asiatic brethren. About ten species of Colobus are generally recognized by naturalists. They are distributed all over the forests of tropical Africa, being, perhaps, most abundant on the west Pp 5 226 THE GEOGRAPHY OF MAMMALS coast. One of them, Kirk’s Colobus (C. kirki) so far as has yet been ascertained, has been only met with in the island of Zanzibar, where, however, it is said to be now nearly extinct. Another well-marked species, the Guereza (C. guereza), is peculiar for its long-haired flanks and tail. It was originally discovered in Abyssinia, but is repre- sented by closely allied forms in British Kast Africa and on the Niger and Upper Congo. The Guenons, or long-tailed Monkeys of the genus Cercopithecus, which we now come to, are likewise entirely confined to the Ethiopian Region, and being exclusively inhabitants of forests, are naturally most numerous in the wooded districts of the west coast and in the great wooded valley of the Congo, They are very numerous in species, as many as forty different kinds having been discriminated, but are mostly confined to small specific areas, not more than one or two species as a rule occurring in the same district. Allied to the Guenons are the Mangabeys (Cercocebus) with about six known species, which has nearly the same area of distribution. Both Guenons and Mangabeys do well in captivity, and are always well represented in the Zoological Society's monkey-house, where they have, in some cases, bred young ones. As many as twenty-four different species of Cercopithecus will be found registered in the Zoological Society's Catalogues, and amongst them are some of the most beautiful and brightly coloured of the Quadrumana, such as the Diana Monkey (C. diana) and Brazza’s Monkey (C. brazze). In the Oriental Region the corresponding form of monkey is the Macaque (Macacus), about fifteen species of which are distributed over Southern Asia and its islands DISTRIBUTION OF MONKEYS AND LEMURS rypat down to Wallace’s Line. Some of these have a wide range, such as the Pig-tailed Macaque (1. nemestrinus) and the Crab-eating Macaque (J. cynomolgus), which occur in most of the large islands of the Indian Archipelago, as well as in the Malay Peninsula, but others are very limited in their specific areas. The Macaques ascend high in the Himalayas, M. rhesus or some of its allied forms going up to at least 10,000 feet above the sea-level. Moreover, two other nearly allied species of this genus are found to the north of the Himalayas, far beyond the limits of the Oriental Region. These are the Hairy-eared Macaque (M. lasiotis) of Szechuen, and the Tcheli Monkey (M. tcheliensis) of Manchuria. The latter inhabits the moun- tains of Yung-Ling, north of Pekin, in latitude 41° North, where the thermometer frequently descends to 10° below zero. An example of this rare monkey, which has been living in the Zoological Society’s menagerie since June 1886, is always kept in a cage in the openair. A third species of Macaque (J. speciosus) is found in Japan, where it is the sole representative of the order Quadru- mana. It is stated to be found all over the island of Hondo or Nippon up to 41° N. lat. and if this be the case, has a higher range north than any other monkey now existing, except perhaps the Tcheli Monkey just spoken of. In the western part of the Palearctic Region, a single species of Macaque is also found. This is the Barbary Ape (M7. inwus), which frequents in the scrubby gorges of the mountains of Morocco and Algeria, and is also a well- known inhabitant of the Rock of Gibraltar. But whether it is an aboriginal denizen of “The Rock” or has been intro- duced by man is somewhat doubtful. At the present time, 228 THE GEOGRAPHY OF MAMMALS being carefully protected by the authorities there, it is said to be increasing in numbers. It appears, therefore, that at least three or four species of Macaque must be considered as inhabitants of the Palearctic Region, while the remainder are confined to the Oriental Region. The series of Catarrhine, or Old World Monkeys, is closed by the Baboons, of which three genera are now usually recognized, one from the Oriental and two from the Ethiopian Region. The Oriental form of Baboons is the Black Baboon of Celebes (Cynopithecus niger)—a feeble representative of its African relatives in the most distant borders of the Oriental Region. The Black Baboon is stated to be also found in Batchian and the Philippines, but may possibly have been introduced by man into these localities. In Africa the Gelada Baboon (Theropithecus), with two somewhat doubtfully distinct species, is re- stricted to the mountains of Abyssinia, while the true Baboons (Cynocephalus) are spread over the greater part of the Ethiopian Region. Of the eleven or twelve species of Cynocephalus usually recognized, the best known per- haps is the Arabian Baboon (C. hamadryas)—the Sacred Monkey of the ancient Egyptians, the likeness of which is of frequent occurrence among the engravings on the Egyptian temples and tombs. Besides the south-west portion of the Arabian Peninsula it inhabits also Abys- sinia and extends into Upper Nubia. Another well- known Baboon is the Chacma (C. porcarius) of the Cape Colony. The proverbial unsightliness of the Baboons reaches its acme in the Mandrill (C. mormon) and Drill (C. lewcophxus) both from West Africa. DISTRIBUTION OF MONKEYS AND LEMURS 229 Section [V.—DIstTrRisuTIon oF New Woruip MOoNkKEYS The Cebidee, or Platyrrhine Monkeys, which we will now consider, are not so numerous as their cousins of the Old World, only from fifty to sixty species being usually recognized, although many of these are not very perfectly distinguished. They are also confined to much narrower limits than the monkeys of the Old World, being entirely restricted to the warmer portions of the Neotropical region, and, being purely arboreal in their habits, to those parts of it which are covered by dense forests. Their northern limit is Guatemala and the adjacent districts of Southern Mexico, the most northern locality for monkeys in the New World positively ascertained being about 23° N. lat., in the State of San Louis Potosi. This, it may be observed, is in striking contrast to the northern range of the Quadru- mana of the Old World, which, as has been shown, extends to 41° N. lat. To the west of the Andes of South America monkeys are only found as far south as the Gulf of Guayaquil, the arid and treeless nature of the whole southern portion of the west coast being quite unsuitable for forest-loving animals. To the east of the Andes, how- ever, monkeys extend all over the vast forests of the valleys of the Orinoco and Amazon and as far south as the wooded districts of Paraguay and the adjoining provinces of the Argentine Republic. Burmeister includes three species of Cebide in his list of the mammals of the La Plata States. The New World Monkeys are usually divided into about nine genera, amongst which the Spider-Monkeys, (Ateles) occupy the highest position. The Spider-Monkeys 230 THE GEOGRAPHY OF MAMMALS number about ten species, which are distributed over the whole area occupied by the family as above described, being most numerous, perhaps, in the great forests of Amazonia. Closely allied to Ateles are the Woolly Spider- Monkeys of the genus Brachyteles, which are confined to the forests of South-eastern Brazil. They have been divided into three species, but the prevailing opinion amongst modern naturalists is, that these are really only varying forms of one species. The typical genus Cebus, which follows next in the series, numbers some eighteen or twenty species, many of which, however, are very imperfectly discriminated. This genus also has an extensive range, extending from Nicaragua to Paraguay, but being most numerously re- presented in Amazonia. With Cebus we close the first and most highly organized group of the New World Monkeys which constitute the first sub-family Cebide. The second sub-family of the Cebide, the Mycetine, contains only the single genus Mycetes with about six species, commonly known as Howlers, from their extra- ordinary voices, produced by a specially modified vocal organ. The Howlers have also a wide distribution in the New World, one species, I. villosus, being found as far north as Guatemala, whilst another, M. niger, occurs in Paraguay. The third sub-family of American Monkeys contains only two genera—the long and bushy-tailed Sakis (Pithecia) and the short-tailed Ouakaris (Brachywrus). Contrary to what is the case in the previous sub-families, the generic area of this group is much constricted, being confined to the basins of the Orinoco and Amazon and to Guiana, which is surrounded by them. ‘The five or six known species of DISTRIBUTION OF MONKEYS AND LEMURS Zo Saki are all inhabitants of various localities within this district. The mode of distribution of the three species of Ouakari is still more remarkable. Each of them, as first shown by Bates and afterwards further explained by Forbes, is limited to a comparatively small tract of forest on the banks of the Amazon and its afiluents. The Black-headed Ouakari (B. melanocephalus), as shown by the accompanying map (prepared by Forbes), which we _ | | nl lel vere x Swe i imi TD a ni ou 5 7 gual Mh hour = os “is =e Foner YS ‘ee, — “Ne BR Ega ox‘Teffo Amazo, | MAP OF THE SPECIFIC AREAS OF THE OUAKARIS. (P. Z. S. 1880, p. 647.) have been kindly permitted to use by the Zoological Society of London, is met with only in a tract traversed by the Rio Negro, the Bald-headed Ouakari appears to be confined to the triangle formed by the union of the Amazon with another affluent, the Japura, and the Red Ouakari to the forests on the north bank of the Amazon opposite Olivenca, and lying between the main stream and the river Ica. Each of them evidently takes the place of the others in its particular district. Of this peculiar ps4 THE GEOGRAPHY OF MAMMALS kind of distribution few instances are known amongst mammals, but many somewhat similar cases have been observed in birds, reptiles, and butterflies. The fourth and lowest sub-family of the Cebide (Nyctipithecinz) includes three genera—the Douroucoulis (Vyctipithecus), the Titis (Callithriz), and the Squirrel- Monkeys (Chrysothriz), and numbers altogether some twenty species. This group, as a whole, has a wide range like the two first sub-families, extending from Central America to Paraguay. But the species are most abun- dant in the centre of the area, where some of them, so far as is yet known, have a very limited range. SEcTION V.—DISTRIBUTION OF THE MARMOSETS Like the Platyrrhine Monkeys the little Marmosets, which constitute the family Hapalidx, are entirely re- stricted to the tropical forests of the New World. The family embraces but two genera generally acknowledged— Hapale with about seven, and Midas with about fourteen or fifteen species. But these small creatures are still little known, and it is probable that many more of them remain to be discovered when the vast forest-region through which they are distributed shall have been more thoroughly explored. The Marmosets do not extend so far north as the true Monkeys, only a single species (Midas geoffroy?) having yet been ascertained to range north of the Isthmus of Panama as far as Chiriqui. Thence, southwards, they are thinly distributed over the South American continent down to the northern provinces of the Argentine Republic, where Hapale penicillata is said to occur in the forests of Salta DISTRIBUTION OF MONKEYS AND LEMURS i and Jujuy. The Marmosets appear to be most nume- rous in the forests of Amazonia, where some of them are confined to very restricted districts. Our best account of these animals is to be found in the journals of the excel- lent observer Bates. SEcTION VI.—SuMMARY AND DEDUCTIONS The subjoined table contains a list of the genera of the Order Quadrumana, or Monkeys, and gives the approxi- mate number of species met with in each of the great geographical Regions. New Sol oie heel ea leesdl sGhnes [reece ere aoa lage Sl) sales 2 7S|HE 5S/82 Bele ' (ps eo ele) sas eS | alee Famity I. | | SIMIIDA. LF Ser ouUn eels AB ee 3 | te | Resimia= 5 9 top| Siler sesis| 1 | 3. Hylobates . 7 || = ri 11 3 8 Famity II. | | | CERCOPITHEUCIDA. | 1. Nasalis . le) | a 2, Semnopithecus 29 | = 28 1 3. Colobus . 10 | 10 ee 4, Cercopithecus . | 40 40 [: 5. Cercocebus . legal 6 sie 6. Macacus . 17 13 4 7. Cynopithecus . 1 | aE 1 8. Theropithecus savisvilre 2 9. Cynocephalus. : . . | 11 11 | Baebes sce mops (PEE IVT SB ioe cose otek 019 al SRR a: ae Gt) 234 THE GEOGRAPHY OF MAMMALS SUMMARY AND DEDUCTIONS (continued). Famity III. CEBID. . Ateles . 11. Brachyteles 12, Cebus . Mycetes . . Pithecia . 15. Brachyurus . Nyctipithecus . . Callithrix . Chrysothrix Faminy IV. EVA PATA DAs, Midas Hapale 19. 20. FAMILIES. T, Simiide:, ~ II. Cercopithecidee Cebidee . . Hapalidee Biol ea te (ete aes 23 | ~2\a5 Se SSlee/ 8/38 ga S/Fs)/ Ae |Re/ Feralas Sey ie) Sol ae ener ears 10 10 1 1 18 18 | 6 6 | 5 5 | 3 3 | a) je ra 1 4 4 BBall sce) BBM ee. beeen ee eee ee 14 || 14 i 7 | ae eer SUMMARY. a 8 Oi Sci Sal vee oe 11 new cp ee 117 ... | 69 43 ae 63 63 |i. Nene hee 21 PAD ae | Lone 212 84 | 72 | 51 | 5 ise) Ou DISTRIBUTION OF MONKEYS AND LEMURS 2 DEDUCTIONS. 1. The Order Quadrumana, or Monkeys, contains about 212 species, divisible into twenty genera and four families. 2. Monkeys are found only in the tropical and sub- tropical portions of the Eastern and Western Hemispheres, and are absent in Australia and Madagascar. 3. The Monkeys of the Eastern and Western Hemispheres are quite distinct in structure,and belong to different families. 4. In the Western Hemisphere Monkeys are restricted to the Neotropical Region, not occurring north of 20° N. lat., or south of 30° S. lat. 5. In the Eastern Hemisphere some few species of Monkeys are met with as far north as 40° N. lat., and in Africa descend to 35° 8. lat. 6. The Monkeys of the Ethiopian and Oriental Regions belong to distinct genera. 7. The Anthropoid Apes are restricted to the tropics of Africa and Asia, the most highly organized of them being the Chimpanzees of Africa and the Orangs of Asia. Section VII.—DIstRIBUTION OF THE LEMURS The Lemurs, which by many recent authorities are united with the Monkeys to form the Order Primates, but which it is, in some respects, more natural to retain as an Order by themselves, number altogether some fifty species—only one- fourth of the number of the Quadrumana. Theyare also very different in geographical distribution, thirty-five out of the whole series being confined to the Malagasy Sub-region, whilst thefew remaining forms are met with onlyin isolated portions of the Ethiopian and Oriental Regions. Besides the family of 236 THE GEOGRAPHY OF MAMMALS true Lemurs(Lemuride) this Order contains the two aberrant types, the Tarsier (Tarsius) and Aye-aye (Chiromys), both of themof family value. Wewill say the few words that are neces- sary about the distribution of each of these three families. The typical Lemurs (Lemwridz), are usually divided into four sub-families, the Jndrisine (Indrises), Lemurine (Lemurs), Galagine (Galagos), and Lorisinx (Slow Lemurs). Of these, as will be seen by the annexed table, the two first sub-families, which contain seven genera and about twenty- four species, are absolutely confined to Madagascar and its adjoining islets. It is, in fact, mainly the presence of these peculiar animals, which constitute altogether nearly one half of the whole mammal-fauna of Madagascar, that renders the Malagasy fauna so very different from that of any other part of the world’s surface, and makes it a moot point as to whether this zoological division should not be more properly treated as a “Region,” than as a “Sub-region.” When we come to the third sub-family—the Galagos—we find the typical genus Galago with its six species dis- tributed over continental Africa, but the three other genera of this family, which contains smaller animals of somewhat aberrant form, are again entirely restricted to Madagascar. The fourth and most aberrant group of the Lemurs, com- monly called Slow Lemurs, from their nocturnal habits and sleepy dispositions, contains four genera, two of which belong to the Ethiopian Region and two to the Oriental Region. Although they vary considerably in structure from the more typical Lemurs, there can be no doubt that the Slow Lemurs possess a true Lemurine structure in many important particulars, so that they must have had a com- mon origin with the true Lemurs. This fact would seem to show that the ancient “Lemuria,” as the hypothetical DISTRIBUTION OF MONKEYS AND LEMURS Zone continent which was originally the home of the Lemurs has been termed, must have extended across the Indian Ocean and the Indian Peninsula to the further side of the Bay of Bengal and over the great islands of the Indian Archipelago. Section VIII.—SumMMaAry AND DEDUCTIONS Table of the genera of Lemurs and of the number of species found in each Zoological Region. eI S =| — 3 z a a eile ome oe more San are SHEARS |ES|SP| be Peles Palle ta) cilia anetl pegy house erste Dal haceeta| hee |) SO ely ee Famity I. | LEMURID&. SuB-FAMILY J.-—INDRISINA. 1. Indris 1 1 2. Propithecus 4 4 3. Avahis . i 1 Sup-Frami.y IJ.—LEMvuRINz.| | Ab, \beiombne 5 e 8 8 5. Mixocebus . 1 i 6. Hapalemur 2 2 | 7. Lepidolemur . | 7 | Sup-FaMILy II].—GaLaGin&. 8. Galago . 6 Genieat | 9. Chirogale . 4 Wield ta | | 10. Microcebus 5 5 11. Otolemur 2 2 | Susp-FAMILY I1V.—LoRIsINn»&.) | 12. Loris. ; iy | y il 13. Nycticebus . i) ere 1 14, Perodicticus 1 || 1 | 15. Arctocebus. Ju | 45 | 238 THE GEOGRAPHY OF MAMMALS SUMMARY AND DEDUCTIONS (continued). | Bl 8) a ik locas g | 2[48/H6)28)e8 -a| 28 hee 3 6 r rs zie) |e c= | 4) | eh) ae nena Famizy II. TARSIIDA. | De Warsi 22 va ao yee Boel) ee ies Poet |e ata ee ee ee Famity III. CHIROMYID. 1. *@hinonmtys ¢ pak ee ee eee Pee eee [eseed\e cn ae 48 SP s6a a4: DEDUCTIONS 1. About forty-eight species of the Order Lemures, belonging to seventeen genera and three families, are known. 2. Of these thirty-six species, representing twelve genera, are restricted to the Malagasy Sub-region, which must be regarded as the special home of the Lemurine Mammals. 3. The remaining forms of Lemurs are thinly spread over the Ethiopian and Oriental Regions, Lemurs being altogether absent in the Palarctic, Australian, Nearctic, and Neotropical Regions. 4, The Aye-aye (Chiromys), forming a family of itself, is restricted to Madagascar. CHAPTER X DISTRIBUTION OF THE ORDER CARNIVORA Section I.—IntTropuctory REMARKS THE true Carnivora are widely distributed over the world, and occur, in fact, everywhere from north to south, with the exception of the Australian Region, where their functions in nature are performed by the flesh-eating Marsupials. The Polar Bear probably ranges farther north than any other species of mammal, while the southernmost point of the South American Continent is inhabited by the Magellanic Dog. In the Antarctic Seas the Carnivora are also represented by several species of Seals, but no land-mammal has yet been found on the Antarctic Continent. The Carnivora are very numerous, nearly 300 species belonging to seventy genera being usually recognized. They are usually divided by modern authorities into four well-marked sections: (1) the luroid, or Cat-like Carnivores, belonging to four families; (2) the Cynoid, or Dog-like Carnivores, consisting of the single family Canide; (3) the Arctoid, or Bear-like Carnivores, with three families; and finally, the Pinnipeds, or Marine Carnivora, with three families. We will take these in order, pointing out the most noticeable features in the distribution of the principal eu best-known forms in each 240 THE GEOGRAPHY OF MAMMALS family, and only alluding to the less-known and more insignificant forms when remarkable for their special distribution or otherwise. Section I].—DIstTRIBUTION OF THE CAT-LIKE CARNIVORA The luroid, or Cat-like Carnivora, among which will be found the largest and most perfectly organized terrestrial animals in the present condition of the world, that have been modified to prey upon their weaker brethren, embrace four families, the Cats, Civets, Aard- Wolf, and Hyenas. The first family, the Felidz, or Cats, contains about fifty species of true Felis, together with the Cheetah, or Hunting-Leopard, which it is necessary to refer to a separate genus. The Cats are distributed all over the main portions of both the Old and New Worlds, failing, however, as already remarked, in the Australian Region, and not extending quite so far north nor quite so far south as the Bears and Dogs. No single Cat, how- ever, has anything like this wide area of distribution, all the species being more or less limited in their extension, and the Cats of the Old and New Worlds being always specifically different, unless it be in the case of the Northern Lynxes. We will shortly describe the specific areas of some of the best-known of these animals. The Lion, the undoubted king of beasts, though it for- merly penetrated far into Europe and even into England, is in these days essentially an African animal, but still ex- tends throughout Arabia, Asiatic Turkey, and Persia into Western India, where it occurs in the Western Provinces. DISTRIBUTION OF THE ORDER CARNIVORA 241 Though the Lion varies greatly in shape and colour, and in other minor features in different districts, it seems im- possible to accept the proposed division of it by some authors into species, or even into geographical sub-species. It is true that an expert, well accustomed to examine Lions, will usually be able to guess correctly the locality whence any particular specimen has been received, but this is by no means absolutely certain. The Tiger, the next noblest beast of prey existing in the present day, and by some writers assigned to a station superior even to that of the Lion, is essentially, no doubt, an inhabitant of the Oriental Region, but has extended its range in many places, probably within a not very distant period, far to the north. The Indian Peninsula may be said to be its focus, where it is found almost everywhere from the Himalayas, which it ascends to a height of 6000 or 7000 feet, to Cape Comorin. Curiously enough, however, the Tiger is not found in Ceylon. Outside India the Tiger ranges throughout Northern Persia, Turkestan, and the southern provinces of China, reaching on the west up to Manchuria and Amurland, where, however, it has learnt to put on a thicker and longer coat in winter to protect it from the severe cold. To the south and east of India the Tiger extends all over the Siamese and Malay Peninsulas into Sumatra and Java, but not, it is believed, into Borneo. Tigers from Sumatra and Java do not, however, attain to the large dimensions of their brethren of India and the north. The third great Cat of the Old World, the Leopard, has a still wider distribution than its two larger brethren. Not only does it inhabit the whole of Africa, including its northern portion, but extends also nearly over every part Q 242 THE GEOGRAPHY OF MAMMALS of the Oriental Region and into the confines of the Palie- arctic. The Leopard is met with throughout Western Asia, Persia, the Indian Peninsula, including Ceylon, the Siamese and Malay Peninsulas, China, and the larger islands in the Indian Archipelago. It varies much through- out its range, not only in size, but in shape and in the character of its markings; yet it is impossible to consider more than one species of Leopard as satisfactorily estab- lished. The fourth great Cat of the Old World, the Ounce (Felis wncia), distinguished at once from the Leopard by its white body-colour, thick fur, and much lengthened tail, is confined to the high ranges of Central Asia, and occurs only at elevations of over 8000 feet. In Gilgit, Hunza, and Tibet, it occurs occasionally within the range of the Indian sportsman, but is more frequent further north, and in the north-west ranges of Siberia and Amurland. After the four great Cats of the Old World, we must proceed to discuss the corresponding animals of the New World. These are two only in number, the Jaguar (F. onca), and the Puma (F. concolor), both inferior in size and organization to the Lion and Tiger, the places of which they are generally supposed to take in the Western Hemisphere. The Puma has a very wide distribution in America, extending over the greater part of both North and South America, from the most northern of the United States and British Columbia down to Patagonia. In the extent of its range from north to south, the Puma seems to surpass every other carnivorous animal. The Jaguar has likewise a very wide distribution in America, though it cannot rival the Puma in this respect. Its northern limit in these days appears to be Louisiana, DISTRIBUTION OF THE ORDER CARNIVORA 243 Texas, and the adjoining southern United States. Hence it ranges throughout Central and South America down to the Rio Negro of Patagonia. As might, however, have been expected from its being essentially a forest-loving animal, the Jaguar does not occur on the western side of the Andes south of Ecuador. Thus we may take it that of the six largest and finest carnivorous mammals of the existing creation, four are found in the Old Continent and two others, generally inferior in structure, in the New World. We need not here go at length into the distribution of the smaller cats, some forty or forty-five in number, as variously estimated. It is sufficient to say that those of the New World are specifi- eally distinct from those of the Old, except in the case of the Lynxes, in which the differentiation of the Canada Lynx (Ff. canadensis) from F. lynx of the Palearctic Region is perhaps somewhat doubtful. The cats of the Ethiopian and Oriental Regions are also, as a rule, different, and many of them are restricted to comparatively narrow geographical limits. The second genus of the Cat family (Felidx) contains only the Hunting Leopard, Cynzlwrus jubatus, which has somewhat the same distribution as the Lion. It is found all over Africa, and extends throughout South-western Asia and Persia into Western India up to the confines of Bengal. Attention, however, should be directed to a sup- posed second species of this genus, the Woolly Cheetah (C. laniger), which has hitherto been only met with in some of the higher districts of the Cape Colony. Next to the Cats we come to the Viverridz, or Civets, a much more numerous group containing about seventy species, usually divided into about twenty-four genera. 244 THE GEOGRAPHY OF MAMMALS In the New World the Viverride are entirely unre- presented, and, as a rule, may be said to belong to the tropical portions of the Ethiopian and Oriental Regions, being entirely absent in the Australian Region. It may also be again remarked that the Malagasy Sub-region is quite peculiar as regards its Viverridx, six well- marked genera of this family being entirely restricted to that anomalous island, whilst the seventh, Viver- ricula, is probably only an introduction from the Oriental Region. In the Palzarctic Region the Viverride are feebly represented by a few scattered species which have strayed from the south into its limits, such as the Common Genet (Genetta vulgaris) and the Ichneumon (Herpestes ichneu- mon), which are both met with in Southern Spain and Algeria. Asa rule, again it may be said that the Ethio- pian and Oriental Viverridz are not only specifically but generically different. But one large genus, Herpestes, has its twenty species divided pretty equally between the two Regions, and the typical Civets, Viverra, are found both in Africa and India. A third family of the Aluroid Carnivora has been necessarily formed for the reception of the Aard-Wolf (Proteles cristatus). This is a hyenoid form well distin- guished by its extremely peculiar dentition. The Aard- Wolf appears to be restricted to the southern and eastern portions of Africa, and constitutes one of the most charac- teristic forms of Ethiopian mammal-life. Finally, closing the A‘luroid series, we have the small family of Hyenas (Hyznidx) containing three well- marked species, which, taken on the whole, must be denominated Ethiopian, as they all three occur within DISTRIBUTION OF THE ORDER CARNIVORA 245 the limits of that Region, and two of them at the present epoch are restricted to it. But the Striped Hyzena (Hyz2na striata) extends from North-east Africa through Arabia and Persia into Northern India, where it is common in the North-western and Central Provinces. It must therefore be registered as a common inhabitant of the Ethiopian and Oriental Regions. SEcTION IIJ].—DISTRIBUTION OF THE’ DOG-LIKE CARNIVORA The Cynoid, or Dog-like Carnivora, consists only of the single family Canidx. Of this the great mass of species (from thirty to forty in number) belongs to the true Dogs (Canis), which, as we shall see, are very widely distributed over the earth’s surface, whilst the three remaining genera are isolated forms, each of one species only, and are re- stricted to narrow geographical limits. The Dogs (Canis) are amongst the most hardy of known mammals, and are spread, as already stated, over the whole earth, being apparently adaptable to all the zones, whether tropical, temperate, or frigid. The ano- malous island of Madagascar is the only land to which they have not penetrated, that is, if we allow the Dingo of Australia to be an indigenous and not an introduced species, which is a point open to question. In the Arctic Regions the Polar Fox (Canis lagopus) extends as far north as any other carnivorous mammal, except perhaps the Polar Bear, and is found far above the Arctic Circle in both the Old and the New World alike. On the other hand, in the extreme south of the New 246 THE GEOGRAPHY OF MAMMALS World we find the Magellanic Dog (Canis magellanicus), which extends to the very farthest extremity of the American continent, while in Southern Africa the Black- backed Jackal (C. mesomelas) is met with in the vicinity of Cape Town, and in Australia the Dingo, now only known in a semi-domestic state, is found over the whole con- tinent. But although, as we have shown, the genus Canis is so widely spread over all parts of the earth, the in- dividual species are in some cases confined to restricted areas. Many well-known members of the genus—such as the Wolf, the Common Fox, and the Jackal—have a very wide distribution. But other species of Dog have limited ranges, and not more than two or three of them are usually met with in exactly the same district. Examples of this restricted distribution are afforded by the Maned Wolf (Canis jubata) of Brazil and Argentina, by the Corsac (Canis corsac) of Central Asia, and by several of the African Fennecs. But as a rule it may be taken that the various species of Dogs are hardy animals with extended areas of distribution. Besides the genus Canis, the Dog family contains three other well-marked genera, each embracing but one species. One of these, the Bush-dog (Icticyon venaticus), is found only in Brazil and British Guiana, the two others, the Hunting-dog (Lycaon pictus) and the Long-eared Fox (Otocyon megalotis) are both peculiar to the Ethiopian Region, where the Lycaon appears to have a considerable range from north to south, but Otocyon is only known from the Cape Colony. DISTRIBUTION OF THE ORDER CARNIVORA 247 SecTION I[V.—DISTRIBUTION OF THE BEAR-LIKE CARNIVORA We now come to the third and last division of the terrestrial Carnivora, which consists of those allied to the Bears and therefore denominated Arctoid. This division embraces three families—the Uvrsidz, or Bears, which have a fairly wide distribution in both Hemi- spheres, the Procyonidx, or Raccoons, which, with a single exception, are confined to the New World, and the Mustelidx, or Weasels, which belong mostly to the Old World with a comparatively few representatives in the New World. The Bears (Ursidz), which head the group, contain, after the Cats, the largest and most destructive of the carnivorous animals of the present day. There has been a tendency of late days, unnecessarily, as we think, to augment the specific forms of the true Bears (Ursus). The species, recognizable by obvious external characters, do not appear to exceed ten in number. Taken as a whole the genus Ursus presents some very interesting features in its distribution. Its generic area embraces the whole of the Palearctic and Nearctic Regions and ° extends into the northern confines of the Oriental. In the Ethiopian Region Ursus is entirely absent, and consti- tutes an important lipomorph. In the Neotropical Region it is represented by a single species, the Spectacled Bear (U. ornatus) of the Andes. In the extreme north of the globe the Polar Bear (U. maritimus) ranges round the Arctic Circle. The next northern species met with is the 248 THE GEOGRAPHY OF MAMMALS Brown Bear (U. arctos), which, under different forms and varieties, occupies the whole Palearctic Region, and is represented in the Nearctic by the scarcely distinct Grizzly Bear (U. horibilis) under various forms. } qa Ay ee | 8 15) 7220 i 1 i! ae 9c. |) AGE) eae eat : 1 eel 1 2, | aeslete| 1 | oe 4 | 1 3 | 1 : 10 | a | 2 ; i 2 “ia ‘ 1 | : 10 1 il 2 2 1 5 illo 2 1 2 1 30 | 91] 31 | 2 on) we) — OND NPNNOrWRRER Wwe | mH Doe be OF DN r Australian. II. Neotropical. 252 THE GEOGRAPHY OF MAMMALS A. TERRESTRIAL CARNIVORA (continued). Ve) Bg) el en ee Se (SESS |ES ES Re rales "a! #/ 3| 2l\"s| 3] =| a | = z i) = = a Famity III. | PROTELEIDA. Bi; PYOLClesin, |e ate etumey ly eedon Meee he fA ree leisescl tac “||. Sc0 Famity IV. HYANIDA. 28; Eby comes, 1c is EMM aD cee le San eSBs feieeel ste ee Famity V. | | CANIDA. | 29. Canis 6 la) Only | <3, Onl aa 30. Lycaon . Se emai MEM Stee NO al| « L vee Sie MOO VON ws chia, ee eee me ee tal ed | 32. Icticyon tel 1 | seu mt 0% 3| 9] 13 Faminy VI. | URSIDA. | Sh ORGS bee uh pera eo ea TWO NM koeeenl| ecalalpeeene ns. 2 4 5 S4elursuse ) eae e are hs MIE cate eel eee Dm essen Sc a5 laropus i ue el qed Se ee | Sa ety een) ||, ..4! 12 1 i 3 4 6 Famity VII. PROCYONIDA. 36. /Alurus . 1 Be es 37. Procyon 2 1 Fe saliby sl 38. Bassariscus 1 il 39. Bassaricyon 2 || 2 Ba 40. Nasua 2 |I 2 1 41. Cercoleptes i.| 1 1 9 || | 9 1 4 | DISTRIBUTION OF THE ORDER CARNIVORA 2 Ot iss) A. TERRESTRIAL CARNIVORA (continued). 42. 43. 44, 45. 46. 47. 48. 49. 50. 51. 52. 53. 54, 55. 56. 57. 58. Famity VIII. MUSTELIDZ. Lutra Latax . Mephitis Conepatus . Arctonyx Mydaus. Meles Taxidea . Mellivora . Helictis . Ictonyx . Galictis . Mustela. Peecilogale . Lyncodon . Gulo. Spilogale a CON PONNRNWH Orr (Je) | NR e I Australian II. Neotropical | | | | Til. Ethiopian. “I Ifa. Malagasy. peje | os o; 4| & a a | | | | SUL CO Md Mes | ieee. Beiliesce re ee i alles Din al be vant ad Sey ile 1 lease Biles eee My 4/10] 15 vaca 2 16 | 24 | 26 THE GEOGRAPHY OF MAMMALS B. MARINE CARNIVORA | 59. | 60. Famity IX. OTARIIDA. Otaria . FAMILY X. TRICHECHID. Trichechus Famity XI. PHOCIDA. . Phoca . Halicherus 3. Monachus . . Cystophora . Macrorhinus . . Ogmorhinus . . Lobodon . . Leptonychotes 39. Ommatophoca Total Species. Te | Arctatlantis. —_ on oc ms 8 II. | Mesatlantis. III. | Indopelagia. Iv. Arctirenia. 2 t DEDUCTIONS divisible into fifty-eight genera and eight families. Vv. Mesirenia. VI. Notopelagia. ciple eee ce peerage ee 1. About 260 species of terrestrial Carnivora are known, 2. They are distributed pretty equably over all the Regions of the earth except Australia, where the Dingo (possibly introduced) is their only representative. DISTRIBUTION OF THE ORDER CARNIVORA 259 5) 3. The only Carnivores in Madagascar are Civets (Viverridx), and out of the seven genera of that family represented there six are peculiar. 4, There are no Bears (Ursidx) in the Ethiopian Region. 5. The Raccoons (Procyonidz), with a single exception, are peculiar to the New World, where they may perhaps replace the Civets (Viverridz), which are confined to the Old World. 6. The Marine Carnivora consist of three families which are divided into eleven genera and about twenty-six species. 7. The Walruses are characteristic of the Arctic Seas and the Sea-lions of the Antarctic, but in the Pacific the Sea-lions are also found in the North. 8. The Seals (Phocidz) are both Arctic and Antarctic, but are represented by quite different genera in these two areas. CHAPTER XI DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS SecTIon I—IntTRopDUcTORY REMARKS THE three Orders, to which it is proposed to devote the present chapter, contain the most difficult and least known members of the whole class. They are very numerous, especially the two latter groups, nearly all of small size, and in most parts of the world have been very im- perfectly studied. Within these last few years large additions have been made to our knowledge of them, especially in the case of the Rodents, and their proper systematic arrangement is still a matter of much discus- sion amongst naturalists. Under these circumstances, and looking to the fact that these small mammals are of much less general interest than their larger brethren, it is not proposed to go very fully into the subject of their geo- graphical distribution, but merely to point out some of the leading and less controvertible facts known upon this part of their history. We will commence with the Insectivores, which are generally allowed to be in many respects allied to the Carnivores, although they present certain points in their structure which appear to show a probability of their original descent from much lower forms. 256 DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 257 Section IJ.—DIstrIsuTION OF INSECTIVORES The Insectivores are for the most part widely scattered over the earth’s surface, but not very numerous in species except in certain localities. In the Australian Region they are entirely absent, their place in nature being there taken by the Insectivorous Marsupials, and in the Neotropical Region, they have only penetrated as far south as certain districts in the Northern Andes. In this case also we may suppose that their functions are performed by the smaller Opossums (Didelphyidx), which subsist almost entirely upon insects. In almost every other part of the earth, as we shall see, the Insectivores are represented either by special types or by members of the widely-spread group of Shrews. As will be seen by the tables given below (p. 260) the Insectivores, according to a moderate estimate, are sup- posed to number about 230 species, divisible into forty-one genera. These genera are grouped in ten families, on the distribution of each of which a few words may be said. At the head of the Order it is best to place the Kaguan, or Flying Lemur as it is commonly called, though its structure is so peculiar and its affinities so little obvious that it might perhaps be more properly ranked in an Order by itself. Of this family only a single genus (Galeopithecus) with two species is known, one found in the Malay Peninsula and the islands of the Eastern Archipelago, and a second, smaller form in the Philippine group. The Galeopithecidx, therefore, may be placed as a characteristic form of the Oriental Region. The second family of Insectivora, the Tupaiidx, or R 258 THE GEOGRAPHY OF MAMMALS Tree-shrews, with two genera and about fifteen species, is likewise confined to the Oriental Region, and forms another characteristic group of that Region. In the third family of Insectivorous Mammals the Macroscelidex, or Jumping Shrews, which, though held to represent the Tree-shrews in Africa, are very different in appearance and are essentially terrestrial in their habits, must be considered as a purely Ethiopian type, though one form (Macroscelides rozeti) has crossed the limits of the Ethiopian Region into Algeria. The fourth family of Insectivora, the Hrinaceide, or Hedgehogs, are rather more widely diffused. The true Hedgehogs (Zrinaceus), of which about fourteen species are known, are spread all over the Ethiopian Region except Madagascar, and are likewise found in the Oriental and Palzarctic Regions. The second genus of Hrinaceidx (Gymnura), of which two species are distinguished, is restricted to the Oriental Region. In the fifth sub-family of this Order the Soricide, or Shrews, we find the most numerous and the most widely extended mammals of this group. Though these little animals are still very imperfectly known and many more species must remain to be discovered, they already number some 125 species. These are spread over nearly the whole earth except the Australian Region and the Neotropical Region, where, however, in the northern borders two or three species are known to occur. The Musk-shrews (Crocidwra) are the most numerous of all the genera, more than eighty species of these little animals having been already described. They are found in Africa and Mada- gascar and are also numerous in the Oriental and Pale- arctic Regions, but do not occur at all in the New World. DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 259 Others of the eleven genera of this family, however, are based on one or two species and have a much more limited distribution, such as the Water-shrew (Crossopus) of Europe and Northern Asia, and Nectogale, the Web- footed Shrew of Tibet. After the Shrews follow the Moles (Talpidz) of which about eleven genera are known containing altogether about twenty-three species. The Moles are specially character- istic of the Palsarctic and Nearctic Regions, to which nearly all the genera and species are confined, but one or two species of true Moles (Talpa) have invaded the confines of the Oriental Region. The seventh family of Insectivores (Potamogalidz) contains only two isolated forms—Potamogale from West Africa and Geogale from Madagascar. This group must therefore be attributed to the Ethiopian Region. Allied to the last-named family, but still more closely to the Tenrecs which follow them, are the two species of the remarkable genus Solenodon, to find which we must go, strangely enough, as far as the West Indian Islands. Here linger the last representatives of this singular group of Insectivores, one species, Solenodon paradocxus, being restricted to Hayti and the other, S. cubanus, to Cuba. The Tenrecs (Centetidx), which form the ninth family of Insectivorous Mammals, number as many as twenty- one species which belong to seven genera, of which the distribution may be very shortly described, as they all belong to the island of Madagascar, and constitute one of the most curious of the primitive forms of animal life that render the Malagasy Sub-region so remarkable. Finally, closing the Order of Insectivores, we find the 260 THE GEOGRAPHY OF MAMMALS Golden Moles (Chrysochloridz), a family containing but one genus, Chrysochloris, with seven species all confined to the Ethiopian Region. Summary and Deductions as regards the Insectivores Table of the families and genera of the Order Insecti- vora, showing the approximate number of species in each of the Regions. Total Species. I. Australian Region. II. Neotropical Region. |TII. Ethiopian Region. Region. - “V. Nearctic Region. VI. Palearctic Region, | IIIa. Malagasy Region. IV. Oriental | | Famity I. GALEOPITHECIDA. if (Galeopithecus:.5 ce. sx fon 22 sivemard| mace litera Veneer meee Famity II. TUPAIIDA, DL UDBIA: 2) sms) | see open PUA Niwas | eee l-oeaulteaesmmilee 3 Ge talOcercuss:- (ep vee eee Ai|iieemel| Seanlll locos l| rece eee 7g ee Ss iss Faminy III. MACROSCELIDIDA. 4. Macroscelides . 10 10 5. Petrodromus . : Od Ii seal! sae 3 G,hubynchocyonice 00.0 ec. | 40)|| ceoal wae tee | 17 ae 117 eae. Faminy LY. ERINACEID. (oGymmnira: tet os tg. we.) 2 2 2 | eh JdabORee 4 4 a oo. a ih até! 4 4 6 4 6 DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 261 Summary and Deductions as regards the Insectivores (continued). We Se tole le lg af 22 | 28|23|38| 38/23) 28 53 || 25|S8|se| S5|Sa| Sa| se BB 28 |e8|"2| 8 | 8 |e | Ae : = = q i > i= A i i =| > FamILy V. | SORICIDA. | 9. Sorex 18 | bes Se || oll 10. Soriculus 4 ss Ay laecnll ates 11. Blarina . 8 Bole pelle Ou (leer: 12. Notosorex . Z a Gand era lerats 13. Crossopus 1 set ae il 14. Myosorex . 4 4 a 15. Crocidura . . 82 AQ) | ZnieoO 10 16. Diplomesodon 1 yet i 17. Anurosorex D) 1 1 18. Chimarrhogale 2 1 2 19. Nectogale . 1 i 25) ees 2 | 44 2 | 36 | 20 | 22 Famity VI. TALPIDA. 20. Myogale 2 eee ee 21. Urotrichus. I | pected 22. Neurotrichus . 1 Ae bee 23. Uropsilus . ee la 1 DA Mal pike, er. oft oars |) LO 1 10 25. Scaptonyx . wee nail see alee 26. Dymecodon I] | Seen 27. Scalops . 3 omer 28. Scapasius 3 3 : 29. Perascalops 1 il 30, Condylura . 1 i PASM easel Mseea leas allan 1 Hi Famity VII. POTAMOGALIDA. | 31. Potamogale Sail iba sa85 eee hi ed en SOmGeOpaleyae cs ete er ell LaiWepen ces Niece |) 2 peo Meal aeace 262 THE GEOGRAPHY OF MAMMALS Summary and Deductions as regards the Insectivores (continued). ol a oe cS | aE eee ecm B= Be = Por =e > Yoel os | cour Id —a@ilee) se) Se/ se) §s/ es] 88 Sexi ko’ AS! sS|/6S|29|/ ao] 8e 53) 2h Sh| Selah) Sh) Se| eh | i) a3 PR Se | Se | es] ee | el | oe |e ea = hen ee te Ls = ian = ze | | ac Famizy VIII. SOLENODONTID. 33. Solenodon . Famity IX, CENTETID. 34. Centetes 1 35. Hemicentetes . 3 36. Ericulus 4 37. Microgale . Sa eae bis ac. Oryzoryctes . 2) | 4 il 1 21 39. Limnogale . 40. Echinops FamIy X. CHRYSOCHLORID. @I, Ohrysoehlorie:. 2 cas uly sthee|| Sze eWaseiil ah DEDUCTIONS 1. The Order Insectivora consists of ten rather isolated family groups, distributed generally over the earth, except in Australia, where they are absent, and the Malagasy Sub-region and South America, where they are very feebly represented. DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 263 2. About 230 species of Insectivores are recognized, arranged in forty-one genera. 3. The most numerous and all-pervading group of Insectivores is the Shrews (Soricidz), of which upwards of 120 species are known. 4, Many of the families of Insectivores are excessively local in distribution, the Kaguans and Tree-shrews being peculiar to the Oriental Region, the Elephant-shrews and Golden Moles to Africa, the Solenodonts to the Greater Antilles, and the Tenrecs to Madagascar. Section III.—DiIstTrisuTion oF Bats (a) Introductory Remarks The Bats, which constitute the Order Chiroptera, are, after the Rodents, the most numerous of all mammals, upwards of 530 species being already recognized, and many others probably awaiting further researches. As regards their distribution, it must be recollected that their powers of flight, which render them capable of passing over tracts of water that could not be crossed by other small mammals, place them in a somewhat different category from ordinary Mammals. At the same time the mode of their occur- rence on the earth’s surface presents many interesting features, concerning which a few words should be said. In these remarks, however, we shall be brief, as the Bats are not creatures of general interest, and as a rule are little studied except by the scientific worker. 264 THE GEOGRAPHY OF MAMMALS (b) Chief Points in the Distribution of Bats The first point to be noticed in the distribution of this extensive Order is, that the members are divisible into two well-marked and easily recognized sections. To the first of these belong the fruit-eating Bats of the family of Ptero- podide#, sometimes recognized as a Sub-order under the title “ Megachiroptera.” This section contains about 110 species, divided into eighteen genera, the whole of which are entirely confined to the tropical and sub-tropical portions of the Eastern Hemisphere, and are quite un- known in the New World. They are spread over the Ethiopian, Oriental, and Australian Regions, the African forms, about five in number, being mostly generically distinct and peculiar to that continent. One of the most remarkable of these is Hpomophorus, distinguished for its large lips and capacious mouth, of which eleven or twelve species occur in the Ethiopian tropics. In the Oriental and Australian Regions Lpomophorus is replaced by Pteropus, with as many as fifty or sixty species. It is a remarkable fact that Pteropus has never reached the African continent, although two of its species occur in the Comoro Islands, only about 200 miles distant from the African coast. Passing onwards to the five families of Insectivorous Bats, we find the two first of these, the Rhinolophide and Nycteridx also absent in the New World, although they are distributed in larger or smaller numbers over most portions of the Old World, including Australia. The fourth family, Vespertilionidx, which is the most nume- rous group of all, embracing some 200 species and twenty DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 265 genera, is found in every Region, and seems to pervade nearly the whole of the earth’s surface, and to extend far north and south. The common Pipistrelle (Vespertilio pipistrellus), ranges into the high north of Europe ; while another species of the same genus (V. magellanicus) inhabits the cold and desolate shores of the Straits of Magellan. The Serotine Bat (V. serotinus) is remarkable as the only species of Bat that is known to inhabit both the Old and New Worlds, being found in North America as far south as Guatemala. It also extends over a great part of the Oriental and Ethiopian Regions. The fourth family of Insectivorous Bats, the Emballo- nuridx, easily distinguished by the tail perforating the inter-femoral membrane or being produced far beyond it, has likewise an extensive distribution, as members of the fifteen genera into which it is divided occur in every one of the six Regions. But these Bats are more abundant within the tropics, and are only feebly represented by a few stray forms in the Nearctic and Palzarctic Regions. Of the two Bats known to occur in New Zealand, one (Mystacops) is a member of this family, and is peculiar to New Zealand. This and another Bat (Chalinolobus morio), belonging to the family Vespertilionide, are the only two indigenous mammals of the New Zealand group, the Rat Mus maorium, often attributed to it, being probably a modern introduction. The last family of Bats, the Phyllostomatidx, or Vam- pires, are entirely restricted to the Neotropical Region, where they form one of the most characteristic groups of mammals, being very numerous alike in species, genera, and individuals. Of the thirty-six genera, with upwards 266 THE GEOGRAPHY OF MAMMALS of eighty species, which belong to this family, the two most remarkable are the purely blood-sucking forms Des- modus and Diphylla, which present an extraordinary modification of their digestive organs, specially adapting them for a diet of blood. These little animals, in fact, are those which have given such a bad reputation to the whole family of Vampires, though it was formerly supposed that the Vampirus spectrum, and other larger forms of tropical America, were likewise sanguinivorous in their habits. But this large Bat is now known, like many other represen- tatives of the same family, to subsist mainly upon fruit, whilst other Vampires feed on a mixed diet of fruit and insects, or on insects only; and others again of the larger forms are said to prey chiefly upon the smaller members of their own order. DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 267 (c) Summary and Deductions as regards the Order Chiroptera. Table of the genera of Bats, showing the approximate number of species found in each of the six Regions. JP eee aceon he 2) eel Sy Sel ees eel Eines g2 || S2|/Se| 25/23) 28) 28) Se a [H8/B IREE |e jes Famity I. PTEROPODID. 1. Epomophorus. OM eel ace tll tre hee tets, lees 2. Pteropus SOM POO ra. eels 0 a0 teem: 3. Pteralopex . 1 LE Weebl | nce el RoBenl NCEE ey Cae Atle oe 4, Cynonycteris . 12 Dl itcett ates lp elas Maat eens | ene 5. Cynopterus PAT oF ikceee | [Bogen |reeanl | lo/e) Mae i] 5b 6. Boneia . ; 2 bs Fie ecisces | Mecca econ | Wee ia leer hace 7. Harpyonycteris . De ec Wetec’ Mectaah eels melon ceemline 8. Scotonycteris . Diprsesce eet) SL x al Petes Ml earns Arete 9. Harpyia ZA a rbd eaeeea beer neal) aco ih ec 10. Cephalotes . ere ae Mates ||| caetecostlleallipese) maa 11. Liponyx DT eset eees ork lnecs Nias 2a |) coe 12. Notopteris . 1 UU coe icaay Mega yl eaanel lt ee 13. Eonycteris . 11 | Boers [eerie Meare eines (eeu Mlle 3 liane 14. Macroglossus . S| ee? TESS Hana coal vee Minscmall abe 15. Megaloglossus Uilewceraliesmet ty ie eieie ete mame tl teeeennet 16. Melonycteris . 1 UN WIRR SN [sneccellerr elton taneee || see 17. Callinycteris . Weil eaieaglienepiltmacseltecocalfemls IMaecialleesine 18. Nesonycteris . 1 1 | Facts |e [Boca Pose Ploas 3p sere Fatty II. DUO Ge es 20M Sade ce las RHINOLOPHID. | 19. Rhinolophus . 26 || 1 8 ihe 4 20, Trieenops A tle, Pal ee 1 21. Rhinonycteris 1 Pee ie Ay ue 22. Phyllorhina 28 || 7 Gilt oles 1 23. Anthops eel ee A a 24, Celops . 1 1 Famity III. 61 || 10 GA es ese 6 NYCTERIDZ. 25. Megaderma Bell 2 26. Nycteris 10 9 268 VESPERTILIONIDZ. 27 28 29 30. 31 32 33 34 35 36. 37 38 39 40 41 42 43 At EMBALLONURID, 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 THE GEOGRAPHY OF MAMMALS (c) Summary and Deductions as regards the Order Chiroptera (continued). Famity IV. . Antrozous . . Nyctophilus . Synotus. . Plecotus . Euderma . Otonycteris . Vesperugo. . . Chalinolobus . . Scotophilus Nycticejus . . Atalapha : . Harpiocephalus . . Vespertilio . Kerivoula . . Natalus . . Thyroptera . Myxopoda . . Miniopterus Faminy V. . Furia ig ee . Amorphochilus . Emballonura . . Coléura . : . Rhynchonycteris . Saccopteryx . Cormura . Taphozous . . Diclidurus. . Noctilio . Rhinopoma . Cheiromeles . Molossus . Nyctinomus . Mystacops . Nearctic Region. | Xe ep: Cd pai* . . . agiaiy Ge VI. Palearctic Region falas. og 39 l@ 18 |2 -|8 lg S383 | 2%|Sm\Sa/Sm Ee S848 | 22/72) ce | 8 [ese Sie EUR ——— eS ee | ope as SWS alesse Se Oe ie oo 1 Bh elec: << 1 |]... | os nee 70 | 41 9/20] 17 90} Bi) All sco ee ele Ayes) 20) eae aie Selecelt So aa os 8 oa | 5 A534) Se) 6) eo Thal) Wa ce) | 10 | 3 || 3 | - | ee ce | 1) ee ae pes 5 || 1 2) 2) God LRA feelin ER aS =| |190|| 19 | 28 | 39 | 5 | 52 | < eee 1 TW kee le MLN ale cocoon are | CAE AR Se Pee A eres Sih sk ale ie leks lee eo 6 Giles 1 || |) oils eee ee TO] va Cs ee 2 || sl Be 2 lisetaSal tet 1 1 etal Pee ote 1 TOV ces ed Opts a 30 || &|-8| 9} 3] 4 il, eae 79: || 22.133) 2154) Salas DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 269 (c) Summary and Deductions as regards the Order Chiroptera (continued). 79. Ischnoglossa 80. Lonchoglossa . 81. Glossonycteris 82. Cheeronycteris 83. Lichonycteris . 84. Artibeus 85. Vampyrops 86. Chiroderma | 87. Sternoderma . 88. Ectophylla. 89. Ametrida . 90. Pygoderma 91. Sturnira : 92. Brachyphylla. .. . OS: ACenbUFIO!. 5) <<) en“ O47 eDesmoduay <5: .4- 4s. 95. Diphylla gi) zs| Fe |22|22| 33| fa] 28 Se) eR SP ae ae] Sh) Se) se Ma| Famity I.' ANOMALURIDZ. 1. Anomalurus rae 9 9 ans WR, Moving. 5 686 o 6 1 Il + 3. Zenkerella. ; 1 il tee 11 1a Famity II, | SCIURIDA. 4. Reithrosciurus .. . 1 opera cee eeoctl leace 1 oe DMPNCLUS Mei, Mie. ie ceirs | Beeaninees Ara lhe eclllbetesraa | Maercr 1 GMOeMTsen eee aie --sn (LOGE \iccsc (ele Wt ses |, GO | 1b 1: °3 ease emia at ose s ere ABOU ct lee kc dleee Ni tee SO Il Baspermopnilus. tyes el 40 Wises ses | Sea | ove |oaee 24 | 16 Se TOMY Ses cy Mo Mcee ithe Wiese | wey la less. Tome Ih ey line OPAL CLOUUVS 2c, Teale ats a |) AON cee sesh ro [esce il aco | 4 |, <6 11. Eupetaurus ee hee ee Ha ed as see 12. Petaurista . 13. Sciuropterus . 14, Nannosciurus. Famity IIT, CASTORIDA. 15. Castor 276 Ig: 18. 19. 20. 21. 22. 23. 24, | 25. | 26. 27, 28, 29, 30. 31. 32, 33. 34, 35. 36. BT 38. 39. 40. THE GEOGRAPHY OF MAMMALS DISTRIBUTION OF RODENTs (continued). Famity LV. HAPLODONTIIDA. 16. Haplodontia . Faminy V. GLIRIDA. Glis . Muscardinus . Eliomys Graphiurus Platacanthomys Typhlomys Faminy VI. MURIDA. Hydromys. Xeromys Celeenomys Chrotomys. Crunomys. . Rhynchomys . Phleomys . Gerbillus Pachyuromys . Meriones Psammomys . Rhombomys . Otomys. Oreinomys. Deomys. Dendromys Limacomys Steatomys . I. Australian Region. II. Neotropical Region a eS WwW CoO bo ee _ CO] ee SS REOFREP BRE DWN OPE HERE =| ined s A Salad =aO|39o aot — 3» oo] SI Be o “Pa | gh _ = = = = = | BON asus te Dr OO: 2 « « Re Ome Bet IV. Oriental Region. | | De eee ett eS V. Nearctic Region. Palearctic Region. | Vat 16 10 DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 277 DISTRIBUTION OF RODENTS (continued). gasy Total Species. I. Australian Region. Region. III. Ethiopian Region. Region. IV. Oriental Region. V. Nearctic Region. Region. IIIa. Mala | | : es Neotropical | | | | ho: Me Palearctic > EO. Ave Malacothrix . . . ./| 1 ORM he io tonel eke) st wy (LOO Bae NGSOKIA) of le i vas} 22 44, Cricetomys 45. Malacomys 46. Lophuromys . 47. Saccostomus 48. Acomys. . 49. Arvicanthis 50. Dasymys 51. Golunda 52. Vandeleuria 53. Chiropodomys 54. Batomys 55. Carpomys . 56. Pogonomys 57. Hapalomys 58. Pithecochomys 59. Lenomys 60. Crateromys 61. Mallomys . 62. Craurothrix 63. Mastacomys 64, Uromys. 65. Conilurus . 66. Lophiomys 67. Cricetus ww oO ce -_ Ore oO oo a ErOOW WH wok OATWHRWeH bo DK Wee: eee eet food by orio ot — Tbe 6 | ees — I Ue. 68. Mystromys : ee 69. Brachytarsomys . 70. Nesomys 71. Hallomys . 72. Brachyuromys 73. Hypogeomys . 74. Gymnuromys 75. Eliurus 76. Onychomys 77. Peromyscus 78. Rhipidomys MOmeR VOM i. at ay eee Sec | yosoed|) Saamll ices S02 Holochilus: -. . ..-. NPE cts, | hat ¢! Sel Pilea Reeth carseat ifort eet tvIR Fat tat tem SSSI ah pear ey ae okie rel SiS o RWS ieee! Msale. Te yee eae ae eae ato me ww FPwWNWOO RE RE POR RP RP ENE DOH HH HP eH He ONDH WH w —_ 278 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. olka 92. 93. 94. 95. 96. 97. 98. 99: 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. Will 112. THE GEOGRAPHY OF MAMMALS DISTRIBUTION OF RODENTS (continued). Megalomys Ligmodon Oryzomys Chilomys . : Reithrodontomys : Eligmodontia Nectomys. Neotomys Reithrodon . Phyllotis . Scapteromys. Ichthyomys . Acodon Oxymycterus Blarinomys . Notiomys. Neotoma . Nelsonia . Xenomys . Hodomys . Phenacomys . Evotomys . Microtus (Arvicola) Synaptomys. . Lemmus (Myodes). Dicrostonyx . Ellobius . Siphneus . Fiber Famity VII. SPALACID A. Rhizomys. Tachyoryctes Spalax . g |8 gi || 28 aE Peas 1 1 fal see 66 | 60 1 1 12 | 3 6 | 6 Roll tf 1 1 5 5 5 5 | 9 2 4 | A 40 || ... | 40 Bees es 1 | ... 1 1 howe 1 SO nant aol Conia teens Gains 19 80 a 4 1 4 5 3 724 | 69 | 162 5 | 3 || 8 16 Ill. Ethiopian Region. IIIa. Malagasy Region. | | | 106! 11 IV. Oriental Region. V. Nearctic Region. Os: Oa: Ww: ae Se TOR QR RrR Ww. 70 e VI. Palearctic Region DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS DISTRIBUTION OF RODENTS (continued). aa A SS (=| S es | @ + Seema ee miner leu loam) Gig re yeh ll es | ee Oo || S$o eS mol ao O5 AS Bem || A a |e. | =o a.O 29 || | S| Ss | | pe | Se efi se|Selgo|ae/o| oe a|| SA} ae | | es | | tl eeretirn latte asl ae |e lS a) il = | = | =! eS i S| : ES ae Faminy VIII. | | GEOMYID. | 113. Geomys Region. | VI. Palearctic HL ILISoproT a) coh AST esata as) | a Aa me Fate Faminy IX. 9 HETEROMYID. | Ibs Dipodomys -- ..-.°+..|| 19 HGS CPeLOdIpUS , aiken | 10 1 710) 117. Microdipodops. . .| {| 1 Ss Rerognathus.-.. -. «| 938 38 moe Heteromys “; .. .. . |) 15 3 12 Famity X. HOa ilies per oua ht st ete BATHYERGID, | | | | n20) Bathyersus: 30.0%... | Pie Georponuss 4 3. )s |) qos |... | 36 122. Myosealops . ... ./| 3/] ... | 3 123. Heterocephalus. . .| 9 | 2 FaMILy XI. IG Bl eee eee ol DIPODIDA. NBL Syeopioyloyiey, 6 5 4 4 AA Mase alll deen Perel erceisc I wieteal wee a EBs “ACTOS 98 eT Sar ane | ete (ata Meee Meal ea a PAG DIGUS Glee tae ed elOs | =i BG PAUACTARAL ne) Gen Xo ul LO! Ili. 128. Platycercomys . . .| 1 || ... 129. Euchoreutes. . . .| 1 |i Faminy XII. SOM eee ll neer beet leh 20 PEDETID&. | ISO, lkawecs Go 3 8 5 i all ae | ee 1 280 THE GEOGRAPHY OF MAMMALS DIsTRIBUTION OF RODENTS (continued). Total Species. I. Australian Region. II. Neotropical Region. III. Ethiopian Region. IIIa. Malagasy Region. IV. Oriental Region. V. Nearctic Region. VI. Palearctic Region | | Faminy XIII. OCTODONTIDA. 131. Ctenodactylus 132. Massoutiera . 133. Pectinator 134. Petromys. 135. Ctenomys. . 136. Acondemys(Schizodon) 137. Spalacopus Se 138. Octodon 139, Habrocoma . 140. Dactylomys . 141. Thrinacodus. 142. Cannabatomys . 143. Loncheres 144, Trichomys 145. Cercomys. 146. Carterodon 147. Mesomys . 148. Echinomys 149. Myocastor 150. Capromys piece | 151. Placiodontia =...) | 152. Pe ae (Aulaco- SETS) We cee gtr thal ARC hier eal) isle ee — —_ EP OH Whee PORE HE DP RHEE ORR DH _ — Pom). etl 0) MOR WORE ER Ore RE DRED tf Famity XIV. HYSTRICIDA. 153. Hystrix 154, Atherura . 155. Trichys FamMIty XV. ERETHIZONTIDA. 156. Erethizon ie ie 2 live aCercolabesies 9) were 9 158, Cheetomys 1 — DISTRIBUTION OF INSECTIVORES, BATS, AND RODENTS 281 DISTRIBUTION OF RODENTS (continued). Faminty XVI. CHINCHILLIDA. 159. Chinchilla 160. Lagidium 161. Lagostomus . Famity XVII. DASYPROCTIDA. 162. Dasyprocta . 163. Ccelogenys Famity XVIII. DINOMYIDZ. 164. Dinomys . Famity XIX. CAVIIDA. 165. Cavia . 166. Dolichotis 167. Hydrochcerus Famity XX. OCHOTONIDA. 168. Ochotona (Lagomys) . Famity XXII, LEPORIDA. 169. Romerolagus. 170. Lepus . _ = =} 4 2/2 2 Sa|og|os BOo|SS |) sg 2p! ow) » &| =o o2 8S “6/246 ) 6 | a Wie AEE | et os | Le | | | 61 i ese 4/10 = nm —_ Sie los eg] asa mo] 29 SS | Ee APl og am oF — = Lond = V. Nearctic Region. DelalOr| de. 20 20 Palearctic Region. [ee 282 THE GEOGRAPHY OF MAMMALS DEDUCTIONS 1. The Order Rodentia is the most numerous of all the chief Mammalian groups, containing nearly 1400 species, which are divided into 170 genera and 21 families. 2. The Rodents are distributed all over the world both in temperate and tropical climes, and are abundant every- where, both in species and individuals, except in Australia where they are poorly represented. 3. The most all-pervading and numerous family of the Rodents is the Muwridx (Mice), which are upwards of 600 in number and are the only members of the group met with in Australia and Madagascar. In Madagascar the Muridxz are represented by a small group of indigenous genera. 4. The Porcupine-like Rodents, of which there are seven families, are specially characteristic of the New World, only the typical Porcupines (Hystricidz) and tive peculiar genera of Octodonts being inhabitants of the New World. 5. Two families of Rodents, the Pocket-gophers (Geo- myide) and the Kangaroo-rats (Heteromyidx), are speci- ally characteristic of the Nearctic Region, the Jerboas (Dipodidx) of the Palearctic, and the Sand-rats (Bathy- ergide) of the Ethiopian Region. : | CHAPTER XII DISTRIBUTION OF HYRAXES, ELEPHANTS, AND UNGULATES SECTION I.—DIsTRIBUTION oF HYRAXES THE Hyraxes and Elephants are nowadays often annexed to the Ungulates, and arranged only as Sub-orders of that great Order, to some members of which they have been shown to be more or less allied by forms of life now extinct. But as, in the present case, we are dealing only with existing mammals, it seems better to give to these two groups their full rank as “Orders,” which they have an abundance of special characters to justify. The Hyraxes, of which, taking Mr. Thomas’s recently published account’ as our guide, about fourteen species belonging to the single genus Hyrax are more or less accurately known, may be regarded as a characteristic form of the Ethiopian Region. As shown by Mr. Thomas’s map (op. cit. p. 58) they are distributed all round the coast of Africa from Senegal through the Cape to Upper Egypt, and also in many places, where they have been searched for, in the interior. Beyond the African continent they extend through Arabia into the borders of Palestine, where the celebrated “coney ” of the Scriptures (Hyrax syriacus) 1 “On the species of the Hyracoidea,” by O. Thomas, P, Z. S. 1892, p. 50. 283 284 THE GEOGRAPHY OF MAMMALS is met with. The Hyraxes are, in most cases, inhabitants of arid rocks, but in other cases are strictly arboreal in their habits. SUMMARY AND DEDUCTIONS AS REGARDS THE OrpDER HyYRAXES =| 3 | . ill Bie Sey ee ies GS -sai/H&| He) S ry ors} || pe! € ||FE| Re Bei oe Belreies = na 2 2) | all I S 8 5 8 un) st oS) 2 a alas 5 | | A) & Te, Mey ras Ys, A ten en | eee ice ieee 14 | aie lips ' 1. The Order Hyraces contains a single genus (Hyraz) forming a single family Hyracide with about fourteen known species. 2. They are found only in the Ethiopian Region, including Arabia and South Palestine. Section I].—DIstRiBuTION OF ELEPHANTS Of this grand form of animal life, formerly much more abundant on the earth’s surface, there remain in the present epoch but two species, one of which is a character- istic form of the Ethiopian Region and the other of the Indian Region. The African Elephant, which, besides its external peculiarities, should be referred from the structure of its teeth according to some authorities to the sub-genus Lowodon of the paleontologists, was formerly found in suitable spots all over the continent of Africa from the Sahara and Upper Nubia down to the Cape. In these days it has in most places been driven by the sportsmen HYRAXES, ELEPHANTS, AND UNGULATES 285 and hunters for ivory far into the interior, but is still to be found in enormous herds in some of the more remote localities of Africa. In the Cape Colony the only spot where it is said still to exist is the forests of the Knysna. The Indian Elephant (Hlephas indicus) inhabits the forest-lands of British India, Ceylon, Burma, the Malay Peninsula, and Sumatra. Its occurrence in Borneo in a wild state has not yet been certainly ascertained. In India, according to Mr. Blanford, Elephants are still found wild along the base of the Himalayas, also in the great forest-countries between the Ganges and the Kistna, in the Western Ghats, and in the forest-clad ranges of Nagpore, but in former times their range was naturally much more extensive. Attempts have been made to separate the Elephants of Ceylon and Sumatra from the continental form as different species, but though there are some grounds for so doing, the distinctions have not been satisfactorily established. The Indian Elephant may be regarded as a characteristic form of the Oriental Region, as the African Elephant is of the Ethiopian. Although we are here only dealing with species of mammals actually in existence, it should be borne in mind that the Mammoth (2. primigenius) has only compara- tively recently ceased to exist on the earth, as is proved by the frozen carcasses of this Elephant that have been exhumed in the tundras of Northern Siberia, and by the enormous abundance of its fossil teeth, which are, even at the present day, a recognized article of commerce. The Mammoth had a very different distribution from the two existing Elephants and was essentially Palearctic in its range, although it appears to have extended across Behring Strait into Alaska. 286 THE GEOGRAPHY OF MAMMALS SUMMARY AND DEDUCTIONS AS REGARDS THE ORDER PROBOSCIDEA || : A r—| es cH S| =| A =) ie | a|.3|.3| 49/3] ei 5 | S$ ||HAlEeo| BS] 4 & SPelce| rs | 6 || SI/Fala el aes|eo| Fa] & is | a ac] + = S 8 al =! 5 | Ss s I o = | = D ~ Ss (=) vA co Ball pa | bo ee ee 1. lephags (4p <2 etl ede le 2 Ailes tal areal ee 1. The Order Proboscidea contains only the genus Elephas (forming a family Elephantidze) with two existing species. 2. One of these belongs to the Oriental and the other to the Ethiopian Region. Section II].—GENERAL DISTRIBUTION OF THE UNGULATES The Ungulates which we now arrive at, and which constitute the ninth order of mammals according to the arrangement here adopted, contain the greater number of the largest and most highly developed forms of the whole class of mammals and embrace in their varied series nearly all the animals (such as the Horse, Sheep, Goat, Ox, Deer, Camel, and Pig) which are most useful to man- kind, although we cannot always recognize the original stocks from which the domesticated forms of these animals have descended. The mode of the distribution of these Mammals over the earth’s surface is, therefore, of special interest, and we must say something about each of the groups into which they are usually divided in classification. The 300 species of Ungulates usually recognized constitute saan el HYRAXES, ELEPHANTS, AND UNGULATES 287 about seventy-one genera which are referred to thirteen families. Some of these families, however, have only one or two species at present existing to represent them, and the great mass of Ungulates, taking them as a whole, belong to the Ox-family, Bovidx, which contains about 200 species. Section IV.—DIsTRIBUTION OF RHINOCEROSES We will begin the Ungulates with what is called by naturalists the Perissodactyle section, which, although abundant in former ages, is represented in the present state of the earth’s fauna only by three distinct types forming so many families, the Rhinoceroses, the Tapirs, and the Horses. In each of these families there remain a few species only, the relics of a vast number of ancestors which have preceded them. Of the existing Rhinocerotidz only five well-ascertained forms are known, two of which belong to Africa and three - to India, although many other species have been suggested upon more or less sufficient evidence. The Rhinoceroses of the Ethiopian Region belong to quite a distinct section of the genus from the Oriental form. These two animals commonly, though not very correctly, called the “Black” and “ White” Rhinoceroses, are best distinguished by the shape of the upper lip, which in the “ Black” Rhinoceros is long, pointed, and extensile, and in the “White” Rhinoceros is short, square, and truncated. The Short- lipped Rhinoceros (Rhinoceros simus), now nearly extinct, has never been met with north of the Zambesi River, which forms the northern boundary of many of the peculiar mammals of the Cape district. On the other hand, the 288 THE GEOGRAPHY OF MAMMALS Black Rhinoceros (R. africanus) extends from the Cape all up the Eastern side of Africa into the plains of the Atbara and Upper Nile. The Oriental Rhinoceroses are three in number. Two of these have only one horn on the nose, while the third is provided with two of these appendages. Of the former the large Indian Rhinoceros (R. wnicornis) appears to be confined to the North-eastern provinces of the Indian Peninsula, whilst the smaller one-horned form (R. sondaicus) ranges from the Sunderbunds of Bengal through the Malay Peninsula down to Java, Sumatra, and perhaps Borneo. The third Oriental species, the Sumatran Rhinoceros (2. swmatrensis), has nearly the same range as the last-named species, but appears to extend rather farther north. Thus we may consider the existing Rhinoceroses as typical forms of the Ethiopian and Oriental Regions, but not to be met with in any other part of the world’s surface. SEcTION V.—DISTRIBUTION OF TAPIRS The second family of Perissodactyle Ungulates, the Tapirs, has a still more remarkable distribution. Out of the five known species four belong to the Neotropical Region, while the fifth, which in some respects is more closely allied to one of the American Tapirs than the American Tapirs are to one another, is an inhabitant of the Oriental Region, being met with only in the Malay Peninsula and Sumatra. This is a good instance of the rare phenomenon of “discontinuous distribution” which, how- ever, may be explained by the fact known from geology HYRAXES, ELEPHANTS, AND UNGULATES 289 that the Tapirs were formerly a prevalent group over a large portion of the earth’s surface, so that in these days we have only to deal with a few scattered remnants of a former considerable series. Of the American Tapirs two (Tapirus bairdi and T. dow?) are found in Central America, a third (7. roulinz) occurs in the Andes of Colombia and Peru, and a fourth (T. americanus) is widely distributed over the South American continent from Venezuela to Paraguay. The Tapirs may be therefore considered as a char- acteristic form of the Neotropical and Oriental Regions, and a “lipomorph” or absent form in all other parts of the world’s surface. SEcTION VI.—DISTRIBUTION OF HORSES The third family of Perissodactyle Ungulates comprises the Horses (Zquidx), now a very isolated group, although allied to the Tapirs by many extinct intermediate forms. The Horses at present known to exist in a state of nature belong to about nine species, of which three may be attributed to the Palearctic Region and six to the Ethiopian. Among the Palearctic species the recently discovered Equus prjevalskii of the deserts of Central Asia is the sole living representative of the typical section of the genus Hquwus with callosities on both the fore and hind limbs. The other eight species all belong to the Asinine section, with callosities upon the hind limbs only. Unless it shall turn out to have been Hquus prjevalskiz, the exact 1 On the species of Horse, consult Sir William Flower’s ‘‘ The Horse” (London, 1891, Kegan Paul & Co.). i 290 THE GEOGRAPHY OF MAMMALS progenitor of our domestic Horse is extinct, but it was in all probability of Palearctic origin. The two Asses of Asia are the Kiang of High Tibet (Equus kiang), which is a larger animal clad with a thick coat of fur in winter, and the smaller, more sandy-coloured and thin-coated Onager (Hquus onager), which occurs in many parts of the deserts of Western Asia and intrudes into the Oriental Region in Cutch. Passing on to Africa we find two members of the Asinine section still wild in the North-eastern part of that continent. These are Hquus teniopus of the deserts of Nubia and £. somalicus of Somaliland. The former of these was probably the origin of our domestic Ass (Hquus asinus). Going farther south- wards into Africa we meet with four distinct species of the beautifully striped Asses commonly called Zebras, viz. (1) the Quagga (HZ. quagga) of the Cape Colony, now nearly, if not quite, extinct ; (2) the Mountain Zebra (Z. zebra) also confined to Africa south of the Zambesi, and now becom- ing extremely rare; (3) Burchell’s Zebra (LZ. burchellv) dis- tributed under slightly varying characters from the Transvaal to British East Africa along the eastern portion of the continent; and (4) Grévy’s Zebra (2. grevii) of southern Abyssinia and Somaliland. The Horses (Hquidz) of the present epoch may, there- fore, be regarded as characteristic of the Palearctic and Ethiopian Regions. Section VIL—DistrisuTION OF BovINE MAMMALS At the head of the great Arctiodactyle section of the Ungulates, which we now enter upon, we meet with the HYRAXES, ELEPHANTS, AND UNGULATES 291 numerous and important family Bovide, to the variety and extent of which in the present day we have already alluded. The Bovidx in fact contain nearly two-thirds of the species of Ungulate animals now existing on the world’s surface, and embrace at least 200 species belong- ing to forty-five distinct genera, amongst which are the Sheep, Ox, and Goat, the animals of which the flesh is mostly used for food by civilized man. According to the arrangement of Flower and Lydekker, the Bovidz are divisible into about ten sub-families, the seven first of which embrace the mammals commonly known as Antelopes. These are mostly met with in the more open districts of the Ethiopian Region, where in former days they roamed about in exuberant multitudes, but have been sadly diminished at the present time by the persecutions of the sportsman and the hunter. We have not space here to go into the numerous and varied forms of Antelopes, but must refer our readers who wish for special information on that subject to “The Book of Antelopes” now in process of publication. It must suffice to say that the roll of Antelopes numbers some 150 species, of which 9 are attributable to the Palearctic Region, 4 to the Oriental, and 135 to the Ethiopian Region. Three of the Oriental species belong to peculiar genera restricted to that Region, and the fourth is a Gazelle, a member of a genus which is also well represented in the Palearctic and Ethiopian Regions. It is therefore evident that the Antelopes, although slightly represented elsewhere, form one of the most predominant and charac- teristic features of the Ethiopian Region. The Rupicaprine, forming the eighth sub-family of Bovidx, and containing what are commonly called the 292 THE GEOGRAPHY OF MAMMALS Mountain-Antelopes, have a very different distribution. They are absolutely unknown in the Ethiopian Region, and are found mostly in the Palearctic Region with stray species in the Oriental and Nearctic Regions. The well- known Chamois (Rupicapra) is the typical form of this group. It is confined to the western portion of the Palearctic Region, and is the single species of the genus. In Eastern Palzarctica it is represented by the genera Cemas and Nemorhexdus, some species of which occur also on the higher mountain-ranges of the Oriental Region. In the Nearctic Region the Rocky Mountain Goat (Haplocerus montanus) is the sole representative of the Mountain-Antelopes. The Goats and Sheep (Caprinz), which follow next, have nearly the same sort of distribution. It should be mentioned that the distinctive differences between the Goats and Sheep, from a structural point of view, are very difficult to define; and that the two forms are so nearly allied that it has been proposed by some naturalists to unite them into one genus. The Caprinx altogether, although by no means satisfactorily worked out at present, may be held to embrace some twenty-six species, of which nineteen are Palearctic, two are Nearctic, five are Oriental, and one species only (Capra walie of the high ranges of Abyssinia) occurs within the confines of the Ethiopian Region. The Caprinx are represented in the Nearctic Region by two, or possibly three, species of Sheep, which extend from Alaska, along the main range, nearly down to Northern Mexico. Finally closing the long list of the family Bovide we have the oxen, or typical Bovinz, embracing about twelve or thirteen species, and thinly distributed over the HYRAXES, ELEPHANTS, AND UNGULATES 293 Palearctic and Oriental Regions, but with certainly one Nearctic representative, and two, or possibly three, in the Ethiopian Region. The true origin of our domestic cattle (Bos), is lost in obscurity, but is usually attributed to Palearctic ancestors, although the progenitor of the humped form, or Zebus, may have more probably been Oriental. The two Bisons belong, one to the Palzearctic and one to the Nearctic Regions, and are very nearly allied. The Yak (Poephagus) is confined to the higher mountain ranges of Central Asia, and must be therefore reckoned as Palearctic. The characteristic Oriental forms of bovine animals are the Bantengs (Bibos), of which sub- genus three species are commonly recognized. The Ban- tengs extend from the mountains of Northern India through the ranges of the Malay Peninsula into the Sondaic Islands. Of the Buffaloes (Bubalus) three species are commonly recognized, one of which is Indian, the parent of the well-known domestic form, while two, or perhaps three, other nearly allied species are peculiar to Africa. Finally, closely allied to the Buffaloes, and hardly distinguishable from them, we have the little Anoa of Celebes, and the lately discriminated Bubalus mindor- ensis of the Philippine Islands. Section VIII.—DistrisuTION oF PRONGBUCKS The second family of Artiodactyle Ungulates embraces only a single species strictly confined to the Nearctic Region, and constituting one of its most peculiar types. This is the Prongbuck (Antilocapra americana), ot the western prairies of the United States, which (as was first 294 THE GEOGRAPHY OF MAMMALS ascertained in this country) carries hollow horns, like those of the Bovidz, but sheds them regularly every year, like the deciduous antlers of the Stags. SECTION [X.—DISTRIBUTION OF GIRAFFES The next family of Arctiodactyles, the Giraffide, consists likewise of what is called a monotypic form, embracing a single species only, and limited, in the present epoch, en- tirely to one Region of which it is one of the most charac- teristic animals. The Giraffe (Girafia camelopardalis) is distributed over the greater part of Africa, from Senegal through the Southern Sahara to Upper Nubia, and thence southward throughout Eastern Africa in suitable localities to the Cape. So far as we know at present it is not found within the great wooded regions of the Congo valley and western coast, but is certainly met with on the Niger and on the Upper Gambia. As has recently been well shown by Mr. de Winton (P. Z. S. 1897, p. 275) the Giraffe presents us with two geographical forms, which may be designated either as species or sub-species. The Northern Giraffe (Girafia camelopardalis typica) which is met with in Senegal and thence across the Sahara to Upper Nubia, Somaliland, and British East Africa is distinguished from the Southern Giraffe (Girafia camelopardalis capensis) by several characters, especially by the great prominence of the third frontal horn which is barely apparent in the southern Giraffe. The Cape Giraffe seems to be met with in suitable localities all up the east coast into British Kast Africa, where it is stated that the two forms meet. More accurate information about the two forms of Giraffe HYRAXES, ELEPHANTS, AND UNGULATES ~° 295 is, however, highly desirable, and the areas of the two sub-species require to be carefully studied. SECTION X.—DISTRIBUTION OF DEER The seventh family of Ungulates which we now come to, though it cannot rival the Bovide, is likewise of importance in an economic point of view, the flesh of nearly all the Deer-tribe forming an acceptable food for mankind. In our survey of the deer we shall follow as nearly as possible Mr. Lydekker’s “Deer of all Lands,” the most recent and best authority on the subject. Mr. Lydekker acknowledges eleven genera of this family, containing altogether nearly sixty species. Of these genera ten belong to the typical Deer and one to the Musk-deer (Moschus). « Deer are found in all the six Regions of the world except the Australian and Ethiopian. Their absence in Australia, as is the case with nearly all the highly organized groups of mammals, can be easily understood, but it is difficult to imagine why there should be no Deer in the Ethiopian Region, when in the New World they have passed so abundantly into South America. We are not aware that any explanation can be given of this anomaly, unless it be that their place is taken by the Antelopes. Commencing with the Reindeer (Rangifer) and the Elk (Alces), we find these two types restricted to the northern portions of the Palearctic and Nearctic Regions. Whether it is possible to recognize more than one species of each of these forms is a matter of doubt. At any rate all the local races of both the genera are closely allied. 296 THE GEOGRAPHY OF MAMMALS The true Deer (Cervus), which we come to next, form the most numerous genus of the family, being about twenty-two in number. They are about equally divided between the Palearctic and the Oriental Regions, with one representative, the Wapiti (C. canadensis) in the Nearctic Region. This occurrence is exactly in a line with that of the stray species of Sheep (Ovis), Bison (Bos), and Mountain Antelope (Haploceros) in the same Region, but is not sufficient, supported though it may be by other similar facts, to convince us of the necessity of recognizing a “Holarctic” Region. It should be stated also that typical Cervus is essentially characteristic of the Palzearctic Region, throughout which (with the above-mentioned exception of the Wapiti) the members of this splendid group of animals are distributed, whilst the sub-genera Awis, Rusa, and Rucervus take its place in the Oriental Region. ‘ The Muntjacs (Cervulus) embrace about five species essentially Oriental, although two of them intrude within the bounds of the Palearctic Region. In the curious form Llaphodus we meet with a Deer with a tufted head which nearly conceals its minute antlers, restricted to the northern portion of the Palzearctic Region, and leading us on to the Water-Deer (Hydrelaphus) of Southern China, in which the antlers are absolutely wanting. Of the Roes (Capreolus) there seem to be three local races, which are often considered as so many species. These allied forms range over the whole extent of the Palearctic Region. We now come to that strange animal the Milou (Elaphurus) which has been sometimes associated with HYRAXES, ELEPHANTS, AND UNGULATES 297 the true Cervi, but which, as has been recently pointed out by Mr. Lydekker, possesses essentially distinctive characters in its remarkable horns, large and spreading hoofs, and long tail, besides other peculiarities. Mr. Lydekker is of opinion that the genus “has nothing to do with any of the living Old World Deer except the Roes, whilst its alliance with the American Deer (Cariacus) seems to be close.” If such be the case its patria is indeed remarkable, for, though only yet certainly known from captive specimens obtained in the Imperial Park, near Pekin, it is said to have been originally brought from Kashgaria, and must therefore be a Palearctic form. The second section of the sub-family Cervine contains the American Deer of which two genera are usually recognized—Cariacus! and Pudwa—containing altogether at least twenty or twenty-one species, which are distributed throughout the Nearctic and Neotropical Regions. As all naturalists agree they form a very natural group, connected by many common characters which separate them from all the existing Deer of the Old World, although the fossil genus Anoglochis, formerly found in Western Europe, seems to have been a closely allied form. As in the case of Cervus the sub-divisions of Cariacus have special areas of distribution, Dorcelaphus being the most northern form, whilst Blastocerus, Xenelaphus, and Subulo take its place in South America. The small Deer 1 We regret not to be able to follow Mr. Lydekker in using the generic term Mazamua for this group. Mazamais a term which has already been most carefully studied and condemned by the late Professor Baird (“North American Mammals,” p. 665) to be rejected for vagueness. Its author, Rafinesque, seems to have been an erratic person whose writings should be held to be of no authority whatever, and it is very doubtful whether any of his names, which were mainly based on the phantoms of his own imagination, should be employed in Science. 298 THE GEOGRAPHY OF MAMMALS of the genus Pudua, distinguished from Cariacus by its remarkable foot-structure, consists of two species, one from Chili and Western Patagonia and the other from the high Andes of Ecuador. The Musk-Deer which constitute a second well-defined sub-family of the Cervidw and should perhaps even be recognized as a different family, contains only the genus Moschus, with two species, which are restricted to the Palearctic Region. SEecTION XI.—DISTRIBUTION OF CHEVROTAINS The Tragulidez, or Chevrotains, consist of a few animals of small size, often known as Moose-Deer, which are intermediate in structure between the Deer, the Camels, and the Pigs. There are only two known genera of these animals at present existing, of which one (7ragulus), with about five species, belongs to the Oriental Region and the other (Hyomoschus), with a single species, is peculiar to Western Africa. The latter form is closely allied, if not identical with the extinct Dorcatheriwm of the tertiaries of the Old World, and is placed by some authorities in the same genus. Other extinct forms of small Ungulates serve to connect the Chevrotains, in former epochs, with the Deer. SEcTion XII.—DISTRIBUTION OF THE CAMELS The forms of the Camel family now existing are two only, the true Camels of the Old World and the Lamas of the New. These are now separate in structure as in HYRAXES, ELEPHANTS, AND UNGULATES 299 locality, but seem to be alike descendants of a group of extinct Camel-like ancestors formerly found in North America. The two species of Camelus now living are the One-humped Camel (C. dromedarius) and the Two-humped or Bactrian Camel (C. bactrianus) both of which are now best known in a domestic state. Indeed the original home of the One-humped Camel has not yet been certainly ascertained, although it is usually supposed to have been Arabia, where wild Camels are said to have existed about - the commencement of the present epoch. We may there- fore, perhaps, class the Arabian Camel as an Ethiopian type. But the true home of the Bactrian or Two-humped Camel is certainly the great deserts of Central Asia, where specimens of the wild species have been obtained both by Russian and English explorers. We may therefore place the Bactrian Camel as a Palearctic animal. The two wild species of Lama—the Huanaco and Vicuna—are found only in the temperate portions of the Neotropical Region; from one or both of these are descended the Lamas and Alpacas of domesticity. The Lamas may consequently be classed as an indigenous form of the Neotropical Region. Section XIIJI.—DIstTRiBUTION OF SWINE-LIKE UNGULATES At the end of the great Ungulate series we come to the Swine or Swine-like mammals, of which the existing forms are usually assigned to four separate families, the Hippopotamuses, Wart-Hogs, true Swine, and Peccaries. Most of these forms, except the Swine, have in these days 300 THE GEOGRAPHY OF MAMMALS a very limited distribution, but in former days the whole series was connected, in locality as in form, by hosts of ancestors now extinct. The Hippopotamide, or Hippopotamuses, formerly widely-spread over the whole world, have now only two surviving species, the larger Hippopotamus amphibius, which is met with in nearly all the great African lakes and rivers, and the smaller Liberian Hippopotamus (H. libertensis) which has hitherto been found only in one of the rivers of Liberia. As regards the existing creation, therefore, this peculiar form of Ungulates must be re- garded as strictly Ethiopian. The second family of Swine-like Ungulates—the Wart- hogs (Phacocheridx)—is also entirely confined to Africa, where two species are widely distributed from Upper Nubia, throughout Eastern Africa, down to the Cape Colony. The true Suidx, or Swine, to which the Wart-hogs are indeed closely allied, embrace three genera—Sus, Pota- mocherus and Babirussa. The typical Swine (Sus) are found in the southern part of the Palearctic and the Oriental Regions, extending from Southern Europe, through Western Asia into India and the islands of the Indian Archipelago. It is difficult in many cases to ascertain what are the real wild species of this group, the domestic forms having varied much under domestication for many ages and having been carried by man all over the world. It is probable that the Swine of New Guinea—the so-called Sus papuensis—and those of other Eastern islands may be descendants of domestic or semi-domestic animals. In the Ethiopian Region the place of Sus is taken by the River-hog (Potamocherus), with a slightly different HYRAXES, ELEPHANTS, AND UNGULATES 301 dentition. Two or perhaps three distinct species of this genus have been generally recognized, but the recent researches of Dr. Forsyth Major (cf. P. Z. S. 1897, p. 359) tend to show that we are by no means yet well acquainted with the species of this difficult group. The third genus of Suid, the Babirussa, remarkable for the peculiar shape of its four tusks, exists only in the far-away island of Celebes and (perhaps introduced) in the adjacent island of Bouru. It must, therefore, be attributed to the Oriental Region, to which, we believe, we have now definitely shown (see above, p. 146) that Celebes must be referred. The fourth and last family of Swine-like Ungulates con- tains only the Peccaries (Dicotyles) of the New World. The two species belonging to this genus range from Texas and the Southern United States down to Patagonia; and must be regarded as purely Neotropical in their geographical classification, although one of the species—the Collared Peccary—undoubtedly intrudes slightly within the boun- daries of the Nearctic Region. 302 THE GEOGRAPHY OF MAMMALS SEcTION XIV.—SuUMMARY AND DEDUCTIONS Table of the genera of the Order Ungulata with the num- bers of species belonging to the great Zoological Regions. a) el" ee alee @ | He|HE/HE SEER) Hel Eg a Z a4 a iG a o Se af [iS |p el a neat Famity I. RHINOCEROTID A. 1. Rhinoceros) t=. ve SO al, Ne 2 ae Famity II. TAPIRIDA. Ls Tapirusis: ts 1 ore eae One| Aas Ole eee ee Famity III. EQUID. My quis: sn... Sh oii a) ae, | sean)» Gol ieee ol acre Famity IV. | | BOVIDA. | Sub-family I. | BUBALIDIN&. le Bnlbalis.c yee cess eel eae oy DeDamaliscus: «tee 3) Sele aod if 3; Connochetes = 3 i agir Ze iliese oa Sub-family II. CEPHALOPHINA. | 4, Cephalophus. . . . | 38 || ... 38 Ay D: MHOMACenOs .) Meee % ae |) Lee ate 1 Sub-family III. NEOTRAGINA. | 6. Oreotragus. 1 1 | 7 Ourebia. 5 5 ', 8. Raphicerus 2 2 9, Nesotragus. 1 1 10. Neotragus . ihe 1 11. Madoqua 6 | 6 bis HYRAXES, ELEPHANTS, AND UNGULATES 303 SUMMARY AND DEDUCTIONS (continued). = E g @| &| 3 | 2 z SE |Re RS SB ESF els = Goh eee ae ate i Mars cS Blea Tea ple silCN aa Sub-family IV. | CERVICAPRIN®. | PeCobuse go ee TOs ts esl 2: HGeRVviCH pra. 4 ses Ae Be Wl css ft ae [8 Wipiclentann Glan re hy tallccral sco OL lees.) |ineee Sub-family V. | | ANTILOPIN &. | 15. Antilope 1 zee 1 ee 16. Aipyceros . 2 2 koe 17. Saiga. 1 1 gol 18. Pantholops 1 | Pos 1 19. Antidorcas. elie il 1 Bt DOmGazellam ss... 6 (025: | ily it 7 21. Ammodorcas . 1 1 22. Lithocranius . 1 1 | 23. Dorcotragus 1 1 Sub-family VI. HIPPOTRAGINA. 24, Hippotragus . 4 4 PoeOrye. 6. 5 5 | 96. Addax . ss if Dele scactiens Ane Sub-family VII. | TRAGELAPHINZ. | 27. Boselaphus il 1 28. Tragelaphus 8 8 Aa | 29. Strepsiceros 2 2 mos 30. Orias ; 3 3 Sub-family VIII. RUPICAPRINA. | 31. Rupicapra . 1 | 1 eoi@emas: .. « 4 heal 3 33. Nemorhedus . 5 Tol iat lees 34. Haplocerus 1 IS ee | 35. Budorcas 2 | 2 304 SUMMARY AND THE GEOGRAPHY OF MAMMALS DEDUCTIONS (continued). Sub-family IX. CaPRIN&#. Capra Hemitragus Ovis . Ovibos . Sub-family X. Bovin&. 36. 37. 38. 39. 40. 41. | 42. 43. 44, | 45. Bos Bison Poephagus . Bibos Bubalus. Anoa. Famity V. ANTILOCAPRIDA. 1. Antilocapra Famity. VI. GIRAFFIDA. 1. Giraffa . Famity VII. CERVIDAi. (a) CERVINA. . Rangifer Alces Cervus . . Cervulus . Elaphodus . Hydrelaphus . Capreolus . . Elaphurus . . Cariacus . Padua (b) MoscHIN&. . Moschus OOODID AP wre — — — ho — Ww Or We dS OO ee bo | | ie Australian. II. Neotropical. Net eal 10] 1 Bh ones 4 e| 22/2] $| m | 8s oes) S Fa. | et & Sissi RS|ae| Fs ral fe 3 m 5 s ° A | 1 Ae ese 500 Bio ee ee — VI. Palearctic. 23 HYRAXES, ELEPHANTS, AND UNGULATES SUMMARY AND DEDUCTIONS (continued). Faminy VIII. TRAGULIDA. 1. Tragulus 2. Hyomoschus . Famity IX. CAMELIDA. 1. Camelus 2. Lama FamILy X. HIPPOPOTAMIDA. 1. Hippopotamus Famity XI. PHACOCHGRID. 1. Phacocheerus . Famity XII. SUID. 1. Sus E 2. Potamochcerus 3. Babirussa . Faminy XIII. DICOTYLIDA. 1. Dicotyles — So re or Australian. II. Neotropical. 1a, Ethiopian. > a ag < = S] S Bae) ae | P = Ss 5 5 500 4 1 006 : 1 Nearctic. 305 VI. Palearctic. U 306 THE GEOGRAPHY OF MAMMALS DEDUCTIONS 1. The Order Ungulata contains about 306 known species, arranged in seventy-one genera and thirteen families. 2. Ungulates are found all over the earth except in the Australian Region. In the Malagasy Sub-region there is but one species, which may possibly be a recent intro- duction. 3. Ungulates are most abundant in the Ethiopian Region, where 156 species occur. Of the great family Bovidee 138 out of 200 known species are restricted to this Region. 4, There are no Bovine Ungulates in the Neotropical Region, and only five (all nearly allied to recent or extinct Palearctic forms) in the Nearctic Region. 5. In the Neotropical Region there are representatives of only four out of the thirteen families of Ungulates— Tapirs, Deer, Camels, and Peccaries. 6. There are no Deer (Cervidz) in the Ethiopian Region. 7. The New World has only 38 species of Ungulates against 270 found in the Old World. eS OU 09 Famity III. 1. Platanista 2. Inia ; 3. Pontoporia . PHYSETERID. PLATANISTIDA. Number of Species. 1 | Tropical and Sub-tropical Seas. 1 | Widely distributed. Dy) | North Atlantic. 2 Widely distributed. 6 | Widely distributed. 1 | Antarctic and Pacific. 3 1 Rivers Ganges, Brahmaputra, and | Indus. | Upper Amazon. . 1 - | 1 Estuary of La Plata. 3 DISTRIBUTION OF THE CETACEANS AND SIRENIANS — 18. 19. . Pro CO CO ~TD rip Ww bO GENUS. Faminy IY. DELPHINIDZ. Sub-family I. DELPHININ«. . Sotalia. Steno . . Tursiops . Delphinus oe Tursio . C Lagenorhynchus . Sagmatias be Deresa~. . < : : Cephalorhynchus : . Neomeris . . Phoceena . . Orcella . Grampus. . . Globicephalus . . Pseudorca . Orca Sub-family IL. DELPHINAPTERIN-. Delphinapterus. Monodon . Number of Species. Dp We Ree ON Of Ob © HH Oe 309 GENERAL DISTRIBUTION. China, Indian Ocean, South America (partially fluviatile) Indian Ocean and South Atlantic. Widely diffused. Widely diffused. Widely diffused. Antarctic and Pacific Ocean. Widely diffused. Unknown. Antarctic Ocean. South Seas. Indian Ocean. | Atlantic and Pacific Ocean. Indian Ocean and Trawnddy River. Atlantic and Pacific. Widely diffused. Widely diffused. Widely diffused. Arctic Seas. Arctic Seas. 310 THE GEOGRAPHY OF MAMMALS DEDUCTIONS 1. The Order Cetacea contains from eighty to ninety known species, belonging to about forty-four genera and four families. 2. Cetaceans are found in all seas from the Equator to within the Arctic and Antarctic Circles, and in some of the larger rivers (the Ganges, Brahmaputra, Indus, and Irawadi in the Old World, and the Amazon and La Plata in the New). 3. The species of Oceanic Cetaceans are mostly very widely distributed, especially the Delphinide, but in some cases are local, some species being confined to the Arctic and Antarctic Seas respectively, and some being peculiar to the Pacific and to the North Atlantic. 4. The Fluviatile Dolphins proper constitute a family of themselves (Platanistidx) with a very singular distribu- tion, one genus being restricted to the rivers of India, and two others to those of South America. 5. Besides the Platanistidze some of the Delphinidz are found in rivers, such as Orcella flwminalis in the Irawadi, and Sotalia tucuwi (with perhaps others of the same genus) in the Amazon. 6. None of the great lakes of any continent is known to be inhabited by Cetaceans. CHAPTER XIV DISTRIBUTION OF EDENTATE MAMMALS Section I.—IntTrRopuctory REMARKS As is the case with most ancient groups, we find the various types of Edentate Mammals strictly limited to certain localities on the earth’s surface, so that the study of their distribution, especially when taken in connec- tion with what we know of the extinct forms of the same group, comes to be a matter of much interest. In the present case, however, we propose to . confine our remarks to the existing Edentates, as we are dis- cussing the distribution of recent and not of fossil mammals, and merely to allude to the extinct forms when necessary. As regards the existing Edentates, as has been well shown by one of our leading authorities on mammals, those of the Old World and those of the New are essen- tially distinct. The two Old World families commonly assigned to this order, are so different in important points of structure from the American families, that it may be even considered doubtful whether they were derived from the same primary branch of mammals. We will, there- fore, take the two groups separately, and begin with the forms of the New World. 311 312 THE GEOGRAPHY OF MAMMALS Section IJ].—DIsTRIBUTION OF THE NEw WorRLD EDENTATES The Edentates of the New World are at present repre- sented by three families, the Sloths (Bradypodidz), the Ant-eaters (Myrmecophagidz), and the Armadilloes (Dasy- podidz), all well distinguished from each other, although essentially modified on the same plan of structure, and more or less united together by other Edentates now extinct. All three families belong entirely to the Neo- tropical Region, although one of the Armadilloes appears to have intruded itself farther north than the generally recognized northern boundary of that Region. Of the Sloths, two genera are well established—the Three-toed Sloths (Bradypus), with about five species, and the Two-toed Sloths (Cholopus), with two species. The Sloths are entirely arboreal in their habits, and are met with only in the dense forests of Central and Southern America, from Nicaragua down to Bolivia and Southern Brazil. The Ant-eaters, of which three well-marked forms are known, belonging to so many genera, each with one species, have a somewhat wider distribution, being not absolutely confined to the tropical forests. The Great Ant-eater (Myrmecophaga jubata), extends as far north as Guate- mala, and from Costa Rica southwards is found in suitable localities all through Venezuela, Guiana, and Brazil to Paraguay. The Tamandua Ant-eater (Tamandua tetradactyla), which varies much in colour and markings, extends even farther north than its larger brother, as specimens have 2 DISTRIBUTION OF EDENTATE MAMMALS cules been obtained in Southern Mexico. Southwards it passes throughout South America as far as Paraguay, where Reng- ger assures us it is common, and widely distributed. The little Two-toed Ant-eater (Cyclothurus didactylus), a purely arboreal form, is also found in Guatemala, Nicaragua, and Costa Rica, but does not range so far south as the two former species, though it extends throughout the great Amazonian valley into Peru. The Dasypodidx, or Armadilloes, which contain the third family of American Edentates, are more numerous and more diversified in their characters than the two pre- ceding families. The eighteen or twenty species generally recognized by naturalists may be divided into four sub- families and seven genera. The general area of their distribution is rather larger than that of the Sloths and Ant-eaters. One Armadillo, as has been already men- tioned, goes as far north as Texas, and Armadilloes are found all over the Argentine Republic down to Patagonia. The most remarkable of the Armadilloes, and one that must form a sub-family by itself, is the little Pichy-ciego of the Argentines (Chlamydophorus truncatus), which is found in the sandy plains of the vicinity of Mendoza, and also, as has been recently ascertained, near Bahia Blanca in the eastern part of the Argentine Republic. 35. net. Psychical Research Society, Proceedings of : ; Vol. I. containing Parts 1, 2, 3, 4. 10s. ” Il, ”? » 5,6, 7. 0s ” Ill. 2» »5 8,9. TOs As av — 58 20, 01s! 8s ” ” ” 12, 13, 14. Ios. 9 VE » 53. Lh sil Osh 7 ne Ose ap WA - ap LO sa Os 2Onmmo ss e \QUI AG pel, 224285 TOs. a Xe nn a>) 245025. cOS ” Xx. ” » 26, 8s ed XI. a Be SOs LOS. 2 ROWE 55 ESO OS: as POET 2G a eS eeoe ok2s: Pulpit Commentary, The (Old Testament Series). Edited by the Very Rev. Dean H. D. M. SPENCE, D.D., and the Rev. J. S. EXELL. Super royal 8vo. Genesis, by the Rev. T. es D.D. ; Homilies by the Very Rev. J. F. MontcomEry, D.D., Rev. Prof. R. A. REDFORD, Rev. F. Hastincs, Rey. W. 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