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and all the more modern geographical text-books give at least some indication of the results obtained. is The facts an attempt to put the main connected with the distribution of animals in a form acceptable to the geographical student.PREFACE known. and has been written by one whose approach the biological side. Director . On the other hand. where they are treated from another point of view. relation between the animals of the various natural regions fully and their surroundings has been much have less still treated by geographers. The facts mostly to be sought in zoological text-books. botanical The intimate relation between the vegetative covering and the physical conditions has been in many cases thoroughly worked out. Special acknowledgement should be of made to Sir Thomas Carlaw Martin. and I am greatly indebted to the various gentlemen named for permission to reproduce their photographs. the more difficult subject of the Within the last few years. and many even of the most recent geographical text-books show uncertainty present book of treatment when dealing with the animals of tropical forest and arctic tundra. to geography was first made from The sources of the illustrations are indicated beneath each. as is well geography has made great strides.

of Keeper the Natural History Collection there.4 the Royal Scottish PREFACE Museum. NEWBIGIN. and Mr. for permission to photograph a the collection. I. Eagle Clarke. number of specimens in For the index I am indebted to my sister. Miss Florence Newbigin. 1913. . I^L^RION Edinburgh.

36 III. IX.CONTENTS PAGE Iniroduction : The Natural Regions of the Globe its .121 . VII.. 9 CHAPTER I. in the Sea . and Fauna ... 208 Appendix. Outline Classification of Animals 226 231 Index .. or Coniferous Forest.. Mountain Faunas -72 The Fauna of the Tropical Forest Tropical Savanas and Deserts . . . The Taiga. 92 VI. .. V. The Tundra and Fauna .. Cave Faunas x. Steppe Faunas and the Temperate Steppes of Asia and North America 52 . Special Features of Island Faunas . its 15 II. . 144 156 182 The Distribution of Animal Life The Animals of Lakes and Rivers. zoogeooraphical regions . IV. .. VIII.

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1. 14. Kiang Bharal Serow Musk-deer ... Congo Tropical Forest Red Howling Monkey American Monkey Black Lemur 102 108 109 . Sage-cock Swiss Alp ... . 31. 23..LIST OF ILLUSTRATIONS FIG. Phalanger 113 . 20. . Virginian Deer Three-toed Woodpecker 48 . 3. 2. African Elephant 28. 4. PAGE 19 . . . 11. 25 39 40 41 . Rain-forest in 22. 98 99 100 101 24. African Flying Squirrel facing 10 BraziUan Tree Porcupine Quica Opossum . .. facing 80 81 Alaskan Wild Sheep Takin 82 83 83 84 84 85 86 16. Snow Leopard Yak . Tundra in Northern Russia Ptarmigan Wapiti Elk . 8. . 9. 17. 7. facing 5. .. Savana 122 . Camels American Bison Prairie 61 facing 62 63 70 74 Marmot . 10. 19. 15. 12.. 18... 13.. 26. 34. Indian Elephant 30. » 111 111 32. 6. 25. Hyrax North Borne 3 21. Puma and Cubs 27. Sloth 33. 109 1 29.. 112 . ...

44. .. 35. Bower Bird 45.. . . 48.. 134 134 . . Seal 42. 38. PAGE Gnu 36... Okapi 47.. 39.. Rookery in Copper Island Seals on shore of Copper Island Kittiwake Gulls „ 43. . 141 40.LIST OF ILLUSTRATIONS no. Penguins Clouded Leopard 46.. Giraffe 37. Kiwi facing ' 150 162 163 41.. . facing 130 131 Jerboa (Central Asia) Jerboa (Egypt) Cactus Wren .. 164 164 facing 218 219 224 225 . Lyre Bird ...

there is very little resemblance between the animals of the grassy plains of South America and those of Africa. The two facts which especially struck him were the two set forth in the chapters on distribution in the Origin of Species. and conversely. These are. On the other hand. as indeed the modern interest in so many problems connected with biology. and it was these phenomena which helped to direct his attention to that line of thought which resulted ultimately in the publication of his theory of Natural Selection. Darwin in his famous voyage on the Beagle was much struck by the curious phenomena of distribution which that voyage broitght before his eyes. that similarity of physical conditions does not necessarily result in a similar fauna.INTRODUCTION THE NATURAL REGIONS OF THE GLOBE The modern of the interest in the facts of the distribution of animals. dates from the publication of the Origin of Species. from the cold deserts of the high Andes to the luxuriance of the Brazilian forest. there is considerable uniformity in the fauna of South America throughout its length and breadth. The second point which struck him was that the resemblances or differences between the faunas of two areas are directly . first. or from the burning deserts of Chile to the grassy plains of the Argentine. even though we find there every variety of physical conditions. despite the similarity in physical conditions. For example.

the small silvery leaves of such plants.. The remarkable features of the animals of isolated regions can be explained on this hypothesis as the result of descent with modification from the original forms. In deliminating the natural regions of the globe the botanists have played a large part. together with the occur- . the the region. the Mediterranean region gives a great deal of information in regard to the region. of the animals of the natural regions of the globe as forming a part of the features of those regions. for they have shown that the existence of the great plant associations depends upon the climate. One result of the direction thus given to the study of distribution has been to the division of the globe into zoogeographical regions. than to their zoological relationships. relief. another and more truly subject. Such facts led him to believe that each species originated in a particular region. and so forth of practised eye a collection of plants from e. and in this case attention is directed rather to the adaptations displayed by the animals. and the other To the devices for husbanding water. 10 INTRODUCTION proportioned to the nature of the barrier which separates the two regions. Similarly. and the study of the faunas make supreme importance. separate two different faunas. geographical point of view is possible. For example. lands separated by a broad and deep belt of ocean may have quite different faunas those separated by a shallow and narrow strait may have almost identical animals. such as the Himalayas or Atlas. and it is this aspect which is studied in most books on the But. We may study of the different regions. Transverse mountain chains. g. obviously. and spread out from that region till it was stopped by some barrier to further distribution.

These are woodland. monkeys. That we find in Brazil. grassland. the Taiga or coniferous forest of Asia and North America has a well-defined assemblage of animals showing some interesting adaptations. and desert. The dependence of the more active animal upon its surroundings is less intimate than that of the passive plant. which roughly divide the surface of the land among them. but as all animals depend ultimately for their food fication upon plants. for example. That Africa has an enormous number of antelopes enormous both as regards species and individuals in that continent of — — suggests the existence and open plains. and so on. animals so thoroughly adapted to the arboreal life as sloths. Such a simple division would help us little in the case of animals. for example. grassland. a consider- conditions of animals which where the winter cold periodically checks plant growth. a modification of this classi- may serve as a framework in the study of animal distribution. as compared with those where no such temperature check occurs. among the various forms of temperate forest. and so forth.NATURAL REGIONS OF THE GLOBE 11 rence of many bulbous plants. suggests at once that dense forest occurs in the region. Thus. speaks to a climate where dry and wet seasons alternate. This gives us a first distinction between the animals of cold or temperate regions and those of tropical regions. . but even here the characters of the animals of a region tell us something of the physical features of that region. and desert occur. able difference in the live in regions There life is. In both of these forest. The botanists recognize three great plant formations. marmosets. but it is possible to pick out certain well-defined types in order to study their faunas separately. so with other regions.

separately. mingled with wood. consider (6) Savanas and This still Warm leaves to be considered the fauna of (7) which show some special features. with its characteristic fauna. which the botanists call Savana. warm or tropical regions where the tempera- is still high. we have (5) luxuriant Tropical Forest. differing according as the forest occurs in Africa. (2) we have to consider (3) (1) the Tundra. Mountain Where the temperature is permanently high. any consideration of the fauna of the deciduous forest which covers so much ground in Such a scheme . is obviously incomplete. whose fauna has many peculiar features. rainfall diminishes the savana thins out into so that we may conveniently Deserts together. and regions. omits. for example. As natural regions with a cold winter climate then. the Taiga or coniferous forest. best (4) developed in the northern hemisphere. The great mountain chains of the earth's surface are peopled by faunas having certain special features. both in Asia and in North America we have vast areas of natural grassland forming well-defined Steppe areas. but the rainfall small and unequally thinning of distributed throughout the year. and also the animals of aquatic habitat. which we must consider Islands. and the vast area of coast-line which fringes the Arctic sea forms a cold desert or tundra. Asia.12 INTRODUCTION Again. the Steppe. or South America. or grassland As the desert. but showing in Again. in ture all cases certain special adaptations. we have a the forest into that type of grassland. or (9) in fresh water. which possess a characteristic fauna. and the precipitation great. according as they live (8) in the sea. in that it It does not cover the whole surface of the globe.

adjust themselves to the altered conditions. and nowhere else. that such regions have for the most part a transitional fauna. made up of units from the surrounding regions. But a consideration of these points would take us too far from our roaches. again. others. some parasites. The difference between the sloth of the of the Amazon forest and the squirrel Siberian taiga. and that further they have been much altered by man's interference. and of the animals of the Mediterranean or scrub forest. in California. zoologist sloths. is a purely arboreal animal. the globe. the common sparrow. We shall find. and prosper under civilization as they never did in former days. for example. it is To the that the a fact of great interest and armadillos should occur in South America. found round the Mediterranean Sea. But the geographer is more interested in the fact that the sloth ant-eaters. It is true that the effects of this interference are often interesting. for the latter . already given. and We fill shall therefore confine our- selves to a consideration of the nine natural regions up whatever gaps this method may leave by a final consideration of the zoogeogra- phical regions into which zoologists have divided the But as the greater part of this book is thus devoted to the geographical aspect of animal distribution. for while some animals die out as civilization spreads. however. while the armadillo. like the rats. fitted only for life in the dense forest. immediate subject.NATURAL REGIONS OF THE GLOBE 13 Europe and North America. zoological and the geographical. and elsewhere. a word or two may be added in further explanation of the difference between the two points of view. shows adaptations to quite other habitats. the cock- and so on. different to zoologist means something very and geographer.

1876) should also be consulted.. A realization of the connexion between the study of the facts of animal distribution and the general theory of evolution may best be obtained from Darwin's Origin of Species and Wallace's Darwinism. London.14 sees in it INTRODUCTION forests. two a reflection of the difference between the which interests the other less. Wallace's classical work on Geographical Distribution (two vols. 1909). References. which gives copious references. and only incidentally. An interesting discussion of the purely geographical standpoint will be found in the chapters on animal geography in de Martonne's Traite de Geographic Physique (Paris. .

absent. In the Antarctic land-mass flowering plants are. and their cessation in relatively low latitudes. Originally applied only to the treeless lands of Northern Asia. roughly speaking. but the climate and the absence of a direct connexion between these areas and the continents have prevented the development of the characteristic tundra plants and animals. Wherever the mean temperature of the hottest month exceeds 50° F. lying to the north of the forest or taiga. its plants its and animals. This limit is. The boundaries of the tundra are well marked. on the continents the southern boundary is the beginning of the forest zone. occurring Old and New Worlds. make the development of true tundra impossible. marked out by the July isotherm of 50°. that treeless area which fringes the margin of the far northern seas. and is characterized by its climate. and to some extent also by topography. with two exceptions. It is true that considerable land-masses occur within the Antarctic area. Associated with this we have a complete absence of land mammals a very striking difference from the northern tundra.CHAPTER THE TUNDRA AND One is I ITS FAUNA of the well-marked natural regions of the world the Tundra.. in latitudes lower than those in which the northern tundra finds its limit. there the tundra alike in the — . and mosses and lichens are only slightly developed. In the southern hemisphere the tapering of the land-masses. As already indicated. the term is now by extension given to all areas of similar character.

The mildness of the Western European climate pushes the tree limit up within the Arctic Circle. In these regions the sea again forms one of the boundaries of the tundra zone. Spitsbergen. lat. ITS FAUNA The result is that the continental tundra forms a band of very varying width. The northern limit of the continental tundra is the polar sea. which extends to about 80° 30' N. but the tundra type reappears on the margin of the land-masses lying within that sea. . Greenland. The other boundary is formed by the presence of eternal ice and snow.. was hunted by the Sverdrup expedition in King Oscar Land. the northern shore of Iceland. The climate throughout this great area naturally Siberia. Franz Josef Land. the New Siberian Islands. varies. Nowhere does the mean temperature of the warmest month exceed 50° F. and even into Newfoundland. Generally it may be said that tundra occurs wherever the meltmg of the snow is sufficient to expose a surface upon which land plants may grow. are all fringed by a band of tundra. Here to the east it revenges itself for its conquest by the forest in Western Europe by pushing the forest zone far to the south. The islands of the American archipelago.16 dies THE TUNDRA AND away into forest.. one of the most characteristic tundra animals. so that a band of tundra extends down the coast-line of Labrador. and reduces the tundra zone to a narrow band to the north of Scandinavia. but the special features are the following. and it is often much below this. Its northern extension may indeed be gathered from the fact that the reindeer. Nova Zembla. Thence it extends through European Russia into and reappears again at the other side of the Bering Strait to extend across Arctic America. Baffin Land. &c.

and therefore the occurrence of land animals. The rapidity of development is of course aided by the virtually continuous daylight. Summer comes with extraordinary rapidity. As the rays of sunlight have a very oblique direction. though slight. local conditions count for much in promoting the growth of plants. bound in winter. springs are absent the rivers are fed only by the melting of ice and snow. so that March or even April may be the coldest month. Elsewhere the mean never exceeds 10" per annum. being frostox'. for Nordenskiold notes that the reindeer are in better condition in the depth of winter than in the spring months. giving rise to the local luxuriance by many explorers. The subsoil is permanently frozen. except south-western extremity of Greenland. clearness strong winds helps to diminish its thickness in places. and may be much The winters especially are remarkable for the and dryness of the atmosphere and the virtual absence of precipitation. and the frequency of less.— THE TUNDRA AND The precipitation is ITS FAUNA 17 in the everywhere small. and fog and mist are common. contrary to what might be expected. and as water. a fact which greatly aids the obtaining of food by such herbivorous mammals as the reindeer and muskDuring the short summer precipitation is frequent. the snow does not form everywhere a thick mantle. and there is thus no ground This renders all erosion mechanical. It is apparently these months which take the largest toll of animal life. and are temporary. There are only two seasons winter and summer and the winter extends far into the months which are spring elsewhere. Thus all the flowers noticed — bloom at once. 1404 Thus a slope where the B ice-cold water . The result is that. and with its coming life suddenly bursts into activity.

and therefore for the curiously local appearance of animals. and so hinder drainage. chemical erosion does not occur. and thus humus accumulates and favours the growth of bog mosses as contrasted with higher plants. When Nansen and his party left the inland ice of Greenland they descended to a little tarn. The picture may be taken as characteristic of the tundra. and the land is imperfectly drained. In their descent. while level ground is very unfavourable. carving up the land into and therefore the process of hill and valley goes on very The slowly. The first of these is the great — . is charac- terized by a very youthful condition of drainage. streams showing a very undeveloped drainage system these are characteristic features. account for the very local development of a covering of vegetation. which. These hold up the water like a sponge. whose water drained by a stream to the distant fiord. aided by the climate. for only here is vegetation abundant enough to supply its needs. As regards topography one must note that the tundra. as the leader notices. imperfect drainage is aided by the fact that the ex- treme coldness of the soil is unfavourable to bacterial life. like the semi-arid lands of other latitudes. A sunny exposure also exerts much influence. As already stated. Lakes fed by melting ice.18 THE TUNDRA AND ITS FAUNA drains away as the snow melts is favourable to plant growth. they found that the main streams had often transverse tributaries. which greatly hindered progress. the rivers are temporary. Without discussing the plants of the tundra in detail we may note those points which are of special importance for animal life. The Sverdrup expedition noted that the Arctic hare is practically limited to the regions where large valleys open out into fiords.

The shows the undifferentiated nature of the drainage. season pasturage is freely available for of the reindeer. and whose buds and seeds are of great importance as part of the food of the ptarmigan and the snow-bunting. summer conditions.. 1. Fig. The Tundra illustration in Northern Russia. or bog moss (Sphagnum). alders. which do not die and are therefore permanently available as pasture for the herbivorous mammals. and shows also that at this herbivorous mammals. B 2 . .THE TUNDRA AND abundance of ITS FAUNA 19 mosses and lichens. birches. Among the higher plants we may note the presence of dwarf willows. whose branches stick up through the snow. It is well known that the lichen popularly called reindeer moss (Cladonia rangiferina) which occurs over wide areas. &c. Other areas are clad in hair moss (Polytrichum). forms a very important part of the winter food down in winter. such as crowberries. In the more favourably situated regions berry-bearing shrubs occur.

and relatively poor in cellulose. and such birds feed in the sea. even of the four mammals named.20 THE TUNDRA AND ITS FAUNA an important part of the available food supply. the habits of all and the conclu- mammals can be said to be permanently dependent upon the land for food these four being the reindeer. on ground enriched by their droppings. Even of the land plants the last statement is partly true.. again forming Is checked by the natural conditions. sion arrived at is that only four of the . and the lemming. Again. All other birds and mammals are either partially migratory or depend to some extent at least upon the harvest of the sea. and are therefore necessarily confined in their distribution to the margin of the sea. &c. as in the case of the Swiss alps (p. the reindeer. which alone can fix inorganic matter in organic form. which makes their nutritive value high. though stunted in appearance. according to Nordenskiold. Everywhere over the globe animals depend ultimately upon plants. only two can be said to be at once permanent residents and wholly dependent upon the land for food. be said. The German expedition collections of to Greenland made large both birds and mammals. 74). With all that can food. the musk-ox. the vegetative growth of the plants whortleberries. but it may be said to be the special . for it is said that the phanerogams are especially abundant about the breeding-places of the sea birds. and observed as closely as possible. These are the redpoll {Linota hornemanni) and the ptarmigan {Lagopus hyperboreus). however. the Arctic hare. and thus. Of the many birds known to visit Greenland. the land offers but little and the vast majority of the tundra animals depend partly or wholly upon the sea. does not disdain seaweed in times of scarcity. they are relatively rich in proteids and fats. Again.

the second large herbivore of polar regions. beginning with the ungulates among mammals. distinction the so-called woodland caribou. but here also seems migratory. The latter. On the continent of America its eastward distribution distribution. which form the basis of their food supply. that it is ITS FAUNA 21 character of the tundra animals. and the smaller forms. it is now limited to the American side of Arctic regions. It occurs also in the suitable parts is where the wolf again absent. but this is impossible with the herds which occur in the Arctic islands. In King Oscar Land and the adjacent regions the Sverdrup expedition found the animal widely distributed but not abundant. bergen perhaps accounts for the greater abundance of the reindeer there. . has a much more limited Found fossil in Europe and Asia. though nowhere very abundant as individuals. rather than terrestrial ones. with larger antlers. seem to migrate southwards towards the forest in winter. Having thus summarized the conditions of life in the tundra we may give some account of the fauna. On the in it continent of Asia the reindeer occurs in summer suitable localities in the tundra region.THE TUNDRA AND of the Antarctic area. of Greenland. the barren-ground forms. The reindeer {Eangifer tarandus) is widely spread throughout the region. which occur in the tundra. is limited by the Mackenzie River. The musk-ox {Ovihos moschatus). but in it is apparently in abundant the northern parts of Greenland. On the continent of America there is a marked between those forms which haunt the forests. apparently on account of persecution by the The absence of this animal in SpitsArctic wolf. like the fewer animals marine plants. and thus not wholly dependent upon the tundra for food.

nor does it seem to migrate in the same fashion. it is. Another important rodent of the tundra is the lemming {My odes torquatus). but in spite of the abundance of that animal in King Oscar Land the Arctic hare is stated to be very abundant there. the lemming. or by nibbling the protruding much smaller animal it is not much bigger than a mouse lives in winter beneath the scraping shoots. a — — snow. ramifying over the surface in all directions. A markedly social animal. It is non-migratory. and in the adjacent regions. In some regions. which seems to feed largely upon it. the herds seem capable of defending themselves against the chiefly Arctic wolf.22 THE TUNDRA AND ITS FAUNA Grinnell Land. however. which is sometimes extraordinarily abundant. it occurs elsewhere on both sides of the Perhaps because of the paucity of Arctic region. finding food throughout the year by scraping away the thin covering of snow. the we should perhaps add the fact that occurs in the tundra in the subfossil condition in Asia and also in Alaska. MTien the snow melts in spring these runs appear. lemmus) is also circum- . which is very abundant in suitable localities. While the other herbivorous animals named find their food in ^vinter either by hare {Lepus away the snow. which attacks young stragglers. this northern lemming does not show the enormous fertility of the Norwegian form. Its greatest enemy is the wolf. Absent from Spitsbergen. The Norwegian lemming {M. this short list To mammoth Rodents are represented in the tundra by the Arctic glacialis). as in Nova Zembla. remarkably abundant. making runs and burrows in the underlying ground. vegetation in its natural habitat. Even in winter it finds food enough in the grasses which project through the snow.

greater intelligence. occurs in Arctic Asia and Arctic America. and refuse. and by some it is believed to store the bodies of these for the winter. shell-fish.. widely distributed in the polar regions of both the Old and New Worlds. as elsewhere. the Eskimo dog. Lemmings are also an important part of its diet where they occur. which follows the lemming to the coasts of the Arctic . like such herbivorous animals as the musk-ox. the catholicity of its appetite may be gathered from the fact that it will eat fish.THE TUNDRA AND polar. therefore. Very insufficiently fed by its master. the presence of large herbivores naturally attracts their enemies the large carnivores. These are the ermine {Mustela erminea). Much more truly Arctic than the wolf is the Arctic fox {Canis lagopus). driven by stress of competition to the unfavourable parts of the surface. but it is constrained to add to this refuse and even seaweed. as a rule. ITS FAUNA 23 but does not extend so far to the north as the other species. but both are immigrants from their natural home further south. when it is often hardpressed. Thus the wolf {Canis lupus). absent in Greenland. necessarily fewer in number. is a very important animal in Greenland and elsewhere. Two other land carnivores occur in the tundra. The carnivores which occur in the tundra. nor are they. Its near ally. seaweed. and usually of As masterful and highly organized carnivores do not as a rule show . do not as a rule form an integral part of its fauna rather are they intruders from other regions. In the tundra. &c. and extends also into such regions as Grinnell Land and King Oscar Land. especially the ptarmigan. Its food consists of birds. animals the large a very close adaptation to one type of country only. in Spitsbergen.

of geese. We may pass next to the birds of the tundra. seals and the walrus. All travellers to tundra regions. whether to the mainland areas or to the lands of the polar sea. of auks and guillemots. where it is found among the drift and floe ice. these birds travel southwards. none of which can be regarded as definitely characteristic of the tundra.24 sea. It is thus not in any true sense a tundra animal. we can only say here that though they breed on the shore they are not in any sense tundra animals. of kittiwakes. THE TUNDRA AND ITS FAUNA and the glutton {Gulo luscus). which. and also probably because they find the long northern daylight an advantage. help to give its characteristic facies to the region. In regard to these it is noticeable that large numbers of migrants occur. although occurring round the margin of the tundra region throughout its extension. 'of skuas. a forest mammal. on the look out for its natural prey. Of the many kinds of seals also. Ave must place the polar bear. though relatively few in number. It is. especially sea birds and those which haunt the vicinity of the shore or of fresh water. though their summer abundance at certain special localities is one of the features of the region. of various forms of ducks. including the eider. and so forth. As its name Ursus maritimus) indicates. Among the mammals which find their food in the sea. speak of many kinds of gulls. These come north to breed in the relative security of the tundra. otherwise with the true land birds. which just reaches the tundra. First . It is unnecessary to give lists of these migratory forms. fleeing from the increasing cold and darkness. As winter approaches. of sandpipers. terns. for it lengthens the period during which the search for food can be ( carried on. petrels. this animal haunts the margin of the sea.

The Ptarmigan. which more than one species occur in the tundra. 2. Northern Asia. and eggs.THE TUNDRA AND among of ITS FAUNA 25 these we must place the ptarmigan (Lagopus).) regions. these birds In the slow . young. sometimes in considerable numbers. showing the stony uplands where the bird breeds in Highland Scotland. In Greenland. {From a apecimen in the Royal Scottish Museum. in Spitsbergen. in Arctic America and the adjacent Fig. with nest. in occur.

and turning southwards in the autumn. nesting in the north. they whiled away their months of waiting for a ship by the help of ptarmigan shooting. and not a few other northern forms the peculiarity of displaying a seasonal change of colour. ptarmigan are not peculiar to the tundra. it was by the help of the ptarmigan and reindeer shot that the final catastrophe was postponed so long. the stoat or ermine. spoken of by most explorers. their crossing of the inland ice completed. the ptarmigan is social. who welcome its There is no doubt that it is largely a migrant. Occasionally. for they extend far to the south of the region. Sverdrup and his companions shot many for food during their Arctic expedition. The only songster of the tundra is the snow-bunting {Plectrophenax nivalis). however. and similarly almost every expedition which has landed in tundra regions has noted the presence and characters of the birds. . except at the pairing season. After Nansen and his companions arrived in safety at the east coast of agony after the Greenland. south in winter than in summer. The ptarmigan share with the Arctic fox. Like many other tundra animals also. a certain amount of migration occurs.26 THE TUNDRA AND of the last ITS FAUNA march of de Long and his companions wreck of the Jeannette. or in stealing unperceived upon their prey. being found in flocks. This is no doubt associated with the fact that food is only obtainable in certain areas. and is twittering song in spring. which is very widely distributed. This change doubtless aids these varied animals in escaping from their enemies. Thus in Greenland they are commoner in the Needless to say. Though in most favourable tundra regions ptarmigan are to be found throughout the year. separated by extensive barren regions.

tundra region lemmings and small snowy owl it not seem be common does to though {Nyctea nivea). The bird has a general resemblance in coloration to a ptarmigan in winter plumage. which inhabits Greenland and Arctic America. a beautiful predominantly white in colour. Except in Spitsbergen and Nova Zembla.THE TUNDRA AND it ITS FAUNA 27 appears to winter in the Arctic. as Zembla and parts of the mainland of Northern Asia. in Nova except where lemmings are abundant. It winters in the tundra. is a very characteristic bird of the Arctic is Another predatory bird the Greenland gerfalcon (Falco candicans). bird. and fresh-water fish are not abundant. upon which the owl preys. but there is nothing characteristic. The third predaceous bird of the tundra is the raven {Corvus corax). It is. and winters in Greenland. As is to be expected from the unfavourable conditions of life. and of the region. there occurs the eats fish in addition to birds and small mammals. Among the invertebrates the insects present some interesting points. and also upon lemmings. however. It may seem . and will eat practically anything from land or sea. reptiles and amphibians are absent. notably in Green- land. which is the scavenger of the region. Representatives of the salmon family occur in some lakes. for it Wherever in the birds are to be found. no exception to the rule that the carnivorous animals of these barren regions need to lay the sea under contribution as well as the land. mosquitoes are enormously numerous. and often lives in the stony regions frequented by that bird. It preys upon ptarmigan and other birds. and is represented by an at least closely allied species in Northern Asia. and form in many regions a terrible plague.

As is weU butterflies usually known. especially about the houses of the Eskimo. after having laid their eggs in the pools. active period of vegetation has a curious effect the development of The very short upon plant-eating insects. when they disappear. but the rapidity of reproduction is certainly not checked.g. except in the coldest regions. but the eggs or larvae remain uninjured. and they are then extraordinarily abundant till the middle of August. and of can be run through in the brief period warmth.. the tundra. &c. e. with the result that the water formed in summer by the melting late in snow tends to accumu- innumerable lakes and pools. and the natural checks. breeding Indeed. in more temperate countries . It is possible that the life-history is lengthened by the cold. These pools freeze or dry up with the onset of winter. but the cause mentioned. In these the mosquito larvae swarm. to hatch or recommence active life with the coming of water in the spring. the bees of the curious parasites known as Strepsiptera. among northern insects. the characters of a semiof the arid region. flies predominate in the refuse thrown out. in the shape of small fish who feed on the larvae. all the other orders of insects are repre- and butterflies fulfilling here their usual role as fertilizers of flowers. as The absence of any run-off in winter gives we have noted. already region.28 THE TUNDRA AND ITS FAUNA curious that they should be abundant in this cold is to be looked for in the uncondition of developed the drainage system. The reason lies apparently in the fact that the larval stage is usually short. must be insignificant. With the exception sented. the percentage increasing as we pass northwards. In addition to mosquitoes a considerable number of other flies occur. From the pupal stage the adults emerge towards the middle of June.

activity and the life-history is rapidly passed through. they must pass a second winter in the larval state. In the spring. early resumed. have considerable direct and indirect importance for the land animals. however. for the land warmer . a few earthworms occur in the parts. or. so that the butterflies appear in summer. and there are a few land and fresh-water shell-fish and some spiders and mites. they winter in a very undeveloped state. In Greenland. in the colder parts. for fuel. Activity does not recommence till at earliest the first week of the following June. In addition to insects. therefore. In general. must necessarily wait in spring until their food plants have made some new growth. This life-history. again emphasizes the baneful effect on animal life of the scanty development and slow growth in spring of the plants of the tundra. the resulting larvae have. As a pendant to this picture of life on the tundra we may add a brief consideration of the animals of the shore. This great lengthening of the lifehistory cannot be ascribed wholly to the cold we must suppose that it is also due to the fact that caterpillars five .THE TUNDRA AND lay their eggs in the ITS FAUNA 29 autumn (except in the case where there are two annual broods). if the larvae hatch. and these hatch at once. Man in these regions is practically dependent upon the sea for food. the butterfly emerging in the following July to lay its eggs and die. and as it cannot be supposed that the animals can pass through larval and pupal life before the onset of cold. only some weeks before winter sets in. which. and for clothing. butterflies do not appear July. and these eggs pass the is winter as eggs. as we have noticed. it may be said that among the invertebrates the insects are the most conspicuous and numerous forms. If till the middle of they lay their eggs.

and toothed whales feed upon fish. littoral A considerable number of large ' Saxicava mussel-like forms. including in the gapers ' Mya and . of the food of the walrus. especially by the minute plants. such as the diatoms. of which the mud-haunting forms are devoured by the walrus. and the two facts are closely connected. but in this respect he resembles all the other animals of the region. so abundant in polar seas. and store up in their bodies reserves of this heat-producing sub- stance. and the fact ' The ' ' ' that they were so utilized of Arctic conditions. Small Crustacea are also devoured by molluscs. which constitute an important part Where molluscs abound. directly or indirectly. the is fairly rich. These largely the small crustaceans caUed amphipods. obtain their supplies of fat larger marine animals almost always. ITS FAUNA commonplace of It is a physiology that in cold climates his need of fat is great. and the floating forms by the whalebone whales. which. The few survivors of the ill-fated Greely expedition kept themselves alive by the collection of shrimps shrimps must have been in the shore-water.30 gives THE TUNDRA AND him almost nothing. by their wealth of small Crustacea and of marine mammals. all of whom seek fatty food. are often extraordinarily abundant seals as individuals. occur. may be taken as emblematic fauna of the Arctic molluscs Apart from Crustacea. The Crustacea feed haunt polar seas. so that the Crustacea form the basis of the marine foodsupply. They are in their turn fed by the more minute forms of life. found in the myriads of small Crustacea which is The food from which the and whose bodies contain much oil. Arctic seas are indeed specially characterized the fish of the polar seas. though of relatively few species. which are enormously abundant in polar seas.

All have. Where seaweed is abundant. however. we must note that if the sea contributes a wealth of food to land animals. ITS FAUNA 31 reach a large which prey upon tliem. seaurchins occur according to Nordenskiold. The poverty of the land is made up for by the wealth of the sea. The ice-foot and driftice also contribute to the rock debris spread over the bottom. and its poverty in land forms of either. bottom. The icebergs which detach themselves from the glaciers carry much rock-waste to the sea. is in Arctic regions carried seawards to help to feed the swarming life of the sea. had terrestrial ancestors. of course. and we may see one justification of the evolution of their adaptations to a marine habitat in the fact just . The characteristic marine mammals of the extreme north are the walrus and the earless seals. toothed and whalebone. the mud which in Central Europe forms such vast fertile areas as the Plain of Lombardy. as is for instance so much of the glacial mud of the Alps. and other less striking invertebrates also occur in numbers. Here. It is this fact which explains the wealth of the Arctic in marine birds and mammals. and the whales.THE TUNDRA AND starfish. and the proximity of the glaciers to the shore in many places must mean that the rock-waste is carried by the turbid streams of summer direct to the ocean. for very many shore animals are mud-eaters. In other words. and size. The mud holds a considerable number of marine worms. instead of being spread over the land. especially at the breeding season. which never voluntarily come on shore at all. west of Nova Zembla they are so numerous as almost to cover the . are also abundant. yet the land also gives something to the sea. and this forms a fine mud very favourable to animal life. all of which spend a considerable part of their life on shore.

is walrus {Trichechus one of the most it is As already noted. — and suitable regions for this purpose. both of whom take advantage of the animal's relative helplessness on land. or at least The massive tusks are used in digging up mussels from the mud. but are very perfectly adapted for swift movement upon fish.32 THE TUNDRA AND ITS FAUNA emphasized the poverty of the land and the relative wealth of the sea in these northern latitudes. whale. which interesting forms. five or six occurring in Greenland. and the fact that they must come on shore to . the is rosmarus). They add shell-fish and is crustaceans. and the many kinds of deer in the pass a considerable part of floating ice-floes. except in the case of man. offering security against attack. and also in helping the animal to clamber upon the ice. feed chiefly upon molluscs. The true feed seals are very clumsy on land. the great enemy of the large mammals. Analogous conditions are presented by the many kinds of antelope which are found in the open plains of Africa. and even seals to these. are to be found in the frigid rather than in the temperate or torrid zones. It seems to its life on shore. Of the Arctic seal-like animals. to which at times they in the sea. but their enemies on land are consisting of the polar bear much more and man. forests of Asia. Another cause of the abundance of the seals and their allies in Arctic and Antarctic waters is certainly to be sought in the fact that these animals must come on shore to breed. stated to fish. but apparently adds swimming on birds. In the sea their great enemy the killer serious. The presence of a number of nearly related forms within a limited area means a very perfect adaptation to the natural conditions. circumpolar. The true seals are represented by a number of species.

avoiding the neighbourhood of land. and lives largely 1404 c . often seek the vicinity of northern coast-lines in spring for breeding purposes. It is a bold species. This is especially true of . The common seal {Phoca vitulina) also occurs in Arctic waters. The very other. and must have relatively open water in its haunts. which fit them for different parts but it is not always of the common environment possible to explain wherein these special adaptations consist. the Greenland seal {Phoca groenlandica) therefore incapable of making a breathing -hole in the which seems ice. though found in practically all seas. The large bearded seal {Phoca harbata) is another form which makes breathing-holes. which does make such breathing-holes. which filter the minute crustaceans and molluscs upon which it feeds from the sea water. On the other hand. and bringing forth its young on ice-floes. and eats cuttles as well as fish. going south in winter to avoid three weeks of their the intense cold of the north. and include some very large animals. is almost nonmigratory. and its elaborate series of baleen plates. The huge rorquals (Balaenoptera). Thus among the whalebone forms we have the Greenland whale {Balaena mysticetus). Among the toothed whales the narwhal {Monodon monoceros) is a true northern form. Many of the seals are markedly migratory. with its enormous head. ITS FAUNA 33 and that the young must spend the first two or life on shore. . apparently a swift curious crested seal (Cystophora cristata) swimmer. the ringed seal [Phoca hispida). is not only migratory but also pelagic in its habits. The cetaceans of the north are numerous. living permanently in the coldest regions.THE TUNDRA AND breed. These two species will thus not compete with each and all the species doubtless show similar minor adaptations.

both of plants and of animals. leads to practical uniformity of distribution. the fact that the salt. Accounts of tundra animals are to be found in almost books dealing with the Arctic regions.34 THE TUNDRA AND is ITS FAUNA upon cuttles and crustaceans. combined with the marked uniformity of physical conditions. generally speaking. are either circumpolar at the present time. and there is almost no resemblance between the desert fauna of Australia and those of Africa or of America. The animals. remarkable ascending rivers for a considerable distance. we may say briefly that the tundra fauna is virtually uniform throughout. whale {Orca gladiator). estranging sea is here more or less ice-bound makes the land areas virtually continuous. The white whale {Delphinapterus leucas) for its habit of similarly an Arctic form. all References. or have once been circumpolar. that is. No such similar uniformity exists in regard of ' ' to the other natural regions of the world. share certain adaptations to forest or desert life. Though the steppes of Asia show a general resemblance in physical conditions to the steppe-like regions of North America. killer is The other hand. No resemblance. and there is no distinction between eastern and western faunas. chiefly occurs in the Arctic in summer. on the a very widely distributed animal. but necessarily desert animals or forest animals. In other words. when the actual affinities of the animals are considered. whatever their affinities. and of the marine forms which occur round its seaward margin. The following may be men- tioned as giving a considerable amount of detail : Manual of the Natural . which If we sum up the facts in regard to the distribution tundra animals. yet their faunas are different. the faunas of the similar equatorial forests of South America and Africa are markedly dissimilar. In the same way. and this.

Geology. E. THE TUNDRA AND History. addition the accounts of their explorations published by Nansen. See also Seebohm's Siberia in Asia (1882). 1897). and gives some characteristic views of tundra life. 1881) Gronland-Expedition der Gesellschaft fur Erdkunde zu Berlin. Kobelt's Die Verbreitung der Tierwelt (Gemassigte Zone). Ernst VanhofEen. Nordenskiold (English Translation. discusses tundra animals in general. 1893. Rupert Jones (London. the Duke of Orleans. The second volume of the last named gives a very full account of the fauna and flora of Greenland by Dr. 35 edited and Physics of Greenland and the Neighbouring Regions. 1911). by A... may be consulted. C 2 . published at Leipzig in 1902. the books dealing with the Jeannette expedition. &c. The animals of polar regions generally In are discussed in Dr. Bruce's Polar Exploration (London. unter Leitung von Erich von Drygalski (Berlin. London. ITS FAUNA . and the same author's Address to Sectibn E (Geography) at the Nottingham Meeting of the British Association. by Professor T. 1875) The Voyage of the Vega. Sverdrup.

Here the climate is especially in the southern portion. and thus prolong the summer abundance into the time of scarcity. Nevertheless here. of heat received. life is caused by the seasons. but in the temperate forest zone. In the forest zone. no less than in the drier the ebb and flow of plant by the varying amounts summer and \^'inter As all animal hfe in these regions ultimately depends upon land plants. OR CONIFEROUS FOREST. with their winter rainfall. not heat. branches of the trees are exposed. it is the alternation of which is of importance. berries. or seeds. vegetative life is practically continuous. however. In subtropical countries. and these twigs and and branches very often carry nuts. In the tropics. more favourable. and the flora far more luxuriant. Further. the twigs the year round. which determines vegetative activity. the forest gives shelter. this means that there are periodic alternations of plenty and steppes. which persist throughout the winter. this is less in the steppe zone. marked than . AND ITS FAUNA is Strikixgly different from the fauna of the tundra that of the temperate forest.CHAPTER II THE TAIGA. and thus diminishes scarcity. it is water. as in the tundra. In the coniferous forests the trees for the most part keep their leaves all However deep the snow. lying just south of the tundra. which forms a wide belt in the northern hemisphere. except where there is a well-marked dry season. that is.

both tundra and steppe animals may seek migration the forest in winter. especially upon the rainfall. limited to the summer months. which are structurally adapted for forest life. on the other hand. and the absence of drying winds in winter. when the coldness of the soil prevents absorption. the covering of dead leaves may protect it from frost hollow trees and dense bushes offer protection to many small the rigour of the winter climate. but which visit the forest for shelter or for food. "'^•^^4^' As a consequence of the fact that the forest onera^' a refuge to animals from other areas. the tree-frog. we find that its denizens may be divided into two groups. In short. where areas abundance even in the summer season alternate with others where famine reigns. and of . and are not at home elsewhere.THE TAIGA AND may shelter the ITS FAUNA 37 The thick branches ground from snow. the woodpecker. We have forms like the squirrel. which show no very special adaptations to forest life. One forest result of this is that on the whole the habit of is less marked in the animals of the temperate than in steppe ones. and especially greater reserves of food. A small rainfall. We have already noted that the northern limit of the forest is determined by adequate summer warmth. and we also find animals. it offers greater advantages in winter. . the lynx. It does occur. and the greater uniformity of conditions in the forest prevents migration occurring on the same scale as in steppe and tundra. the tree-creeper. while the forest does not offer the rich pasturage of the steppe in summer. animals. the dormouse. favours grass rather than trees. but. and so forth. Its southern limit depends upon much more complicated factors. like some kinds of deer and wolf and fox.

taiga as a whole Lianes and creepers are t3rpicaUy absent. or carnivores. a belt of forest of very similar characters occurs. being Its important trees are the Siberian fir . crowberries.. with many minor forms. eagerly relished Cembra pine. The important mammals of the taiga belong to three orders they are ungulates. The species of conifers are different. and its inhabitants show a general resemblance to those in the Old World. or rodents. especially abundant in clearings.38 THE TAIGA. and on the whole arboreal adaptations are few. whortleberries. . but the general aspect of the forest is the same. such as cranberries. &c. therefore. markedly uniform in character. this ditions. mountain ash. we shall describe the inhabitants of the coniferous forests of both the eastern and western hemispheres. Under the name of taiga animals. and in areas devastated by forest fires. Very many of the inhabitants of the taiga possess the power of climbing without showing any great modification of structure. such as juniper. forest. its &c. OR CONIFEROUS makes conifers their slow rate of transpiration more tolerant than broad-leaved trees of unfavourable con- The consequence is that in Asia we find a band of predominantly coniferous forest separating the tundra from the steppes of the central region. The result is that we have no animals of purely arboreal habit. larch and the Siberian the for its edible seeds. which is called the taiga.. fruits. and the is much less dense than the tropical forest. . bird cherry. important in that they bear edible In addition the taiga resembles the tundra in great wealth of berry-bearing bushes. especially in the eastern part of Canada. In North America. important by many animals . such as occur in the equatorial forests.

S. AND ITS FAUNA 39 There are in addition a few bats and a few insectivores. often miscalled Rocky Mountain elk '.FOREST. Except in South America. we may say that the ungulates of the taiga consist of deer. 3. especially in South America. Deer are typically forest-haunting animals. We shall begin with the ungulates.A. U. as including the largest and most obvious of the inhabitants of the forest.) ' which just penetrates the Asiatic taiga from the south. Fig. but the predominant forms belong to the orders named. really mountain animals. though some of them feed on the outskirts of the forests. The Wapiti of the American forests. where large hoofed mammals are few. {Photo by the Biological Survey. With the exception of the wild boar {Sus scrofa). and excluding from consideration such forms as sheep and goats. and a few species inhabit open plains. the place of deer on grassy plains is taken by .

The Eurasiatic continent. belonging to a group numerously represented further south. In the Asiatic taiga the Virginian .40 THE TAIGA. but many of these are confined to tropical and subtropical regions. When ungulates occur in tropical forests they are mostly small animals. Deer are not only big animals. with narrow compressed bodies. with closely related forms of similar habits. In both hemispheres there are taiga varieties of the reindeer. especially altogether absent. America has in addition the . Though these forms do not occur in Canada. canadensis) is a nearly related species. which allow antlers them to glide through the brushwood. their place tends to be taken by sheep. leaving relatively few to occur in the taiga proper. the wapiti (C. where deer are High up on mountains also. is rich also in species of deer. Some authorities regard these antlers as evidence that deer have only recently taken to wooded regions their presence at least excludes the animals from very dense thickets. but the broad. In the Asiatic taiga. spreading of the male would make rapid movement through a dense wood impossible for him. such as the maral stag (C. deer {Cariacus virginianus). maral). The taiga fact that deer are characteristic animals in the in itself sufficient to enable us to is draw certain conclusions in regard to the nature of the region. OR CONIFEROUS abundant in Africa. goats. the red deer {Cervus elaphus) occurs. and distinguished by the great development and curious structure of the antlers. and both have elk or moose (Alces machlis). which is rich in antelopes. So far as these forms gc there is a close resemblance between the eastern and western halves of the continent. and allied forms. forests.

U.S.) . {Photo by the Biological Survey. The Elk or Moose of Alaska. 4.Flo.A.

«5 P^ .

near the upper Limit of the forest. upon trees and bushes and short neck prevent it is .FOREST. except where the grass very long. on the Sayansk divide. nuts. 85). In winter it leaves the swamps for the dense forests on the higher ground. and there feed upon reindeer ' moss The elk '. it depends largely upon mosses and lichens. compelled to feed by the fact that its long legs from grazing on the ground. &c. twigs and leaves both and deciduous trees. as outside of it. Carruthers' expedition to Mongolia. though its The last is an aberrant form. berries. In the Asiatic taiga some interesting observations on woodland reindeer have recently been made by the. which are swept bare of snow by the wind. almost hare-like in its habits. perhaps because it is better fed. In the winter the animals seek the open hill-tops. both very characteristic of the taiga. Here. of coniferous above mentioned are at least That is. AND ITS FAUNA musk-deer (Moschus) true is 41 small roedeer {Capreolus caprea) also occurs. the members of the expedition found traces of a forest-haunting form which spends the summer in the rhododendron scrub. bulk largely in their diet. members of Mr. and much persecuted for the sake of the scent gland of the male (see p. and grass is relatively less important.. and the home lies further south. Most of the animals largely forest-feeders. found occasionally. The most northerly forms of those mentioned are the woodland reindeer and the elk. in both the Old and New Worlds is a true forest animal. and chiefly haunts marshy ground. In North America the woodland reindeer is larger than the tundra form. Within the forest.

the striped squirrels or of the various kinds of squirrels. The chipmunks feed on nuts and seeds. showing progressive adaptation to arboreal striped squirrel or of a chipmunk {T. The American taiga also lodges the Canadian porcupine. both birds and mammals.42 THE TAIGA. an arboreal animal. The squirrels form a very interesting group. which obtains in winter by pawing away the snow. though they hibernate. asiaticus) is ground form which occurs in North America has other species in addition. The Siberian an example both North America life. in living mostly on the ground. a great fondness for beech mast. . home is rather further taiga are the The most important rodents (Sciurus). sharing with it many other animals. and Asia. It extends into Canada. the musquash (Fiber). where they often excavate holes. but spends a good part of its time upon the ground. which extends though it is not common there. and the (Castor) . and. stage is represented by the American red is squirrel or chickaree {S. such as the true squirrels chipmunks (Tamias). and the American variable hare in addition to forest region. OR CONIFEROUS The red deer seem to feed chiefly on the outskirts of the forest or in the open glades. the Polar hare. forest produce. The next like the true squirrels. but its true south than the taiga proper. but the American wapiti eats a considerable amount of leaves The Virginian deer also eats a large and amount of twigs. but are not nearly so agile as the true squirrels. hudsonianus). flying squirrels (Sciuropterus) the beaver into the the Polar hare. a swamp animal. They differ. Our . where it sometimes excavates holes. of which they store large quantities for the winter. They can climb trees. however. which a graceful and agile climber.

Mexico. The food consists of leaves. known. widely distributed more northerly parts It of Asia. twigs. by means of a descent to terra firma. beech mast. Though chiefly inhabitants of the tropical forests of Asia. if water-lilies. though it does not possess the same adaptations to this mode of life as are exhibited of by its southern allies. While the porcupine of the inhabits rocky country. not identical with the Eurasiatic one. one species occurs in tem- perate Asia and one in temperate North America.. was once widely distributed over the cooler parts of the northern hemisphere. which haunts the streams flowing through wooded regions. &c. tend- ing to be darker in colour in the colder parts of its range in summer. though the nest is usually made on the ground among stones. and has great powers of shows interesting colour variations. the tree-porcupines South America. It will eat almost any forest product. The final stage in the mastery over the forest is represented by the which are furnished with a parachute which they can take long leaps from one tree to another. of such plants as species. the porcupine of warmer parts of Asia Canada is a purely arboreal animal in habits. is at Beavers feed upon bark and twigs. and turning white in winter where the climate is severe. &c.FOREST. It spends the greater part of its time in the trees. The American least very nearly allied. and thus obviate the risks due to flying squirrels. and also upon the roots. and other nuts. The Canadian form {Erethizon dorsatus) follows the taiga to its northern limit. The beaver. its and also follows the western coniferous forest to southern limit. in the AND ITS FAUNA 43 common British squirrel {S. and the . and hibernates during winter. bark. leaping. vulgaris). as is well rarely descends to the ground.

minks. In Siberia. is the skunk {Mephitis mephitica). The wild cat {Felis the wooded parts of Europe does not occur in the Siberian forest. and the only one which ranges far north. the special features of the northern forests are the relative poverty in true cats. although it is common in the coniferous forest of the catus) of Rocky Mountains. and are true forest animals. so that it must find its food largely on the ground. a member of a group characteristic of South America. The bears also are characteristic of temperate forests. The lynxes are represented by the common lynx and by a closely related species {F. and badgers. The common woK is not {Felis lynx) in Siberia. abundant in tropical forests generally. There is a close resemblance between the carnivores of the American and Siberian taigas. s. the common fox. weasels. found lynxes.^ specially fitted for forest life . the glutton.44 THE TAIGA. stoats. They are expert climbers. Thus in both are so common wolf.. the tiger is sometimes found. the especially in the south. As regards the carnivores. while it is altogether absent from America. and take there the place of the true cats and civets elsewhere. OR CONIFEROUS animals are stated to defoliate completely the trees they attack. it lives either in the . which are most abundant in temperate latitudes. but with no Old World representative. bears. and the wealth of arboreal animals of the weasel family. An animal found in the Canadian forests. canadensis) in the woods of Canada. while in the Canadian taiga the puma {Felis concolor) does not occur. though not confined to them. martens. and among less purely forest forms. but the Canadian form is stated to feed chiefly on hares and birds of the grouse family.

and climbs the pines is in search of well as seeking to rob the peasants' stores. It occurs throughout the greater part of North America. while the fisher marten {M. of similar habits. Closely related is the American marten {M. though both descend to the ground upon occasion. Both the black and brown bears eat fish and other aquatic animals in addition to warm-blooded forms. and. is also largely arboreal. In the taiga of replaced by the black bear Ursus ( americanus). it takes a large amount of vegetable food. in spite of its name. but only because in the woods it in the open. It is ( taiga. as in the stated to be particularly fond of the edible seeds of the Cembra pine. which is of similar habits.FOREST. as Like its North America bilis) is it is very fond of honey. americana). haunting mountain regions in the . The fox with finds the necessary shelter and abundant food. as well as open country. in the form of the Siberian wild dog (C. It can climb. AND ITS FAUNA 45 wherever food can be easily its burrowing habits is more of a forest animal. pennanti). relatives also it them. The grizzly {U. The brown bear Ursus arctos) occurs in the Siberian woods of the temperate region of the Eurasian continent generally. like all the bears. with a more northerly range and a more valuable coat. which apparently haunts the high Alpine forests. nor do they disdain insects. Both are thoroughly arboreal. horri- only found in the western forest. woods or obtained. though it is not so agile as some of its relatives. zihellina). the pine-marten [Mustela martes) and the more valuable sable {M. In the Siberian forests two kinds of martens occur. alpinus). It has no special adaptations to life there. member In addition to wolf and fox Siberia has another of the genus Canis.

Among especially abundant in northern or especially adapted to life there. and the Siberian form {M. is chiefly found within the forest. the other hand. erminea) as already indicated. we may species are . vulgaris). eversmannia) occurs in Siberia. though a burrowing form. OR CONIFEROUS warmer parts of its range. forms which found there in summer time. a fierce and voracious carnivore. Passing now to the birds. though they are found there as well as elsewhere. birds. taxus) is found in the Siberian forests. we find that the forest is adapted forms. the American form [Taxidea americana) is chiefly a prairie mammal. and the complete absence of monkeys as well as of the man^ carnivorous or egg-eating animals which haunt tropical forests. and that in addition there are large numbers of migrants who spend part of The forest gives relative security their time there. and feeds largely upon the prairie marmots (see p. occurs On the other hand. and are forest animals to the extent that localities suitable to their habits often occur within forest areas. where it On preys upon mammals and disabled deer. found both in Northern Asia and in Northern America. and the stoat or ermine (M. are not definitely forest animals.46 THE TAIGA. The polecats. and help to account for the number of forests. vison) of North America. the common weasel {M. The absence or paucity of snakes in the northern forest also. the wolverene (Gulo luscus). siberica) haunt streams and lakes. during the dangerous period when brooding is taking rich in specially place and the young are being reared. increase the value of the northern forest from the birds' standpoint. of which one {M. and. 64). all manner of small and even attacks weakly or The common badger (Meles chiefly in woods. The mink {M.

Woodpeckers are very widely distributed. and also long. AND . viscid. The woodpeckers fitted for climbing. Related is the curious wryneck (lynx torquilla). often called ruffed grouse {Bonasa umbellus). a few only ranging into the northern woods. The and the most highly tails. of both Old and New Worlds. while the . jays. worm-like tongue enables them to capture the insects thus exposed.FOREST. and the spotted woodpeckers (Dendrocopus). In the Asiatic taiga we find the black grouse tetrix). is specialized forms have stiffened process. FAUNA 47 note the grouse and their alHes the grosbeaks and their relatives the crossbills the woodpeckers and wryneck . {Lyrurus the larger capercaillie {Tetrao urogallus). found in summer in Northern Europe and Asia. which aid in the climbing The beak powerful and chisel-shaped. is an important factor in their distribution. while in the is and the hazel grouse Canadian forest found the Canadian grouse {Canaa partridge. and their allies. The abundance of berry-bearing bushes. which is without the stiff tail and climbing habits of the woodpeckers. Among these are the green woodpeckers (Gecinus). for these berries constitute a considerable part of their food. Grouse belong to the same family as the ptarmigan. {Tetrastes honasia). of which we have already spoken. and are especially characteristic of northern latitudes. enabling the birds to excavate holes in wood. and of the appreciation in which they are held to lever off bark. the nutcrackers. and also the chites canadensis). are birds showing remarkable feet are well adaptations to an arboreal habitat. ITS . We have spoken of the edible seeds of the Cembra pine. found in the temperate regions of the Old World. both in the tundra and in the taiga.

A. . Note the and especially the position of the tail.) of food products. The Three-toed Woodpecker of North America. and form a much sought after food supply.48 THE TAIGA. characteristic attitude. OR CONIFEROUS by many inhabitants of the Siberian taiga. 6. U. yet they also are edible to many In animals.S. Though the seeds of other conifers have not the same wealth Fig. (Photo by the Biological Survey.

Among amphibia. young water-birds. the latter represented in the eastern taiga by the Siberian jay {Perisoreus infaustus). the former especially. The forester is constrained to apply heat to obtain the seeds without damage. but the crossbills have the beak converted into an instrument by means of which the seeds are readily prized out. In the Eurasiatic continent the 1404 common tree-frog {Hyla D . with soft fluffy plumage of a reddish colour. an important form being the large bull-frog {Rana catesbiana) of North America. so that their extrication is a matter of bullfinch.FOREST. devouring fish. as in Siberia. with their usually greenish colour and the curious adhesive pads at the ends of the toes. AND ITS FAUNA 49 both the Siberian and Canadian taiga they form a very large part of the natural food of the pine grosbeak {Pinicola enucleator). The tree-frogs (Hylidae). may be seen its breaking the cones of the Cembra pine to pieces with powerful bill. . More slashing are the methods of the nutcracker {Nucifraga caryocatactes). and during the ripening process the seeds are carefully protected by the closing of the scales.. a gorgeous bird related to the and of the crossbill {Loxia curvirostra) also with bright-coloured plumage. Related to the nutcrackers are the magpies and jays. &c. and derives its common name from its noisy croak. The western coniferous wood of North America is inhabited by another species. difficulty. by means of which they keep their foothold. frogs and toads haunt the pools and swamps. which is compared to the roaring of a bull. which is exceedingly voracious. which in the Alps. Pine cones. Reptiles and amphibians are few in number in the northern forest. ripen slowly. have a much wider distribution in America than in the Old World. it will be remembered.

sometimes in countless numbers. parasitic forms prey upon Many of the beetles especially are important pests from the point of view of the forester. In Asia wild honey bees reach the Arctic circle. according to Marshall.50 THE TAIGA. the wood-wasps (Sirex). Europe just reaches the south of the taiga. who rob the nests . where it has been introduced with pinewood. The adult female has a boring apparatus. they scarcely pass latitude 50° N. while in North America. In insects the northern forests are perennial vegetation feeds for the many herbivirous forms. the forms which store up honey extending much further north in the Old World than in the New. by means of which she bores a hole in trunks of pine trees in order to lay her egg at the bottom of the The wood-wasps are preyed upon by excavation. so that the resultant larvae may prey upon the wasp larva. Another form which feeds in the pine woods is the pine saw-fly (Lophyrus pini). of which a large species gigas) often appears in summer-houses or other wooden erections in Britain. Wild bees also occur in the woods. There is here a singularly delicate adjustment between host and parasite. comparable to the relation which exists between the length of the proboscis of certain butterflies and the length of the tubes of the flowers which they visit. when they insert their long ovipositors and lay their eggs close to the wasp's egg. ichneumon flies. The honey is greatly prized by bears. As examples of wood-eating forms we may mention {S. whose larvae feed upon the needles. whose delicate senses enable them to find the burrows of the wasps. and many carnivorous or these. North America have a more rich. OR CONIFEROUS arborea) of but the tree-frogs of northerly extension.

Kobelt's book. gives a full account and descriptions will also be found in Brehm's From North Pole to Equator (in translation.FOREST. inhabitants of the taiga are mostly those characteristic of the temperate them will be found books mentioned at the end of chapter x. As the of the taiga animals. 1896). regions of the Northern Hemisphere. London. already mentioned. from which Refeeences. 51 without regard to the stings of the bees. Most descriptions of journeys through Siberia also give some account of the taiga animals. full accounts of in the D 2 . AND ITS FAUNA their thick coats doubtless protect them.

is it. of soil or local variation of some favouring condition make this possible. characterized by the absence or scarcity of trees.CHAPTER III STEPPE FAUNAS AND THE TEMPERATE STEPPES OF ASIA AND NORTH AMERICA To the south of the coniferous forest of Asia hes a belt of varying width. Further. or the warm desert of the rainless regions to the south. This belt is of of steppe land continually merging into desert. grasses and by the periodical growi:h and other herbaceous plants. wherever the banks of a water-course. for indeed the tundra is but a special type of steppe. and chiefly confined to the summer period. very hot in summer and very cold in winter. and as it is remote from the sea. This very brief description the main features may be said to sum up temperate steppes. though the rainfall is always slight. there is some reason to believe that it is subject to great variation. The region is also continually swept by very severe storms. perhaps cyclical in character. the forest sends long feelers into or climate. the cold desert of the high uplands. a country of As Central Asia generally high mean elevation. Similarly. injurious alike to plant and animal life. the climate is continental in character. Some of the conditions which we noted in the tundra are here repeated. Thus in both we have a seasonal and highly local abundance of of regions of . and the peculiar conditions cause them to be inhabited by special types of animals.

and far more diverse. If the scarcity reigns. As the safety of the species depends in this case upon the strength of the male. the tundra animals then. But as the wealth of the steppe is greater than that of the tundra. are far more exposed in the open steppe than in the forest. alike in time and space. altitude being one of the most important of these conditions. necessarily the majority. as . always. to move about in flocks from one region of abundance to another. For this reason there are frequent alternations Like of famine and plenty. and it is comes past . to prevent death steppe animals are mostly swift. with special adaptations to ensure rapidity of movement. in any case as is. resulting in a great abun- dance of food for the herbivores of the region. soon as the favourable season the height of that alike in summer and in the depth of winter. we have seen. they attempt to escape unfavourlocal conditions by migration. 53 and local scarcity. the steppe animals tend to be social. But produce minor variations in climate. or have singularly keen senses. they are far more numerous. Like the tundra animals as a whole. and therefore they are either burrowers. as with the wild is here of great aid. the able conditions must be rapid.STEPPE FAUNAS OF ASIA AND AMERICA food. and as these migrations by starvation occurring between one region of pasturage and the next. we find that he must win and keep his post as leader and defender by his strength. and that he is The herbivores. the strength of the male enables him to protect the females and young from attack. If the rains of early summer are abundant in the steppe there is an extraordinarily rapid awakening into life on the also a seasonal and part of the steppe plants. asses. The social instinct also In some cases. enabling them to perceive danger from afar. rains fail.

or kill Now no natural barriers separate steppe regions from the neighbouring areas of forest. such as the rodents. Similarly. social animals are less forest. In Central Asia buran or hurricane may practically the dreaded exterminate all life within a given area. We have already seen that this fertihty of steppe animals is associated with strong migratory instincts. deposed by a younger male so soon as his powers begin to Again. of semidesert. risks of storm. seen alike in the steppe rodents and in This is drought and consequent lack of food. As such storms are frequent. the social instinct is an aid in another way. risks associated with winter cold and summer heat.54 STEPPE FAUNAS AND TEMPERATE fail. In the where food is more uniformly distributed and the natural shelter greater. the fact that to overrun the region. as a special character they tend periodically of subsistence within their of steppe animals. and led to its being forest-clad . except such animals as can find a refuge underground. The more helpless forms. must out large numbers of animals. of desert. and so forth. of steppe animals is their great fertility. a season risks of the natural habitat risks of ' — associated with the ' a series of such seasons. these sentinels giving warning of the approach of danger in time for the party to seek safety in flight or underground. frequent. Another marked feature insects like the locusts. of deficient rainfall. We find then. The direction of the winds and the position of Europe on the western border of a great continent give it a moist climate. great natural fertihty is necessary to repeople the devastated regions. means own and therefore to flow out into the neighbouring regions. usually appoint sentinels when they feed.

But the animals of the steppe can only find in burrows and in sleep protection against want of food and extremes of temperature. at least at times. and the shelter and presence of food even in winter in the forest makes it relatively infrequent there. occur in many other temperate regions besides Asia and South-eastern Europe. and their quiescence is reflected in that of ^ = * many of the steppe animals. A great area of land of this type extends from the border of the Canadian forest southwards through the United States in the region west of the Mississippi river. nearly to the Gulf of Mexico. just as the fir-tree itself becomes active whenever the temperature permits. and clothed. while others. rapidly died out. Similar areas occur in the Argentine and in Patagonia in South America. and . though the steppe region of Asia is continued into it through Southern Russia and part of Hungary. with grasses and herbs.STEPPES OF ASIA AND AMERICA 55 almost throughout till man interfered. like the saiga antelope. like the hamster. Another character steppe is of the animals of the temperate the prevalence of hibernation among them. have kept their hold. By extension the term steppe can be applied to all these regions. in South Africa. The squirrels in the fir woods wake whenever the temperature rises in winter. Large areas of land almost devoid of trees. so that here both aestivation and hibernation may occur. and in the south-eastern part of Australia. some of which. The tundra is too cold for hibernation to be practised to any great extent. But the plants of the steppe disappear beneath ground so soon as their brief period of activity is over. We find then that since the close of the glacial period Europe has been continually liable to incursions of steppe animals from Asia.

but their place in nature was taken by enormous numbers of rodents. with trees scattered among the grass. We may then say generally that the steppe regions of the world are the regions which form the natural home of the ungulates most valuable to man. In North America the steppe lands. and such land has some of the characters of a steppe. Again.56 STEPPE FAUNAS AND TEMPERATE there are other large areas in Africa and in the north of South America where open park-like country occurs. within the human period at least. but the fact that ungulates introduced by the white flourished apace man into both countries for us to have suppose that any natural obstacle to their presence there existed. but of the white. displaying the general characters of a steppe. In South America. That South America has few native ungulates and Australia none are two of the most interesting facts in geographical distribution. espeAsia and Africa. No horse. prior to the immigration were almost absent. makes it unnecessary . large ungulates and the natural pasture was utilized by marsupials or pouched animals. in Australia no ungulate whatever occurred. they are the natural home of of the great ungulates. no relative of cattle or sheep or antelope cropped the herbage of the great plains. within the human period. The great cially in interest of the steppe areas is that. which reached here a size not attained elsewhere. Again. of which the most important grasseating form is the kangaroo. but of these few the bison was represented by countless numbers of individuals until it was exterminated by the white man. have been inhabited only by few species of wild ungulates. all the great deserts of the world are fringed by semi-arid areas. the very many specialized of most highly which are steppe animals.

we find that it is in the steppe regions that the cultivation of cereals is being most vigorously pushed at the present time. and the plateau of Tibet. with special conditions of climate. found in the steppes of temperate Asia. Further. Steppe conditions reign over that great area of Central Asia which is practically ringed by mountain chains. has a peculiar fauna of own. and is separated from the forest region to the north by the great area of elevated ground in which the Siberian rivers arise. Excluding Tibet and the mountain we have a great steppe and desert area which extends from the eastern base of the Pamirs to the Khingan Mountains. is the biological equivalent of the flocks of rodents and the flights of locusts which the steppe sent out in earlier days. but the fauna throughout this area is not uniform. and so forth. and from the its chains then. the region to which the following description specially refers. as well as to the west of it. horses. which pours into the civilized world. The mountain chains themselves have their own characteristic animals. of meat. . as steppe regions do not require to be cleared of trees. and therefore were once forest regions. Formerly these watered lands were liable to be periodically flooded by cattle necessary for the the overflow of the fertility of the steppes. and. climatic and other. often produce abundant crops. and of the hordes of wandering nomads which came later. and wants of civilized communities.STEPPES OF ASIA AND AMERICA It is the steppes of the 57 more remote parts of the world which now support the vast herds of sheep. of wool. where the rainfall is sufficient. To this general account we may add a few words on the special conditions. and the steady stream of wheat. The regions of dense population are mostly regions of abundant rain. tide is now This great being gradually regulated.

. Thus the snows of the Pamirs water Russian Turkestan through the Amu Daria and the Syr Daria. by mountain this area. which makes the growth of trees virtually impossible. is continued through Southern Russia in Europe into Hungary. ditions do not reign throughout this great area. and the small precipitation. thus Sven Hedin found temperatures of — 17° F. The frequency dry east winds in winter is an important feature in checking the growth of perennial plants. and is bounded to the south-west by the plateau of Persia. giving a dry winter and a dry summer and autumn. Further to the east much lower winter temperatures die down to the ground. and the river here is frozen for three . while the mean annual precipitation is only 17". the hot summers and the bitterly cold winters. Climatically. and the mean July 70° F. the general features of the steppes of temperate Asia are the great range of temperature. those characterized by the absence of severe winter cold. of the climate.58 STEPPE FAUNAS AND TEMPERATE regions of China to the south-east wooded chains. but no absolutely dry month. and this westerly region as a whole has a much milder winter climate than the easterly region. But uniform con- which the general name may be given. to Through the Dzungarian Gate is of Desert of Gobi or Shamo continuous with the extensive band of steppes which runs through Russian Turkestan to the Aral and Caspian seas. Tarim basin. permitting of the extension into it of animals really peculiar to the southern steppe and desert regions. which shows many alternations of fertihty and of aridity. as much we have seen. with a June maximum. which tends to occur in late spring or early summer. occur in the . except such as of As indications of the nature we may note that at Orenburg the mean January temperature is 4° F..

ungulates is the saiga antelope {Saiga tartarica). In the historic period it has been gradually retreating eastwards. sub- which extends to the Gobi desert. In giving a brief account of steppe animals. the so-called Persian gazelle (G. will begin with the ungulates. the gazelles are more .STEPPES OF ASIA AND AMERICA and a half 59 Daria. is frozen for about a month every year. especially during the rainy season. The steppe animals are thus subjected to very unfavourable natural conditions. The of the steppe Perhaps the most characteristic antelopes are also represented in the steppes gutturosa). The animal occurs in thousands in the Kirghiz steppes. characteristic of the warmer deserts to the south. and the . and the Mongolian gazelle {G. means a great intensity of life for a short period. an ungainly animal with rather short legs. than three kinds of horse-like animals haunt the Asiatic steppe the tarpan or wild horse [Equus less No — caballus). and. but the high summer temperature. as well as in Belgium and Southern France. Formerly it had a much wider distribution to the west. gutturosa). the male having a peculiar swelling on the face. prejevalskii). we shall limit ourselves chiefly to the steppes of Central Asia just described. by two kinds of gazelles. Prejevalski's horse {E. here especially important. thus losing the ground which its ancestors conquered in the ages when steppe conditions were more widely spread than at present. Even the Amu in a western region of relatively mild winters. and the fact that its yellowish coat becomes almost white in winter suggests the severity of the climate in its home. as before. months every year. generally speaking. which makes it appear hook-nosed. a somewhat larger animal but. its remains having been found even in Southern England.

and will drink saline and brackish water. Such a dun tint is common in steppe and desert animals. It is almost by scraping away the snow from uniform in coloration. The kiang (see Fig. so that it is a typical inhabitant of the salt steppe. above. and of Prejevalski's horse is of a similar colour almost white underneath. Its appetite moreover. coat is so thick as to form a kind of fur. Still and occurs at great heights in Tibet. as is the one-humped camel for the sandy deserts to the south. 16) inhabits the higher parts of the a dun colour. The animal feeds largely upon the bitter saline plants of the steppes. legSj which with its long hair. all sorts according to Prejevalski. and camel as representawe may pass on to the consideration of the rodents. horses. hemionus). which are numerous . and traces of striping on the limbs. In winter its. it will devour of animal matter if vegetable food fails. and it obtains food the ground. now limited to the regions of the steppe most remote from human influence. Like its allies it goes about in small troops under the leadership of an old male. for. is another group of ungulates represented in the steppes by the Bactrian or two-humped camel {Camelus hactrianus). tives of the ungulates of the steppes. but is steppes. and is doubtless protective. and has extraordinarily acute senses and great swiftness. remarkably catholic. Taking antelopes. The two humps allow for the storage of fat when the animal is well fed. and thus permit tion. it to withstand periods of semi-starvais. moderately short and hard feet is as well fitted for the hilly and rocky regions of the steppes.60 STEPPE FAUNAS AND TEMPERATE The tarpan is kiang or wild ass {E. instead of being striped as in the African zebras. In the kiang there is a dark stripe down the back. but the body is of a uniform darkish colour.

This is the more remarkable in that there is no resemblance in regard to the ungulates of Fiti. Their chief ungulate was formerly the bison (Bos americanus). no wild horse nor ass. while the bison of the temperate regions of the Old World was a forest-dwelling animal. the two regions. but not in the steppes of Asia. an animal related to the antelopes. The prairies of North America have no antelope. A point of very considerable interest in regard to is the steppe rodents that there is much general resem- blance between those inhabiting Asia and those of North America. found in Europe.STEPPES OF ASIA AND AMERICA 61 both as to individuals and as to species. but differing markedly in the fact that the . 7. no camel. and often show some curious adaptations to steppe Hfe. The other important ungulate of the North American prairies is the prongbuck (Antilocapra americana). Transport Camels in the Australian Desert.

and markedly contrasted with the long tail and large tufted ears of the common squirrel.62 STEPPE FAUNAS AND TEMPERATE trast horns are shed annually and are branched. hibernating in winter. Li the first place. taking small birds and their eggs when occasion offers. we shall refer chiefly in the following descrip- tion to the rodents of the Asiatic steppes. Like the squirrels. but storing in autumn a large collection of roots. berries. both adaptations to the terrestrial and burrowing habit. which has a double significance. and also small rodents belonging to other genera. we may note that marmots occur in both regions. with representatives both on mountains and in the steppes. for winter use. but the common steppe form {S. they are not averse to a certain amount of animal food. and that the jumping mouse of the American and their aUies in the Old World. or recently exterminated. The conbetween the living. just as the tundra forms a steppe because it is too remote from the Equator for tree growth to occur. ungulates of the Old and New Worlds is the more remarkable when we find that America has many fossil horses. though no living ones. &c. that the sushks of Asia are represented by the forms called gophers in America.. The susliks (Spermophilus) are related to the squirrels. citillus) differs in the very short tail and the minute ears. We shall see later that it is not unusual to find and mountain animals. relationships between steppe a fact . In proof of the statement just made as to the resemblance between the steppe rodents in the Old and New Worlds. which belong to the same genus. As the adaptation to steppe life is prairies takes the place of the jerboas similar. The marmots (Arctomys) form an interesting genus. seeds. Like other steppe rodents the sushks are social and burrowing animals.

3 a u •c <I> a Ph .s P-t <a -^ a o O >^ (S a a o .


so common a sound in the mountainous regions of Switzerland in summer. the result of these excavations is . Pyrenees. and tunnel the ground in all directions with their burrows. but also brings back a time when much of Europe was in the condition in which the steppes are to-day. the burrows being extensive. with the gradual improvement of the climate. The food appears to be entirely vegetable. after the retreat when the climate was unsuited to trees. Bobacs occur in very large colonies. If this be so. bodies. were driven up the mountain sides to the Alpine regions. On the other hand. not only recalls the conditions of the Asiatic steppes. of the fore-hmbs here gives greater purchase in burrowing. the bobac {A. c> The shortening ^ According to Kobelt.STEPPES OF ASIA AND AMERICA the elevation prevents forest growth. The shortness of the hmbs must greatly Kmit the range of vision. clifmsy animals with heavy and usually short tails. the characteristic marmot of the Asiatic steppes. and Caucasus. eastern frontier of Germany. then the shriU whistle of the Alpine marmot. extends into Europe as far as the ing to a wide extension of steppe conditions. and the return of forest-haunting forms. The Alpine marmot is now limited to the Alps. and the animals have very markedly developed that habit of sitting up on the hind Umbs to take a wider Marmots are rather view which is common in relatively short-legged animals. which. it is believed that in those parts of the world recently sub- was a period. short limbs. jected to glaciation there of the ice. lead- This brought in its turn a wide extension of steppe animals. and the animals hibernate much more profoundly than the susliks. hobac). the time being far past when it also inhabited the low ground. 63 so the upper parts of mountains form steppes because Again.

however. The jerboas show a curious form of compromise to these varying needs. though no less perfect. if the legs are elongated to give speed. shows also a general resemblance in habits. A different. Thus the prairie marmot {Cynomys Ivdovicianus). generally parallel to that shown by the kangaroo. In feeding.64 STEPPE FAUNAS AND TEMPERATE soil. fore-legs The animal therefore progresses by leaps. In the jerboas the fore-Kmbs are short. as is seen in the giraffe (Fig. of the large leaves of these plants and the presence among the lyme a very obvious feature. the not touching the ground at all. 37 and 38). has similar habits. on the other hand. though not very nearly related. then feeding on the ground steppe Hfe and drinking become difficult. to modify the condition of the so that special '. 9). and also modifies the herbage in the regions of its occurrence. In speaking of the marmots we noted that for rapid and efficient burrowing. marmot gardens which can thus be recognized from afar. and its speed cannot be great. On the other hand. not a burrowing animal. which is. but the hind-legs are enormously long (Figs. if the hind-legs are also short. and the tail also long and capable of helping in the support of the body. The interesting point is that rodents of similar habits seem to inhabit steppe regions in general. but. The hemp-leaved nettle plants appear in the ' and wild rhubarb occur especially grass (Elymus) of the steppes is here. short fore-legs are necessary. while the viscacha of the pampas of Argentina. a related form inhabiting the prairies of North America (Fig. thus permitting the animal to burrow. the animal's range of vision is small. 36). . while the loosening of the soil by the burrows makes ' progress for horsemen very difficult over the gardens '. adaptation to is shown by the jerboas or jumping mice.

while on the other hand the bobacs at once seek their burrows on an alarm. and is an adaptation to swift movement over a relatively hard surface. Another peculiarity of the hind-foot. for have only one toe on each foot. the five-toed jerboa as it is called. and not wholly of grass. which it shares with other jerboas. and make underground tunnels to the feeding-grounds.. it is animal only measure some it cannot be overtaken by a horse. carried its allies The commonest jerboa of the steppes is Alactaga decumana. Though the head and body together seven inches. As a result a very strong bone called the cannon bone is formed. As in rodents in general. forming of this stated that 1404 v\ . as in the kangaroo and in most ungulates.STEPPES OF ASIA AND AMERICA 65 a crouching position is adopted. Like the other steppe rodents the jerboas are social. Another very interesting structural point in regard to the jerboas is the reduction of the number of toes from five to three on the hind-foot. is the fusion of the bones which form the sole of the foot in man. similar reduction. berries. This also is an adaptation to ensure swift movement over firm ground. because the lateral toes are present on the hind-feet. the food being carried to the mouth by the fore-Hmbs. quite comparable to the cannon bone in the antelopes and their allies. The curious elongation of the hind-legs gives such speed that the jerboas can escape their enemies by flight. so that they are never far from an open burrow. though it is formed in a different way. the food consists largely of roots. though they are minute and functionless. the horse and Among the ungulates we find a however much further. seeds. for it gives the necessary rigidity during the taking of the long leaps. &c. so extraordinarily rapid is its speed. This reduction gives speed. that is of the metatarsal bones.

they are (cf. notably D. but some species. According to Prejevalski. They are remarkable for the size and elaboration of their burrows. extend into the steppes. however. for winter use. essentially in- habitants of arid regions p. They form a very important part of the food of the carnivores of the high steppe. These animals are smaller than a rabbit.66 STEPPE FAUNAS AND TEMPERATE insects burrows in companies. these haystacks may weigh 10 kilogrammes . sagitta. tailless hares or whistling hares as they are variously called. the genus which includes the picas. like extent. replacing here the lemming of the tundra. Another characteristic genus of the steppe is Lagomys. But though originally doubtless steppe animals. In MongoHa. though to a less have attached themselves to man.. especially of the steppe lark. In the steppe region are also found the hamsters (Cricetus). and at the approach of winter stores up large quantities of hay near the openings of the holes. than to depend upon the precarious vegetation of the steppes. in which they store quantities of corn. the hamthe bro^^Ti and black rats. they are found on lower ground. ogotona occurs in large colonies. with a western extension into Europe. sters. Kving high up among rocks and stones in the mountains of Central Asia. Though the habits of the members of this genus are generally similar to those of the preceding in their coloration and in their intolerance of rain or damp. In Siberia. and are for the most part mountain animals. The jerboas of the genus Dipus chiefly inhabit the hot deserts further south. &c. L. 133). mmgle With the vegetable food they and the eggs and young of birds. and even extend into the tundra region. and find it more profitable to plunder his fields and gardens.

rhea. about the size of the common cat. perhaps in more than one variety. emu. but. E 2 a great . and can Hve in localities far from water. Here as elsewhere the ubiquitous wolf also occurs. the corsac fox (Canis corsac). The special cat of the steppe is. and so dispense with the trouble of storing fodder on their own account. Its main food consists of the steppe rodents. 67 sufficiently numerous for it to be Mongolians to bring their cattle to feed upon the pica's stores. examples of birds without any power of flight. though as a tropical animal. thus showing parallelism with the mammals. it does not disdain the rodents. In regard to the birds of the steppe it is noticeable that they tend to acquire swiftness in running. and apparently alhed to the wild cat of Europe. usually regarded to the north. a beautiful animal with a bushy tail. Like some other of the steppe animals.. a steppe animal. As to the carnivores of the steppe. extends far and occurs in steppe regions. which also feeds chiefly upon the steppe rodents. where no rain nor dew occurs for months and are worth while for the at a time. offer safe places upon which to rest. As the is birds are exposed to danger so soon as they alight. like the forest. &c. the tiger. The reason for this tendency is possibly that the steppe or desert does not. unlike it. When nobler game fail. where we have in ostrich. but of great fleetness of foot. the manul cat {Felis manul).STEPPES OF ASIA AND AMERICA (22 pounds). fleetness advantage to them. notably the picas. special fox. the picas are remarkably resistant to thirst. and also of The steppe has also its the ground-nesting birds. This is carried further in the birds of the warm deserts of the world. however. which is necessarily upon the ground.

tetrax) and the ruffed or Macqueen's bustard {Hubara undulata) also occur. The plant is also reUshed by horses and camels. are fleet of foot. and having the habit. In addition to the form mentioned.68 STEPPE FAUNAS AND TEMPERATE As examples of steppe birds. but Pallas's sand-grouse {Syrrhaptes parais characteristic. This is necessary in birds whose eggs are laid in a mere depression scratched in the loose soil. The young are precocious. to increase the range of vision over the plain. of squatting down when undisturbed. we may name first the bustards. just enter the region. the latter preferring running to flying. the ostrich of the Saharan desert. the great bustard has been compared in build and habits to Bustards. both the young and the adult showing a delicately patterned type of coloration which is eminently fitted to conceal them in their natural habitat. but stretching itself to its (considerable) height on an alarm in order {Otis tarda). common among steppe animals. and show a preference for open country. the little bustard {0. though good fliers. which are bush-haunting birds. reaching even the British doxus) . so as to take full advantage of its protective coloration. Pheasants. They do not seem to drink. when it makes incursions on lands adjacent to its normal habitat. being able to run as soon as hatched. and its seeds are used as food by the nomads as well as by the sand-grouse. and it assembles in large flocks in places where this plant of the salt wastes is abundant. In the steppes the bird is said to feed chiefly on the seeds of a member of the Chenopodiaceae called AgriophyUum. The sand-grouse shares with many steppe animals the peculiarity of being periodically very abundant. Flocks of it have appeared at various times in Europe. of which the largest form. and take both animal and vegetable food.

and They haunt the desert regions where bushes of saxaul and tamarisk grow. Reptiles in the steppes and adjacent deserts are not numerous in species. sometimes called chough-thrushes. or squat escaping notice. is 69 The bird differs from the bustard in that water it. as means of In the steppes of both the Old and New Worlds such forms as curlews. plovers. preferring among the herbage. but the individuals are often very abundant. they only fly with reluctance. feeding upon insects and seeds. and so on. geese. lizards belonging to the genera Phryno- . Fig. just as Pallas's sand-grouse feeds Asiatic steppes phasianellus). snipe. are abundantly represented in the breeding-season. but do not show any very notable adaptations to steppe life. Siberian lark {Melanocorypha calandra). ' brush ' {Artemisia tridentata) in the western states. In the American prairies game birds are represented by the cercus related to the grouse {Tympanuchus americanus). 10). with many of the smaller singing-birds. Though capable but run of flight. and do not seem to drink. alliance. Very characteristic of the Asiatic steppes are the birds belonging to the genus Podoces. a form also by the sage-cock {Centrourojphasianus. which are members of the crow their though exact position is doubtful. necessary to and it makes regular daily visits to particular drinking-places. ducks. nesting in bushes or sometimes on the ground. which feeds upon sageprairie-hen . while the more grassy regions of the steppes often swarm with the swiftly. . when hard pressed. In that part of the desert of Gobi which is called Alashan. to run along but only rise the ground.STEPPES OF ASIA AND AMERICA Isles. upon Agriophyllum in the and by the prairie-chicken {Pedioecetes All these can fly rapidly and powerfully.

70 STEPPE FAUNAS AND TEMPERATE life. They are also greedily consumed by birds of and even by such mammals as wolves and foxes and domesticated dogs. (Photo by the Biological Survey. 10. Turkestan there occurs a curious gecko {Teratoscincus scincus). and so forth. Sage-cock. Among snakes we find the small sand-snake (Eryx jaculus).) prey. which enable it to run over the loose sand. in power in of burrowing. cephalus and Eremias are very abundant.A. g. and show special adaptations to desert swiftness.S. So abundant are they summer that they attract flocks of the demoiselle crane {Grus virgo). and has acquired curious fringes. in the in coloration. which feeds upon Fig. In the steppes and deserts of them. U. a harmless . e. which has lost the disks on the digits by which its alhes chmb.

by Lull (1904). 1889). as well as it buries itself in the sand vipers and others. six vols. by Shimer (1903). insects we need only note that grasshoppers and locusts are very abundant. and is 71 interesting from the way in which means halys. dangerous to by and the poisonous Trigonocephalus cattle but not to man. e. . and Tossorial Adaptations in Mammals'. mention may especially be made of Prejevalski. See also the books mentioned at the end of chapter iv. Refeeekces. which extends into Greece. especially 'Cursorial Adaptations in Mammals'. g. Amphibians are limited to the regions in the vicinity As regards of water. A series of articles in the American Naturalist. will be found interesting in this connexion. Wissenschaftliche Resultate der nach Centralasien unternoinmenen Reisen (Leipzig. and offer no special features. of its snout . edited by Shipley and Harmer. open grassy plains being the natural home of these insects. the relevant volumes of the Cambridge Natural History (London). The works of give accounts of steppe animals. from which the locusts make periodical raids upon cultivated lands.STEPPES OF ASIA AND AMERICA form.. edited by Lydekker. where details will be found of the different animals mentioned. Kobelt and Brehm already mentioned and reference should also be made to any of the natural histories. Of the various books published as the result of travels and giving accounts of the life of the steppes. or The Boyal Natural History (1893-6).

we may note that elevated ranges may be divided into three regions. At a point which varies with the exposure. and there comes a zone with steppe characters. The width of this band and its nature vary greatly. g. with its Alpine plants plants e. but with tufted leaves and other indications of checked vegetative growth. Many of these plants reappear in the Arctic tundra. g. In e. on the hills of Scotland or Norway.CHAPTER The some IV MOUNTAIN FAUNAS animals of mountain and plateau regions show make it worth while Taking mountains first. each of which presents pecuhar biological interesting conditions which to consider them separately. Senecios. In tropical regions. it is characterized by the conifers predominating as growth of relatively small herbaceous plants. this steppe area is replaced . whether they occur in the Alps proper. often with large and gorgeously coloured flowers. the mountain regions of Europe. where . The lowest region is usually clad in forests. features. by a band where the show desert characters the Lobehas. In all regions where the mountains attain sufficient elevation. and tree-heaths of the upper zone of Ruwenzori and Mount Kenya form good examples. above the steppe or desert zone comes that of eternal snow. even on a single mountain. we ascend. the slope. and other factors. or in the Caucasus. in the Himalayas. but the fact that they are specially abundant in the Alps has led to them being called Alpine plants. tree growth ceases. in Africa.

where the surface is narrow alpine zone on. &c. the temperature that it we may note diminishes steadily as we ascend.MOUNTAIN FAUNAS and lichens. due to inversions. much above the zone of maximum precipitation. In tropical regions the snow-line is pushed far upwards. such as the great plateau of Tibet far less extensive plateau of Spain. the low temperature increasing the need for xerophytic characters by diminishing the power of absorbing water. Plateaux of great altitude and great horizontal extension tend to show some of the characters of a cold desert. but gives to those possessing such agility a certain .g. 73 the exposed rocks arc at most scantily clad in mosses in detail the characters of moun- seems probable that the upper limit of the forest corresponds roughly to the zone of maximum precipitation. Plateau regions. but the conditions tend to be less which reign extreme than in small and isolated areas of great elevation. With certain modifications... thus giving us the special conditions in Tibet. which demands special agility upon the part of the animals inhabiting them. Without discussing tain climates. e. beyond which it begins to diminish. the Alps clad in plants of definitely drought-resisting type. Plateaux differ from other steppes or deserts in the peculiar nature of the relief. The is relatively due to the fact that the vertical distance between the zone of maximum precipitation and the snow-line is short. may be said to correspond to the steppe zone on an ordinary moun- and the tain. and the wide interval thus produced between the forest and the snow allows for the development of a broad band of gradually increasing aridity. In ascending a mountain the rainfall increases up to a certain point.

in certain localities at least. of Fig. according to some observers. 11. the presence of a close low bushes (mostly berry-bearing). . yet there is pasturage enough to spare to feed the domesticated animals upon which the community largely depends.74 security. In Tibet. The richness of the pasture is suggested by the number of cows short turf and particularly nourishing fodder plants. MOUNTAIN FAUNAS due to the rugged nature of the surface. together Further. probably with the rapidity of mechanical erosion. the pecuHar cHmatic conditions. though wild herbivores exist in flocks of thousands. promote the growth. of The absence of trees. are very characteristic of the upper regions of the Alps as well as of many other mountain and plateau regions. A Swiss Alp. A large part of the pastoral industry of Switzerland is due to the pasturage obtainable on the high mountain shelves or alps.

and they seem able chffs. In the case of sheep. and as the trees in their native habitats are mostly xerophytic. — — acter. the equally agile goats depend largely upon the shoots. and twigs of bushes and small trees. Like steppe animals in general. domestication has produced so many changes in character. especially ungulates. so sheep and goats are the typical inhabitants of elevated regions. wild sheep are and Kke many such animals they seem mostly Their powers of to scale to appoint sentinels while feeding. leaping are very great. apparently inaccessible While wild sheep. the difference in diet the goats usually occupy ground which is more scarped and rocky than that favoured . From tastes. social. or spiny plants native to high steppes. and antelopes typically steppe or savana animals. but in the Old World at least the two genera of sheep (Ovis) and goat (Capra). like domesticated forms. Just as deer are typically forest animals. at least. which appeared late in geological time. The ungulates of mountains belong to more than one group. hairy.MOUNTAIN FAUNAS The relatively 75 relative rich pasturage. and the account for the fact that mountains and plateaux have a considerable number of peculiar herbivores. it is noticeable that the goats have very catholic not disdaining the resinous. and their other special adaptations to Hfe in mountainous security. The thick coat protects them and the fact that the young necessarily few in number in animals with such a habitat are able to follow the mother anywhere within a very short time after birth is an important adaptive char- and elevated against cold. chiefly graze upon the ground. regions. that it is somewhat difficult to realize the extraordinary agility of the wild animals. leaves. include animals which are especially adapted for mountainous districts.

As a general rule the latter seek to capture their prey by ruse rather than by direct pursuit. where. There is evidence that the herbivores acquired swiftness as the carnivores acquired intelli- gence and strength. other types of ungulates also show adaptations to mountain life. For convenience' sake we shall take a rapid systematic survey of the animal kingdom in studying mountain animals. and there the carnivores can hardly be said to have followed them. Just as the by two latter types of animals are of relatively it is recent origin. doubtless. hke the ungulates. As will be explained later. To escape them. The characteristic . there being few large flesh-eaters at great elevations. and they have apparently even greater powers of chmbing and leaping. are represented high up in the region of coniferous forests. and in the open steppes and plains their task is relatively easy. some herbivores sought the heights. so probable are the adaptations to mountain life shown by other mountain ungulates. especially in those regions which have no wild goats or sheep. the Primates or monkeys. but it may be noted first that as forest. they find relative security from the great carnivores. For example. apparently. steppe. no less than to utilize the rich mountain pastures. always conspicuous in such regions. in the Himalayas we find that those truly arboreal animals. and tundra are all represented on mountains. Indeed the fleeing to the mountains appears to be the latest phase in the long struggle between herbivores and carnivores. it is natural that animals from all these regions should occur there. Indeed the chief flesh-eating animals of the mountain ranges of the globe are the vultures and their allies.76 MOUNTAIN FAUNAS sheep.

a group of slender-limbed. and has a curious tiptilted nose. there can be no doubt that the Tibetan form has found its way into China from the south. and the number of forms found on the lower ground in India and the adjacent region proves that its the acquisition of the mountain habitat in this case must be recent. which lives in the forested regions of the east of Tibet. The particular species is Semnopithecus schistaceus. and its allies while sharing with arboreal to life the usual adaptations to —to life be described later — it show any special fitness for mountain is does not seem life.000 feet. . especially in the vicinity of Lake Kuku It is found at Nor. elevations of over 10.MOUNTAIN FAUNAS monkey of the 77 Himalayas is one of the langurs. leaf-eating forms widely distributed over South-eastern Asia. but never descends to the region of cultivated land. Better adapted for at high elevations the Tibetan langur (S.000 feet. and migrates to some extent according to the seasons. All who have seen it in native home speak of the peculiar effect of the animal among the snow-covered pines and deodars there. unless it has any- thing to do with facilitating respiration at these high prized The hair is very long. The animal is stated to live chiefly on fruits but no doubt the seeds of the mountain trees are also important. The Tibetan langur is somewhat stouter in build and shorter in the limbs than the other forms. As the other monkeys of China are species of Macacus. related to the common The Himalayan form is very closely sacred monkey of India. and also extends into China. and the skin is greatly by the Chinese on this account. roxellanae). through the mounaltitudes. ' '. the district plundered so frequently by the low ground forms. whose use is unknown. and the animal occurs at heights up to 12.

quite dififerent from that of Tibet proper. As this food is much more portable than leaves. have a damp chmate. The . which seems to occur in much the same region as the Tibetan langur. where it . notably the two monkeys named. and also extends into China proper. in Sechwan and the south of Kansu. The macaques differ from the langurs in their diet. they have cheek-pouches in which it may be stored. MOUNTAIN FAUNAS Its presence is thus an indication that mountains do not necessarily form a barrier to distribution. The macaques also include one moimtain form. The narrow and broken mountain chains of this region. which have large sacculated stomachs. The Himalayan and Tibetan langurs and the Tibetan macaque appear to exhaust the members of the Primates which haunt mountains. None of the New World forms seem to be specially fitted for life at high altitudes^ and though some of the African baboons inhabit rocky regions at some elevation. none seem to be pecuHar to high mountains or plateaux. even to animals reputed to be so sensitive to cold as monkeys. for they eat insects and other forms of animal food in addition to fruits and seeds. The insectivores have various mountain representatives. Alpine shrew {Sorex alfinus) inhabits the upper parts the of the lofty mountain regions of Central Europe Himalayan swimming shrew {Chimarrogale himalayica) inhabits the lower slopes of the Himalayas. and mto the valleys the monsoon forest insinuates itself. according to recent observers. these being absent in the langurs.78 tain barrier. comparable to the stomachs of some ungulates. Macacus tibetanus. some showing peculiarities of distribution. taking with it part of its characteristic fauna.

79 haunts streams and. Of the cats the most characteristically mountain form is the ounce or snow leopard {Felis uncia. greyish above.000 feet in summer time. battle of life. and there is nothing remarkable in the fact that peculiar forms are found in mountain ranges. also into the —may extend a beautiful animal with very thick fur. the bear alliance being that with most mountain representatives. and small fish the desman {Myogale pyrenaica) of the Pyrenees inhabits the banks of streams in that mountain range.000 feet in winter. In Ladak it is said to ascend as high as 18. are not very abundant. and is stated to prey upon the Bighorn sheep. 12). and not to descend lower than 9. attains a considerable height in the Rocky Mountains. It attacks wild goats and sheep.MOUNTAIN FAUNAS their larvae. seeming to be intermediate between moles and The insectivores are. Fig. mole-shrew {Uropsilus soricipes) is a small cursorial animal. as forest or steppe we have seen. feeds upon aquatic insects and tadpoles. however. Carnivores. Of other cat-like carnivores extending into mountainous regions. and also rodents. but peculiar forms are not numerous. regions The forms which inhabit the neighbouring mountain regions. and pure white below. . though not specifically a mountain animal. 26). In the New World the puma {Felis concolor. with black spots. Fig. when we remember the shifts the members of the group have been put to in order to survive in a world where the mammals as a whole have better developed brains and greater differentiation of structure than they can boast of. we may note that the lynx of the . but its habits are not well known. losers in the shrews. and spends a conthe Tibetan siderable part of its time in the water .

in America contain another mountain shape of the spectacled bear {Ursus ornatus). That forested region of Eastern Tibet which lodges the Tibetan macaque and the Tibetan langur has still of South the . of which several occur in mountain regions. &c. also Its very indiscriminate appetite it makes it possible for to exist on many different kinds of ground. though also descending to the In the wooded regions of the Himalayas occurs the black bear (Ursus torquatus). into the plateau of Tibet. the packs hunting down wild sheep. It is known short is and differs from most bears and thin coat. antelopes. We come next to the bears. the common fox is replaced by a variety with thick fur and a very large brush. Africa south of the Sahara has no bear. which in Tibet. instead of to forest-covered ground. considerable elevations. when the animal is very fat a means of providing for the winter time of scarcity. in the Himanot in the layas does not voluntarily quit the forest. The Indian wild dog {Canis deccanensis) shows a similar extension of habitat from the forests of India into the wastes of Tibet. Thus the Himalayas and Tibet have special varieties of the brown bear. It is a social animal. like bears in general. is very resistant to cold on account of the thickness of its coat. In the Himalayas and Tibet also. especially in autumn. which. related The Atlas Mountains have a closely form. The Andes bear. chiefly a vegetable feeder and an excellent climber. The necessary resistance to cold apparently obtained by the great oiliness of the skin. and in the Rocky Mountains the large grizzly bear {Ursus horribilis) occurs at and powerful plain. where it shows some adaptations to rocky and barren country.80 MOUNTAIN FAUNAS northern forests extends in a special variety. or species {Lynx isabellina).

as fs* .

^S' Ph j^id^ai .

sometimes. the Bos grunniens — — — — of zoologists. once beheved to be related to Aeluropus. occurs at considerable elevations in the south-eastern Himalayas and feeds chiefly on vegetable matter. on account of its great endurance. especially bamboo shoots. we may note that just as one form the muskox is admirably adapted for life on the timdra. the animal is used as a beast of burden in Tibet. so another the yak of Tibet is perfectly fitted for life at a great elevation. and by great agihty and hardiness. and it is stated to be an excellent climber. heat. As is well known. The latter feature is remarkable in view in tastes.MOUNTAIN FAUNAS . A curious little animal called the panda {Aelurus fulgens). Beginning with the cattle. the abimdance of ungulates in mountain regions. has a very thick close fur. seems to be purely vegetarian bamboo thickets which in this region ascend the damp valleys up to a height of the fact that the animal It is said to inhabit the The young shoots of bamboos form the greater part of its food. and while impatient of and is capable of thriving on the coarsest of herbage. as we have seen. which are forest animals. and is now believed to be most of nearly 10. This animal. its is characterized by its long hair. somewhat fox-like in appearance. 81 another remarkable form in the parti-coloured bear {Aduropiis melanoleucus) It is curiously marked in black and white. The panda is an arboreal animal. Travellers have often 1404 V extraordinarily resistant to cold. .000 feet above sea-level. and enormously powerful jaws. nearly related to the carnivorous raccoons of America. The comparative rarity of large and fierce carnivores helps to explain.000 feet. is It ascends to a height of 20. ascending to a considerable elevation above sea-level. as in the case of the forms called coatis (Nasua).

while the other {Ovis hodgsoni) occurs at great heights on the Tibetan plateau. like the reindeer of the tundra. Of the remaining two one. the animals in winter seek exposed positions where the wind sweeps away already alluded may add . the Indian urial. reaching the size of a small donkey. The and have been to. They present a remarkable instance of adaptation to very unfavourable conditions. one. the Bighorn. and. so closely related to the wild sheep of species. form is found is in the Rocky Mountains. The yak is the only member of the cattle group with mountain life. To the preceding description we a brief note on the wild sheep called argali. all only the urial extending south of the Himalayas. Both are large animals. eight inhabit various mountain chains and plateaux in Asia. but in this connexion it is important to remember the great extent of the plateau of Tibet. are generally similar.82 described the MOUNTAIN FAUNAS way in which the animals push their way through snow. but this. The food consists of grass. and having short and close hair. Passing to the sheep we find that there are eleven well-defined wild species. but also that the home of is the mountainous regions of Central Asia. the mouflon. and its great uniformity. mingled in winter with mosses and lichens. the Barbary sheep. occurs in mountain regions in Sardinia and Corsica. or through icy glacial torrents. extending to relatively low ground in the Punjab and Sind one special adaptations to . Of these. Kamchatka as to suggest a former land connexion across the Bering Strait. We are thus justified not only in saying that sheep are definitely the genus habits of mountain animals. and the other. of which one form {Ovis ammon) is found at moderate elevations in Mongolia. is found on the southern slopes of the Atlas Mountains in Africa. or across glaciers.

(Photo by U.S. a form closely related both American Bighorn and to the wild sheep of Kamchatka.A.) the Biological Survey. . to the The Alaskan Wild Sheep (Ovis dalli). 14.Fig.

Photo by Ca/pt. -Tli^r-ai-". Bailey. 15. ( Tlic iuryu lateral hoofs are a special feature. M. Half-wild Kiang. F. 16. F.t^^ldk^PSM^ --T!^:>y^-^ Z FiG. Tibet. near Gyantse. Bailey.- A young Takin in captivity in Tibet.) . Fig. The type of country is very {Photo by Ca/pt. characteristic. M.

Like agile. with long spirally twisted horns in the male. such as Hemitragus. all wary and The goats are mostly distinguished from the sheep by the presence of a beard on the chin. Some ten species of wild goat occur in the Old World. In Europe there is a curious tendency for the (isolated) goats. except that there is no true American goat. and no less than three species of wild goats inhabit the Caucasus.MOUNTAIN FAUNAS the snow. two species. while the takin of Eastern Tibet is a large form with curiously shaped horns. Of the Asiatic species the most attractive is the markhor (Capra falconeri) of the Himalayas. their allies. which seem to connect the goats with the antelopes. enter Africa. there being thus a curious separation between the range of the two species. now exterminated as a wild animal. the ibex of Abyssinia and the Arabian wild goat. The interest of these genera is merely that they emphasize what has F 2 . an ibex the Alpine ibex. sometimes . which extends into Upper Egypt. The gorals (Cemas) and serows (Nemorhoedus) are Himalayan and Tibetan forms. the animals are very while the female has smaller and thinner structures. and of these. 83 In the male the horns are large and massive. called mountain chains to have peculiar species of Thus there is a Spanish wild goat. their distribution showing a remarkable analogy to that of the sheep. and fringes of hair on the chest and shoulders in addition to the beard. and another species found in the Nilgiri Hills of Southern India. including the tahr of the Himalayas. and by the strong odour of the males. is peculiar to that chain. The remainder occur in the mountain regions of Europe or of Central Asia. They also are mountain animals. Closely related to the goats are a series of smaller genera.

a rather clumsily built animal with short hooked horns. and adjacent regions. notably the Tibetan gazelle which ranges up to 18.84 MOUNTAIN FAUNAS been said already as to the number of kinds of ungulates found in Tibet. It is widely but discontinuously distributed over the mountain regions of Europe. shy than most gazelles. is and less Siberian steppes. which apparently give it great sureness on the rocks. notably the rhebok {Pelea capreola) of the south and east. In the distended nose it somewhat resembles the saiga of the {Gazella picticaudata). and the animal seems to be very resistant to cold. Tibet has also a peculiar antelope. an animal in which the male has a curious swollen nose and long horns. Both its Dutch and Latin names refer to its jumping powers. the Himalayas.000 feet. which has the habits of a chamois. but is found fossil on the low ground also. Of the true antelopes many of the gazelles ascend to considerable elevations. the chiru {Pantholops hodgsoni). This form occurs at the same heights as the gazelle. often called the Rocky Mountain goat. stumpy tail. and having very small feet. . and feet especially fitted for scrambling about rocks. there occurs in the Rocky Mountains a white ungulate.000 or 9. frequent Of the vast number of African antelopes some hill regions. Nearer the antelopes than the goats is the chamois {Rwpicapra tragus). found at heights up to 8. Though America has no true goat. about the size of a large sheep. sometimes in very large herds. although it also descends at times to sea-level.000 feet in Abyssinia. Its hair is very long. and apparently related to the serow. Another agile and chamois-like African form is the klipspringer [Oreotragus saltator). spending its time near the upper limit of the forest.

M. (Photo by Capt. F.Fig. Bailey. 18. one Note the curious markings on the of the wild sheep of Tibet {Ovis nahura). Bailey. M.) . 17. F. The Serow of Tibet. The Bharal or Burhel.) Fig. legs. (Photo by Capt.

canines of the male. Note the large lateral hoofs. the coarse coat. 19.) . the promment {From a specimen in the Royal Scottish Museum. Musk-deer.Fig.

000 feet in this is partly the hotter parts of their range.000 to 8. The kiang (p. At the wet season they go high up the mountain sides. The vicuna has a more limited extension. the hair being long and coarse. and the animals seem never to descend below 6. South America is singularly poor in ungulates. or 7. which are no bigger than hares.000 feet above sea-level in Ecuador and Colombia. they avoid equally rocky regions and snow and ice. ascends to an elevation of 7. due to intolerance of heat seems indicated by the fact that the guanacos come down to sea-level in Patagonia. which apparently assist it in obtaining foothold on hard snow or rocky ground. One of the American tapirs also {Tapiriis roulini). showing any special adaptations to mountain life. apparently. In the Andes of Chile and Ecuador also occur the tiny deer of the genus Pudua. These animals have very soft feet. and giving much protection against cold.000 streams. From the Himalayas the musk-deer extends northwards into Central Asia and Siberia. the guanaco and the vicuna. The coat is thick. 60) occurs in Tibet as well as in the Mongolian steppe. As already explained. It has imusually large lateral hoofs. and feeds largely on twigs and buds. but in the Andes two members of the camel family occur. That . being usually found in the birch forest above the zone of pines. being confined to the district between Southern Ecuador and Central Bolivia. but when the vegetation there dries up at the approach of the hot season they descend to valleys watered by springs or perennial The long coat affords protection against cold. and though they ascend to the highest ridges. without.MOUNTAIN FAUNAS 85 Of the deer we may note that the somewhat primitive musk-deer {3Ioschus moschiferus) occurs at considerable elevations in the Himalayas.

those attaining a great elevation seem to differ chiefly by flight.000 to 11. in which they feed. cannot escape and cannot burrow. only two possible means These are to haunt rocky country where the fallen stones and blocks offer natural shelter. They have four welldeveloped toes on the fore foot. breed. and resembling most helplessness and want of those animals also in their of swiftness. . show a marked difference in habit in that they live entirely among trees. where it pops in and out of the crevices as a rabbit pops in and out of its burrow. of escaping their enemies remain. and Being relatively hardy they occur at great sleep. with the spreading toes. called hyraces (Procavia) as regards adaptation to mountain All are small animals. and three on the hind. AU the hyraces are confined to Africa and the adjacent regions. e. or to seek the trees. very like rodents in appearance and habits. without any notable difference in structure.86 MOUNTAIN FAUNAS The very aberrant and somewhat primitive ungulates show some interesting features life. g. though frequenting suitable ground elsewhere in addition. and the size of the feet. of Kilimanjaro. The common hyrax of Syria gets its name of rock coney from its habit of frequenting rocks. As rocky country is more frequent in mountainous regions than elsewhere. in their thicker coats. enables them to scramble about rocks very easily.g. WTien small herbi- vores cannot defend themselves actively. we find that the ground hyraces usually occur in elevated districts.000 feet in the forests In both the ground and tree forms. at from 7. e. Both methods are adopted by the hyraces. elevations. Among the true rodents we find. The tree hyraces. Syria and Arabia. many mountain species. In the consideration of steppe faunas something was said of marmots (Arctomys). as already indicated.



with its soft thick fur. inhabits the higher parts of the Andes. South America. in particular localities. natural that they should occur also in the steppe region mountain chains. It has been found on the Finsteraarhom at a height of 12. curious Alpine or Alps and Pyrenees there occurs the very snow vole {A. even above the snow-line. run into varieties. The Himalayan marmot.MOUNTAIN FAUNAS and as these are typical denizens of lofty is 87 it is of cold steppes. lemming. resembling in habits the Alpine form. though none extend to the south of the Himalayas. It differs notably from these forms in its habits. for it lives high up the mountains. allied to the bobac of the steppes. Thus the common chinchilla {Chinchilla lanigera). has a considerable number of peculiar mountain rodents. and displaying marvellous agility. which is sometimes pure white. In Europe the Alpine marmot and the Caucasus. or species. the usual elevation being 12. but on account of its habits is rarely seen. living in rocky ground. to the barren wastes which most nearly resemble the home fou/id in the Alps. A larger form {Lagidium cuvieri) inhabits the . though it once occurred on the low ground also. nivalis). the genus showing a remarkable tendency to feet. but is structurally very near the Thus in the continental field vole and the bank vole.000 to 13. Other mountain species occur in the Himalayas and Tibet. The voles (Arvicola) are well represented in mountain regions. is limited to the region above the tree limit.000 feet. making runs beneath it after the fashion of the Arctic in its search for vegetation. with its paucity of ungulates. In Central Asia many kinds of marmots occur. It appears to live actually beneath the snow. that is. of the bobac. the Pyrenees.000 In North America we have similarly a Rocky Mountain marmot.

which consists of rodents. on the other hand. found in the Alps as well as in Asia and North Africa. Among birds we find that many kinds of birds of prey haunt mountains. which lives in large societies in the upper part of the Andes. thus illustrating the marmots. found in the mountain ranges of Europe. the young of ungulates and game birds. which inhabits barren mountain sides. kills its own prey. we have already noted that the picas or tailless hares occur at great elevations same in the Himalayas in addition to frequenting the Siberian steppes. related to the Arctic hare already discussed 22). and the magnificent condor {Sarcorhampus gryphus) oi the Andes of South America. tunnelling the ground in all directions with its burrows. has become virtually a bird of . though they are not necessarily confined to them. and Central Asia. and has similar habits. The even more widely distributed Golden Eagle {Aquila chrysaetus). and in this connexion it is curious to note that the New Zealand kea {Nestor notabilis).000 feet. same conditions The mountain hare of Europe is as the closely (p. which does not descend below 14. Other forms which may be mentioned are Bonelli's hawk-eagle {Nisaetus fasciatus). Some of the cavies also ascend to momitain regions.000 or 15. the most interesting being Lepus hypsibiiLS. notably the Bolivian cavy {Cavia holivensis). Of wide distribution is the laemmergeier {Gypaetus harbatus). which apparently feeds largely on carrion and animals not killed by itself. In tropical countries some of the parrots range up moimtain sides. Tibet has several species of hares. The pacas of the Brazihan woods are represented on the mountains by a smaller type {Coelogenys taczanoivskii). North Africa. As to the hare-like rodents.88 MOUNTAIN FAUNAS regions.

found by probing in rocky. Both in the Alps and in Central Asia is found the curious wall-creeper {Tichodroma muraria). At first confining itself to dead sheep. smaller than the common form. Thus in the Alps we find the ptarmigan. In the mountain ranges of South Europe generally the Greek partridge {Caccabis saxatilis) lives. and having a yellow instead of a red beak. which lives on insects and spiders. especially in the Tyrol. while high up on the Alps is found the beautiful black and white snow-finch {Montifringilla nivalis). allied forms of which also occur on the mountains of Central Asia. its and that the great peculiar game birds.crevices. To the pheasant family belong the snow-partridge {Lerwa nivicola) of the Upper Himalayas. In regard to other t3rpes of birds. In shape and flight this bird has a remarkable resemblance to a large butterfly. a few words may suffice. as it hovers over the rocks.MOUNTAIN FAUNAS 89 prey since the introduction of sheep into that area. and it is also found on the other mountain ranges of Central Europe. on most of which the black grouse or capercaillie is found. and the very large snow-cock {Tetraogallus Jiimalayensis) of the Himalayas and Tibet. As is to be expected from the conditions. but these sufficiently indicate the two essential facts —that some of the game forests birds of the tundra and pine extend to the mountain ranges of the as it interior. the bird is stated now to kill living animals with blows of its powerful beak. occurs the citril finch {Chrysomitris citrinella). Other examples might be given. In the Alps the commonest member of the crow family is the Alpine chough {Graculus alpinus). elevated area in Central Asia has has its peculiar ungulates and its peculiar rodents. In the pine forest of mountains. using its long . game birds are frequent on mountains.

it is obvious that ice-free water for the regions of the mountain sides. there is some evidence that the mountain habitat tends to produce darkening of the colour. and unspotted. Similarly the Alpine salamander {Salamandra atra) is black in colour. which fit them for progression over rough ground. but then this hardy animal also extends into Siberia. Of the summer visitors to the Alps. As the animal ascends to an elevation of 9. This form shows a peculiar adaptation to life at considerable elevations. though not at such a great elevation. thus differing markedly in colour from its allies. a brightly coloured bird.90 sharp its bill MOUNTAIN FAUNAS to pick out its prey just as a butterfly uses proboscis to suck up nectar. extending up into the barren In the Alps and Pyrenees also the Alpine accentor {Accentor collaris) is common. most of the accentors are mountain-haunting birds. height of 10. and these pass through the gilled stage within the body of the mother. It is remarkable that the common viviparous lizard {Lacerta vivipara) ascends to a. having strong feet and legs. in that it only produces two young at a time. so that at birth they are lung-breathers like the parents. we may note the Alpine swift {Micro'pus melba). and it has a similarly wide range.000 feet in the Alps. The common viper {Pelias herus) also occurs in the Alps. As regards amphibians.000 feet. and not gill-bearing tadpoles. . Thus the common frog {Rana temporaria) occurs in the higher parts of the Alps in a dark-coloured variety. With the exception of the hedge-sparrow. Reptiles do not appear to show any special adaptations to mountain life. is In the Alps another frequent bird the rock-thrush {Monticola saxatilis). and the same thing is said to be true of the common toad {Bufo vulgaris). which is larger than the common form.

1900) . Kobelt's book has a good chapter on mountain animals. and Tschudi's Thierleben der Alpenwelt (ninth edition. Etherton's Across the Roof of the World (London.MOUNTAIN FAUNAS relatively long period of larval 91 development would not be easy to find hence the adaptation described. P. T. and Travel on the Tibetan Steppes. Among these may be mentioned Demidoff's After Wild Sheep in the Altai (London. 1899). An account of Alpine animals will also be found in Anderegg's Schweizerische Alpwirtschaft (Berne. reaching — a length of over five feet. during the short period of regions. Leipzig. Adventure. 1911). 1911) . There is a marked general resemblance between the insects of mountains and of polar regions. In regard to the lower forms of life we need only note that with the great development of flowering plants in most mountain development of insects. doubtless on account of the coldness of the water. 1872) should also be consulted. summer we have a great References. Fergusson (London. especially butterflies. There are a number of works on travel in Tibet and the neighbouring regions which give accounts of various of the characteristic animals. In mountain streams in China and Japan occurs Siebold's giant salamander {Cryptobranchus maximus). : . Sport. due doubtless partly to the similar physical conditions which prevail in the two regions. Relatively few species of fish occur in mountain streams. by W. N.

makes possible the occurrence of all many fruit-eating animals. Thus the anthropoid apes are fruit-eaters the fruit-bats have the same diet. of which temperature also The constantly high temperature.CHAPTER V THE FAUNA OF THE TROPICAL FOREST Just as the temperate trees is tropical forest as contrasted with the its remarkable for wealth of species of and for the luxuriance of its undergrowth. makes a warm coat unnecessary. The absence of seasons. fitted for this habitat alone. seasons. at day. in the sense in which they ditions We may occur in higher latitudes. and some orders. roughly speaking. like the Primates among mammals. as the name indicates For such parrots eat seeds and fruits. so its fauna is remarkable for the number of special tjrpes. begin by a few words upon the special conwhich reign here. The same condition makes reptilian life abundant. results in abundance insects. forms life in colder climates is impossible. for fruits occur at . again feed many other animals. To begin with. food is abundant all the year round. for the heat The high of the sun is available to hatch the eggs. are. for there is no seasonal check to vegetation such as occurs in temperate climates. and so on. As the tropical forest is characterized by a very . for relatively few of their members occur outside this region. . least during the so that animals producing valuable furs are relatively rare. Adaptations to life in the tropical forest occur in almost all groups of animals.

The true lemurs do not possess parachutes. and by great density. and having as their sole function the suspending of the body from the branches of the trees. ing lizard {Draco volans) . also of flying lemur. together with some other animals. as in the sloths of two conditions Somespecialized forms. which is so inserted that it can be opposed to each of the other digits. South America. differing in several respects . flying phalangers . but in general the monkeys and their allies. or at least of taking long Thus we have flying frogs (Rhacophorus) a fly. incapable of being separated from one another. This is the modification of one digit on both hands and feet. and enables them to leap from one flying squirrels. This is true of the South American whose feet are quite unfitted for terrestrial pro- gression. and it is notable that very many different kinds of arboreal animals have some form of parachute. This condition is suggested also in some of the thumbless monkeys.FAUNA OF THE TROPICAL FOREST 93 luxuriant undergrowth. we find not a few arboreal forms which never voluntarily put foot to the ground. producing . monkeys. show a character which gives as great security of grip. &c. partial powers of flight. tree to another. some and so of the forth. g. As regards the structure occur in the more highly times. giving them leaps. sloths. of forms like the Where the forest is less dense than usual. while conducing to greater freedom of movement and greater agility. and toes are converted into hooks. such purely arboreal forms must at times find it difficult to pass from one tree to another. the fingers of the feet. Galago) have the ankle greatly elongated. but some forms (e. which gives them a very froglike appearance. African from the forms found in temperate climates the so-called flying lemur.

a gripping hand and foot is produced by certain digits being birds. in the kinkajou. and so forth. Again. As great freedom of movement . in some of the South American ant-eaters. But this condition of opposable to the other is In the opossums and phalangers. in parrots among one (or more) digit being not confined to the Primates. in opossums. the lower forms having pointed fingers. the thumb being absent or minute. with nails instead of claws. Like the hookshaped hand it tends rather to occur in relatively sluggish forms than in those whose agility is sufficient to enable them to recover easily from a false movement. and the chameleon among lizards. This condition is only gradually acquired in the Primates. primitive mammals whose young are bom in a very undeveloped state. Thus Brehm notes that the * five-handed ' New World monkeys are not nearly such good climbers as the four-handed monkeys of the Old World. the great toe is also opposable. digits At the same time the extremities of the tend to be flattened. which never have prehensile tails. Some minor adaptations to the arboreal life may next be touched upon. Still another adaptation to arboreal life is seen in the prehensile tail. and with the flattening of the finger-tips a nail tends to replace a claw. opposable to the others. As already suggested also. the hand in certain Primates tends to become hook-like. giving more gripping power. Only in man among the Primates is the great toe not opposable. We find it in the chameleon. This also occurs in widely separated groups.94 THE FAUNA OF thumb and the condition described as the opposable great toe. in the South American tree-porcupines. in the New World monkeys.

. this is very noticeable. as in these are not down or to the body by will A comparison of most quadrupeds. in quadrupeds. such animals as dog or cat with monkey man make this point clear. in order to give a rigid support to the body but in most arboreal animals they are separate. as the arms are of very great importance in arboreal animals. It is the presence of the clavicle which permits of the upward movement of the arm. the two bones of the forearm and the corresponding bones of the foreleg tend to fuse. for only thus can the young be carried without loss of balance by the mother.g. To this account of the chief peculiarities of arboreal animals we may add a few words upon the tropical In the tropical forest considered as a source of food. and in a monkey. arboreal forms have often only one or two young at a birth. easily seen on comparing the position of the fore-limbs in. This gives such mammals a characteristically broad chest. and notably in the swift ungulates. which tend to be lost in quadrupeds. Finally. In the sloth. but preventing the animals from acquiring any great speed on the ground. a sheep. . With the loss of other arboreal characters man has also lost his long arms. e. we note that skin. so necessary in climbing. Again. in the monkeys and lemurs. we find that they are proportionately verylong. Another common feature of arboreal mammals is the presence of collar-bones. even when their fingers and toes make it possible for them to attempt to run. a movement impossible to pure quadrupeds like antelopes and deer. and the mammae are frequently pectoral in position. giving much greater freedom of movement to wrist and ankle.THE TROPICAL FOREST of the limbs tied is 95 necessary. Again. these being proportionately very much shorter than in the apes.

. The tropical forests of New Guinea yield almost no food for man. The fact that all the more intelligent of the forest animals soon learn to rob man's plantations. and it has doubtless also its effect on the wild animals. Elsewhere there are few human groups indeed who manage to subsist solely on the products of the tropical forest. They do not reach a high state of civilization. but we note among them that life the physical conditions most favourable to plant are not those best suited to the mammaUa at least. and those who do. If there is one Cembra pine loaded with ripe seeds in the taiga. the Congo pygmies and the Phihppine negritos. no seasonal check and therefore no marked variation in the amount of vegetable food available. and yet to sufifer from chronic starvation. Further. and have at times run almost the same risks to vegetation. the enormous number of plant species in the tropical forest diminishes the chance that a particular useful species will occur in numbers. of starvation as explorers in the tundra. suggests that the wild mammals are no better off than forest-dwelling man. e. But there is much evidence to show that the total amount is less than might be expected.96 forest there is. The reasons why the conditions should be unfavourable in little relatively the tropical forests are perhaps a obscure. so that recent expeditions have had to import every particle Consumed. g. THE FAUNA OF as already indicated. or to prey upon his flocks and herds. even when armed with modern weapons. seem to be few in number. there will almost certainly be many in the vicinity if the squirrels find one hickory tree in the American woods whose nuts are . trees of The fact that commercial value only occur in isolated specimens is a great barrier to the exploitation of the tropical forest by man.

animals are specially exposed to danger when feeding. specialization showitself chiefly in brain development. for large quantities of vegetable food must be taken. One point of structure is interesting.THE TROPICAL FOREST ripe. On the other hand. food can be temporarily stored in 1404 G . In most of the monkeys. Most herbiing vorous. Many of the ungulates have an arrangement whereby the hastily swallowed food can be stored within the alimentary canal. their generalized teeth. has to hunt for individual camphor trees in the Formosan forests. If we consider in systematic order the groups of animals showing adaptation to life in the tropical forest. in some instances their prehensile tails. there will also 97 be many others. or for individual rubber trees in the Brazilian forest. their strong clavicles. the number the reduction in the — of fingers of the limbs. careful mastication taking place later. show their descent from relatively primitive ancestors. On the other hand. on the other hand. This is perhaps part of the reason why the tropical forest seems to the explorer to be deprived of animal life save insects and reptiles. part of the reason why the forest mammals mostly occur in relatively small numbers. whose members just as man display practically described. and number of young at a birth all these fit them for life among the trees. their able thumbs and great toes. and in the fact that the hands are progressively suited for many uses. all those characters which we have oppos- Their long arms and short legs. their pectoral mammae. in addition to their prime function as aids in climbing. so must the fruit-eating wild mammals hunt through the forest for the scattered trees whose fruits supply their hunger. and toes present. the condition of the bones and other characters. we naturally begin with the Primates. or partially herbivorous.

with its very long arms and its covering of long reddish hair. eats some animal food in addition to fruit. As. The chimpanzee. which has a very limited distribution. the gibbons from South-East Asia. all Primates in New World monkeys and are denizens of the tropical forest. The way in which ' ' . cheek-pouches. not nocturnal. shar- ing a taste for and also some fruits with the negroes of the consuming some which the negroes do fact that it not care for. the orang in Sumatra and Borneo. Like man himself. of which there are four living kinds the gorilla and chimpanzee from West Africa. Apart from man. and is an extraordinarily powerful all the members of the order here — . at least at times. and of the Tibetan and Himalayan forms already mentioned.98 THE FAUNA OF later. but it is nevertheless common. Wild plantains and palm cabbage are freely eaten. Very different in appearance from the gorilla and chimpanzee is the orang. with the exception of baboons. it is impossible to name a few examples only can be picked out. to be chewed and swallowed This all arrangement the is far from universal — ^it is absent in some Old World forms. animal. The gorilla. is a dweller in dense forest regions. and Uke most Primates except the lemurs. like many mammals. the anthropoid apes are diurnal in habit. the highest living forms are the anthropoid apes. It is purely vegetarian and fruit-eating. even though the animals appear to be very few in number. The gorilla and chimpanzee resemble man in the region which they inhabit in having dark skins and hair the body is of course much more hairy than in man. The robs human plantations shows that food is sometimes scarce. region. which extends into Central Equatorial Africa apparently.

Whereever this type of forest occurs it shows similar density (see Fig. Figs. a fact which explains the existence of so many purely arboreal animals (cf. Lomas. The Margin of the Equatorial Rain-Forest iii ]S'orth Borneo.) . - »^^ <^£y ^ -* *.41. 23 and 32) which spend their whole lives among the trees. (Photo by H. 22). -I Fig. 21.M.

22.) . The Congo Tropical Forest (Belgian Government.Fig.

The animals are indeed excellent climbers. which are also leaf -eaters. is where the tropical forest best developed. . is also inhabited by the monkeys of the genus Cercopithecus. With the other egg-eating animals of the tropical forests they have probably been one factor in determining the migration of so many birds every year to the northern forest. and the diet is very G 2 . are long-tailed. in the upright its allies. already mentioned (p. though not rapidly. and walk. but differ in that they can plant the foot flat upon the ground. widely distributed over South-East Asia. They do not appear to come voluntarily to the ground. In West Africa the langurs are replaced by thumbless monkeys belonging to the genus Colobus. to which the monkey usually carried about by organ-grinders belongs. Though the arms are relatively short. the region. The gibbons are very much more active in their movements so active. but are somewhat deliberate in their movements. where they can rear their young in comparative security. 77). especially eggs and nestling birds. They mingle with their vegetable food a considerable amount of animal matter. The gibbons have even longer arms than the orang.THE TROPICAL FOREST the knee turns outwards makes the upright position than it 99 even less fitted for but the resultant inward position of the sole of the foot makes it even more skilful in cHmbing. making their way through the trees with great rapidity. as to be able to catch birds on the wing. it will be recollected. the animals are exceedingly agile. All have cheek-pouches. and are fitted for dense primaeval forests. West Africa. indeed. and are remarkable for their long silky hair. leaf -eating monkeys. position. Of other forms we may note that the langurs.

Very different from all the Old World monkeys are those which haunt the equatorial forests of Brazil. which have a mixed diet. and southwards to lat. they have long slender limbs. none has an opposable thumb. The leaf -eating howling monkeys (Mycetes. Though characteristic of the equatorial forests. The spider monkeys (Ateles) are comparable to the thumbless monkeys As their of Africa in the total absence of the thumb. In the absence of cheek-pouches they carry away part of their spoil in their hands or under their arms. the long and markedly prehensile tail playing a great part in the process. and are very skilful climbers. The baboons of Africa are much more widely dis- tributed through that continent than most genera of monkeys. name indicates. THE FAUNA OF In Asia the macaques are similar forms. the arms not being indispensable in climbing. being chiefly found on rocky ground. Examples are the various kinds of capuchins. at which they do not appear to be skilful. which is carried on largely by the tail. the tail is usually prehensile. but this is associated with the fact that they are not forest animals in the true sense. with very large cheek-pouches and a most indiscriminate appetite. and being practically quadrupeds. and share with some Old World forms the habit of plundering cultivated land. these New World monkeys extend northwards into Southern Mexico. animal food entering largely into the diet. 30^ S. but a considerable number of other genera occur. 23) may also be named.100 varied. Fig. some without prehensile tails. All these animals are small. and constitute the Platyrrhine or American forms. and the partition between the nostrils is broad. little Of much more limited distribution are the . The shortness of the arms is a drawback in climbing.

) prehensile tail. an American form with a {From a specimen in the Royal Scottish Museum. . The Red Howling Monkey.Fig. 23.

fSH .

Existing lemurs occur in the tropical forests of Africa. The lowest members of the Primates are the lemurs. Here also the animals reach a relatively large size. except the great toe. they do not grip the branches and swing freely from one to another. which contains many strange and primitive animals. In the absence of the special adaptations shown by their allies. but stick in their claws and climb along the branch on — The diet consists of fruits and insects. In Africa and Asia they are few in number. and are always small in size. as well as of structure. which occur only in the tropical forests of South and Central America. the animals climb like squirrels rather than like monkeys that is. In Madagascar there are more than thirty species. they show adaptations to varying conditions of life. or by the attacks of carnivores. in the forests of the south-eastern region of Asia. the largest being about two feet in length. the hind legs are larger and better developed than the front ones. apparently in individuals as well as in species. and show much fewer adaptations to the arboreal life than the higher Primates. have pointed claws instead of nails. and in the Old World have apparently been pushed far to the south by the development of better organized forms. The number of species and differentia- . The tail is nonprehensile. and the and finally in the island of individuals are abundant in every wood. as the number of species suggests. and the great toe is very small. They are entirely absent from the New World. Thus the fingers and toes. and. in spite of the fact that they spend their lives among the trees. all fours. which show some marked pecuharities of distribution. an area for all lemurs are very helpless animals. These animals are very squirrel-like in appearance.THE TROPICAL FOREST 101 marmosets. Madagascar.

We have not hitherto spoken of bats here. and where the carnivores of the cat family are absent. It is therefore the interesting to find that. but they are no less definitely adapted to arboreal hfe. the common fruit or fox-bat of India measuring four feet from tip to tip of the wing. and is even better developed in the httle tarsier (Tarsius) of the Malay region. or from one end of a branch to the other. though not to New Zealand. because those forms which occur in temperate regions show no special adaptation to any particular type of habitat. Their mastery of the are virtually absent. but in the east extend southwards to the island continent of Austraha and to Tasmania. so in which have abandoned the arboreal gascar Mada- we find the ring-tailed lemurs.102 tion of structure THE FAUNA OF seem to be associated with the fact that the lemurs here are protected by isolation. The same peculiarity occurs in the mouse-lemurs (Chirogale) of Madagascar. The fruit-bats are larger than the insect-eating forms. They are entirely absent from the New World. which Hve among We . and much less highh' differentiated. which progresses by leaping from one branch to another. gives them the power of leaping Hke a frog. It is otherwise with the large fruit-bats. members of the less differentiated civet alhance taking their place. where they are chiefly found in forests. Lemurs are less intelligent than monkeys. just as more baboons are monkeys hfe. foxy-looking rocks in regions where trees have already spoken of the fact that the galagos of West Africa have an elongated ankle which. Hving as they do on an island where there are no true monkeys nor apes. animals. which are practically limited to the tropical regions of the Old World. though the mechanism is a httle different.

the female being monkey-lilie. Though the general shape is Fig.male only The brown with a white rufif. (Figs. 25. {From a specimen in the with those of Royal Scottish . is black. Note the opposable thumb and great toe. Male and female of the Black Lemur. 23 and 24). the foxy faces should bo contrasted monkeys Museum. Madagascar.


In the forests of tropical America there occurs an interesting family of bats. Further. some of them eat Others eat fruit and insects. but the way in which they raid the orchards of the fruit-growers shows that fruit is greatly preferred when it can be obtained. In Australia. Similarly the index finger is not reduced to the rudimentary condition seen in other bats. where wild fruits are somewhat rare. Fruit-bats spear their food with the claw of the thumb. We have already pointed out that insectivores are intelligence of the peculiarly helpless animals. but. they have taken advantage of man's fondness for fruit to extend their range to regions where his plantations can be robbed. as the fruit-bats feed in trees they are more arboreal in habitat. and by the strongly hooked thumb of the hand. attaching themselves by the hind feet. From the diet the animals must necessarily have been once confined to regions where wild fruits were abundant. but nevertheless fruit only. without the which enables many Primates to escape from . The vampire bats belong to the insect-eating section (Micro- chiroptera) of bats. some of whose members present a curious analogy to the fruit-eating bats of the Old World. like some members of the order Primates. or vampire bats.THE TROPICAL FOREST air is less 103 complete than in the insect-eating forms. and have teeth so modified as to allow them to crush pulpy fruits. and is here usually furnished with a claw. while others again are purely blood-suckers. the arrangement of the tail apparently preventing them from giving those sharp turns so necessary in an animal which feeds on insects caught on the wing. where it is useless. being able to scramble about the trees. the Phyllostomatidae. the fruit-bats seem to devour the flowers of the eucalyptus.

though smaller than most. as Galeopithecus is to flying squirrels. tail. The membrane how- ever. with sharp claws to allow the animals to cling to the rough bark. Borneo. and resembles the flying squirrels of temperate able that various forms should life here. together with the sHghtly enable the animal to swarm up trees. but extending also into the mainland of India and Burma. the socalled flying lemur (Galeopithecus). and adjacent regions. strongly clawed. of which two species occur. The animal altogether life. The fingers and toes are short and and they. and the other in the PhiUppine Islands. example of adaptation to forest In the same region as the flying lemur. and the animals use their forefeet like squirrels in feeding. The feet are naked beneath. sitting up are on their hind-legs and holding the insects or fruits in . forests in having a parachute of skin extending from the fore to the hind limbs. upon leaves. better developed. for it cannot fly in the sense of ascending against It feeds and has strongly cusped offers a curious front teeth. The tree-shrews very like squirrels in appearance. presumably for the purpose of nibbling the leaves. we find the little tree-shrews (Tupaia). one in the Malay Peninsula. tropical forests offer great possibilities in the the consequences of their The way of shelter and protection. the tail is long and bushy. in that it involves the tail as well as the limbs.104 THE FAUNA OF own incapacity for defence. which enables flying leaps it to take is. from tree to tree. The flying lemur is about the size of a cat. Sumatra. prehensile gravity. and it is therefore not remark- tions to show special adaptaThe most striking of these adaptations are those shown by a very aberrant animal. which so far as adapta- tions go may be said to be comparable to ordinary squirrels.

It shows. They haunt the the other large cats. The clouded leopard or tiger (Fig. It is apparently a bad climber. however. On the other hand.THE TROPICAL FOREST the forepaws. but this may be partly because it is very impatient of great heat. with the exception of Madagascar and Australia. less successful Their limited distribution shows that they have been much more much have many to acquire than squirrels. This is even more true of the South American jaguar. being much more a denizen of open plains or districts with thick reeds and grass. the widely distributed leopard climbs well. being capable of running up a straight-stemmed tree. and the tropical forests of aU regions have their full share of species. On the other hand. The puma (Fig. is much more frequently found in wooded regions than elsewhere. 45) is an arboreal form found in SouthEast Asia. the tree-shrews are very different from the squirrels. The habits of the civets are . which in swampy regions may be almost purely arboreal in habitat. like tion. the less differentiated forms known as civets (family Viverridae) are limited to the warmer parts of the Old World. and preys largely upon monkeys. are universally distributed. though occasionally found in forest regions. The tiger. considerable adaptability^ being found in savanas as well as in forests. though their it less necessary for them purely arboreal habits than for more helpless animals. so that the resem- blances are due to adaptation. of course. forest for food rather than for protecOf the large forms the lion is not a forest animal. The powerful and dominance makes intelligent carnivores representatives in the tropical forests. in India especially. The smaller cats. 105 Structurally. 26) does not appear to haunt dense forests.

forests of Of the raccoons. though it is a clumsy climber. represented in the tropical jungles. but among the arboreal forms we may note the palm-civets or tree-cats (Paradoxurus) of India and West Africa. however. though perhaps the majority of the tropical Asiatic deer tend to haunt relatively open country. In Africa. the coatis (Nasua) are found in the South and Central America. As a general rule ungulates are not abundant in the tropical forests. where deer are absent. despite their total absence from Africa south of the Sahara. but the animals do not predominate here as they do in the north. with a prehensile tail. are well represented in the tropics. which have naked feet. Though not a few of the dog alliance extend into wooded country. It feeds fruits. their bulky bodies not fitting them for life amid dense and tangled undergrowth. birds and their The widely distributed weasel family has representatives in the tropical forests. forests of South-East Asia contain the Thus the small black Malayan bear {Ursus malayanus). a common feature in arboreal animals. and the bear-cat (Arctictis) of South-East Asia. and the tropical forms show no special features of interest. On the other hand. naked feet. The deer of temperate woods are. none can be said to have any special adaptation to forest life. a cat-like animal. and upon small mammals. which is essentially arboreal and frugivorous in habit. and strong claws. some of the .106 THE FAUNA OF varied. and their general preference for temperate and upland" regions. In the forests of India the sloth-bear {Melursus ursinus) is common. and the kinkajou {Cercoleptes caudivolvulus) of the tropical forests of America. and lives in forest regions. has as special arboreal characters a prehensile tail. the bears and their allies. eggs.

where it inhabits swampy regions. and are inhabitants of the swampy parts of forests. In tropical Asia the sambar and its allies are the chief deer of forests. wooded country. and their slender legs and bodies enable them to glide through the jungle. The wild pigs of India and the adjacent regions. We find. and the African sable antelopes (Hippotragus) are found in tropical forest also In the African found the rare and little-known okapi. rivers or swamps. occurs on the west coast of Africa. therefore. ing those of a zebra (Fig. In the Asia occur various members dense jungles of South-East of the genus Tragulus. their In a densely forested region. called chevrotains. that such parts of the forest have special animals. 107 This is especially true of the little duikerboks (Cephalophus). almost rodentlike animals. but with stripes recallis wooded country. are examples of forms which haunt the damper parts of forests. whose slender bodies enable them to glide through the undergrowth. and thus to persist despite their generalized structure. An allied form. true pigs. of primitive structure. The African harnessed antelopes (Trage- laphus) also. a relative of the giraffe. with special types of vegetation. the bush pigs of Africa. Africa has also its hideous wart-hogs (Phacochoerus) while the islands of Celebes and Buru lodge in their forest the curious babirusa with its gated tusks. 46). but the muntjacs (Cervulus) also haunt regions. obviously afford advantages to animals not fitted for arboreal life proper.THE TROPICAL FOREST antelopes have become modified for forest life. choosing upland They are the smallest of living ungulates. including small. enormously elonbut the slendersimilarly All these . being only about a foot in height. America has no limbed peccaries replace them. the water chevrotain.

the squirrelsrand flying squirrels. like rhinoceroses of South-East Asia and the common rhinoceros of Africa. in the way in which these toes spread out to prevent the animal sinking in the mud. are found in dense jungles. in regard to habitat. which is regurgitated and pig-like animals differ thoroughly masticated later. As in the case of the pigs it is probably partially determined also by the need for finding relatively soft and succulent food. The great enemy of the American forms is the jaguar. for while The rhinoceroses show somewhat different conditions some haunt more or less the Javan and Sumatran open plains. The elephants. forest animals. but in both cases this seems to be largely on account of the animals' great intolerance of heat. both the Malayan form and those occurring in South America. both Indian and African. on the other hand. when the animals feel themselves relatively safe from pursuit. Among the rodents. are found in forested country. haunting much the same regions as do the pigs. for they emerge into the more open regions both at night and during the cooler parts of the day. others. and in the absence of the elaborate arrangement which enables a sheep or an antelope to swallow large quantities of partially masticated vegetable food.108 THE FAUNA OF from those ungulates which live on hard ground in the greater number of functional toes. . and the fact that when attacked the animals endeavour to reach thick cover suggests that the forest habitat is partially determined by the need for protection. for the teeth of pigs of structure and tapirs have not the elaboration found in the more highly differentiated ungulates. are forest animals. and Horses are not subsist largely on leaves and twigs. Among the odd-toed ungulates the tapirs.

00 ^^ ID <U -f^^J dj o C a-^ Pm .

{From a specimen in the Royal Scottish Museum. The African Elephant.) . 27.Fig.

which has overlapping horny scales on its under surface near the base. Elephant in teak forest. are also found maximum number in the Malayan region. genus Pteromys. India and the adjacent regions have large flying squirrels belonging to the Fio.THE TROPICAL FOREST in tropical ones. which are very different Note especially that the long. 28. straight not bound down by skin as in the horse.) thigh is from those of cow or horse. and the true knee appears to be very low down. and thus . the squirrels reaching their 109 already described in temperate forests. The position brings out the peculiarities of the hind-limbs. stated to be used in obtaining a hold on the bark when the animals are climbing trees. Upper Burma. but Africa has forms which belong to quite a different family (Anomaluridae). Couchman. {Photo by Gol. The Latin name is given on account of the peculiar tail.

belonging to the genus Bradypus and Choloepus. 47). no adaptation among which they pass their The fact thai^they possess practically . and that their place in nature is taken there by rodents there is. We have already repeatedly emphasized the fact that ungulates in America are few. The great toe is absent. resemthey show a blance to the chevrotains of South-East Asia. therefore. This is represented in the tropical forests of porcupines of is Mexico and Brazil by the treethe genus Syntheres. or by acquiring burrowing and concealed habits. and the hind-foot is so inserted as to make it easy for the animals to grasp the branches. nothing surprising in finding that the rodents called agutis (Dasyprocta) haunt the forests of Brazil and and have become adapted for life in the dense jungle. perhaps the most truly arboreal of all mammals. With their slender limbs and bodies close. and probably do not descend voluntarily from the trees (see Fig. of the tropical forests. The animals are stated not to drink. which present some interesting adaptations to arboreal life. though entirely superficial. the trees for they are quite helpless if removed from whole lives. With the Edentates we come to a primitive order. but its place is taken functionally by a fleshy lobe which can be bent inwards so as to be partially opposed to the toes. beyond them either by retiring to the depths elsewhere. 30). have already spoken of the Canadian porcupine (p. The paca (Coelogenys) has similar habits. In the forests of the Amazon occur the sloths. . pecker We (p. The tail markedly prehensile. whose members have kept their hold in a world which has grown. 43) in the offering THE FAUNA OF some analogy to the stiff tail of the woodwhich has the same function.

Though not nearly related the animals show a superficial resemblance to one another. 29. |The characteristic on the under-surface of the tail cannot be seen in this specimen. With this figure should be compared that of the Flying Phalanger (Fig. is well shown. which helps to support the parachute. 33).Fig. scales An African Flying Squirrel {Anomalurus). . but the bony structure projecting from the elbow. {From a specimen in the Royal Scottish Museum).

to J^ .300 o 5 o £ SJ .S 12.

THE TROPICAL FOREST to permit 111 them to pass from one is tree to another shows that their natural habitat tree is the dense forests where bound to tree by lianes. or do not require to use the tail as a balancing organ after the fashion of squirrels tree-shrews (cf. &c. the individual hairs appearance of a lichen-covered branch. but terrestrial. Not a few of the marsupials have acquired arboreal Thus the tropical forests of New Guinea and Queensland are inhabited by tree kangaroos. ing back The usual position hang- downwards from the branches of trees. We and have already spoken of the elongation of the forelimbs in the sloths (p. most are found in rocky country. are converted into The fingers mere hooks. while the little though some of these are partially arboreal. Among the special features we may note the nature having a fluted surface. which have . The South American ant-eaters are also forest animals. habits. both in the structure of its limbs and in appearance. where they burrow in the ground. has a curious adaptive resemblance to a sloth. the tail is a stump. anthropoid apes. The hairy anteaters of South America are replaced in India and Africa by the scaly ant-eaters or pangoHns (Manis). the large form called Myrmecophaga jubata being strictly two-toed ant-eater (Cycloturus didactylus) is arboreal. and so give the animals the of the coat. The food consists of leaves and fruits and is the animals do not drink. 95). and except that it has a prehensile tail and lives upon insects. 32). bears. many helpless forms they are active only at night (see Fig. a common feature in such arboreal animals as have not prehensile tails. and the animals sleep rolled up in a ball with the head tucked Like between the arms.). on which algae lodge.

In North Australia and in the islands of the Indo-Malayan region is found the spotted cuscus {Phalanger maculatus). and a slightly developed .112 THE FAUNA OF apparently recently taken to the trees. and presumably. life closely related animals. In the forests of Western Australia occurs the little Tarsipes. In other kangaroos the forelimbs are very short. in these forms they are nearly as long as the hind ones.inches. and the tail. great flying phalanger {Petauroides volans) of Queensland and Victoria has both a flying membrane and a prehensile tail the squirrel flying phalanger or sugarsquirrel {Petaurus sciureus) has a bushy squirrel-like tail. The feet are naked beneath. not prehensile. There are several species. though not prehensile. phalanger {Acrobates pygmaea). The pygmy flying It occurs in Eastern Austraha. mostly about the Just as a cat. but like of the forest animals will also take birds and size of other small creatures. Much more trees as definitely arboreal are the phalangers. tail is very markedly pre- so many Cuscuses feed upon leaves and fruit. and a well-developed parachute. . a shrew-like animal with a prehensile tail. has a long tail with the hairs arranged at the sides to imitate the vane of a featner. so among the The 33) we have also flying types. which sucks honey from flowers with its worm-like tongue. As in the other members of the phalanger family the great toe can be opposed to the others. among insectivores. the rodents we have flying squirrels among and flying shrews phalangers (Fig. is said to be used in climbing. and the long hensile. in which the total length of head and body does not exceed about 2^. obtains insects also in this way. but as well fitted for in the most kangaroos are for life on the ground. like humming birds. for they are stated to be slow and clumsy climbers.

fe .

Fig. 33. Australia. The Lesser ilyiug Piialaiiger the (From a specimen in of Eastern Royal Scottish Museum.) .

They must be added to the very considerable list of peculiar arboreal mammals possessed by South America. and doubtless aided by its feather-like tail. a fact well illustrated by the gorgeous birds of paradise. the great toe being opposable to the others. Curiously enough. Birds are so abundant in the tropical forests that we can only pick out for special remark a few which illus- In spite of the gloom of the jungle brilliantly coloured forms are common. can be opposed to the other three. which are absent from the South American continent (see Fig. but is also stated to descend to the ground in order to dig for roots. it possesses cheek pouches like an Old World monkey. as often happens in such cases. the trate points of interest. are represented in the United States by the widely distributed common opossum {Didelphys marsupialis). as in other phalangers. the koala or native bear {Phascolarctos cinereus) from its allies in adaptations. for they do not occur outside of the American continent. On the foot. Another member of the phalanger family. which. and they take the place in nature elsewhere occupied by insectivores. 31). and shows some curious It is sluggish in its movements. The majority spend their lives in trees. The last of the marsupials to be considered are the opossums of South America.THE TROPICAL FOREST parachute. 1404 H . It is stated to 113 agility in show marvellous is leaping from branch to branch. bear. there are five toes. and. though chiefly South American. sloth. The animal feeds chiefly upon leaves.). though some spread to the pampas of Argentine. the tail has disappeared In the hand two of the fingers (cf. is very different appearance. &c. small insect-eating animals with opposable great toes and prehensile tails. the condition which occurs in the chameleon.

that is one in which the first and fourth toes are directed backwards. of which the upper hinged as well as the lower. The climbing foot however best developed in the parrots. as of the fruit- mammals. and the toucans with enormously developed bills and bright colouring. The little love-birds are African. which are for the most part thoroughly arboreal. but they extend much further to the south of the tropic of Capricorn than to the north of that of Cancer. to whose branches they cling easily with their strong feet. is used in cHmbing as well as in cracking hard shells and kernels. Parrots are chiefly tropical. Of almost eating all parrots it may be said. Of the parrots. and almost all these are arboreal birds. fruitshort.114 THE FAUNA OF forests of most brightly coloured of animals. handsome birds found in the South American forests. which haunt the New Guinea and the neighbouring regions. green predominating. the crested cockatoos are Australian. As is well known. They include the widely distributed woodpeckers. and the gorgeously coloured macaws with their long tails South American. is hooked bill. The parakeets are found in Africa. while the second and third point forwards. though the birds seem to dwell entirely in the trees. as it often does among arboreal. their colouring is often very striking. and the portion is eating birds. that they show a marked preference . They spend much of their time in the trees. as is also the familiar grey parrot with a red tail. India. The purely frugivorous plantaineaters of Africa differ in that the fourth toe can be turned backwards or forwards at will. found in tropical no less than in temperate forests. strong. The legs are short. the jacamars. and Australia. Among Picarian birds a whole division possesses the climbing foot.

Quite similar conditions occur among the game birds. after the fashion of a fruit-bat. preferring to trust to the thick cover unless danger approaches too closely. Among reptiles the crocodiles and their alHes haunt the lakes and rivers of forest regions. the spur-fowl of India jungle-fowl.THE TROPICAL FOREST for robbing 115 man when they can. through which their wedge-shaped bodies enable them to travel easily. and so Very remarkable is the condition presented by the hoatzin (Opisthocomus) of South America. and extending southwards to Australia. We may mention especially the fruit-pigeons. The hoatzin is believed by some but is ornithologists to be related to the game-birds. Such birds fly with considerable reluctance. A very curious arrangement of the breathing organs enables them to drown their prey by holding the H 2 . Not a few of the pigeons are arboreal. which they use in climbing about trees. through which their slender bodies permitted them to force their way. where the young are hatched with claws on both the thumb and index finger. Among the mammals we found that in addition to the truly arboreal forms there were a number like the chevrotains and agufis. Among such forms we may mention the wood-partridges of the Malay forth. where various members of the pheasant family live in dense thickets. the guinea-fowl of West and Ceylon. the Africa. where they lie in wait for the forest animals as they come down to drink. The bill is distensible at the base to permit the birds to swallow large fruits whole. which took advantage of the shelter afforded by the dense forest. rather than hmiting themselves to wild products. found in SouthEast Asia. region. a very primitive form.

but habit of dropping into the water with a splash. and the hands and feet conThe limbs are long and slender. fourth. and the wings when not in use are folded close to ' ' the body. to all. and extending into Europe in Spain. specially characteristic of Africa. and feed upon birds. which are outside.116 THE FAUNA OF animals caught under water till death takes place. with prehensile chiefly tails. The animals are insectivorous. upon The mechanism is thus entirely different from that seen in any fljdng mammal. and their parachute is a lateral fold of skin supported the posterior ribs. the third. and their power of colour-change is famiHar All have prehensile tails. They haunt especially trees overhanging water. They occur in the Indo-Malayan countries. and fifth forming the outer group. The body is greatly depressed. if alarmed. America and the West Indies show no special adaptations to this mode of life. Like their allies the pythons and . the chameleons. and are said to have the curious of tropical The iguanas are also arboreal forms. stitute perfect grasping organs. In the foot the inner bundle is made up by the first two toes only. two being opposed to the other three. On the fore-limb the first three fingers form an inner bundle opposed to the fourth and fifth. however. The fingers and toes form two groups. the forelegs especially so. which are greatly elongated. The most purely in arboreal of the lizards are. Of the lizards the flying forms (Draco) are interesting. some- times from a great height. Of the snakes the tree boas of South America are forest-haunting forms. but also found India and Ceylon. the crocodile being meantime protected from choking by the nature of the internal nostrils.

including the most poisonous forms. they are non-poisonous. whose members occur alike in Madagascar and in tropical America. shooting out their heads in order to capture their prey. flying frogs of the A number of other frogs arboreal habits. and also throughout South-East Asia. the show more or less well-marked commonest adaptation being the of adhesive disks at the development of some form end of the toes. which occur in Madagascar. like the trees among which Of the amphibians we have already mentioned the genus Rhacophorus. The huge anaconda is {Euneces murinus) of the Amazonian forests aquatic. tint. with a markedly prehensile and a generally green it lives. whose secretion is used in South America as an arrow poison. An exception is the climbing tree-viper {Trimeresurus gramineus) of tail. and some species at least have poison glands in the skin. crushing their prey their coils. The true boas occur both in Madagascar and in tropical South America. The beautifully coloured and marked wood-snakes (Herpetodryas)of tropical America represent there the tree snakes (Dendrophis) of the Old its World. to permit of the animals fixing themselves to the branches of trees. In India the very agile whip-snakes (Dryophis) twine their slender bodies round branches of trees. This is seen in the family of Dendrobatinae. Most other snakes. India and Burma. Such an associa- . and inhabit the dense forests. all of which are arboreal in habit. lurking in pools till semi- prey approaches the water. through which they can glide without difficulty. often inhabiting dry and sandy places. It feeds largely on peccaries. They are beautifully and often vividly coloured. The American forms feed upon lizards and young birds.THE TROPICAL FOREST by the weight of 117 true boas. are ground animals.

In regard to fresh. None of these animals voluntarily quit the water. The typical tree frogs of the family Hylidae are widely distributed over the globe. Other amphibians are not adapted for the arboreal Hfe. peculiar. The can anterior fins are curiously modified. found in the streams of the tropical on its double-breathing seaward margin. Even more curious are the habits of the mudskippers (Periophthalmus). in the tropical streams of Africa by Protopterus. are represented in the tropical streams of Queensland by Ceratodus. They have suctorial disks at the ends of their are usually protectively coloured in shades of green. which haunt the mangrove swamps and mud-flats on the shores of the Indian and Pacific These are Oceans and off the coast of West Africa. and in those of South America by Lepidosiren.water forest. or fish we may note one or two peculiar forms. The very peculiar fishes. the tints varying with the surroundings. and thus obviate the necessity for seeking ponds to Their breeding habits are often moisten the skin. bony fish which are partially adapted for Hfe on land. so that the fish cUmb about the supporting roots of the mangroves. which are animals furnished both with lungs and gills. but the fact that the two last named can breathe when buried in the mud as well as when swimming in water adapts them for life in tropical regions. where there is often a well-marked alternation of wet and dry seasons. but are absent from Africa south of the Sahara. . toes.118 tion of THE FAUNA OF bright is colouring with poisonous or noxious quahties of common occurrence in animals. from Arabia. and from India. and have glands upon the under surface whereby they can absorb water from the damp leaves on which they rest.

protrusible. serve as transmittors of the tropical diseases most fatal to man and beast. either solely or in of food. which are often blood-suckers. some of which species. are furnished with distinct eyelids. and thus permits a great wealth of individuals to occur. It has a special accessory breathing organ. many reptiles some carnivores. which are wandering forms. The eyes are very prominent. the equal temperature allows the insects to go on breeding the year round. as in the case of mosquitoes and tsetse flies. tropical butterflies. Further. while others from their of all numbers and ferocity are the enemies other animals. and second the ants. which are very tadpole- appearance. The driver ants of West Africa. and voluntarily leaves the water to do so.THE TROPICAL FOREST and it 119 can also leap and skip over the mud. feed upon insects. a freshwater form which travels long distances over land by hitching its pectoral fins round plants. eat combination with other types all insects. of the marsupials. another fish is found which can live on land. The fish is also stated to be able to climb trees. which enables it to breathe air when out of the water. the edenand amphibians and so on. a special feature of the tropical forests is the great wealth of insects. and are like in The fish. is the climbing perch {Anabas scandens). In India and the south-east of Asia generally. the insectivores. In regard to the invertebrates. while the special conditions favour a great wealth of genera and Apart from the size and beauty of the most impressive of the insects are first the flies. are stated to devour every living creature in their path which cannot make its . many tates. and then may. That this must be so will be clear when we reflect that This some of the forest-dwelling primates. the are extraordinarily destructive to plant life.

give good accounts of South and Central American animals... serve for the transmission of deadly diseases. 1908) should also be consulted. like flies. Wallace's Malay Archipelago (London.120 FAUNA OF THE TROPICAL FOREST and will escape. and ticks. or so-called white ants. even attack human beings if from any cause these are rendered incapable of defence or flight. 1897) Lydeker's Game Liberia (2 vols. We may add that. just as a few fish in tropical climates have taken advantage of the greater opportunities which the land offers as compared with the life upon the of other sea to acquire partially terrestrial habits. London. . Bates's Naturalist on the Amazons (5th ed. which has a marked organisms in forest areas. London. predatory forms which often show a marked resemblance to the vegetation among which they live. Johnson's works give very full details. British Central Africa (London. special As most of these forms do not. including the large bird-catching form of South America.. 1884). however. For details as to the animals. to which they return for breeding purposes. In regard to the animals of the African forests. 1906). members of the same order form the beautiful stick and leaf insects. reference should again be made to the natural histories already mentioned. The termites. 1888). show it adaptations to forest life. . is sufficient to merely effect indicate their abundance. so various crabs have become partially adapted for life on land. obvious in the dense forest than in more open Other but some species do occur there. with many figures see The Uganda Protectorate (2 vols. London. spiders. 1869) gives a fascinating account of the animals of that region. . London. and Belt's Naturalist in Nicaragua (2nd ed. . H. and the same author's Tropical Nature (London. 1902) Anitnals of Africa (London. which. and wander about in those forest regions which are at no very great distance from the sea. As well as with insects the tropical forest swarms with scorpions.. 1878) may also be mentioned. References. are as a rule less country. Sir H.

through the intervention of the savana wood. like the similar conditions in the steppes. a characteristic park-like appearance to the landscape. and the savana passes into the tropical rain-forest. or inland from the Australian bush to the central desert. and there is a well-marked dry season. The rain comes in the hotter season of the year. migration This. South America. have a highly The typical savana climate is tropical. where the rainfall is abundant the trees increase in number. and developed and characteristic fauna. separated by regions without trees. Where the rain is deficient the savana passes into desert. On the other hand. The period of greatest vegetative activity accompanies or follows the rainy season. and Australia. Further. forming the so-called gallery forests. giving or shrubby vegetation.CHAPTER VI TROPICAL SAVANAS AND DESERTS In the tropics the steppes of temperate latitudes are represented by what are called savanas. which are regions where grass occurs. Such savanas occupy large stretches in the tropical regions of Africa. as we see in passing northwards from the Sudan to the Sahara. and the dry season may witness an almost complete cessation of activity. . involves a certain amount of among the inhabitants of the region. with its numerous trees. Elsewhere the trees follow the courses of the streams. . mingled with arborescent Sometimes we have in these regions scattered trees only sometimes clumps of trees occur.

Lloyd. Various kinds of grasses occur. It stores water in its thick trunk. with scattered acacias. whose foliage is reHshed by the giraffe. i . {Photo by Capt. Other important trees are thorny acacias.122 TROPICAL SAVANAS fires frequent. while thorny and xerophilous plants are of course frequent.''^#'lk'~^s>««' ."''~^'^s»ji. forming dense thickets. are The type of vegetation naturally varies in the which savanas occur. on the margin of the desert in Kordofan. Savana. and so forth. different regions in j#.) and becomes leafless in the dry season. In the savanas of the other . which offer much shelter. In Africa the mighty baobab [Adansonia digitata) is conspicuous among the trees. bush and often have devastating effects. palms. 34. owing to the great power of the sun. Fig. various species of cactus-like Euphorbia. and often grow to a great height.wm^mf^ Mim::^S^:^i^^m>^^^^a^:p^^mA ' v"'-^if i| i'-'T~.

AND DESERTS is 123 continents. The Kalahari. conditions Vegetation also gradually diminishes as the become less favourable. Of the great savana regions is of the world the African especially rich in ungulates. ' ' . 32° S. were relatively poor in mammalian types in recent mammalia till savana the advent of European man. pass into a savana region further north. the general facies the same-^tall grasses. cactus-hke and generally plants presenting which protect them against periodical while allowing drought. this being especially true in South America. In the desert proper few organisms can live. typical savana regions insects as locusts. while the savanas of South America. the mammahan fauna diminishing first. It should be noted. however. One proper of the special climatic features of the is the periodic abundance of rain. and more uncertain. and the rain becomes less. indeed. but the less unfavourable regions carry a reduced savana fauna. No hard and fast line separates savana regions from steppes. and the fauna becomes impoverished. especially of insects and reptiles. a country which has witnessed an extra- ordinary destruction of geological time. the Australian in herbivorous marsupials. cactuses or forms. notably in antelopes. As we pass from the savana to the desert the dry season increases in length. thorny acacias. where the treeless pampas or steppes of the Argentine south of lat. that in addition to the characters already given. is said to serve as a reservoir for such which breed there until starvation forces the swarms to sweep outwards to the better watered lands on the margin of the desert. though the species differ. notably kangaroos and wallabies. them to take advantage of the special features periodic tropical downfalls.

shoots. To this rule. however. where it feeds upon fruits. and seeds. which they use in routing about for the insects on which they feed. buds. and move in a series of leaps. mostly fitted by nature for hfe in the tropical forest. insectivores we may note especially jumping shrews. as we have already had occasion to remark. the large cat-hke forms. 64). for. as we have already seen. the baboons form an exception. however. The Arabian baboon {Papio hamadryas) occurs abundantly in the Sudan. suggests that it was once a desert animal. The type genus is Macroscelides. and finds the necessary shelter in rocky ground. being mainly but not exclusively tropical. and feeds largely upon that the strange coniferous plant Among the known as Welwitschia. In Africa it is abundant in the Kalahari desert. and has several obvious advantages which have been already discussed (see p. they are quadrupeds. They have also curiously elongated snouts. anuhis) inhabits very dry country. is The elongation of the hind-limbs interesting. as we have seen. we may note that as a general rule the primates do not occur in such regions. The uniform colouring of the lion. and show a curiously close resemblance to the jerboas among rodents. for it is frequent in steppe and savana animals. In Angola another species (P. hving in open rocky country.124 TROPICAL SAVANAS are distinguished from the typical steppe region already described in the absence of a severe winter. show no very special adaptation to one habitat. The jumping or elephant shrews have greatly elongated hind-legs. which are confined to Africa. Proceeding now to consider the animals of the various savana regions in systematic order. being. Of the carnivores. and the animals are widely distributed in Africa. but also extends into savana and steppe .

and of following its natural prey. An interesting desert species is the . where the savanas pass into steppes. which store water in their underground parts during the time of drought. a burrowing animal. when the nobler animal has As a rule the members of the dog alliance inhabit open country. or the smaller ungulates. upon which it is partly dependent. Like most of the smaller mammalia of open country this animal is a burrower. which seems to feed largely upon termites. the spotted hyaena of Africa being often found in the same districts Somaliland.AND DESERTS regions. We may note especially the African jackals. and inhabits open country throughout the southern half of Africa. the mongooses are ground animals. making deep holes in the sandy veldt with its sharp foreclaws. usually haunting open country. Of the smaller cat-like forms. the meerkat (Suricata tetradactyla) haunts sandy ground in Cape Colony. kill '. with weak teeth. in that ' it obtains the remainder of the satisfied its hunger. man. A considerable number occur in the tropics. which hunt in packs and feed upon rodents. where as the lion. with a northern limit apparently in The hyaenas it has been once found. Another adaptation to life in this region is shown by the fact that it feeds chiefly upon the bulbs and roots of the veldt plants. Another steppe or savana form of wider distribution is the aard-wolf {Proteles cristatus). its distribution being probably 125 determined by its per- the double need of avoiding the proximity of sistent enemy. or sick or wounded members of the larger species. and avoiding dense forests. as the common habit of hunting in packs suggests. An allied form. the larger ungulates. are also inhabitants of savana country. and may therefore be included in the fauna of deserts and savanas.

. small birds. and is burrowing and nocturnal in habits. It is widely spread throughout Africa. and sets forth in search of insects. are absent from Africa south of the Atlas. is tawny-coloured like the desert sand. At dusk it becomes active. Though the animals hve in holes. The animal is in company. It would appear that the dogs have little or no burrowing power themselves. This little animal has very large ears and great acuteness of hearing it desert. where it inhabits open country. which has a curious and unexplained resemblance to the spotted hyaena. or fennec. So swift are these dogs that they are said to be able to overtake the swiftest antelope. it is clear that the animals are unfitted for life in such regions. Not a few other dog-like animals haunt the African savanas. even fruit if obtainable. burrows being made A very much fiercer animal is the Cape hunting dog [Lycaon pictus). the young being born underground. The allies of the bears.126 TROPICAL SAVANAS long-eared fox. which inhabits the Sahara and is called Canis zerda. as in we have already seen. the these may serve as types. Bears. As these two countries have the best-developed savanas. but lizards. and that in the Andes. their holes being either natural or obtained by ejecting the original occupant. They are indeed absent aUke from temperate steppe and tropical savana and desert. being said to appear to sink through the sand. which are borne down by sheer weight of numbers. In South Africa the flocks of the white man form a favourite source of food. It burrows with great rapidity. such as the raccoons and coatis. and one species only occurs South America. rodents. are also . or partially social. preying upon the ungulates. yet on an alarm it seeks safety in flight rather than in the burrow like the fennec.

and the weasel group is not well Among the few weasel-like forms we may note the South African weasel (Poecilogale) and The latter has a somewhat wide distribution in Africa. and is in mountain ungulates (cf. and never more than one calf at a birth. The animals feed upon small mammals and birds. however. It has a peculiar resemblance to the American skunks. tarily quit the vicinity of water. is This habit speaks to always obtainable. and like them is protected from possible enemies by its very offensive odour. which is replaced in the regions richer in trees by the short-horned buffalo {Bos pumilus). and are found in rocky the Cape polecat (Ictonyx). the diversity of species being as remarkable as the number of individuals. As we have repeatedly stated. It is when we come most to the ungulates. p. 75). it is the antelopes which characterize specially the savanas of Africa. In the reedy swamps throughout the greater part of the continent occurs the so-called Cape buffalo {Bos coffer). and differs from martens and polecats in that it is unable to climb. It is hidden by the mother among long grass. there Both live in herds. bathe.AND DESERTS represented there. In several points of structure antelopes are more primitive than hfe in regions where cover striking distinction to the habits of the . that we find the specialized and abundant of the animals of the African savana. its birth the mother separates from the and remains within easy reach of her young. 127 absent from savanas. in and do not volunwhich they love to The young are born in the warmer season. till man interfered. which she visits at short intervals. and for about is ten days after herd. and frogs. where the young must be able to travel with the herd very shortly after birth. districts. lizards.

96). while in the tropical forests. moving back to the grassy plains to feed and drink. and are by the presence of horns in both sexes. and where possible spend the hours of greatest heat in the shelter of woods. the addax from the deserts of North Africa and Arabia. and the gemsbok {Oryx gazella) from the deserts of Southwestern Africa. usually on very rough ground. they prefer open plains with scattered timber. Typical savana animals. but unlike the buffaloes they can tolerate thirst. preference for bush or thicket-covered country is even more marked in the harnessed antelopes (Tragelaphus). but extend into the desert on the one hand and into the open savana wood on the other. social plants are also infrequent (cf. as well as characters. Kudu (Strepsiceros) are distinguished from eland by by other They haunt thickets and country covered with bush. On the other hand. and in the Kalahari desert apparently obtain sufficient water by eating water-melons and similar succulent fruits or plants. which we have already mentioned as being found even in the tropical forest. &c. Eland are the largest African antelopes. Their speed is not very great. Where possible they drink daily. and they appear to be older. where social animals are rare. Like most ungulates they live in herds.128 TROPICAL SAVANAS oxen or sheep and goats.) are social also. Like not a few distinguished tropical animals they are intolerant of the hot sun. p. and occur in small parties. The the absence of horns in the female. and the student of botany will notice that the fodder plants upon which these animals depend (grasses. are examples of forms which inhabit . They were formerly common over the whole of Eastern and Southern Africa.

and is stated to rest its forefeet against the trees. which resembles a chamois. perhaps because it had few enemies till man appeared on the scene. and remarkable and for its power It prefers the proximity of water.AND DESERTS 129 very arid country. &c. all being inhabitants of open country. Among of we may notice the pala antelope {Aepyceros melampus). by the The rhebok. also small animals. has a South African member in the springbok. other species of the genus Oryx occur in different parts of Africa. and appear capable of going for long periods without tasting water. giving it a resemblance to a miniature giraffe. Of the little klip- we have already spoken steinboks. Other African species also occur. They are remarkable for their speed and their close resemblance in colour to the ground. which inhabits rocky districts. hilly and inhabited (p. the latter seeking the swamps when pursued. The gazelle group. North Africa. 1404 I . but not to be swift. The horns are very powerful. but avoid hilly country. An allied form of interesting habits is Waller's gazelle {Lithocranius walleri) from East Africa. and the gemsbok is stated to be able to beat off the lion by means of them. In addition to the gemsbok. where thorn-jungle occurs. among the acacia scrub of taking long Southern and South-Eastern Africa. and has a remarkably long neck. in order to reach the foliage. It feeds chiefly upon twigs and other forms leaves. 84). found for its great swiftness. In swampy regions where reeds are plentiful the reitbok and waterwater or active buck antelopes occur. occur in open country or savana wood. largely represented in the more arid regions of Asia. mountainous springer districts On are the other hand. once enormously numerous on the margin of the Kalahari desert. leaps..

The wild antelopes (cf. and seek safety there. as e. the strength and powerful horns of the bucks. though one member of the genus extends into Syria. frequent the swamps. and some. thorny jungle. the hartebeests. with some resemblance to oxen. negro children and malaria) are apparently tolerant of the parasites of the various diseases. hke the water-buck. like the duikerboks. and frequent grassy plains. a few. especially those which have so great an extension in Southern Africa. and the bless- The \vildebeests (Connochoetes) are ungainly looking animals. Generally. hke the gemsbok. The virtual absence of wild ungulates in South America when it was colonized from . while not a few extend their range up the mountains. All the three mentioned are exclusively African. The genus Bubahs includes the somewhat stag-like hartebeest. the blessbok. into the tropical forests. for these diseases form the greatest obstacles to the introduction of domesticated ungulates into the great plains of Southern Africa. The resistance which these wild forms offer to the diseases carried by tsetse-flies and ticks must have been a factor in their persistence. rocky regions. which affect them but httle. and the bontebok. by means of which introduced forms may be infected. extend into the desert proper. Less powerful and slower forms must haunt country which offers some form of shelter. group of African antelopes includes the large wildebeest or gnu. the klipspringer being a typical example. and their swiftness. Those which inhabit open plains jSnd security in their numbers. They thus serve as reservoirs of infection.130 TROPICAL SAVANAS last The bok. which inhabit open country in South and East Africa. we may say that very many of the African antelopes are typical savana animals. g. but some. or swampy districts.


Fig. 36.


Giraffe drinking, showing the great difficulty
in spite of the length of

has in reaching the ground,
specimen in

the neck.

which the animal {From a

Royal Scottish Museum.)



Europe meant the absence of such reservoirs of disease, and accounts for the greater ease with which domesticated ungulates were introduced there.
ungulates from South America
It has


suggested as a reason for the disappearance of so

abundant there


parasitic disease.

for they were once they were killed off by some this were so, then with the dis-


appearance of the hosts the parasites must have died also, leaving the field clear for reintroductions later. In Africa we have to notice that the tolerance of minute blood parasites a tolerance doubtless originally ac-

quired at the price of a fearful death-rate
of protection of the existing

^is a means forms against the intru-

sion of

the trypanosomes carried

new and competing forms. The resistance by the tsetse-fly has even

to to

some extent protected the antelopes from their great enemy man, for European hunters are constrained to avoid regions much infested by the tsetse on account
of the difficulties of transport.

Another very beautiful ungulate which frequents all the open parts of Africa south of the Sahara is the
which shows a pecuHar adaptation to life in the The great elongation of the neck and of the forelimbs enables it to browse upon the leaves of


the acacias and other trees of the scrub. Despite the length of the neck, however, drinking presents great

the animal being compelled to straddle


legs apart before it

can reach to the ground.


similarly difficult,

and is

rarely practised.


inhabits arid country, notably the Kalahari desert, and,

sandy plains where the scrub on which it It can apparently go many months without drinking. The animals go about in herds, and are capable of great speed. Only a single young one is

feeds occurs.

I 2



produced at a

and when three days old


can trot

by the

side of its mother,




from the

attacks of carnivores


large eyes

by kicks with her powerful legs. and the long neck give the giraffe a

very wide range of view, very necessary in an animal which frequents open country. Deer are usually absent from open plains, and are entirely absent, as we have seen, from Africa south of the Atlas. But in South America, where antelopes are totally absent, the deer extend their range to the plains and swamps. Thus the pampas of the Argentine and Paraguay, together with similar regions further south, are inhabited by the pampas deer (Cariacus campestris), which finds the necessary shelter among the long pampas grass. This deer occurs in pairs or small parties, and the female is particularly ingenious in protecting her fawn. The latter makes off through the grass on an alarm, and then cowers down, while the mother takes herself off in another direction in a slow

and hmping manner. We have spoken of the camels of the steppes of the Old World, and have seen also that their aUies, the llamas of the New, are intolerant of great heat, so that
the tropical savanas, whether in Africa or in America,

have no members of this family. Of the odd-toed ungulates the rhinoceroses, as already mentioned, extend into the plains, this being especially true of Burchell's rhinoceros, a grass-eating form. Of the striped horses of Africa Burchell's zebra and the quagga are (or were) both inhabitants of the open grass-covered plains, but both are chiefly extra-tropical, inhabiting the vast plains of the extreme south of Africa. The African wild ass is similarly an inhabitant of the arid regions of North Africa. While Burchell's



zebra and the quagga formerly occurred in vast herds, the wild ass, an inhabitant of a region where food is

always scanty, occurs in smaller parties. Its sober tint harmonizes with the uniform colour of the desert, and the fact that the domesticated form will eat dry and prickly food points back to the days when its ancestors contented themselves with the thorny herbs of the desert. The stripes of the zebra and quagga apparently harmonize with the light and shade playing through the open bush country of their native haunts. It will be noted that the presence of the various forms of wild horses on the plains to the north and south of Africa,


their limitation chiefly to elevated regions in the

tropics (Grevy's zebra), speaks to a relative intolerance of

heat, a not


feature in the larger ungulates.


to the rodents,

we note

that in Africa those of

the savanas are relatively unimportant, while in South

America, with the paucity of ungulates, there are many large and important rodents. We have already spoken of the jerboas, which extend from the steppes of Asia into the deserts of North

and have noted

their special adaptations.


the southern half of the continent the place of the
is taken by the so-called jumping hare (Pedetes very common at the Cape, but extending northwards into Angola and Mozambique. While related to the jerboas the animal resembles the common hare in size and colour, but it has a long hairy tail, and elongated hind-legs. It feeds upon all-fours, but escapes from its enemies by taking long bounds, stated sometimes to cover as much as thirty feet. The animals are burrowers and nocturnal, and frequent arid districts. True hares also occur in Africa, but offer no special






Among the various mole-like forms of Africa mention may be made of the little sand-rats (Heterocephalus)

—which are no bigger than a mouse—are nearly naked,

of the

sandy deserts

of Somaliland.

These httle animals

perhaps as an adaptation to life in sandy soil. They resemble moles in having no external ears, and in their almost functionless eyes. Of the rodents of South America the majority have a wide distribution, ranging from the true savanas southward to the pampas of the Argentine and the plains of Patagonia. The viscacha {Lagostomus trichodactylus), according to Mr. W. H. Hudson the most typical inhabitant of the pampas, extends beyond their Hmits, though scarcely into the savanas proper. On the other hand the mole-hke tucotucos (Ctenomys) are burrowing animals, which occupy sandy regions from Brazil and Bolivia to the extreme south. The coypu (Myopotamus), an aquatic form haunting the banks of rivers and lakes, has an even wider distribution. Many species of cavy also occur, some being partial to moist situations, while others dwell in rocky regions. The largest living rodent, the capivara or capybara {Hydrochoerus capivara) has a very wide distribution, and is aquatic in its habits, frequenting the margins of lakes and rivers, but also feeding on the open plains. Of the South American Edentates most of the armadillos inhabit open country within the tropics, though some extend into extra-tropical regions. Most prefer
arid regions,

and feed on



carrion, together

with some vegetable matter. The tiny pichiciago or fairy armadillo (Chlamydophorus) is found in sandy regions in the western part of the Argentine where the vegetation consists of cactuses and thorny brushwood. It burrows with great rapidity, and is very sensitive

38.) . 37.) Fig. (From a specimen in the Royal Scottish Museum.J Jerboa (Dipus) from Central Asia. The animal is of a greyishsandy colour which renders it very inconspicuous in the desert regions in which it lives. (From a specimen in the Royal Scottish Museum. Fig. The Egyptian Jerboa (Dipus).


It aids the animal in maintaining the upright position. though it has no armour It seems to feed chiefly on as the armadillos have. and therefore necessarily of the neck to permit feeding to take place. without corresponding elongation of either forelimbs or neck.. termites. In addition to the protective covering of bony plates. the type genus Dasypus are widely distributed. which they reach by standing in the upright position. 135 The members of Another species is found in Bolivia. are ground forms. being especially common in the Argentine. The kangaroos and wallabies. We hind-limbs.AND DESERTS to cold or wet. far is an adaptive peculiarity which is from uncommon in animals of the open plains. the aard-vark or ant-bear (Orcyteropus) occurs in open regions. another curious edentate. or even in the case of the larger forms. and is also of burrowing nocturnal habits. In the ungulates the other type of adaptation. &c. of trees. The tail is long and strong. In Africa. that which consists in the elongation of both pairs of limbs. where armadillos are completely absent. and also in running that is. on leaves. is the usual one. Of the marsupials the kangaroos and their immediate allies are exclusively confined to the Australian region. and in the fact that they are mostly nocturnal. they find security from their enemies in their burrowing habits. feeding not only upon grass but also upon shoots of bushes and shrubs. the so-called white ants. and there they formed the natural fauna of the savanas till the advent of the European with his flocks and herds. in — leaping. of have already noticed the essential points structure in the kangaroo the elongation of the . Just as among the rodents . with the exception of the tree-kangaroos already mentioned.

or as the long-legged antelopes and giraffe have been evolved from the ancestral short-legged of the on all fours. That the same need produces the same results is obvious when we note that the kangaroo has four toes on its hind-foot instead of five. and that of those four two are rudimentary. and in the structure and shortness of its hind-feet approaches the phalangers. to a fusion of hmb bones. so among the kangaroos we have forms like the rat-kangaroos (Potorous) in which the elongation hind-hmbs is much less marked. Uke the useless side toes in antelope or deer. just as the jerboas on the Asiatic steppes have been evolved from ordinary rodents. but we may repeat that if an animal of the open plains cannot burrow. for they are said to be employed in cleaning the fur.g. . We saw also in connexion it must have acute senses. it must have a wide range of vision in order to note their advance. we have various stages in the elongation of the hindand therefore in the development of the power of leaping. or defend itself passively in some way. e. In the scrub of Queensland the fivetoed kangaroo {Hypsiprymnodon moschatus) occurs. it must be swift to escape its enemies.. and so on. The nature been already emphasized.136 TROPICAL SAVANAS limbs. in order to give the necessary strength and rigidity for rapid movement. showing that the true kangaroos have been evolved in response to the special conditions which prevail on the open plains. the giant kangaroo's means of defence. though in the kangaroo the rudimentary toes have apparently a subsidiary use. with the ungulates that there is a marked tendency to ungulates. in response to similar conditions. and which run and cannot dehver the powerful kicks which are so important a part of. of these special conditions has reduction in the number of digits.

Further. It however always returns to it on an alarm. The mother shows great devotion to her young. and the young one is born in a very imperfectly developed condition. In the case of the kangaroo con- ditions are very different. Others regard the action as equivalent to an abandonment of the mother. and soon the young one is able to move and leave the pouch. who. It is placed by the mother in the pouch with which she is provided. as — capable of flight. the young ungulate. but has apparently been justified in the by the higher survival rate of we may note readily among domesticated sheep. however. The period of gestation is very short. We have noted that among the swift ungulates the tendency is for the young to be born very fully developed. and is not able to care for itself till it is some eight or nine months old. by the action of certain of the muscles of abundant food. growth is rapid. the kangaroo shows a very striking contrast to the ungulates. wiU return to pick up her joey '. when suckling. able after a very short period to run by the side of the parent. to this ' Owing . but if hard pressed by dogs she will throw the young one out of the pouch into the fern. The consequent long period of gestation is dangerous to the mother. if she escapes. According to some authorities the chances are that the dogs will neglect the young one and continue to chase the mother. and attached to the teat. Milk is then pumped into its mouth. thus relieving herself of a weight which diminishes her speed. The action has been variously interpreted.AND DESERTS 137 In one respect. incapable of sucking and quite incapable of voluntary movement. is not allowed by the mother to satisfy its appetite completely it must never be so satiated as to be instruggle for existence the young.

has been justified in the struggle for existence as against the marsupial method. take both vegetable and animal food. long hind-legs. that practised by the ungulates. The jerboas and jumping mice of other desert regions are represented . tions. the hind only one of . the animals are markedly gregarious. and do not attempt to defend themselves nor to hide. Only one kangaroo is produced at a birth. TROPICAL SAVANAS In any case the fact that. suggests that the placental method. because a reduction in the number of young is frequent in animals which must escape with their young by flight when threatened. A similar tendency is well marked in the higher ungulates. primary purpose of the elongation of the limb is to give swiftness. For example. remarkable in showing an ungulate-like reduction in the number of the toes. capable of kiUing a dog. yet secondarily. despite the greater risk to the mother before birth and at birth in the former method. Like so many denizens open country. e. though the helpless.138 the young.g. But it must not be supposed that the kangaroo is entirely In the ungulates we find that. The fore-foot has the animals two functional toes. with rare excepmarsupials have only succeeded in surviving in the Australian region. these adaptations exhibiting a curious convergent resemblance to those which appear among the placental. In regard to the other marsupials we need only note that Australia contains various forms shomng special adaptations to open grassy plains or desert regions. Similarly the kangaroo can use its long hind-legs to deHver heavy blows. This is an interesting point. a small animal vnth. the powerful hind legs can be used as weapons. throughout the greater part of Australia the open grassy plains are inhabited by the so-called pig-footed bandicoot. in the horse.

Among the inhabitants of savana regions. and is found in sandy regions where ant-hills occur. but they drink freely if water is available. Like many attacked. the gregarious habit. and found also in Syria. the length of the hind-legs and the reduction of the number of toes to two gives them speed (cf.AND DESERTS in 139 Australia by small marsupials of similar habit. and use their hind-legs as weapons when The young are active almost immediately after hatching. as in the ungulates. ostriches can go without water for a prolonged period. . giraffe). Arabia. Again. a bushy tail. The males are bigger and stronger than the females. In South America the place of the ostrich is taken by the rhea. the banded ant-eater {Myrmecobius fasciatus) shows the long extensile tongue common to ant-eating forms. again the giraffe). must be made seen in adapted to these the running birds. being associated with polygamy. The birds are chiefly found on the pampas. The habits are somewhat similar. again recalling the precocious young of the ungulates of the steppes and savanas. African distributed throughout Africa wherever the open sandy regions suitable to its habits occur. savana animals. mention of the districts. notably by Antechinomys laniger. Like the large antelopes with which the ostriches associate in the southern parts of Africa. which correspond rather to steppe regions than to savanas in the strict sense. and the habit of progressing by jumps. a little jerboa-like animal with long hind-legs. the ostrich is a social animal. which are specially The adaptations are well ostrich (Struthio). and Mesopotamia. which has three toes instead of two. The great length of the legs and of the neck gives the animals a wide range of vision (cf. and some other structural differences.

is an inhabitant of W'ooded Another interesting group of birds which inhabit the grassy plains of South America. but possess the power of flight. suggests that the ground-hauntmg habit is a recent acquisition. are the tinamus. The curious secretary bird (Serpentarius secretarius) also. haunting the plains and open country. on the other hand. The flight is swdft. Its structure do not shows that it is allied to the birds of prey. though extending also into forest regions. and rise with reluctance. and are for long unable to run upon their elongated legs. it is capable of flying well and strongly. it prefers to run along the ground. The nest is a huge structure and is often placed in a mimosa bush. rarely venturing out into the open. The emu. with its long legs. where it eats a large amount of grass. like the cassow-ary. like the ostrich. which resemble the rheas in having three toes. but they fly nearly so well as the game birds. being popularly called partridges. of which many species occur in Africa. just as the common form occurs in the steppe regions of Asia. preferring to skulk among the long grass. but cannot be kept up for long at a time. but of these the cassow'aries are forest birds. and the fact that the young are very helpless. which show certain resemblances to ostriches. The remaining living member of the group. They strongly resemble game birds in outward appearance.140 TROPICAL SAVANAS In Australia and the neighbouring regions two other types of running birds occur. districts. is a savana bird. is Though very characteristic of the African savanas. . these being the emus and cassowaries. the kiwd of New Zealand. Among other birds characteristic of savana regions mention may be made of the bustards.

39. U. it The Cactus Wren (Gampylorhynchus brunneicapillus). but showing a curiously close nature .the cranes. (Photo by the Biological Survey. birds apparently related to. a characteristic bird of the desert regions of California and Texas.AND DESERTS 141 Fig.S.A.) In South America the place of the secretary bird in is taken by the seriemas. where builds a complicated nest in cactus bushes.

and has the curious power of being able to absorb water through its rough skin. Somewhat similar in appearance are the homed (Phrynosoma) of the desert regions in the toads warmer parts of the United States and in Central America. In the sandy districts of Western and Southern Austraha occurs Moloch horridus. which are admirably adapted for desert life. show any very marked adaptations. The presence of the hzards named. and are said to appear actually to sink into the sand. together with many others. but of the numerous hzards which occur there one or two may be named as showing interesting pecuharities. They also have a rough and prickly skin. as desert and savana regions. and is a cursorial bird.142 external TROPICAL SAVANAS resemblance to the secretary birds. This is presumably an adaptation to permit the animal to avail itself of the rare showers which fall in the desert regions where it hves. speaks to the abundance of insects in These insects do not. or so-called white ants. and in colouring resemble the sandy soil in which they hve. and possess remarkable powers of burrowing. as being . There is nothing very characteristic about the reptiles of the savanas and hot deserts. a hzard covered with spines and tubercles. The animals are assisted in the process by a curious fringe at the sides of the body. The Brazihan seriema (Cariama cristata) haunts open districts with scattered bushes. but we may a mention the termites. In the hot deserts of Northern Africa ' ' and the adjacent regions occur the skinks (Scincus). though it roosts in trees. their bodies enabling them to move through the soft sand as fish swim through water. They feed upon a variety of insects. It Hves upon ants. rule.

Mention may also be made of W. References. All the books 143 very characteristic of the savana regions of Africa and mentioned in the previous chapter as giving accounts of African animals. H. and of Tristram's Flora and Fauna of Palestine (London). end X . 1892) . describe the inhabitants of the savanas as -vvell as those of the forests. especially The Naturalist in La Plata (London. describe among others the savana animals. as giving accounts of the zoological regions. Hudson's books. The books mentioned at the of Chapter also.AND DESERTS elsewhere.

Even to flying animals a large expanse of sea may constitute a formidable barrier. the climatic conditions are similar. but no such . The question whether a particular animal does or does not occur in an island is merely a question of observation while that as to whether a form found on a mountain does or does not constitute part of the mountain fauna is a much more delicate one. it is On the other hand. We have already seen that mountain areas have special types of animals. The special conditions in their case include both climate and topography. to be noted that while . and in whatever region of the globe the mountains occur. by the slow migration is possible in the case of islands. if they are greatly elevated. because they form isolated regions cut off by the physical conditions from the their neighbouring districts. Islands are regions cut off from neighbouring regions sea. save in the case of those with exceptional powers of flight.CHAPTER VII SPECIAL FEATURES OF ISLAND FAUNAS Islands present some interesting features as regards animal life which makes them worthy of special study. and can be studied with much more ease than the animals of any other natural region. involving an elaborate process of reasoning. which forms a barrier to the passage of all but the marine and the flying animals. The consequence is that island faunas form a whole. In the one case a gradual migration with concurrent adaptation is possible from plain to mountain or vice versa.



mountain animals, like those of steppes or forests, are exposed to virtually uniform conditions of climate, &c., no such uniformity exists in the case of islands. We
speak, it is true, of insular climates, meaning that the proximity of the sea has a certain moderating effect, more especially if there is a large expanse of ocean all round, or if the prevailing winds blow from such an expanse. But even within the same island there may be great variations of climate, and there may be almost all types of surface mountain, wood, steppe, tundra, and so forth. Thus, while the inhabitants of islands may be readily counted and classified, and form a statistical unity, they do not form a biological unity in the sense that the animals of the other natural regions do, and there are few general statements which can be made of them as a whole. The special features of island faunas may best be illustrated by a few examples, and we may begin by a consideration of the Galapagos Islands, an interesting group which has been studied by various naturahsts, including Darwin. This archipelago consists of a group of islands lying some seven hundred miles off the coast of Ecuador, and on the equator. They are of volcanic origin, the surface being covered with extinct craters, and are placed upon a submarine bank, which rises steeply from the depths of the Pacific. The climate is singularly cool for the position, this being due to the fact

that the islands are washed

by the cold Humboldt

In the lower regions of the islands the rainfall is markedly deficient, rain only falling in small amounts in the earlier months of the year (FebruaryMay). Here the vegetation is scanty, and xerophytio in character, the most important plant from the





economic point of view being the archil lichen from Darwin which the dye called Htmus is obtained. describes this lower zone as uninviting in the extreme,
the scanty brushwood leaving the black basaltic lava
exposed, and giving the district the appearance of an
iron-smelting region in the

black country




But the clouds hang


about eight hundred feet

and above an elevation of not only is there abundant

rain during the rainy season, but even at other seasons

a constant mist keeps the vegetation fresh. The plants of the two regions differ fundamentally, and in the upper cultivated plants from many cKmates thrive. Of the native vascular plants, about one-haK are pecuUar
to the islands.

The group

consists of five principal

and eight small islands, with a total area of nearly The hills range in height from 3,000 square miles. 3,000 to 4,000 feet. The more conspicuous bushes and trees are species of euphorbia, cactus, and acacia, so that the islands generally resemble savanas and
desert regions in their vegetation.

Palms are



Turning now to the fauna, we find that mammals are There are two mice, but of these one belongs to the widely distributed genus Mus, and the other belongs to has doubtless been introduced a South American genus (Hesperomys), and has also possibly been introduced. That there is nothing in the conditions of the islands to prevent the existence of even large herbivorous mammals is, however, shown by
practically absent.

the fact that goats, pigs, horses, &c., introduced by

man, have become


and have multiplied exceedinteresting features.


birds present







have been



obtained in the islands, and of these forty-one, or considerably more than half, are peculiar to the islands,

not being found elsewhere. Most of those which are not peculiar are birds with considerable powers of Of the flight, capable of travelling great distances. true land birds all but one are peculiar, and this one is the common rice-bird {Dolichonyx oryzivorus), which occurs throughout the whole of the American continent. This bird is markedly migratory, spending its winters in the West Indies and Central America, and travelling northwards in vast numbers to breed and spend the summer in the northern United States and Canada. There is, therefore, nothing improbable in the supposition that stragglers from the migrating flocks reach the Galapagos Islands from time to time, and so prevent an island type from establishing itself. The other land birds of the islands show, as Wallace points out, all gradations from close resemblance to forms occurring elsewhere to generic difference. Thus the archipelago contains a special owl Asio galapagensis but this is very nearly related to the widely distributed shorteared owl {Asio hrachyotus). On the other hand, among the finches peculiar genera occur, e. g. Geospiza, related to genera of restricted range found on the continent of South America. Again, though the islands are relatively so near together, a peculiar genus may be


by three



on as many

different islands.

well exemplified in the case

of the honey-creepers of the genus Certhidea, the three

species of whicli occur


different islands.

The reptiles of the islands are especially remarkable. The giant tortoises have long been famous, and give the islands their Spanish name. Such giant tortoises
occur only on islands, beiag found in certain islands in





the western Indian Ocean, and in the Galapagos archipelago. In addition to their large size the animals have
as special pecuHarities the fact that they

individual variation,

and tend


show much to spht up into

species, different species occurring

on adjacent


small tortoises occur on the islands containing these

A final peculiarity is that the bony carapace tends to be reduced in thickness, perhaps because tiU man came the animals had few enemies. The diet
large forms.

exclusively vegetarian,

and the animals occur

in both

the upper well-watered region and in the lower arid

The fact, however, that they drink greedily, and are fond of wallowing in muddy water, suggests that they are not native to arid regions. Darwdn gives an interesting description of the long journeys made by

the low-ground tortoises to the springs of the upper
region, in order to obtain water.

The cactuses

of the

arid regions



form a favourite food plant. The exact species is a matter of doubt, owing to the

individual variability already mentioned.

In addition to the giant tortoises, the Galapagos possess both lizards and snakes. Two of the lizards are very large and of very peculiar habits. One {Conolophus svbcristatus) is found only on the central islands of the archipelago, and occurs both in the upper and lower regions, but more abundantly in the latter. The forms from the arid districts, however, do not travel

upwards to the springs to drink, and are far more numerous than the upland forms. As the diet of these lizards is quite similar to that of the tortoises, it would seem that originally the lizards inhabited the arid ground, and the tortoises the damp upper regions, but each has become partially adapted to the territory of the other. The lizards reach a length of a yard, and

liave stout





are very partial to

cactuses, but also climb the acacia trees to feed


the foUage.
Closely related to Conolophus, and therefore like it one of the iguana family, is the marine lizard Amhlyrhynchus cristatus. It also is entirely peculiar to the islands, and is the only Kzard known which is marine
in habitat.
It inhabits the shores of most of the islands, and feeds upon seaweeds which grow below tide mark, and, as Darwin proved, it can tolerate prolonged immersion without any ill effects. It will be recollected

p. 116) that the typical

iguanas are forest animals.

to which the forms in the Galapagos Islands are exposed has made it necessary for these to adopt modified habits. The family Iguanidae is further represented in the islands by various peculiar species of the genus Tropidurus, other members of which occur on the continent of South America. Various species of geckoes belonging to the widely distributed genus Phyllodactylus also occur, most of these being pecuhar to the islands. There are also two snakes, both being nearly allied to South American forms. Amphibians are entirely absent. Of the invertebrates the insects are few in number, but show the same characters as the other kinds of animals. In other words, most of the species and many genera are peculiar, and the forms tend to have a Hmited distribution among the islands, very few being found in all the

The exceptional conditions









feature in island insects.

It is not necessary to describe the fauna of other

islands of the

in detail, for the general characters are the

same type as the Galapagos archipelago same in

It is noticeable that the power of flight tends specially to diminish in animals found in islands much exposed them to hurricanes. and the presence of the large tortoises in the Galapagos Islands is held by some to indicate a connexion. due apparently to the isolation of the stock. a fact readily explained when we recollect that both the eggs and adults are very intolerant of salt. especially insects and land shells. while those which flew little or not at all would be more likely to breed and continue their own type. This is very marked in some birds. animals readily introduced by man. dentally Wallace. and are separated from the nearest continent by deep water. flying forms tend to lose their wings or to possess only limited powers of flight. and thus destroy them. and therefore could not be readily transported across sea-water by floating logs of wood or any similar means. that is. In such islands generally the commonest land forms are birds and invertebrates. at a remote period. do not lie upon the same continental shelf as the continent. thus the flightless dodo. for here even short flights might expose the animals to strong winds. . believes that the ancestors of these forms were acci- introduced.150 all. possibly on floating timber. with the continent of America. are no indigenous land These in all cases tend to run into local races. which would sweep out to sea. In such islands there mammals. SPECIAL FEATURES Islands which do not he very near a continent. Further. and so forth are all inhabitants of islands. Reptiles are not very frequent in islands separated by deep water from continents. the solitaire. on the other hand. the kiwi of New Zealand. show great general similarity as regards their fauna. often with the doubtful exception of mice and rats. Amphibians are also absent.



As contrasted with . &c. we find that its fauna is poorer even than that of the larger island. As Ireland was apparently cut off from the continent before Great Britain. islands oceanic islands we have the which are obviously merely separated portions of the adjacent continent upon whose continental shelf they stand. quoted from Wallace. most of which occur in lakes. Great Britain 40. called by Wallace. That the red grouse (Lagopus scoticus) of the northern regions is pecuhar is generally admitted. The differences are. at least to a large extent. A good example is furnished by the British Isles. but shows impoverishment in several respects. We have also some peculiar fresh-water fish. for the paucity of sources of animal food tends to make — their human inhabitants cannibals. though it is nearly allied to the willow grouse of Scandinavia. not very striking. .or ISLAND FAUNAS oceanic 151 Generally the fauna of islands like the Galapagos. and Ireland only 4. will help to render the above statements more precise. Great Britain 13. apparently as a result of the glacial period. however. and at a stage when many of the animals driven south by the ice had not had time to recover lost ground. and Ireland 22 Belgium has 22 species of reptiles and amphibia. or by mechanical transportation on floating wood. whose fauna is very similar to that of the adjacent continent. The following figures. is such that its members may be supposed to have been. accidentally introduced by storms carrying winged animals out of their usual course. Incidentally we may note that the absence of mammals on oceanic islands has often important human consequences. Scandinavia possesses about 60 species of mammals. The question as to the number of pecuHar forms in the British fauna has been actively debated.

In this case we shall take Madagascar. There are few rodents. which belongs to the civet group. which is of the continental type. The fauna is ably rich. the fish of one loch interbreeding with those of another There are also a considerable number but nothing comparable to the differences which separate the animals of the Galapagos Islands from those of the adjacent mainland. in contrast with the oceanic type. which is so in the . and is as markedly characterized animals it includes as by those of which it is Thus the island has no monkeys in its wide remark- by the devoid. about the size of a common cat.152 SPECIAL FEATURES fact that the chances of and are no doubt due to the are very small. The total area of peculiar species or varieties of insects. and no true cats. Insectivores are fairly numerous.500 square miles. Suchislands as the British Islands are called by Wallace continental. and the enormous wealth of large ungulates. and there is much forest. but in addition to eight civets there is a relatively powerful animal called Cryptoprocta.000 square miles of the Galapagos Islands. To add point to the contrast we may note that the area of the British Isles is in round numbers. 121. of this large island is nearly twdce that of the British Isles. One more example of an island fauna may be given. tropical forests. There are not many carnivores. and include in the tailless hedgehogs (Centetes) primitive forms whose nearest aUies appear to be some shrews West Indian Islands. constitute half which the mammahan population of the island. but is separated from the continent of Africa by a channel so deep and wide that it must be supposed that the connexion between the two was broken at an extremely remote period. as contrasted with the 3. but has some thirty-three species of lemurs.

and the presence of primitive or peculiar forms. notably the lemurs. Among both snakes and lizards we find the peculiar feature that American forms are represented among the lizards by two Iguanidae. Both the species and genera of mammals are strikingly peculiar . Saved from the competition of higher forms by isolation. forms of &c. The beetles. have reached a high . and consequently the original mammalian and other fauna of Madagascar was primitive in type. Without considering in detail the various questions snakes. Australia. many forms. for that important order is only represented by a river-hog and a subfossil pygmy hippopotamus. raised by these pecuHarities. and favoured by the abundant food. thus it appears that of the twenty-four of mammalian genera Africa. had penetrated Africa. the higher ungulates. but they also show.OF ISLAND FAUNAS characteristic of the continent of Africa. otherwise peculiar to America. is 153 here scarcely suggested. other lizards are mostly peculiar famihes. again.g. The . show affinities with those of India. for they present affinities both with the Indian region and with South America. the absence of enemies. which abundant in Africa. though to a less degree. but are members is of African a special feature the number of chameleons. the same feature of the absence of the African forms which would be expected. Madagascar only two occur in The birds are not quite so pecuhar. it may be sufficient to say generally that Madagascar Africa. was formerly connected to to India. There are no poisonous The amphibians are remarkable. and the Malayan region. and the great forest area. the monkeys. and the absence of the monitors. and perhaps also through a chain of islands The connexion was broken before the higher life.. e.

others beheve that the presence of giant land tortoises For is in itself a proof of a former land connexion. or before they had had time to become widely distributed. or (1) no reptiles. has made it difficult to apply. or (2) reptiles of such as could be accidentally carried by floating wood. or has disappeared. &c. this must have been prior to the deposition of the lower Tertiary beds. and have many different directions. because the pro- gress of geology. it is sufficient : to recognize the follow- ing three conditions When an island has never a continent. St. g. e.154 SPECIAL FEATURES own lines. the Sand- wich Islands. a very primitive type. for at this . we may note first that Wallace's degree of specialization along their evolved in distinction into oceanic and continental islands is not of very great practical importance. &c. no amphibia. the Bermudas. The most important members of its fauna will be birds. insects. It involves a conception generally accepted. many coral had a connexion with and volcanic islands. Helena. or has only had such a connexion at a geologically remote period (Galapagos?) before the higher forms living at the present day had originated. which is not now Thus. its fauna contains no mammals. since the classification was laid down. fitting them for their habitat. Examples of such islands are the Azores. and land shells. Among flying forms the conditions favour those in which the power of flight is moderately developed. Where such islands have had a continental connexion. If we sum up the more important general points in regard to island faunas. while Wallace names the Galapagos group as a characteristic oceanic group. Owing to the isolation the animals will tend to (3) exhibit special pecuharities. of the permanence of ocean basins our purpose then 1.

2. which a minor extent in the same author's Darwinism. When the separation of an island from the adjacent continent has taken place within geologically recent times. including representatives of all the chief classes and phyla. Java. it will tend to have a rich fauna. e. Nat. all the fauna will be in essentials similar to that of the adjacent land mass. but the more modern classes will be represented by primitive forms. On Some Peculiarities in the Geographical Distribution and in the Habits of certain Mammals inhabiting Continental and Oceanic Islands {Ann. 3. 1884). See also Dobson's. ' ' the classical work dealing with the subject. e. may occur.OF ISLAND FAUNAS time 155 mammals were evolving rapidly and spreading widely. When an island has been separated from adjacent land masses for a moderately long time. Japan. xiv. since the middle of the Tertiary period. i. 1895). &c. their primitiveness depending on the length of the period during which the island has been isolated. the most satisfactory classification of islands from the point of view of their fauna depends upon the approximate length of time since which they have been isolated. London. Of such islands Madagascar and the adjacent islands of the Indian Ocean are good examples. in the Pleistocene or Post-Pleistocene period. References. 1843). Borneo. A very interesting account of the Galapagos Islands will be found in Darwin's Journal of Researches into the Natural History and Geology of the Countries visited during the Voyage of the Beagle (London. In other words. Hist. Wallace's Island Life (second edition. 1892) is is also discussed to . i. The subject is also fully treated in Beddard's Text-hook of Zoogeography (Cambridge. though minor differences Examples are the British Isles.

and savana wood into forest. The two great conditions which influence the type of animal to be found within any part of the sea are. for land- mud only accumulates off shores. All these organisms agree in that. as we have seen into one another in the previous chapters. the presence or absence of hght. we have the httoral area. the presence or absence of a substratum. In the open water occurs another group of organisms. A plaice. At the margin of the sea.CHAPTER VIII THE DISTRIBUTION OF ANIMAL LIFE IN THE SEA While. so shaped that it and where hght penetrates. and the minute algae must have light before they can five. fixed to a rock and containing symbiotic algae. if less apparent. Thus a sea anemone. and. they show adaptations to the two striking features of the area. is adapted for lying on is a sandy surface and bearing well-developed eyes. — much for here the divisions are relatively sharp. the natural regions of the land more or less grade savana. But the dependence of a mud-haunting animal feeding upon minute algae upon httoral conditions is no less real. first. similarly fitted. derived . in structure or in habit. it is veryeasier to separate the oceans into natural regions. peopled by littoral organisms. passing into savana wood. second. is obviously adapted for life in a region where a substratum is present. where the light penetrates to the bottom. for example.

or pelagic stages in their Hfe-history. and usually bottom Finally. each of which we must consider separately. and (3) the abyssal. able to take advantage. where the littoral Shallow littoral animals are replaced by the abyssal ones. we have forms entirely independent of sunlight. each class showing adaptaThese three groups are tions to a natural region. (1) great variations in the physical of food are here three : The basal sources (2) (3) the waste of the land. in the great depths. of the many currents which influence the surface waters of the sea. and conditions. example. the fixed algae of the shore rocks. contain only and pelagic animals. the members of the different groups do not necessarily restrict themselves throughout their whole It is. very common to find that littoral forms have pelagic larvae. seas. for life history to one of the natural regions. but directly or indirectly dependent upon the presence of a substratum. These are the abyssal animals. (1) the littoral. but for the most part display sensitiveness to it. whose existence was first fully demonstrated by the Challenger expedition. the minute floating algae or phyto- . The littoral zone extends outwards from the (1) margin of the land to the edge of the continental shelf. As we shall see. is Throughout its extengreat sion the littoral area its characterized by its wealth of food. or roughly to the one hundred fathom line. such as the North Sea. (2) the pelagic. We can thus divide the inhabitants of the ocean into three well-defined classes. Beyond this line the sea-bottom usually slopes rapidly in the Continental Slope to the great depths.ANIMAL LIFE IN THE SEA constituting the 'pelagic forms. which require 157 no subof stratum. demand hght. for purposes of distribution. at the the ocean.

which in their turn feed many fishes to many and so forth. At high tide this zone is covered with a sufficient depth of water to be available as a pasture ground even for the larger fish. coelenterates. and is not available as a source of supply either for the . such estuaries are specially favourable to many marine to shell-fish like mussels and oysters worms. The Laminarian zone. molluscs. The fixed algae can only grow where some firm sur- face to which they can attach themselves exists. . and so forth perfectly adapted to life among the long brown fronds and about the branching roots. has always been known as a rich coUecting-ground. and here therefore shore animals cluster. especially. . worms. If the water be fairly clear. or by the waste of the land. which lies just beyond low-tide mark. Diatoms. In their habits and adaptations most of these are pelagic. They are therefore limited to the margin of the lands. that is apparently independent of the presence difference but the fact that there is considerable between the phytoplankton of the shore water and of the open sea. pelagic or for the abyssal animals. which also feed fish This land-waste has only a limited seaward extension. but there form rich feeding-grounds. in the shore waters minute algae swarm. suggests that these forms are in some way indirectly affected by the proximity of the sea-floor. which are eagerly consumed by many of a substratum. Finally.158 plankton. It contains many small forms Crustacea. THE DISTRIBUTION OF The importance of the waste of the land may if we think of the number of marine animals be realized which find food in the mud-deposits which accumulate round the mouths of rivers and estuaries. small Crustacea. . and the — — fisherman reaps rich harvests. and where the sunlight is strongly felt.

but there they degenerate. Many shore fish. Tides and currents. and thus keep the water sweet. strong to stem the tide. powerful forms like porpoises and dolphins and even whales there are times when the strength of these avails them nothing tide or current. whose influence is manifest in several ways. in aU the open seas. and yet. it may be asked. but they no more form a true part of the life of the pelagic area than does that shoreborn weed which floats at the surface of the Sargasso Sea for a period. and bring constant supplies of food and oxygen. the shore naturalist may coasts he is sure of a rich harvest knows that off many when wind enforces beach at certain seasons is strewn with flotsam and jetsam. It is this constant movement which helps to give the shores of the oceans so much richer a fauna than the shores of lakes. The constant movement brings with it also a danger. But. are represented in the shore waters species 159 by many more than in the open ocean. or of enclosed seas like the Mediterranean or Black Sea. into the ocean.ANIMAL LIFE IN THE SEA animals. if these algae have no attachment to a substratum. Of the physical peculiarities of the littoral area the constant movement of the waters is characteristic. how do they contrive to remain in the shore waters. notwithstanding their strength. their be to serve as food for the pelagic animals of the open seas. and probably do not destiny Produced in the shore waters. for the — . Some of the shore forms are swift swimmers. the strong cuttles among molluscs. ensure that no stagnation shall occur. in spite of currents and tides which must tend to sweep them seawards ? The answer —by is apparently that myriads are so swept out live long. ere it perishes and is replaced by new fragments torn off by the currents.

. the range is considerable. can the ranges of temperature in the sea be compared with those on land. the latter Speaking generally. the are rapidly killed by them. &c. that the range of by the variations in the temperaby latitude. though not so obvious. however. the limpet among molluscs. have a heatregulating mechanism. and they are found dead or stranded on the beach. of course. Between the two. In polar waters. Such a mechanism is absent in all sea animals except those (e. and is influenced by local causes. while the range similarly small. but the shore waters especi- — — ally are greatly affected ture of the land.). According as they can or cannot tolerate variations in the temperature of sea water. but cling tight and allow the danger to pass. The weaker abandon the attempt to struggle against the waters the lumpsucker and the gobies among fish. The temperature of the oceans varies much. that very many land animals.).).160 against the THE DISTRIBUTION OF movement of the waters. temperature in temperate regions. g. who are descended from terrestrial ancestors. the acorn shells and barnacles among Crustacea these do not attempt to resist the tides. owing to the high specific heat of water. The former are tolerant of great variations. which the shore forms must guard against is the constant change of temperature and of salinity in the sea water. In tropical waters temperature is small (about 10° F. Another danger. while the mean is neither high nor low. quite as real. whales. sea animals may be divided into eury thermal and stenothermal forms. for instance all the warm-blooded ones. Nowhere.). which is characteristic of the shore. We find. whereby they can adjust themselves to changes of temperature. the mean low (about 28° F. and is the mean temperature is is high (about 80° is F.

the temperate zone must form a barrier between the animals of the two other zones. Something must now be said as to the characters of shore animals. and to that account some more general statements may be added. each one seeks out a special spawningground. having its characteristic depth. and Hjort states that of the seventeen species of the cod family found in the North and Norwegian Seas. for though many show a perfect mastery of the water. Of the marine mammals we must regard all the seals and the walrus as littoral forms. at least at certain seasons. and salinity. outside the Arctic area. as sudden changes are always more dangerous than gradual ones. and more or less prevent their mixing. and thus are dependent on a substratum. which occur in the north. temperature. though this perhaps affects pelagic animals more than littoral ones. In addition to the seals already mentioned. Further. a fact which may be partly due to greater susceptibility of the larvae than of the adults to minor variations in pressure. and salinity.ANIMAL LIFE IN THE SEA littoral 161 animals of temperate latitudes must be eury- thermal. we have the true fur-seals (Otaria). temperature. Variations in the salinity of the water seem to have similar effects. 1404 and the many rossi). weddelli). while those of tropical and of polar seas may be stenothermal. seals of the Far South. In the chapter on the tundra we mentioned some of the common forms to be found in Arctic regions. all come on shore for breeding purposes. As the actual temperature in the case of many stenothermal animals seems to matter less than its constancy. such as Weddell's seal {Leptonychotes Ross's seal {Ommatophoca the crab- L . the contact of a cold current with a warm one seems often to lead to a great destruction of animal life.

swimmers. while the whalebone whales haunt relatively shallow water in high latitudes. and are pelagic.162 eating seal THE DISTRIBUTION OF {Lobodon carcinophaga). of ice-pack. two orders of mammals whose members are entirely aquatic. which ninus). because of the abundance of small marine organisms there. and are still bound to There are. which are more frequent in Since polar seas than elsewhere. and the manatee (Manatus) which lives in the estuaries of the great rivers M^hich flow into the . ever. howterra firma at the breeding season. the last named being a fur-bearing animal found off rocky coasts in the North Pacific. their young being born under water. are all carnivores which have taken recently to the aquatic life. The fact that aU these different kinds of seals tend to occur in cold regions cannot be ascribed wholly to deliberate choice on their part. and the animals seek out places as free as from molestation. the sea-leopard all occur in the region fur-seal [Otaria jubata) occur {Stenorhynchus leptonyx). or many species are being driven further and further back into the inaccessible parts of the polar have been virtually exterminated.Antarctic seas as well as elsewhere. the dugong (Halicore) found in the Pacific and Indian Oceans. and extending southwards to the shores of Australia. ' The Sirenia include only two living animals. man joined their enemies seas. Their only connexion with shore is that the greater abundance of fish in shallow water brings many of them to the vicinity of submarine banks '. while the sea-elephant {Macrorhinus leo- and the southern in sub. for the possible young of all are born on the land. The seals. and sea-otter. and the Sirenia The former are powerful or sea-cows. These are the Cetacea or whales and dolphins. walrus.

{Photo by Prof. haunt the smooth ground at the back of the rookery.) Fig.of Copper Island. Thompson. The shore The seals prefer the . unable to obtain a foothold there. rocky portion of the beach close to the water. D'Arey W. in the Bering Sea. and only the weaker forms. 41. showing a Rookery of Fur-seals.

O ^ £«: ^ .

puffins. Quite comparable in habits. and are also used in pushing food into the mouth. gannets. and could therefore find no food in the open sea. Others. though pelagic for much of the year. which present the same features as the seals in that. make it a perfect paradise for the penguins. Of the littoral birds perhaps the most interesting are the flightless penguins. though not related. which was similarly incapable of flight. auks. and having a perfect mastery of the water. and have been described by all the Antarctic expeditions (see Fig. 44). and fell an easy victim to man during the breeding-season. breed on the shore.ANIMAL LIFE IN THE SEA tropical part of the Atlantic. The hind-legs are entirely absent. who resort to its rocks and beaches in millions to breed. The animals are not nearly such good swimmers as the Cetacea. which are suckled above water. but the fore-limbs may touch the ground as the animals swim leisurely along the bottom. was the extinct great auk (Alca impennis) of the shores of the North Atlantic. and are helpless outside it. guillemots. apparently takes a large variety of aquatic plants in addition to algae. for the purpose. like and so forth. choosing cold or inaccessible regions Of the various adaptations to such L 2 . 163 and in shallow bays or Though both spend their whole life in the water. it Of the other birds which feed in the sea and frequent it. yet both are definitely littoral. they must come on shore for breeding purposes. and their heavy bodies and massive bones fit them for life near the bottom. and in carrying the young. forsake at the little breeding-season for inland regions. some. The absence of any carnivorous land mammals in the Antarctic area. The manatee lagoons. like certain of the seagulls. and its inaccessibility. for they feed upon the large algae.



breeding-places which they show the


pointed egg, which does not

roll ofE

the rocks readily,

while the eggs themselves are frequently protectively

Fig. 43.

in the Firth of Forth.

Kittiwake Gulls nesting on basaltic pinnacles on Majj^Island, {Photo by Mr. George' West.)

coloured, except where, like those of the puffin, they

are concealed in holes.

Of the

reptiles relatively

few are marine animals,

the habit being most frequent


the turtles and



tortoises (Chelonia).


The very curious leathery
apparently found in

(Sphargis coriacea)




it is

either very rare or its habits



infrequently seen and caught.

It is a large animal,

sometimes measuring six and a half feet long, and feeds exclusively on fish, molluscs, and Crustacea. The limbs are turned into paddles, the anterior ones being very long, and the animal is apparently pelagic in habit and a swift swimmer, though it is tied to the land by the need The eggs are laid on sandy shores, of breeding there. especially on the eastern side of tropical America.

The green

or edible turtle of

commerce {Chelone
it is

also marine,

but the fact that


upon algae and other aquatic

plants, binds

though the animals are good swimmers. As usual the eggs are laid on sandy beaches. Another species of the same genus is the hawksbill turtle {Ch. imbricata), which supplies the tortoiseshell of commerce, and is carnivorous, feeding upon fish and molluscs. Both these forms have a wide distribution
closely to the
seas, while the loggerhead {TTialassochelys into the Mediterranean and Bay of extends caretta) Biscay. This is a carnivorous form, and occurs hundreds of miles from land, floating on the surface. Again, however, it must approach the land at the breeding-




We have already spoken of the marine lizard of the Galapagos Islands (p. 149), and there are also seasnakes (Hydrophis) similarly adapted for life in the sea. These animals, which are especially found off the coasts of the Indian Ocean, have compressed swimmingtails,

and feed upon



in the case of so


forms which have secondarily returned to the aquatic life, their dependence on the land is especially shown



at the breeding-season, for the gravid female must come on shore to produce her young. For this purpose she visits the shores of low islands, where she may be found

up among the



fact that amphibians

through which
intolerant of its

have unprotected skins, can pass very easily, makes them presence, and we have thus no marine

Littoral fishes are many, and range from forms like the lumpsucker, the gobies, the sea-horses, the pipefish, and so forth, which inhabit rocky pools within or close to tide marks, through forms hke plaice, dabs, turbot, skate, fishing frogs, &c., which haunt the sandy bottom, their shape fitting them for Hfe here, to forms Hke cod and haddock, whose chief adaptation to littoral life is that they are ground feeders, depending upon animals like hermit crabs, marine worms, shellfish, &c., which only occur on the bottom. It is a curious illustration of what has been already said as to the tendency to change from the pelagic to the Httoral habitat during the life-history, that the httoral cod and haddock should have floating or pelagic eggs, and should in their early hfe haunt the
surface, often seeking shelter within the bells of the

life is



the herring, which in


a pelagic

feeding on free-swimming Crus-

tacea, has

heavy or demersal

which sink to the



tunicates or sea-squirts mostly attach them-

have pelagic free-swimming larvae. Of the molluscs a vast number are littoral forms. Many cuttles cower down on the sea-bottom waiting for their prey, or clamber over the rocks by the aid of
selves to the sea-bottom, but

however. or they are attached or tuft of silky threads like the by a byssus common mussel. to forms which can only walk like the crabs. Indeed. not only have most of them pelagic larvae. Curiously enough. the Friendly Islands. others. The most striking example of this is the palalo worm of various islands in the Pacific. which off Fiji. many very interesting adaptations to their relatively sedentary The majority relatively of the annelid worms are httoral. simul- taneously in thousands of worms. we can trace among the shore Crustacea the gradual acquirement of the ground-haunting habit. are adapted for life on the bottom. This worm Uves among the coral reefs. such as periwinkles.ANIMAL LIFE IN THE SEA their sucker-bearing arms. 167 The flat creeping surface which forms the foot in most gastropods. whelks. and elsewhere. but at certain seasons a portion of the body containing the sexual is elements Hberated. these showing hfe. however. The liberation of these sexual portions takes place at night. have pelagic larvae. or in sand like cockles. like the copepods or water-fleas. so that the whole surface of the water becomes filled with the animals. . hke crabs. like Mya and Lutraria. leading of swift by various stages from a form with the power swimming like the lobster. often live buried in the mud. few being adapted to the pelagic Hfe. before death occurs. indicates the dependence The bivalves of these animals upon a substratum. Many of these forms. and so forth. or they bury themselves in the substance of the rocks Hke Pholas and Saxicava. but certain forms become temporarily pelagic at the breeding-season. and leads a brief free-living life at the surface. While many of the Crustacea are pelagic. and various forms of prawns.

a common feature of shore animals of simple structure. and thus finds protection from its tions to shore hfe. Very many of these annelids and it is their abundance in the shore waters that attracts so many are greatly prized as food fish there. and manifest obvious adaptations to this mode of hfe. Starfish and brittle-stars similarly clamber over the bottom. and so forth. which possesses symbiotic algae. . A similar process British annehds. which builds up tubes from the sand. which lives inside the shell inhabited by a hermit crab. can only name the sea-mouse Hves buried in mud Serpula. which secretes Hme from the sea water and fashions a tube. is in the coral reefs regenerlater the process and a year repeated. Perhaps the most interesting forms are. a pretty example being the turbellarian called Convoluta. while the holothurians either live buried in the sand or attach themselves by their tube feet. the Coelentera. or else chmb about the rocks by means of their suctorial tube feet. fishes. and implants them deeply in the sand so that it is safe from the wash of the waves a Nereis. The sea-urchins burrow in the sand or mud. The fronds of the shore . enemies. however. Not a few unsegmented worms are littoral. We (Aphrodite). and is is suggested in certain a device to ensure that the eggs are carried to a distance from the parent. by Very many of the echinoderms also are httoral. which often show a double adaptation to httoral and to pelagic life. The littoral annelids show many beautiful adapta.168 are eaten THE DISTRIBUTION OF by the natives as food. The portion of the worm which remains behind ates the missing segments. which it attaches to rocks or to the hard parts of other animals or to weed Terebella. which .

sea-pens. like Others are pelagic remain permanently larvae which have never grown up. But there are no very great obvious adaptations to littoral life. — occurrence among the however. No swimming-bell floats away from the coral as it does from the sea-fir colony. throughout. A number occur in shallow water. For example. while still others attached to the rocks. lie loosely on the surface of . Those delicate bells which we call Ctenophora. save in and are fixed save in those very early stages. which stratum. sea-anemones. many of the corals of seas.ANIMAL LIFE IN THE SEA the sea-firs. great jellyfish are shore-born. and so forth. 166). ' ' fir special cups may be These buds drop out and float away. while some abyssal forms the ooze. tunicates already mentioned The sponges. Between the two extremes all stages exist. On the other hand. colonial 169 algae are often covered with the dehcate tracery of coelenterates of sedentary habit. have no littoral stage at all. that frequently one of the two modes of life is emphasized at the expense of the other. all these being attached to some solid body. so that the fertilized egg may find a home far from the parent stock. the generations. spending a considerable part of their lives attached to rocks. becoming swimming-bells sexual forms which carry eggs and sperms to a distance. Somewhat similar conditions occur in the (p. save perhaps that shallow-water sponges seem to be always attached. warm their earliest stages. or comb-bearers. are all require the presence of a subeither littoral necessarily or abyssal. We find. is of common coelenterates. This. are littoral. but are pelagic throughout their life. and their allies. which the biologist calls alternation of Among the branches of the seen in which buds form.

but present no special adapta- here. e. in the Norwegian Sea. saHnity. in the Red Sea and in the Mediterranean. which cause the surface waters to move in great whirls.170 THE DISTRIBUTION OF are as well represented on the shore as The protozoa tions to (2) life in other parts of the sea. the ocean currents. . as has been already suggested. is currents are very important. Now. where there no substratum. but where many at least of the organisms are fully exposed to the action of light. and so on. amount down from the sun upon the the difference in the rise to differences in of heat well known. it occurs also off the eastern coast of North America. in contrast to the littoral area. which streams surface of the sea gives temperature and density. and there is a junction layer where a sudden change in temperature. the salinity is greatest in warm regions exposed to the drying trades. Such a meeting of warm and cold currents is found. is We come next to the pelagic region. and therefore to we have also differences of Roughly speaking. where the cold Benguela current meets the warm Agulhas one. exposed to different types of winds. and least towards the poles. for instance. When this occurs the warmer water floats on the surface of the colder. such regions . and specific gravity occurs. As the surface in the different latitudes is differential evaporation. tides On As the other hand. With regard to the movements of the water here. where the Polar drift meets the warm currents of that sea . where the Labrador current meets the Gulf Stream it occurs off the Cape of Good Hope. salinity. we must notice first that. are of Kttle importance. In not a few parts of the globe we find that currents bringing cold water of relatively low salinity meet warm currents of highly saline water.g.

they can vary their specific gravity in harmony with — — variations in the specific gravity of the water. and Dr. By a series of very ingenious investigations he has shown that. not only does such a junction layer between the surface and lower layers of water occur. and helps to account for the abundance of fish. light water is floating on cold. But there is something more than this. that such an accumulation of pelagic organisms in an area of sudden change of density occurs in the power of ' — vicinity of aU the great it fishing regions of the world. Very many possess certain powers of adjustment that is. in many parts of the ocean. They must be able to float without effort that is. The delicate organisms which float in the surface waters of the sea are necessarily very sensitive to changes of density. which thus forms what Dr. following this line of change of density. Hjort has shown that many of the littoral fish at certain seasons leave the shore. When warm. their specific gravity must be that of the water in which they float. the . a region where living and dead plankton animals accumulate. where food accumulates where it is as abundant as it is There seems some reason to believe near the land. but that this layer is a region of great wealth of life.ANIMAL LIFE IN THE SEA are regions of 171 pelagic great mortality among the organisms. it is apparently impossible for the more dehcate forms at least to pass through the junction layer of the two. prolongs the plenty of the shore out into the open. for the stenothermal forms are Idlled by the sudden changes of temperature which occur. It is a false shore-line in mid-ocean. as were. and thus. and swim out into the open ocean. As to the actual temperature of ocean water. Hjort calls a false bottom '. dense water. this But adjustment has its limits.

lat. and here At 900 fathoms they remain we reigns. but another point has also to be considered. however. must suppose that absolute darkness These statements are true for middle latitudes. . In practically all parts of the ocean the temperature at depths exceeding 1. Obviously. so that we — — of life in the must add to the above consideration of the conditions open some account of the pelagic algae. at about 270 fathoms depth. In 50° N. we shall see directl3% the intensity of light greatly influences the bathymetrical The basal food-supply open as off depth) distribution of marine animals. and varies but little throughout of littoral waters hold — — the year. at about 430 fathoms depth. for photographic plates are slightly affected here. Dr.172 THE DISTRIBUTION OF made above in regard to the temperature statements good here also. lat. and therefore they will penetrate to different depths in different latitudes. because. as (i. at about 100 fathoms depth. An important point. absolutely unaffected after long exposure. is that nowhere not even under the Equator does the warmth of the surface layer extend far downwards. Some of the violet rays tion. is : This is important. At 300 fathoms the red and green rays have disappeared. are stiU present at 500 fathoms. The conditions as to light demand detailed consideraVery careful experiments show that all raj's of light are present at depths of 50 fathoms. the direction of the rays is not the same in all latitudes. In 67° N.e. lat. in the of marine animals the shore consists of algae.000 fathoms is permanently low. Hjort gives the following in- figures in regard to the depths at tensity of light which the same found in different latitudes In 33° N. but blue and violet rays remain.

The next point to be considered is the distribution of pelagic animals.ANIMAL LIFE IN THE SEA 111 173 the first place it is noticeable that the larger algae are unrepresented in the open sea.' and there is reason to believe that then the fish of the shore may follow them. The dead forms. swarms of the weed is. as light- demanding organisms. as we have already suggested. Those found in greater depths are resting stages or dead forms. though in diminished quantity. are limited to the surface layers of the water. but. There is. currents may carry the shore forms out to the ocean. They occur. water of the open ocean seems to be poorer in algae than coastal waters. and can live exception. There is a tendency among many writers. The basal food-supply float at the surface. at least temporarily. animals which live among the floating masses of weed in the Sargasso Sea show what an important addition to the food-supply of the ocean there for a time. no real justification for this use. however. the phytoplankton. therefore. for this weed is torn off the shores. to confine the term pelagic to those forms which live sufficiently near the surface for light to penetrate. the minute algae which Necessarily these algae. as has been already suggested. and though it proliferates vegetatively in the open. at least tacitly. being especially abundant in the uppermost 25 fathoms. tend to accumulate. the increase of density occurs. An animal living at a depth of 2. but it is The it is not a truly pelagic form. especially off the Bahamas. of the pelagic animals is. floats at the surface of the Sargasso The weed which Sea is an apparent apparent only. at levels where a sudden As regards quantity. down to 50 fathoms.000 fathoms is as truly pelagic as one living at ' ' .

beginning with the plankton. taken in conjunction with some previous observations. We must now consider some of the characters of pelagic organisms. and such forms as the sunfish. or were merely caught as the net a depth of say 2. were abyssal fish. large eyes or no eyes. A sponge with an attaching stalk. But if a net which has been dragged along the bottom at when examined. feeding on the are great.000 fathoms proves. bodies. and have usually flattened and expanded Sometimes. found about the light limit. believes that the recent (1910) expedition of the Michael Sars. for it was attached to the bottom. Plank- ton animals are those able to float in water udthout exertion. however. obviously. . most abundant in the upper 100 fathoms or so (2) mesopelagic or he is right. then pelagic animals may be divided into at least three groups (1) surface forms. some protozoa. intermediate forms. (3) bathypelagic forms. if it can be proved that neither has any dependence on the proximity of a substratum. and having a bizarre form and colouring. as in the very abundant copepods. taken in deep water. of which they . which varies with the latitude occur below the limit to trate. oil makes the body hght. how can we tell whether these fish were truly abyssal forms. has yielded evidence which proves that many of these curious fish If are truly pelagic. Hjort. bottom.174 THE DISTRIBUTION OF a depth of 50 fathoms. which which the rays of light penebut yet not upon the sea-floor. was hauled up ? The difficulties in coming to a decision and until recently there has been a tendency to assume that all fish obtained in deep water. Dr. But here. and so on. is obviously an abyssal form. to contain fish. Hving at or below the light limit. are independent. a globule of as in At other times. a difficulty occurs. : .

different kinds of gasteropods are adapted to life in the open sea. surface jellyfish are blue or violet or almost colour- while in deep water bright-red forms occur. so that it is invisible as it floats in the clear water. there are enormous numbers of examples. Surface pelagic forms are so numerous that no useful purpose would be served by giving a list. the sea-butterflies (Pteropods) being good Some other less familiar groups are also unrepresented in the open sea. a pelagic tunicate. curiously modified gasteropod in Salpa. With one exception. appear in Sagitta. way in which they appear in unThus while the shore annelids are is brightly coloured. so it will equally be invisible when seen from below against the sky. a The same peculiari- ties worm of uncertain affinities in Phyllirhoe. a . but it may be interesting to note those groups which have no pelagic representatives. less. Tomopteris. in some pelagic Crustacea. though their larvae are often pelagic. colour- and has an expanded body. We see this in the protozoon called Noctiluca. 175 is due to the presence of air or another In the surface forms the body is often transparent and that colourless. in the beautiful tunicate called Pyrosoma.). very many types of Coelentera are of the pelagic there are no pelagic alcyonarians (sea-pens. and shell-less and so on. &c. Many forms of the surface are phosphorescent. and there are no pelagic sponges. On the other hand. As young animals have often delicate . though many pelagic. There are no pelagic bivalve molluscs.ANIMAL LIFE IN THE SEA the lightness gas. While. there are no pelagic echinoderms. as already suggested. is That these peculiarities are adaptive suggested by the related forms. while in other cases it is blue or violet. to be considered later. a pelagic form. The less. pelagic larvae. and so forth.

Wings are completely absent. and the body is covered with a greyish pubescence. in the open sea a number of powerful swimmers (nekton) able to make headway against the currents. but was similarly for long regarded as an independent organism. because its relation to the parent was for long unknown. The insects appear to be perfectly pelagic in that even the eggs are produced in the open. The related bottle-nose (Hyperoodon). and is found especially in the North Atlantic. for what reason is not known. and the young have been found on floating soHd bodies in the sea. Some fifteen species of Halobates are known from the warmer parts of the oceans. but they may also approach shores. Thus the huge cachalot or sperm whale {Physeter macrocephalus) is found especially in warm seas.176 THE DISTRIBUTION OF translucent bodies it would seem as though they could be very readily adapted to the pelagic life. called Leptocephalus. which also lives on cuttles. The animals breathe air. related to the forms which skim over the surface of fresh-water ponds. The whalebone whales of these are the . occurs in colder seas. where it finds the large cuttles upon which it feeds. whales and their whose distribution is tied to that of the animals upon which they prey. and belong rather to the surface of the ocean than to its waters. As two interesting forms mention may be made of the transparent larva of the eel. which shines in the sunlight. The most important allies. we have In contrast with these plankton or drifting forms. a kind of bug. Very remarkable is the presence in the open sea of the members of an insect genus. which is the larva of the rock lobster. and the glass crab (Phyllosoma). Halobates. The female carries the eggs for a time attached to her own body.

Provisionally. the bonito {Thynnus pelamys). &c. which is also pelagic.. the fish-eating dolphins and porpoises especially frequent shallow waters. though widely distributed. but varying M .) feed on the floating plankton. however. All these forms. except when some floating wreckage. the forms which occur between the surface layers of water and the bottom. related to the mackerel. the beautiful dolphin (Coryphaena). however. near the light limit.ANIMAL LIFE IN THE SEA on sea-butterflies (Pteropods). found practically no surface fish. at least in the surface is apparently that food near the not sufficiently abundant. the following statements may be made. while cruising in mid. The above forms may be regarded as characteristic of the upper 100 fathoms or so of water. waters. surface. &c. especially and are very common and cold waters. The reason surface is with a burden Crustacea. the Prince of Monaco's yacht. these being based especially on the results obtained by the Michael Sars surface forms may — expedition. involving the use of comphcated methods not hitherto used at very great depths. purely pelagic in cool fish are not very abundant. another flying form. With the exceptions already given. despite their obvious adaptations to pelagic life. where the fish on which they prey are most frequent. 177 (Balaena. such as drew them to the ' ' Among the be noted the flying-fish (Exocoetus). are more abundant as the land is approached than far away from land. and its allies. The study of these is necessarily a difficult matter. At depths between 200 1404 to 300 fathoms. of attached invertebrates. On the other hand. Professor Bouvier notes that the Princesse Alice.Atlantic. We have next to consider the mesoplankton and mesonekton that is.

178 THE DISTRIBUTION OF with the latitude for the reasons aheady discussed. of red deeper. Sehastes norvegicus being an example of the former and Goniostoma elongatum of the latter. occur forms in which the colouring is black or red. the only pelagic echino- derm (Pelagothuria). A very curious point about it . of which the fish have been chiefly investigated. environment where a considerable amount of sunhght penetrates. which fives in the Atlantic at depths of somewhere about 200 fathoms at night. Large which perhaps catch the last rays of sunhght. and live in the same depths. It has huge jaws. when the fight conditions are the same near the surface as in the depths where it habitually dweUs. a common feature in these deep-water fish. especially blue and violet rays. The last-named fish is purpfish-black. and doubtless other forms also The fish hving at these depths inhabit an occur. In the absence of any red rays both these colours must look the same. at depths of from 300 to 500 fathoms. They have often large eyes. though medusae. light organs are present. Another form with the same colouring and characters is Cydothone signata. bright -red prawns. and red or black fish. such as Acanthephyra muUispina. lives at a Still depth of about 300 fathoms. Its young have the same colouring as their parents. . and either will protect the animal. which sometimes protrude at the end and are then called telescopic. this fish has been found within some 80 fathoms of the surface. with small fight organs and small eyes. An example is the silvery fish Argyropelecus affinis. At these depths occur deep -red or chocolate-coloured medusae. where aU trace and green rays have completely disappeared. which of short stalks. there seems to be a fauna of silvery or iridescent animals.



that in the daytime
it rises

it lives

at about 300 fathoms,

but at night

to within 80

fathoms of the surface.

young are

crystal clear,

and are surface forms

upper 80 fathoms. On the other hand, some members of the genus Alepoeephalus live as far down Their as 1,100 fathoms, and have no light organs.
in the

young are the same black colour as the parents, and
live also in the great depths.

To sum

up, so far as

we know

at present, forms

live in the surface

waters of the ocean are crystal

clear or pale violet or blue. Those which live in layers where only blue and violet rays penetrate freely tend to be silvery and to have large eyes and large light organs. Still deeper, where no red and no green rays, even in some instances no rays at all, penetrate, the colour becomes red or black, and light organs are small or disappear. That there is an intimate relation between the characters and colouring of the organism and the


of light present is

shown by the

fact that


the larva inhabits the same depth as the parent
the same colours.


has inhabits another zone it

takes on the normal colouring of the inhabitants of

Further, the fact that the forms from deeper water occur in different layers of water in difEerent latitudes the level corresponding to the degree of penetration of light in the particular latitude, so that the animals live higher up in high latitudes

that zone.

than in low

—suggests that



of the animals is

strongly influenced

by the amount

of light present.

One other point
of these depths.

interesting in regard to the

pelagic larvae out to the open sea, so

Just as the littoral fishes often send some of the

abyssal fishes seem to have bathypelagic larvae,

larvae living in the open water, but at considerable




Thus the members
of the

genus Macrurus,

believed to be true abyssal, that

ground-liAnng, forms,

have apparently such eggs and larvae. The abyssal hermit crabs, also, have bathypelagic larvae, just as the Uttoral forms have larvae which form part of the
surface plankton.
(3) The abyssal region of the oceans is contrasted with both the other regions in that its waters must be permanently calm. It is in harmony with this calm that we find, as already stated, some deep-sea sponges which have no attachment to the substratum, upon which they seem to lie loosely. The pressure must be enormous, for the animals of the depths support a weight of several miles of sea-water upon their bodies. Though there are no plants, oxygen seems to be abundant. But this is a consequence of the welldeveloped circulation of the great oceans, and is not a feature of enclosed seas. Thus the Black Sea has no animals in its depths, and the waters there are impregnated with sulphuretted hydrogen, and contain but

In the Mediterranean the conditions are Light is scanty. The water is, of course, entirely absent in the depths. is permanently cold. The basal food-supply must consist of the dead animals of the upper zone, which In fall downwards through the water as they die. consequence we find that many of the abyssal forms



extreme, but the deep-sea fauna



To what extent

the floor

of the

remains a difiicult question. ocean As aheady noted, Dr. Hjort beheves that genuine abyssal fish, in the sense of grovmd forms, are few, most of the so-caUed abyssal forms being bathypelagic and having been obtained in the process of hauling in the dredge. But there can be no doubt that many

depths, especially over regions covered


invertebrates do occur on the sea-floor in the greatest

by Globigerina and echinoderms preponderate, molluscs and crabs among Crus-


these invertebrates sponges

Of the sponges the old-fashioned forms are numerous, and calcareous sponges are absent. A few corals occur. Among the echinoderms all the classes are represented, and the oldtacea being rare.

fashioned crinoids, rare in shallow water, are relatively



the sea-urchins the special feature

the abundance of old-fashioned irregular forms, which

Annehds occur especiIn harmony with the great uniformity of the conditions over wide areas, the abyssal animals are widely distributed. Further, as the conditions of temperature, and during the long Arctic night the conditions regarding illumination, show resemblance in polar and abyssal regions, there is apparently some resemblance between polar animals
occur so abundantly as ally in the red clay.

and abyssal animals.
References. The forty-two volumes of the Challenger' Reports ( 1880'

More 91) give the results of the epoch-making cruise of that vessel. popular accounts of its work are to be found in Mosely, Notes by a Naturalon the Challenger (London, 2nd ed., 1892), and Wyville Thomson, The Depths of the Sea (London, 1873), and Narrative of the Voyage of the ''Challenger'' (London, 1885). The following may also be consulted: Ortmann, Grundzuge der marinen Tiergeographie (Jena, 1896) Chun, Aus den Tiefen des Weltmeeres (Jena, 1900) Marshall, Die Tiefsee und ihr Leben. For the more recent work discussed in the present chapter, see Hjort, The Michael Sars North Atlantic Deep-sea Expedition {Geographical Journal, April and May, 1911); Hjort, Die erste Nordmeerfahrt des norwegischen Fischdampfers Michael Sars im Jahre 1900 (Petennann's Mitteilungen, IV, 1901), and The Deptlis of the Ocean, by Sir John Murray and Dr. Johann Hjort (London, 1912). Two interesting popular articles on the work of the Princesse Alice are Les Vertibris de la Surface and Im Faune Pilagique des Invertebres, by Professor Bouvier
' ' ;






{Revue Ginirale des Sciences, 1906).

The water
from that
of salt.
of rivers


of very


lakes differs

of the sea in the absence of

any considerable

quantity of




water contain a special fauna, contrasted with the fauna of the ocean, may which, as but in addition to this be described as impoverished negative character, this fauna possesses also some Certain positive ones, which we shall consider later. other lakes, either because they had a former connexion with the sea, or because they have no outlet, contain saline water. Such masses of water maj^ contain some animals with distinctly marine affinities, e. g. in the Caspian are found a seal and a kind of herring. But the Caspian is far from being as salt as the sea, and in addition to animals of marine type it contains some typically fresh- water fishes. On the other hand, masses of very salt water, such as the Great Salt Lake of Utah, contain pecuhar brine-shrimps (Artemia) not found in the sea. A still further compUcation is introduced by the fact that certain lakes, though their waters are perfectly fresh, contain animals of distinctly marine Thus Lake Baikal, though its waters are not facies. salt, lodges a seal aUied to the Caspian seal, and a Lake Tanganyika, in Central Africa, marine worm contains several animals of marine type, and so on. In consequence we cannot sharply separate freshwater animals from marine ones, in the sense of imply-

Such bodies






ing that


all the latter live only in the ocean, and that and rivers contain only the former nor can we assume that all aquatic animals can be divided into marine and fresh-water types, for this would exclude



the brine-shrimps,



it is

and brine

But, at the same time,

convenient to recognize a general distinction between the animals of the oceans, the marine forms, and those

and rivers, which are for the most part fresh- water animals. Using the terms in this general sense, we may note that the poverty of the fresh-water fauna is due to the
characteristic of lakes
relatively unfavourable conditions.


these un-

favourable conditions

we probably need not


the absence of considerable amounts of salt dissolved
in the water, for the presence of

an appreciable amount

of salt, e.g. in the Caspian, does not give rise to a


increase in the fauna as compared with freshwater lakes, and the presence of a large amount of salt, e.g. in the Great Salt Lake, is distinctly inimical to animal life. Before proceeding to discuss the especially unfavourable conditions, we may note that it is generally believed that life originated in the sea, not in fresh water, and that therefore all the animals now found in rivers and lakes must have originated in one of two ways (1) they may have been derived directly from marine forms, or (2) they may have been derived from land animals which have reacquired the aquatic habit of their remote ancestors. Of the first group, fresh-water fishes may be taken as examples. They have certainly had marine ancestors, and in some cases, e.g. the eel and the salmon, the existing forms have retained the power of living in

to consider the groups life But when we come which are richly represented in the sea. the . and the — The . the water-spider. true spiders. though they live in water. No cuttle hunts its prey in the depths of the great lakes. respiratory organs when ters. nor creeps along river bottoms. . in fresh Most of the higher Crustacea hve in the sea. The otter which haunts the salmon rivers has lungs no less than its distant relative the polecat the water-tortoises have lungs Hke the land forms the water-beetles and the many insect larvae found in pools. 176). and their peculiar would be inexplicable if we could not assume that their immediate ancestors Hved on land. the water-mites. most amphibians have a fresh-water and some pass much of their life in water. no (cf. seaurchin. Similarly. brittle-star. It is otherwise with most of those numerous forms which. echinoderms are exclusively marine no starfish. no on. are yet adapted for breathing air. and so larval stage. and the few crayfish and prawns which live water are certainly descended from ancestors which Uved in the open ocean. are all air-breathers. the bivalve the sea as molluscs of the lakes and streams have certainly had marine ancestors. the tale is different. which has hardly any insects air-breathing moUuscs. The polychaete worms are almost exclusively marine.184 THE ANIMALS OF LAKES AND RIVERS weU as in fresh water. their far-off ancestors acquired terrestrial charac- Therefore in possessing some of these forms the fresh-water fauna is not poorer but richer than the sea. no hving amphibian is ever found at any period of its Again. nor sea-lily is known from fresh water. p. while in the sea. In these cases there is no reason to beheve that the animals have ever sought the sea since the time pond-snails.

though they include some peculiar forms. where also there are but few sponges. This fact.THE ANIMALS OF LAKES AND RIVERS higher Crustacea largely 185 Coelentera so. and be represented by a series of stagnant pools. The great group of the very scantily represented in fresh water. a vertical circulation is produced which carries down oxygen to the deeper layers of the water. This makes Ufe in the deeper layers possible. as the ice melts in the former case.water fish also are few in number as compared with their marine is allies. and before it is formed in the latter. the rivers may periodically cease to flow. and the however. pike. where the surfacewater periodically cools below the point of maximum density of water. Fresh. like the ganoids sturgeon. lakes In the general case this applies.g. and at least one ganoid (Polyp terus) have accessory breathing organs which enable them to breathe air. as in many parts of the and inland globe. as well as gills. taken in conjunction with the relatively large numbers of air-breathing dipnoi or double-breathers. In the summer the heating of the surfacerise to a stratification into two layers. to seas rather than to rivers.). (e. interesting to note that of these pecuUar fish all the three living dipnoi. when the warm oxygenated surface-water lies upon a water gives . of course. suggests that one difficulty in colonizing fresh water has always been its relative deficiency in oxygen. But where a marked dry season occurs. &c. whose water is oxygenated by its motion. invertebrates. In the lakes of temperate regions. bony It is. though some American experi- — layers occurs only in spring ments suggest that the oxygenation of these deeper and autumn. whose waters become very fetid conditions highly unfavourable to animal life. at least at certain seasons.

which would form a deposit on the surface of the breathing organs of animals Hving in them. the deep-water a deep-water fauna. This relative paucity of oxygen naturally checks life in the deeper parts of the lake. whose oxygen and transporting agents. Rivers are powerful eroding deeper. then. because of the absence of a well-marked circulation of the water. and greatly retard respiration. increases towards the Equator. The second unfavourable condition in fresh water is its frequent turbidity. Now if an increase in the amount of mud in the water kills estuarine animals. There is much evidence to show that its waters are becoming muddier year by year. which are naturally tolerant of some suspended matter. lakes are great the course of rivers of . and leads to a virtual absence of In brief. The Firth of Forth is a great estuary swept by the tides. fauna of lakes is always poor. filter-beds on necessarily. once inhabited by great numbers of marine animals tolerant of estuarine conditions. colder layer. the waters both must often contain fine particles in suspension. and the deficiency in the amount of oxygen. therefore. apparently because drainage of the mosses in the upper regions of the river has made the run-off there more rapid. and that there is no spring melting of ice. even a small amount of suspended matter will prevent the ' ' — . In tropical lakes the fact that the surface-water does not cool in the same way in autumn. with the resultant paucity of life. notably by oysters. As a result of the increasing muddiness many animals have disappeared the oyster-beds long ago ceased to be productive. and other forms also are diminishing. The importance of this may be illustrated by an example.186 THE ANIMALS OF LAKES AND RIVERS is only renewed very slowly. makes the vertical circulation much less marked.

160). which are almost permanently frozen. may render the water unsuitable animals. the land is This statement not in- consistent with that already made. that the waste of an important part of the food of the seais a prime necessity which precedes even the need of food. and must be a great bar to the passage of marine animals up estuAlmost any coast-Une will illustrate aries and rivers. the only . This is an extreme but it suggests the adaptations necessary before animals can thrive in small masses of water. The shore collector knows that to get the rarer and more deUcate animals he must seek rocks or beaches remote from the mouths of streams. But it is not only the suspended matter in the water of lakes and rivers which is inimical to animal hfe. selves. of Victoria Land.THE ANIMALS OF LAKES AND RIVERS 187 growth of the more dehcate animals. and will yet hatch out if the temperature rises and the ice melts. however. case. p. or their offspring. Low temperatures are not.water forms must be eurythermal (cf. and few animals can take in oxygen if their respiratory organs are choked with mud. These variations are normally so great that they render the water absolutely unsuited All fresh. and temperate and polar forms must have special means of protecting them- to stenothermal animals. for oxygen to certain forms. The presence of large amounts of hme or manganese salts. the same thing on a smaller scale. or of humic acid. Still another point of great importance is the enor- mous variation in temperature to which lakes and rivers are subject. for at the immediate outlets of these only is the hardiest forms occur. against the cold of winter. the naturahsts of Sir Ernest Shackle ton's expedition found rotifers which can apparently In the ' lakes ' tolerate being frozen into the ice for years.

The amoebae in a waterspout can be dried to dust. but within considerable limits. so that thousands of tadpoles die because the drought comes before they are ready for their land life. One other unfavourable feature of rivers and lakes is found in the strength of their currents. high temperatures and low. and must also be provided against. the lung-fishes He in their mud cases. but the tadpole avoids the risk of drought by adjusting its time of metamorphosis to the time of drought. For example. with resultant desiccadangerous. are just as that where the problem of seasonal adaptation cannot be directly faced. and the spout once again becomes suitable for their active life. are risks which every fresh-water organism must face.188 THE ANIMALS OF LAKES AND RIVERS High temperatures. In the swamps of the Paraguay or of the Central African rivers. waiting till the wet season again permits them to become active. it may be avoided. such currents cause Httle harm to . Every summer one may find cases where the adjustment has not been sufficiently delicate. life in the temperate and frigid zones is markedly seasonal. and the Httle frogs return again to them to pass the winter in their mud. and with the exceptions already of the ocean are (p. given 161). so that it is ready to leave the water at the time when the pond would naturally dry up. In these. and it is interesting to note danger. those shallow bodies of water which are favoured by frogs as spawning-places are very apt to dry up in full summer. tion. as on dry land. and the lessons learnt in the conflict with them are stamped deep on all the inhabitants of stream and lake. In autumn the ponds fill again. The waters swayed also by currents. and will lie unharmed within their protective cysts until the rain comes. Drought and freezing.

whose purpose distribution along the shore. and are sometimes filled with water rendered very salt by evaporation. a number of marine forms occur. a kind of sea-fir (Hydromedusan). it is relatively easy for them to progress. is nearly fresh. to another to which it has no adaptations. showing that it is not its freshness alone which prevents such forms from colonizing fresh water. or free-swimming stages. 35). while at the rainy season the water In spite of these variations in salinity. Now sea-anemones produce usually free- Museum. connected with the sea. (1907) p. In the nature of things littoral animals must have always had special faciUties that is. including a seaanemone. and estuaries. and sometimes shut off from it. but the regions into which they are swept may be more suitable than those they have left in the case of the littoral animals the currents are the great distributors. facilities in regard to space ^for conquering fresh water. some artificial ponds at the mouth of the Ganges a very curious fauna has been described {Records Indian These ponds are sometimes i. Tides and currents sweep the members of the plankton hither and thither. in to the colonization of fresh water. from the shore up But we have already seen that one of the great peculiarities of Httoral animals that they tend to produce free-swimming young. it is to ensure and the existence of these free-swimming stages or young offers a serious obstacle For example.THE ANIMALS OF LAKES AND RIVERS 189 marine animals. for it means the passage from a speciaUzed environment to which the animal has become adapted through progressive variation. . So far as their — — — relations in space go. But for a lake or river animal to be swept out to sea must in the general case mean death. and a polychaete worm. rivers it is actively or passively.

it may be said. for these forms would not have a free-swimming stage.190 THE ANIMALS OF LAKES AND RIVERS called swimming larvae planulae . with rare exceptions. . though they avoid regions where the water is in rapid movement. hatch as miniature adults. the planulae and the trochospheres would tend to be swept out to sea by the stream. Uke land snails. which makes Hfe in ponds and streams easy for them. and so can be peopled afresh from it. This perhaps gives us a second reason why so many animals with a terrestrial ancestry have succeeded in colonizing fresh water. But. So real is this danger that. but if we suppose that at some future period the connexion with the sea were to be reduced to a swift-flowing stream. can resist moderate currents. fresh-water snails. all delicate forms incapable of swimming against currents. fresh-water organisms have no free-swimming stage in their hfe-history. Thus while most marine gasteropods have freeswimming larvae. or which managed somehow to survive until this free-Uving stage could be suppressed by progressive variation. how do the lake . like many insect larvae. Hydromedusae produce free-swimming medusoids polychaete worms have free-swimming larvae called trochospheres. powerful swimmers. for the medusoids. in a large lake the need for distribution is as great as in the sea. and there would be thus no young forms to replace the parents when death took place. Only those marine animals could colonize fresh water which had no plankton stage in their Hfe-history. could we suppose that the animals would persist ? Obviously not. These arguments of course only apply to those deUcate organisms which are inRelatively capable of swimming against currents. These particular ponds have at intervals a connexion with the sea.

She then discharges the young. In most of these cases the modifications are . remarkable are the conditions in the fresh-water mussel (Anodon). and so on. and develop into mussels like the parent. With this preliminary account of the conditions of Hfe in lakes and rivers. constitute the fauna. we may proceed to consider the animals which ing adults. These shelled eggs More are doubtless easily transported by currents. which may produce currents of unusual violence. Here the mother retains the young within her body till sticklebacks approach the mud in which she is living. and so distributed. . Thus we have many kinds of otters. the curious duck-mole or Ornithorhynchus of AustraHa. and are thus carried about for a time. such as the common water-shrew and the Russian desman various rodents. such as the beaver and the water-vole a monotreme. We may take two examples of familiar forms. In addition there are some curious special adaptations. being assisted by sudden freshets. and of the resultant adaptations. . which have webbed feet and feed upon fish not a few insectivores. or. . . and produce fresh hydrae in spring. capable of carry- more likely. who fix themselves to the fish. The fresh-water hydra as contrasted with the marine seafirs has no free-swimming stage. resting eggs to a distance. A considerable number of mammals show at least a partial adaptation to fresh -water life.THE ANIMALS OF LAKES AND RIVERS 191 animals succeed in spreading from one part of the lake to another if they have no free-swimming stage ? The answer is apparently that the distribution is largely passive. Later they drop from the stickleback to the mud. While the weather is warm it reproduces itself by buds towards autumn it produces coated eggs which survive the winter.

and are apparently primitive forms which. used in swimming. Brahmaputra. Of mammals with more profound adaptations to aquatic life. Amazon almost to their sources. Baikal seal also occurs in Lake Oron. and a broad flattened tail used as a rudder. This suggests that in most cases the modification is relatively recent. especially the Amazon. We have already spoken of the Lake Baikal seal {Phoca sibirica) and the Caspian seal {Phoca caspica). Among the more frequent modifications are webbed feet. as high up as the depth of the water permits. and in addition to its seal Lake Baikal shows certain marine features in its molluscan fauna. . not sufficient to cause the animals to diverge very markedly from their terrestrial alhes.192 THE ANIMALS OF LAKES AND RIVERS not very profound. direct or sea of Tertiary times. The explanation is perhaps that the lake had once ' ' a connexion. Other fresh-water dolphins occur in the large rivers of South America. both closely The related to the Ringed seal of northern waters. Among with the great inland the Sirenia the manatee largely fresh-water. but it has never {Platanista gangetica). ascending such rivers as the But these animals thrive equally well in the ocean. been seen in the sea. These dolphins feed upon freshwater fish and river prawns. on the course of the river Vitim. close dense fur which prevents the water reaching the skin and so chilhng it. is indirect. but The one which is most definitely adapted for this habitat is the blind dolphin of the Ganges. few occur in inland waters. This animal occurs from tidal waters to the upper reaches of the rivers. like Ganoid fish. Cetaceans are exclusively marine. and Indus large The certain dolphins occur in fresh water. large rivers requires and their presence in no special explanation. have found a refuge in fresh water.

ponds. breathing animal the modifications which fit an animal for life in the sea must necessarily be more profound than those which fit it for life in fresh water. abundance of food there. in the water. as tions to the aquatic among mammals. marine than in fresh-water forms. 1404 If . a form reaching a considerable size. which are but little modified. which usually occurs in relatively small masses. Among the Chelonia a considerable at least a great part of their time in the water. Such typically marine mammals as seals. fresh-water rodents like water-voles and beavers. have only slightly modified limbs. however. Marine turtles have their limbs turned into paddles fresh -water Chelonians swim by limbs which are only slightly modified as compared with terrestrial forms. by the relative live Such forms. Cetaceans. It has strongly webbed feet and is carnivorous. This is quite parallel to the conditions which occur in mammals. Such fresh-water carnivores as otters. 193 very large number of birds haunt the vicinity of especially at certain seasons. and rivers throughout the greater part of North America. being attracted to it. which is found in lakes. fresh-water in- hke water-shrews.THE ANIMALS OF LAKES AND RIVERS A fresh water. the adaptalife are much more marked in . and where sectivores the animal inhabitants often retain considerable depen- dence upon the land. As might be deduced from its limbs. and Sirenians have their limbs (when present) turned into paddles. on the surface rather than interesting to note that In regard to reptiles it is among the Chelonia. retaining many features found in terrestrial This is equivalent to saying that for an airtypes. number spend As two examples we may take the American Snapping Turtle {Chelydra serpentina).

194 it THE ANIMALS OF LAKES AND RIVERS leaves the water. except at the breeding-season. None to life of the snakes in fresh water as show such definite adaptations do the sea-snakes to life in the in search sea. is that the young live in the water and breathe by gills. while the adults live on land and breathe by lungs. which has also webbed feet. and migrate from one pond to another if the first shows signs of drying up. or in order to migrate from one region to another. especially true of the natrix). its life in which sometimes spends water. should want of food is or any other cause render this necessary. Throughout much of North America. But there are some interesting exceptions. in which it swims curious conditions life. in a way that would be impossible for a marine turtle. The amphibians show some very in connexion with adaptations to the aquatic The normal condition. which when adult is normally terrestrial. and is of a brownish colour. and capable of (awkward) progression on land. as already suggested. with yellow spots and . but certain forms enter the water freely This is of food. or to lay their eggs. These little creatures are fond of coming out of the water to bask. either to bask in the sun. The living crocodiles are still so far bound to the land that they leave the water voluntarily. which. seem to seek the water for the sake of the food to be obtained there. there occurs a salamander {Amhlystoma tigrinum). common grass- snake {Tropidonotus the greater part of easily. The same tale of partial adaptation to aquatic life can be told of the various kinds of crocodiles. like the forms just mentioned. and especially in Mexico. In Central and Southern Europe the ponds and swamps are inhabited by the pretty little pond tortoise {Emys enropea).

at times. never grow In up. The same statements may be made are of Siren lacertina. have large and a tail fringed with Now. This animal uses both its lungs and its gills for breathing purposes. these larvae as axolotls. This brief account of the air-breathing vertebrates found in lakes and rivers enables us to draw some general conclusions in regard to these. originated from fresh-water forms (see p. called Necturus maculatus. the mud-eel of parts of the United These examples of the reacquisition of the exclusively aquatic habit by animals whose ancestors had in water. which sometimes breed while retaining the characters of the tadpole. but for causes not clearly understood. known external slender limbs. become adapted for life both on land and Amphibians. as In a more advanced form the same condition occurs it were. which possess gills at some stage in their ' ' — : N 2 .THE ANIMALS OF LAKES AND RIVERS blotches. Obviously we (I) The can classify them under three headings amphibians. larvae. in certain other amphibia. conditions. swamps of the eastern part of the Missisand in some of the Canadian lakes. life. 195 The gills. That is. As a much rarer phenomenon this may occur in the common newts. and while retaining larval characters. with three pairs of external gills and a fringed tail. there occurs an amphibian of about one foot in length. of the Amblystoma eggs laid in a lake some may give rise to the salamander-like adult. 202). it may be noted. have probably been derived from fishes of the ganoid type that is. while remaining in the water. while others may breed while retaining tadpole characters. the lakes and sippi basin. but remain permanently in the tadpole stage. under natural may become sexually mature and lay eggs. which. a swimming membrane. and retains its gills throughout States.

but driven from the sea by the competition of more advanced forms. or because .. life is safer there for relatively helpless animals (3) the fresh. the fresh. ters. are of more ancient origin. on the other hand. the freshwater mussels have had time to become greatly modified.water crayfish. water-tortoise and crocodile among reptiles. The fauna of swift rivers must in the general case consist only of powerful swimmers like fish and freshwater crayfish. more or less accidentally cut off from their natural habitat. because enriched by many forms. The .water hydra. The seals found in Lake Baikal and the Caspian Sea are truly marine forms. pelagic. the manatee. but the fresh. includand invertebrates. which retain some dependence upon the land.water habit by animals adapted to life in the sea. &c. or of animals which can so fix themselves as to avoid the force of the currents. illustrate the transition from the fresh-water to the terrestrial life. there is a greater wealth of life. But of by far the largest. the fresh-water dolphin of the Ganges became fluviatile at ing fishes a period which is geologically but of yesterday. show land . either animals in the process of reacquiring aquatic characbecause food is easier to get in water than on land. some forms showing a tendency to revert to the aquatic life (2) such forms as otter and desman among mammals. and have had more time to become fundamentally modified. like the larvae of insects. The manatee is still a marine animal.water dolphin of the Ganges. exemplify the acquisition of the fresh. these groups the littoral it is is and abyssal groups. or quitting it voluntarily in search of food.196 THE ANIMALS OF LAKES AND RIVERS life-history as well as lungs. and it is possible. as in the sea. to divide this life into littoral. The remaining animals of lakes and rivers. In lakes and ponds.

The fauna of the lochs of Scotland has been studied in great detail by the members of the Scottish Loch Survey. 440 species of animals were found. many species or is cf . but the salmon family. in the Caspian. g. near the margins of the lochs. e.g. so that it is possible to make a considerable number of general statements in regard to it. however. and Loch Ness of over 750 feet. Here.THE ANIMALS OF LAKES AND RIVERS abyssal fauna rarely is 197 sometimes virtually absent. exclusive of vertebrates and of insects and their Most of the Loch Morar has a maximum depth of over 1. owing to the paucity of oxygen. rendering an abyssal fauna possible. We know. and if its members in ever show the seen the animals of the ocean abysses. lochs studied are shallow.000 feet. which run into varieties . water-beetles and water-boatmen. The members of the Survey studied in all 562 lochs. specialization e. Of the invertebrates the majority may be described as littoral in that they occur in shallow water. which peculiar to that lake. and. and therefore there is no time for a special abyssal fauna to develop. including the large mussels and the small fresh-water snails. As considerable differences exist in the fauna of different lakes. Small . This seems to be partly because lakes are necessarily temporary phenomena. too. are found nearly all the few molluscs. Here are found many insect larvae. and not a few adult insects adapted for life in the water. from other sources that the special feature is the number of members of larvae. the Loch Leven trout. destined to be filled up after a longer or shorter period. it seems better to give some account of a few types rather than to make general statements in regard to lakes as a whole. The members of the Survey did not especially investigate the fishes.

the animals which constitute the fresh-water plankton but cosmopolitan. and four protozoa. and the forms from deep water show no special feature. is the presence here of some Arctic Crustacea. twelve rotifers. Of these thirty species most are very wddely distributed. and believed to form the community of organisms on the earth (Sir John Murray). the lochs are mostly shallow. The special feature. together with such small animals and so forth. especially those forms popularly called water-fleas. only in Loch Ness were successful deep-water dredgings made. a marine crustacean caUed the opossum shrimp (My sis) occurs. and the absence of certain forms found in other European lakes. however. three worms. small Crustacea. an insect larva. leaving a small group of animals. In certain of them. Of the two deep lochs. In the Scottish lochs the species of Mysis found is that .198 THE ANIMALS OF LAKES AND RIVERS from their Crustacea are numerous. especially those which lie but little above sea-level. three Crustacea. Only some thirty species are typically pelagic and thus constitute the plankton. water-bears (Tardigrada). both no doubt being all ' oldest ' due to the northern position of the lochs. These comprise fourteen as water-mites. and a few infusoria. As has been just stated. Before leaving the fauna of these lakes. so that it would be more correct to say that a few only of the littoral forms can live in depths greater than 300 feet than to say that a special deepwater fauna exists. Many rotifers occur. jumping movements. Here it was found that at depths greater than 300 feet the majority of the httoral species disappear. including one mollusc (a small bivalve called Pisidium pusillum). These occur also in the httoral region. and therefore an abyssal fauna is rarely developed. one interesting point may be noticed.

being about 475. to indicate their supposed former connexion with the sea. Its fauna has been subjected to detailed investigation. Reliktenseen).. and the lakes in which they occur are called relict lakes (German. we find that here the fauna forms a unity. Necessarily there is no deep-water fauna. . The area of this lake is about 250 miles. but its average depth is only 10 feet. another variety or species occurs. a large shallow body of water in Hungary. and in addition two forms of special interest. As its shallowness makes Lake Balaton a mere pool. The fish show some There are no trout. pike. With the Scottish lochs may be contrasted Lake Balaton or Flatten See. &c. called Mysis relicta.THE ANIMALS OF LAKES AND RIVERS which also occurs round the coasts water. but the lake contains such common fresh-water forms as perch. connected with the Danube drainage system. especially those in Denmark and on the North German plain. it is But in certain continental lakes. Such organisms are said to constitute a relict fauna. but have been long separated from it. but exclusive of insects surface. eels. interesting peculiarities. gudgeon. and is it is not possible to discriminate accurately between littoral and pelagic everywhere so close to the of species does not differ greatly from that of the Scottish lochs. roach. including 38 kinds of fish. The modifications which this form displays are believed to indicate that the bodies of water in which it occurs were once connected with the sea. in salt 199 and brackish and its presence is doubtless to be explained on a recent migrant from the the supposition that sea. gobies. and the maximum 36 feet. One is the catfish called ' Wels ' by the Germans {Silurus glanis). faunas. as the bottom The total number and air-breathing vertebrates. tench.

The wels is found only Eastern Europe. J. Moore and others. and is a member of a small group of old-fashioned fishes which used to be called ganoids. which once lived in the sea. for the ganoids are not a homogeneous group. As a third example of a lake fauna we may take that Lake Tanganyika in tropical Africa.200 THE ANIMALS OF LAKES AND RIVERS sterlet {Acipenser ruthenus). The name has been abandoned in more recent classifications of fish. Tanganyika has an area of 12.700 square miles. It of . but not in Scotland. but their interest for us is that they are fish of a primitive type. On the other hand. and the other the cat-fish constitute Tlie fish a large family of fresh-water in especially characteristic of the equatorial region. which has been the object of careful study by Mr. though they are absent from the lochs of Scotland. S. Similarly there is a greater wealth of moUuscs. Of the invertebrates it is sufficient to say that of its family in member with considerable general resemblance to those of the Scottish lochs. and such forms as the fresh-water crayfish (Astacus). Thus. but comparisons of this sort do not profit very much. found in England. and is the only European waters. but have been driven into fresh water (though the sturgeon also occurs in the sea) by the competition of the more highly organized bony fish. are present here. the Arctic types of Crustacea are absent. the rotifers are far less abundant than in the Scottish lochs. they display certain minor differences. The sterlet is a near ally of the sturgeon. E. a considerable number of genera of more or less southern facies being here represented. as it is difficult to be sure that the investigations have been conducted along exactly similar lines in the two cases.

In the place the lake contains a species of Protopterus. the species of Protopterus have a wide distribution in the lakes and rivers of the middle portion of possess. All the dipnoi both lungs and gills. owing their perthe acquisition of specialized breathingTheir discontinuous distribution over the ' ' At this period. The fish normally inhabit regions where a periodical dry season occurs. as the African continent. but the two sets of breathing-organs are not in use simultaneously. fauna show some curious features. In the case of Ceratodus the lungs are used when the water is too foul for the gills to be of any use.100 feet have been recorded at various localities. breathing by their lungs till the organs. but the animal does not make mud nests. As well as occurring in Lake Tanganyika. The two other living genera inhabit the one (Lepidosiren) the rivers of tropical America. but depths of 1. and the other (Ceratodus) those of Queensland. The seen. when the water in which they live either becomes very foul or dries up. Lepidosiren and make themselves nests of mud. therefore. many fossil forms having been found. surface of the globe is another point of interest in .624 feet above sea-level. Protopterus they lie water returns. The first fish The elevation is 2. Thus the dipnoi must be looked upon as the fresh-water survivors of a highly primitive group sistence to of marine fishes. Their ancestors were certainly marine. in which dormant. the second in that of the two other genera. All three genera show certain very primitive characters in combination with this specialized mode of breathing. one of the three living genera of dipnoi or lung-fishes. first we have condition is frequent in the natural habitat of the Queensland Ceratodus.200 to 2.THE ANIMALS OF LAKES AND RIVERS 201 has never been systematically sounded.

It seems as if the limb of Polypterus could more easily give rise to that of a terrestrial vertebrate Protopterus. common with old-fashioned animals. We have already considered several cases of land animals re- acquiring modifications of structure which life fit them for gesting one but Polypterus is in which land ar^imals arose from aquatic ones. Its ancestors were certainly marine. for the organ which forms the air-bladder in many other fish seems here to be used in respiration. Another very peculiar fish-type in Tanganyika is Polypterus congicus. however. the species Polypterus markedly from those of Protopterus. Into these points we cannot go here. in the water. it is a land vertebrate than that of Protopterus. but it may be differ say that many zoologists are of opinion that they show that terrestrial vertebrates arose from a stock common to them and to Polypterus. another example.e.202 THE ANIMALS OF LAKES AND RIVERS Such discontinuity is regard to them. a member of a ganoid genus. but were driven into fresh water by the competition of better-organized forms which appeared in the sea. whose waters become periodically points of foul. which ditions. In very of many structure. Like Protopterus ' ' this fish has an accessory respiratory organ. which is restricted to tropical Africa. owe their local persis- tence to their adaptation to special environmental con- The tapir in the swampy forests of the Malay is region and in South America. perhaps because the fish inhabits the same regions as Protopterus. Though the lung of the dipnoi is much more like the lung of sufficient to than is the air-bladder of not believed that dipnoi are near the line of descent of terrestrial vertebrates. but nowhere else. rivers i. largely because of the structure of their limbs. interesting as sug- way .

show the same we find a com- bination of forms found in other African lakes. very usual inhabitants of equatorial lakes. which retain many archaic . and. As we shall see directly the invertebrates general pecuHarities as the fish —that is. We have already noted. and these apparently led the way tion of purely terrestrial characters. the result being an unusually large fauna. Moore shows that the lake had once a direct connexion with the sea to the west. He places the rupture of this connexion so long ago as the period called by geologists to give the lake believes that this the Jurassic. and of quite peculiar forms of primitive type. The lake characterized by very large number of fish. Another peculiarity of the is fish fauna of Tanganyika its the enormous number is of fish belonging to the family Cichlidae. there are a considerable number of cat-fish. more remarkable still. that it is fish in lakes. fish of the lake are for the most part similar to those which occur in the remaining African For instance. Of these. nearly a hundred different species having been described. These in course of time evolved into the peculiar types now found in the lake. of the fifty-eight species of this family described in the lake only one is known outside the lake. The other lakes. in the allies of the ancestral Polypterus. through what is now the Congo basin. in speaking of the Scottish lakes. not unusual to find peculiar varieties or species of but the fact that many peculiar genera occur in Tanganyika speaks to long isolation.THE ANIMALS OF LAKES AND RIVERS The relatively unfavourable conditions rivers 203 in which exist made it necessary to develop accessory breathingto the acquisi- organs. and believes that at the time of the rupture the lake was peopled by Jurassic forms. more than half belong to the family Cichlidae.

the ordinary fresh-water fauna of characters. as already stated.g. The great point of interest is the occurrence of invertebrate animals of defuiitely marine type. &c. Some of these five at depths of 600 feet and upwards. e. the first two genera being cosmopohtan and very old. and it is of course very inaccessible. with strong marine hgJohmnic affinities. hope to have the detailed information in regard to its fauna which is available for the Scottish lochs or for Lake Balaton. for. We have already mentioned their occurrence in other lakes. Unio. notabty of a fresh-water jellyfish. Tanganyika is an enormous lake when compared with the lochs of Scotland. which is in itself an exceptional feature. but they seem to be especially numerous and peculiar in Tanganyika. It has never been systematic ally sounded. Planorbis. This hypothesis has not been universally accepted. Lake Baikal. Invertebrates of marine type " ' are of course not confined to in Tangamdka. therefore.204 THE ANIMALS OF LAKES AND RIVERS Later. giving the present- day ininghng of special types with marine affinities (haloHmnic forms) and ordinary fresh-water types. No less than fifty species of molluscs have been described from Lake Tanganyika. But in addition there are a number of gastropods of pecuhar and primitive characters. and. We cannot. " ' . so that it is not possible to distinguish between the faunas of the different depths in detail. As the figures already given suggest. they coexist there with ordinary fresh-water forms.water elsewhere. Africa gradually reached the lake. Thus we have species of Limnaea. and the evidence upon which it is based depends upon detailed anatomical points which cannot be considered here.. Of these a considerable number belong to the genera commonly represented in masses of fresh. Bithynia. constituting part of Moore's fauna.

and in Lake Nyasa no life has been found at depths greater than 100 to 150 feet. organism can be said to have solved completely the problem of adaptation to life in the brine.water jellyfish has been described from Rhodesia. 617). vol. This is the is Brine-shrimp {Artemia salina). and also two crabs and two prawns. three fresh-water sponges. xvii. The occurrence of deepwater forms in Lake Tanganyika is perhaps associated with the antiquity of the lake. Only one ordinarily unfavourable to animal life. and indeed those of lakes in general. which in excessively abundant (cf. but none of these is well known. The Crustacea include the usual small copepods of fresh water. which has given time for the evolution of a special deep-water fauna. The protozoa of Tanganyika seem to resemble generally those of other African lakes. and another apparently very similar form has been found in the tributaries of the Krishna in India. and is thus another example of the deep-water fauna. (1901) p. To these accounts of characteristic lake-faunas. of variations in the salinity of the water. all pecuhar to the lake. summer Scottish Geographical Magazine. A closely related fresh.THE ANIMALS OF LAKES AND RIVERS as 205 we have already explained. One of these crabs extends downwards from 500 to 600 feet. and the jellyfish (Limnocnida) already mentioned. a word or two may be added on the fauna of the Great Here the conditions are extraSalt Lake of Utah. and the fact that the eggs can develop parthenogenetically without previous fertilization enables the animals to reproduce themselves very rapidly when the conditions — — that . Among the other interesting forms are a peculiar polyzoon. there appears to be a deficiency of oxygen in the depths of equatorial lakes. It is very tolerant of cold and is.

Curiously enough. An and interesting example of a cave animal is Proteus anguinus. instead of showing the deep-red or black coloration found in many animals which live in the ocean near the light limit. Some of the of abyssal forms therefore reappear here for example. also occur in the shore waters. and perfectly black specimens can be produced by gradual exposure. we we find that cave animals tend to be white. Cave Faunas. though ters may be low. and reaches a The skin is without pigment. Otherwise it is singularly barren of animal life. where as have seen (p. something may be said here of the more characteristic animals found in them. Cave animals live in conditions which in many respects resemble those prevailing in the ocean abysses. 172) certain rays penetrate deeply. Green charac. As caves usually contain a considerable amount of water. But as the light conditions are quite different from those which obtain in the ocean depths. larvae of two is also During the warm season the a Tipula and a Chironomus. In the caves of Texas there is an allied form {Typhlomolge gilled length of about a foot. is remarkably uniform. This animal is one of the permanently- amphibians already described. The animal remains permanently within the water. also. flies. and the eyes are below the skin.206 THE ANIMALS OF LAKES AND RIVERS all are at favourable. are necessarily absent. Dalmatia. and a water-boatman (Corixa) found in the lake. it and the temperature. . plants. found in the caves of Carniola. Thus there is often complete darkness. though it rises to the surface to breathe if the water is not quite pure. if the white body is exposed to hght pigment develops in the skin. the eyes tend to disappear or become minute. Carinthia.

Archinotis. and a few molluscs. Many of the general works already mentioned also discuss fresh-water faunas. is To make up for the loss of the eyes furnished with a great number of tactile Of other cave animals the most important are Cruswhich has no eyes.g. 1906). by Dr. of South America in Archhelenis u. 1891). direction of Sir The first volume of the scientific results of the Bathy- metrical Survey of the Scottish Fresh-water Lochs. especially isopods. the organisms found in the Scottish Lochs. and has concealed eyes. See also articles on the river detailed lists of mussels. Other references to the same subject will be found in the bibliography mentioned above. by Hermann von Ihering as they affect (Leipzig.g. The same type adaptation also reappears among the cave fish. Animal Life (London. Kobelt's book (cf. e. and insects. See also Semper's Conditions of Existence fish of the Kentucky . and to them may be added Forel's Handbuch der Seenkunde (Stuttgart. 35) has an interest- ing chapter on cave faunas. Otto Zacharias and others (Leipzig. 1901). by J. 1881). Adelops and Bathyscia. A full account of the fauna of Lake Tanganyika is given in The Tanganyika Problem. Titanethes albus. John Murray and Laurence Pullar (Edinburgh. e.und PJlanzenweU des Sllsswassers. notably beetles. as mcII as gives also a detailed bibliography of limnological literature. Lausanne). This volume The account of the fauna of Lake Balaton given in the text is based upon the Besultate der wissenschaftlichen Erforschung des Balatonsees. One of the most interesting found there is Amblyopsis spelaea. E. the tendency to lose eyes and the pigmentation of the skin being well marked. 1872). which is white in colour. &c. 1903). Moore (London. p. herausgegeben von der Balatonsee-Commission der Ungarischen Geographischen Gesellschaft (Vienna. 1892-1904. contains several articles dealing with the fauna of lakes. The blind cave are discussed by Putnam {Amer. myriapods. which have been especially studied in the Mammoth Cave of Kentucky. also some spiders. References.CAVE FAUNAS raihhuni) of 207 of similar characters. and Die Tier. conducted under the 1910). the head papillae.. Nat. and the same author's large work on Lake Geneva (Le Leman : Monographic limnologique. tacea. 1907).

In other words. and so forth. and parts of the adjacent . for example. and the sloths. These have long since become extinct. certain kangaroos have taken to the trees and become arboreal in South in Madagascar lemurs swarm in the forests America particular kinds of monkeys. people the and so on. and the adaptations which these show to their particular habitats. found wherever the forest But in New Guinea. as well as other animals. called the is adaptation has worked raw material upon wliich not the same in all parts of We have found that animals inhabiting dense forest generally show certain arboreal adaptations. is The only possible explanation of these conditions the hypothesis that the speciahzed forms in each separate region have been evolved from pre-existing unspeciahzed forms inhabiting the region. course of these descriptions it In the that obvious has become what may be the globe. or other area. occurs. and the existing sloths have apparently been saved from a hke fate by the acquisition of marked arboreal characters. has its own types of adapted animals. the opposable thumb and great toe. selvas. . the opossums. which protect them from many possible enemies. and have noted their more important occupants. In the continent of Australia.CHAPTER X ZOOGEOGRAPHICAL REGIONS We have now surveyed those natural regions of the earth described in the Introduction. such as the prehensile tail. each isolated forest. In South America we find fossil representatives of ground sloths.

Living marsupials also occur in South America. and less diverse. In order that a particular stock differentiation to isolation may give rise by specialization and many such groups. and .ZOOGEOGRAPHICAL REGIONS regions. they are few in number. as in possible enemies. any forms inhabiting a natural region. Lemurs also occur in Africa where they have to face the competition of the more intelligent monkeys. practically absent in AustraUa. and have thus protected themselves from the competition of groundliving forms no less than from the attacks of some part savana animals. kangaroos are abundant. then. 209 and are for the most But where tropical forests occur. in Madagascar the lemurs are very numerous and very diverse. there certain forms have taken to the arboreal life. more or less perfectly adapted to of habitat the possible varieties which the region affords. For example. for otherwise new incoming stocks may colonize the vacant places before the first stock has had time to become differentiated. small in size. that terrestrial all stock will tend to give rise to specialized groups. 1404 O India. and is there that animals reaches a isolated from any cause. The same statements might be made animals. North Queensland and New Guinea. Similarly. and the attacks of more powerful carnivores than any which are found in Madagascar. but there. but there they are represented by a very limited number of types. generally. of of desert of savana animals. or. because they have had to face the competition of placentals. being adapted for different modes of life. We may say. in Australia almost all habitats suited to terrestrial mammaUa are occupied by marsupial animals. some degree of seems to be necessary. when any stock of new region.

and similarly how the numerous deer of the north are shut out from barrier Africa by the. how scantily represented in temperate are the abundant antelopes of Europe and Asia Africa. for both the mountains and the deserts prevent intermixmg. and by isolating much of Africa on the one hand and the peninsula of India with Further India on the other. The animals of temperate Europe and Asia show a marked difference from those of tropical Asia and of tropical Africa. impassable desert. to favour evolution in these two areas. Now there is much evidence to show that there has also largely deserts been a progressive desiccation in geologically this region within recent times. One effect has been to favour the development of the many steppe and desert animals of the region. To some extent this difference is doubtless due to the existence of transverse mountainchains. To the zoogeographer. Necessarily these barriers must be supposed to have a definite date of origin. for example. to them. We have seen. but it is As an example due to the existence of a great band of and wastes across Asia and Northern Africa. but another has certainly been to separate an originally homogeneous fauna into three parts. many divisions as the result of the of the origin in time of such a barrier let us take a progressive climatic change.210 ZOOGEOGRAPHICAL REGIONS isolation of portions of the earth's surface is Such caused by the existence of barriers to the distribution of terrestrial animals. however. and this origin must always be supposed to be later in time than that of the ancestral stock of the group under consideration. such as the Himalayas and the Andes. save on a very limited scale. the greatest . now pro- tected from the incursion of northern forms. but to precede the differentiation of the stock into isolation.

has geographers with the idea that great changes in the distribution of land and water have taken place on the surface throughout geological time. Without such a consideration. as in most of the works on the subject. we cannot hope to explain in its entirety the distribution of animal animals at the present time. Now to consider this distribution in its relation to life and evolution from the Cambrian period onwards would obviously be a Herculean task. This is especially true when the subject is treated in detail.g. to be carried passively host. In dividing up the globe in this way the first point to be emphasized is that. and in this concluding chapter we shall consider briefly the classical regions into which the globe has been divided by zoologists. or resistant enough. on the basis ' ' especially of the distribution of birds and mammals. e. no such distinction o 2 . however. it is of interest to note the But without attempting main features in the Intro- of the distribution of the higher forms of Ufe in the existing continental masses.ZOOGEOGRAPHICAL REGIONS is 211 undoubtedly the sea. save those small enough. At the same time for historical reasons. to do this. by Arldt. especially in Geological research. Here. if for no other. the subject is too important to be entirely omitted. though it is one which has been attempted. though the geographers are accus- tomed to lay great stress upon the distinction between the American and Eurasiatic continents. familiarized recent years. As we saw duction. obviously. for a broad channel offers an insuperable barrier to the passage of most land animals. between the Old and the New Worlds. such a division of the globe into zoogeois graphical regions primarily of interest to the zoologist rather than to the geographer. by winds and waves or an animal we approach difficult questions.

We thus have to consider as forming one great of the earth's surface the Realm whole Eurasian con- tinent with parts of the Malay Archipelago. and though they are now functionally separated from the great land-mass in the eastern temperate and frigid zones regions.212 exists ZOOGEOGRAPHICAL REGIONS between the animals. by belts of deserts and by tralisverse mountain-chains. there was apparently a time when it was physically possible for land animals to spread from the far north of Europe and Asia to the far south of Africa and India. as already suggested. as well as possibly across parts of the North Atlantic. . Further. was in communication. the con- tinent of Africa with the isolated region of Madagascar. as we would expect from its great extent. before the progressive formation of these belts of desert. and North America. both geological and zoological. to show that not long ago there was free land communication between the Old and New Worlds in the northern hemispheres. and that only as we travel southwards does differentiation appear. though parts of what are now the continents of Africa and of India were. There is much evidence. In this great land-mass evolution. though it seems to have been leng separated from South America. probably separated by stretches of sea from the land-masses to the north. across what is now the Bering Strait. in early Tertiary times. yet. and south-eastwards to parts of the Malay Archipelago. at least of the northern We have already pointed out that the tundra animals of the east and west show very marked resemblances to one another that similarly there is considerable resemblance as regards both the plants and animals of the belt of coniferous forest in both hemispheres. free With this great eastern land- mass North America. has been .

Where the tropical forest is dense a few primitive forms linger. and some also have been saved by isolation. A mingling of faunas certainly occurs in this region. where a boundary line has to be drawn through that mass of islands which stretches between Further India and the northern shores of Australia. For this great area various names have been proposed. and no monotreme. earlier age. however. and any hard some doubt upon . In two regions. like the Hatteria of New Zealand. Wallace drew the line which separates his Oriental (or Indian) region and the AustraHan region between the islands of Bali and Lombok. because of its mainly northern position. Recent detailed research has thrown this statement. points us back to an Here and here alone do we get the highest from man the anthropoid apes and the dog-faced baboons.g. 213 Here we find the highest mammals and the most speciaHzed birds.ZOOGEOGRAPHICAL REGIONS rapid. difficulty occurs. Here and here alone do we get the highest and most specialized of the ungulates. Thus it has been called Arctogaea. and so on. or the Northern World. e. No primitive reptile. In regard to the boundaries of this realm it may be said that over much of the area they are formed by the sea. One is in the south-east. A narrow but deep strait separates these islands. though Africa extends far to the south. No marsupial (save one opossum in North America) now lives within this great area. and Wallace of the primates apart — believed that all the islands to the east of this strait (Wallace's line) possessed faunas with a distinctively Australian facies. in Madagascar. But as a general rule the primitive forms have been crushed out of existence by higher forms. while those to the west had faunas of Indian type.

It is enough for our purpose to know that such a transition exists. at a not very remote period. g. Here again the difficulty is usually got over by the erection of a transitional region to include a considerable part of the south-western United States. New Guinea. a certain occurs. The other area where a fine is difficult to draw is in the American continent. The rest of the globe is sometimes included in one Realm. amount That this mingling is not greater than it is is doubtless due to climatic differences. and the increasing probability that they were connected. or Southern World. absence of the highly differentiated forms named . minghng . The Notogaeic realm so defined is characterized by the presence of monotremes (in Austraha) and marsupials by the presence of running birds and dipnoi . seems to be justified by the increasing proofs of the affinities of their faunas. and so on. The difficulty is munication. therefore. despite their considerable separation in space. and by the above. (these being characters shared with Africa). forming the Notogaeic Realm. and on the other the continent of AustraUa with Tasmania. There are many striking differences between the faunas of North America and South America. to the occurrence of desert regions. e. and some of the islands of the Malay Archipelago. as weU as the scattered groups of Polynesia.214 ZOOGEOGRAPHICAL REGIONS fast line and must necessarily be of a more or less sometimes got over by the erection of a transitional region. This union of Australia and South America. through the Antarctic continent. of Necessarily. and including on the one hand Southern and Central America. with Mexico and the West Indian islands. the islands of New Zealand. in Mexico and the west. but the two regions are now in free comartificial character.

This classification may be summed up in tabular form as follows : : ARCTOGAEIC REALM. including India south of the Himalayas. including the whole of temperate Europe. Malagasy region. and (6) the eastern or Australian. in the characters already mentioned. Asia. we may note that some authorities separate it on this account from the Arctogaeic realm.ZOOGEOGRAPHICAL REGIONS Now it is 215 obvious that these two realms are large divisions. (2) (3) (4) Holarctic region [sometimes divided into an and a western (Nearctic) sub- Oriental region. and approaches the South American and Australian regions. To these many would add a fourth region for the island of Madagascar. Without pursuing this point further. and parts of the Malay Archipelago (3) the Ethiopian. including Africa south of the Atlas and Sahara desert. Again. is strikingly dissimilar. we find that it may be subdivided into the following regions (1) the Holarctic. Africa. the Holarctic region. (1) eastern (Palaearctic) region]. . . and North America (2) the Oriental. Oriental region shows many resemblances to that of turn subdivided into regions. the differences being largely attributable to differences of climate. whose fauna is so strikingly difl^erent that it may well form But while the fauna of the (4) the Malagasy region. Further India. : . the Notogaeic realm obviously falls into two regions (5) the western or Neotropical. Ethiopian region. and unwieldy They are therefore in their Taking the Arctogaeic realm first.

their place being taken by the hang-nests or Icteridae. To the east. but both are rich in carnivores. Thus North America has no true starlings. (5) (6) Each may now be considered in a Httle detail. Australian region. but not in the west. while the Nearctic section shares the skunk and raccoon with the Neotropical region. In' both divisions rodents are very numerous. p. In the Palaearctic section the sheep and goats are well represented. however. absent in the New. where it extends into the Oriental region. rather than orders or families. we find that the western or Nearctic section has no primates. Elephants do not occur in either division. while deer are abundant. Beginning with the Holarctic region.216 ZOOGEOGRAPHICAL REGIONS Neotropical region. camels and horses occur. The bird faunas of the two regions differ from each other somewhat markedly. but this is partly because both regions receive many seasonal immigrants from the hotter land-masses lying to the south of them. but this is a fact which has already been sufficiently discussed. The differences also chiefly affect genera or even species. 82). Ainong the special forms it is noticeable that the badger is peculiar to the Old World. to which the rice-bird or bobolink (Dolichonyx) belongs. The . of these regions NOTOGAEIC REALM. Among the insectivores the mole (Talpa) is a form peculiar to the Old World. 77-8). The antelopes are scantily represented in both. but to the west they are few (cf. and the eastern or Palaearctic only those species of Semnopithecus and Macacus already mentioned (pp. of the Two familiar families Old World are. Both sections have oxen.

which is remarkable for its development of salmon.ZOOGEOGRAPHICAL REGIONS flycatchers (Muscicapidae) of the Old 217 also World are absent in the New. peculiar to the island of Celebes. pike. —a very interestmany fruit- The Oriental region is rich in bats. sticklebacks. and the gibbons of the region. already mentioned the peculiar proboscis monkey (Nasalis) of Borneo. including two species of the very peculiar genus Tarsius. Beginning with the Primates. The animals of the Oriental region show so marked a general resemblance to those of the Ethiopian. certain welldefined differences in addition to basal resemblances. perch. &c. their place being taken by the tyrant shrikes (Tyrannidae). There are four lemurs. also been said of the fresh-water fish fauna. There are. with some peculiar genera. scantily represented. that some naturalists would unite the two as a Palaeotropical region. belonging to the genera Macacus and Semnopithecus. The genus Cynopithecus includes only one species. the so-called black ape. the majority Something has of which are peculiar to the region. the orang of Sumatra and Borneo. We have usually included in the Oriental region. Reptiles are somewhat but we have already touched upon the abundance of tailed amphibians. Among the insectivores the very peculiar flying lemurs (Galeopithecus) of the Philippine Islands are . carp. and in Madagascar about thirty-six species ing contrast. There are also many kinds of monkeys. In the Malay continent of Africa there are eight species. we find that the Oriental region has two peculiar kinds of anthropoid apes. having representatives both of the insect-eating and the eating forms. There is nothing specially worthy of note as regards the invertebrates. however.

green bubuls (Phyllornithidae). especially squirrels and mice. and are very numerous. Carnivores are numerous. the abundance of deer. another point of contrast with Africa. and include peculiar genera. . Very striking also are the bulbuls (Pycnodontidae). as are also the tree-shrews (Tupaia). On the other hand. com- pared with their abundance in the Austrahan region. and the fact that wild asses just reach the region. Among the ungulates we note the presence of oxen. the region being especially characterized by the number and beauty of its pheasants. snouted forms of the latter (Gaviahs) are restricted to the region. Rodents. of rhinoceroses. the pangolins. the paucity of antelopes. does not reach Ceylon. True cats of the family Felidae are also numerous. many and an elephant. Birds are very numerous. A good many of the peculiarities of the Ethiopian region have been alluded to incidentally in the fore- .218 ZOOGEOGRAPHICAL REGIONS confined to the region. The babbling thrushes (Timehdae) have their headquarters in the region. and bears. we have already spoken of There are many reptiles The longthe flying Kzard (Draco) and the crocodiles. where there is one species in the Ganges. the grant into India. are abundant. whose presence in India affords region. represented elsewhere only in the Ethiopian region. and is believed to be a relatively recent immiLike the Ethiopian region. The Edentates are represented by members of the genus Manis. among is the interesting the presence of contrasts with the Ethiopian region beheved to have originated within the Oriental The tiger. pigs. of of a tapir. Oriental one is rich in civets and their allies. parrots are relatively few. the presence of chevro tains. and the hill-tits (Leiotrichidae).

Asia.E.Fig. Clouded Leopard of S.) . (From the picture hy W. 45. Walh.

46.) .Fig. The Okapi. {From a specimen in the Royal Scottish Museum.

In regard to the rodents a peculiar feature is the absence of the true flying squirrels and their replace- ment by the peculiar family of Anomaluridae (cf p. and with one exception so are true pigs of the genus Sus. but rhinoceroses. and . river hogs. The region is almost equally characterized by the forms which are present. two peculiar forms confined. Thus we have jumping-shrews. several kinds of horses occur. as we have seen. though wart hogs. the aquatic potamogale and the golden mole. as well as the curious hyrax and pecuHar forms Uke the giraffe and okapi antelopes are abundant. So much has been already said of the ungulates (p. The birds are not so abundant nor so beautiful as . chevrotains.) that a few general statements may suffice. Insectivores are peculiar and primitive. Among the Primates it possesses the gorilla and the chimpanzee. and by those which are absent. but some further details may be added. 129 et seq. Among the carnivores an interesting feature is the presence of two peculiar dogs. belonging to the two genera of Lycaon and Otocyon. Fruit-bats are fewer than in India. 109). and is noticeable that the weasel scantily represented. bush pigs. Goats and sheep are practically absent. elephants. There is no tapir and no camel.ZOOGEOGRAPHICAL REGIONS 219 going description. as well as by pangoHns. to a relatively small part of the conMonkeys are represented by five peculiar tinent. We have already spoken it is of the absence of bears. The Edentates are represented by the curious aard-vark (Orycteropus). the latter beheved to be alhed to the curious tailless hedgehog or tenrec of Madagascar. the latter having very primitive alliance teeth. genera with many species. . and the hippopotamus occur. lemurs by two or three genera and some eight species.

regions. The insectivores are somewhat numerous. which they somewhat resemble in beauty of plumage and in habits. Some other peculiarities of the fresh. The other two are hippopotamus (with one extinct species) and Potamochoerus. the river hog (with one species) . ' ' these differences testifying to thirty-six its long isolation. and the curious ganoids known as Polypterus and Calamoicthys. both helpless forms. The Malagasy region or sub-region (p. 152) shows some very striking differences from the mainland. and among the fish we have to note the presence of the dipnoan genus Protopterus. and the genus is one of the three which Madagascar shares with the adjacent mainland. and shows many primitive characters. there are no anthropoid apes nor monkeys. though not confined to the region. that of the tenrecs or Centetidae. Of the insectivores all save one species of musk-shrew (Crocidura) belong to peculiar genera.220 ZOOGEOGRAPHICAL REGIONS The plantain(Coliidae) those inhabiting the Oriental region. of Africa. eaters (Musophagidae) and the the peculiar families. for there are over twenty species. Reptiles are numerous. The musk-shrews are widely distributed in the Old World. all belonging to peculiar genera. Parrots are not very numerous. is very characteristic. and most belong to a peculiar family.water fauna have been already alluded to. but species of lemurs Thus some occur. which is entirely confined to the island. The true ostrich. colies are among Though the beautiful sun-birds (Nectarinidae) are represented in the Oriental and Australian teristic they are especially charac- where they take the place of the humming-birds of South America. must be associated with the paucity and small size of the carnivores. The relative abundance of these and of lemurs.

ZOOGEOGRAPHICAL REGIONS Rodents are represented by ing to the five peculiar 221 genera belongis mouse section . The bats are very peculiar. The absence of the true cats of the genus Felis. thus showing a curious parallelism with the lemurs. and it teristic the non-fljdng mammals of will be noted that they form the plantain-eaters a striking contrast with those of Africa. Here are no anthropoid apes. and another of small. Here only (with a slight extension into North America) do we find the vampire bats. which is both highly pecuHar and very rich in species. while the fruit-bats of the Old World are completely absent. of camivora are peculiar to the island. but on the other hand a peculiar family of broad-nosed (Platyrrhine) monkeys. of which the largest the cat-Hke animal called Cryptoprocta. believed to be related to the Centetidae of . is a remarkable feature. e. though in the West Indian Islands there occur two shrews of the genus Solenodon. no dog-faced (Catarrhine) monkeys. both families differing in a number of respects from all the Old World forms. and no lemurs. but are more abundant in the island of Madagascar than anywhere else. These lizards occur in Africa and in India. furry forms called marmosets. also broad-nosed. Turning now to the Notogaeic Realm we may begin with the Neotropical region. all The above include Madagascar. elsewhere so widely distributed.g. The chameleons present some interesting features. as well as another family called the horse-shoe bats (Rhinolophidae). The characand colies. are similarly absent. carnivores by some seven All the genera members of the civet family. African birds. while the reptiles are very remarkable in that some show affinity with those of South America rather than with those of Africa. Insectivores are practically absent.

which are of great interest because they are beHeved to belong to the herbivorous or diprotodont section. p. 140). the motmots. that of which the kangaroo is the most familiar representative. and like the skunks are common to this region and the Nearctic. llamas Old World camels). sheep. We have already spoken of the number and peculiar nature of the rodents. the tanagers. goats. but these are beyond our range. In South America there occur also other marsupials. Among the important famihes are the humming-birds. though. is it. the jacamars. the toucans. pecuhar pigs (pec- caries). tapirs (shared with the Malayan region). the macaws (Conurinae). The birds of South America are almost as pecuhar as the of mammals. Of the carnivores it is only necessary to say that bears are represented by only one species. Very remarkable conditions are exemphfied by the ungulates. but no antelopes. the chatterers. therefore. oxen. . and nothing need here be added to what has been said on pp. nor horses. There are many peculiar fossil forms. and the presence of many kinds of opossums.222 ZOOGEOGRAPHICAL REGIONS Madagascar. The discovery of these Uttle animals in South America is of great theoretical importance. because it was formerly beHeved that (aUies of the the diprotodont marsupials had originated within the Austrahan area. another link in the increasing chain of evidence which points to a former connexion between South America and AustraUa. while raccoons are plentiful. the tinamus (cf. &c. as was to be expected. the curassows (Cracidae). many the characteristic forms extend also into North America. The pecuhar nature of the Edentates has also been pointed out. and had never occurred outside Their existence. known as selvas (Coenolestes). 56 and 87. We find pecuhar deer.

the dingo or native dog and the pig of New Guinea. a few rodents of the mouse section. in Queensland and New Guinea. we must region. and a peculiar feature is that the more of Australia.. and is the negative characters by several species. which is probably an introduced animal. occur is to the south. and New Guinea. whereas the more primitive forms. and the lizards of the family Iguanidae. the region further includes the only living monotremes. and Proechidna in New Guinea only. occur towards the north. Among we may note the absence of crows and ravens. rather than from the north. especially tree-frogs (Hylidae). consider the Australian markedly peculiar Here the higher or placental mammals are absent. the certainly modern tree- kangaroos. As already explained. Tasmania. Echidna in Australia. which do not extend south of Guatemala. The marsupials are extraordinarily numerous and very diverse. &c. . Among the fish the dipnoan called Lepidosiren Finally is important. while there is a large number of toads and frogs. We need not consider further the marsupials of the region beyond noticing that there are forms adapted to almost every kind of Australia received south. in the island of Tasmania.ZOOGEOGRAPHICAL REGIONS The so-called 223 is American ostrich (genus Rhea).g. through the islands of the Austro-Malayan region. the Tasmanian wolf and Tasmanian devil. Ornithorhynchus being found in Australia and Tasmania. As in Australia there are many parrots. e. with the exception of bats. the boas and anacondas. represented charac- teristic. Among the reptiles interesting forms are the rattlesnakes. link in This another the chain of evidence which suggests that its original marsupials from the through a connexion with South America. specialized forms.g. e.

the marsupial mole (Notoryctes). 118). New casso- Zealand has the curious Apteryx. eaters (Meliphagidae) are peculiar to the region. no woodpeckers. Zealand has a very primitive lizard (Hatteria. The birds of the region are interesting and peculiar. New Guinea is especially remarkable for its beautiful birds of paradise. which has become carnivorous. The type genus Hyla occurs all over Australia. and no vultures. There are no true finches. while the related cockatoos are almost peculiar.224 habitat. with many peculiar birds. and the presence of many members of the family in the Austra- lian continent is the more remarkable in view of its complete absence from Africa and India (cf. ZOOGEOGRAPHICAL REGIONS and showing curious adaptive resemblances to the placentals. as well as the moa (Dinomis). Very extinct striking also are the little Running birds. An is interesting form. The fish fauna of the streams and lakes is somewhat . no pheasants. has some remarkable parrots. 228) and no snakes. p. preying upon sheep. with a curiously close resemblance to the typical mole Old World. and the nocturnal owl parrot (Stringops) with a degenerate keel on its sternum. Mound-turkeys and lyre-birds occur both on the mainland and in the and the parrots are exceedingly numerous and very characteristic. represented on the mainland chiefly by the bower-birds. discovered not many of the years ago. and but little power of flight. Pigeons are also very abundant. and include the most brightly coloured members of the order. p. Australia the emu and wary. New Zealand. Queensland and New Guinea share with America reptilian fauna is less peculiar. The but New an abundance of tree-frogs of the family Hylidae. notably Nestor. The honeyislands.

m p .

Lyre Birds with nest and eggs. {From a specimen in the Royal Scottish Museum.Fig.) . 48.

1896) Beddard. and the extraordinary differences between its fauna and that of the rest of the world. See also Geoffrey Smith. The Geographical subject of this chapter . by Bartholomew. and Geological Distribution of Animals (London. La Oiographie Zoologique (Paris. Still another curious relic is the mountain shrimp (Anaspides). A Text-book of Zoogeography (Cambridge. Clarke & Grimshaw (Edinburgh. lived in the far-off the marsupials in suggesting the long isolation of x4ustralia. Die Entvncklung der Kontinente und ihrer Lebewelt (Leipzig. A Naturalist in Tasmania (Oxford. plates.ZOOGEOGRAPHICAL REGIONS fish 225 poor. 1907). a primitive type of crustacean found only in the mountain region of Tasmania. cussion of the origin of the Australian fauna. and apparently closely related to forms found fossil in Carboniferous and Permian beds in Europe and North America. now found only off Australian coasts. Geographical Distribution of Animals (London. and is one of the curious rehcs of the AustraHan continent. will be found in the Atlas of Zoogeography. 1890) Heilprin. but the presence of the very primitive dipnoan Ceratodus is important. The following are the : more important books on the Wallace. 1876) Sclater. A Geographical History of Mammals (Cambridge. 1887) . but once abundant in the Jurassic and Cretaceous seas of Europe. 1899) Lydecker. . This fish. Arldt. Refebences. with \ 1404 . Another is a mussel called Trigonia. 1911). for a dis- A very elaborate series of a considerable amount of text and a full bibliography. now found only Devonian period in the seas of the northern hemisphere. . The Geography of Mammals (London. 1909). 1895) Trouessart. These old-fashioned forms are perhaps in some respects even more striking than in the rivers of Queensland.

or the more systematic Natural Histories. .APPENDIX OUTLINE CLASSIFICATION OF ANIMALS Readers who are not well acquainted with modern zoology may ence. find the following outline classification useful for refer- Fuller details will be found in any of the zoological text-books. including warm-blooded hairy animals whose young are nourished by milk after birth. A whose members are of structure. whose young are nourished before birth and are therefore including nine orders. The Royal Natural History (London). the highest and most intelligent manmials. with three sub-classes : Sub-class I. Thomson's Outlines of Zoology (Edinburgh). including monkeys. Class Mammalia. and have their respiratory organs. is This phylum divided into a : number of smaller units or classes as follows PHYLUM VERTEBRATA. as well as . and are believed to have been descended from a common ancestor. the word phylum phylum includes a group of animals characterized by a common ground-plan originating from the anterior end of the food-canal. by an organ 1.g. whether lungs or gills. e.g. apes and man. possess a tubular nervous system. Of the terms used is in important. e. Thus we distinguish the members of the 'phylum of vertebrates from all the varied phyla of invertebrates by the fact that all have at some period of life a dorsal supporting rod called the notochord. called the allantoic placenta. Eutheria or Placentalia. modern classification. born with the form of the adult Primates.

affinities : shrew. and The elephant and the rock-conies (Hyrax) of Syria show certain Rodentia. including life in only two living forms. including the opossums and the Tasmanian wolf. or whales Rabbits. Examples 5. &c. and having by means of which they obtain their food. Chiroptera. may be the only one present. Sub-class II. bear. Carnivora. Examples mole. 7. characteristic chisel-edged incisor teeth. Cetacea. or flesh-eaters. and e. usually small. sheep and cow. generalized and usually small animals. In one sub-order (Perissodactyla) the third or middle toe tends to predominate. usually feeding on insects. including mammals whose young are born imperfectly developed. with strong teeth. absent in other ungulates. 227 the more primitive lemurs. hedgehog. 4. Metatheria or Marsupialia. rats and mice are examples. mammals very perfectly adapted 8. Edentata. ant-eaters. and are carried by the mother after birth in a pouch. with the power of 3. all brains. armadillos. forms with usually opposable great toe. with both with primates and with marsupials. or marsupium comprising two orders: 1. herbivorous animals. &c. with numerous incisor teeth. g. insect-eating or fruit-eating forms flight. 6. or sea-cows. e. also adapted for 9. primitive characters. the fore-limbs being greatly modified. an old-fashioned heterogeneous order. horse . adapted either for or in water. Polyprotodontia. an old-fashioned group. Sirenia. having four to five toes. Insectivores. Ungulata. in the other sub-order ( Artiodactyla) the third fourth toes are equally developed. or bats. the gnawing animals. a large order of often powerful life : animals. including with relatively simple sloths. mostly arboreal in habitat.. carnivorous forms.g. on land cat. with a reduced number of toes. the water. . and dolphins. for life in water. and teeth adapted for a vegetarian diet.APPENDIX thumb and 2. dog. and having cusped teeth. P 2 . often of large size.

without tails in adult life. the Sub-class III. &c. having a keel of flight. Monotremata. Lacertilia 5. Sharks and skates. Carinatae. &c. g. with one living order. and usually the power 2. including forms and toads. The classification is a matter of difficulty. and many extinct ones 1. Chelonia or turtles and tortoises. cassowary. and the two spiny ant-eaters (Echidna and Proechidna). with 3. which contains only three mammals.. permanently aquatic forms with gills and scales. forms with a completely bony skeleton and a swim-bladder used as hydrostatic organ. Prototheria. OUTLINE CLASSIFICATION OF ANBL\LS Diprotodontia. long-legged forms. mammals. herbivorous forms. but sharing with the Elasmobranchs certain primi- . Crocodilia or crocodiles and alligators. with only two incisor teeth in the lower jaw. and those like newts and salamanders. including only a few living forms. The living forms may be divided into two divisions living kinds of : 1. RhyncJiocephalia. cold-blooded scaly animals with five living orders. on the sternum. emu. or flying birds. no keel on the sternum and rudimentary fore-limbs. and have a cartilaginous skeleton. or lizards. Class Reptilia or Reptiles. Ganoids we may group a few living forms. including kangaroos. very different from each other. Class Aves or Birds. including warm-blooded forms clothed with feathers. the ostrich. New Zealand lizard called Hatteria. including only the 4. gills in Class Amphibia or Amphibians. : Ratitae.228 2. 2. like frogs owing to the vast number of fossil forms. with many primitive characters. are Elasnwhranchs. egg-laying primitive wombat. or running birds. with many orders. and lungs in the later. Class Pisces or Fishes. It is sufficient to note here that most living fishes are Teleosteans. smooth-skinned forms with the early stages. the duck-mole (Ornithorhynchus). Ofhidia or snakes. &c. the living forms being the As specialized remnants of a large and primitive group. which keep the tail throughout life. e.

there are the many phyla of invertebrates not nearly related to each other. forms like oyster and mussel. active 2..g. The phylum vertebrata includes in addition some other small groups. and so forth. and gills. is PHYLUM The highest MOLLUSCA. Though we commonly use the term invertebrate as a convenient contrast with vertebrate. the Dipnoi or double-breathing only three living forms (Ceratodus.APPENDIX tive features. the polypterus of the and so forth. including snails. millipedes. marine forms in which the shell is usually lost. &c. having a loose fold of skin called the mantle. gills. though they have a swim-bladder like the Such are sturgeons. 229 Teleosts. e. but almost tadpole-like when young. degenerate when adult.. usually sedentary in habit. and carnivorous in habit. called the foot. Protopterus. The very large PHYLUM AETHROPODA includes animals which some at least of the segments of the body are furnished with jointed appendages. with a double shell. the snail creeps. the Tuni gates or Sea-squirts. 2. periis winkle. in which the shell present or absent. Finally. There are three important classes 1. 3. and centipedes. usually furnished with a limy shell. Most live on land. . and having one pair of feelers or antennae. Nile. &c. fish include and Lepidosiren). Lamellibeanchiata or bivalves. The Crustacea. such as the Cyclostomes or round-mouths (hag and lamprey). mostly aquatic forms. bony pike. and when present has but one valve. There are three main in classes 1. and the curious worm-like Peripatus. all breathing by air-tubes. interesting because they possess both lungs and gills. The Cephalopoda or cuttles. on which. slugs. Gasteropoda or univalves. including forms breathing by and a muscular protrusion. Antennata Tr ache ATA. including large forms like crab and lobster. including insects. with a hard coat. two pairs of antennae. and the small copepods or water-fleas.

many of marine animals. large jellyfish and related forms. usually marine. and the breathing organs are very diverse. . and the minute PROTOZOA. number of and of great diversity. all aquatic and mostly marine. of minor importance so far as distribution is concerned.230 3. as well as parasites. because of the delicate combs by means of which they swim. including spiders. and such free-living forms as Rotifers. often of great beauty . all of somewhat more complicated structure (ScYphozoa) and finally those delicate iridescent bells which float in the open waters of the ocean. Among them we have sea-firs and their allies. of radiate form. Among worm-like types we have also many parasitic forms. scorpions. star- (Asteroids). corals. often with limy skeletons. sea-cucumbers (Holothuroids). alcyonarians. The sponges. and constitute the Ctexophora. brittle-stars (Ophiuroids). .. including both the sea-worms and the earthworms. &c. form phylum of very simple animals. constitute the PHYLUM PORIFERA. The fish living classes comprise the sea-urchins (Echinoids). OUTLINE CLASSIFICATION OF ANIMALS Arachnoidea. organisms. The PHYLUM ECHINODEEMA includes a great number crab. The PHYLUM COELENTERA importance in that a large many parasitic forms are included. the king- They have no antennae. The Phylum of of ANNELIDS includes all the different kinds segmented worms. and sea-lilies (Crinoids). of great includes a great aquatic. and the fresh-water hydra (Hydromedusae) the seaanemones.

40. 213. sea. Acrobates. 119. Alaska. seal. 175. 213. 23. 98.3. 123. 218. 213. Bactrian camel. Alces. Agulhas. 142. Amblystoma. Arabian. Aelurus. a3. 1. &3. 129. Arctic America. 114. 59. 60. 98. 223.23. South America. 215. 69. 128. . 32. 122. Circle. 119.27. 106. 5)0. 55. North. 128. 196. Ants. 194. 54. 118. 205. Adansonia. 166. Eastern. Ateles. 112. 178. 204-5. 210. Antilocapra. 119. 105. 78. 107. Acorn shells. 49. 21. 98. Agutis. 52. 86. 129. Agriophylluni. Sea of. 7. 26. 122. 215. 179. 11. 106-8. Addax. driver. Andes Mts. 55. Armadillos. 62. 151. Acacias..i3. 84. 101.INDEX Aard-wolf. 168. Anodon. 12. Australia. vark. :M. Amoebae. 119. 1. 21. 214. 88. 217. 154. 65. 34. 126. 136. 115. 139. 128. 142. 98. 3. 178. KiS. 199. 2. 1. 207. 110. South. 80. tropical. 192. 181. 214 . Bali. 128. 213. 200. wild. 46. 119. 111. 133. 182. 100. i. Acanthephyra. Antarctic area. 162. 220-5. 1. 200-24. 124. South-western. 81. 86. 5(). 87. 217. 129. 40. 15. 179-81. tropical. Baobab. 40. 111. 118. 125. 11. of lakes. 192. 147. 9. 62. of. 173. Anabas. great. Alps. 163. 124. 44. 224. continent.16. 32. 10. 188. 146. 210. .^3. 132. 122. 111. 56. 117. 86. 165. 47. Anaconda. Accentor. 142. 210. Anaspides. 112. 115. 72. 106. 9. 161. 82. 111. 182. 13. 219. Anomaluridae. 173. 88. AJepocephalus. 88. 87. saiga. 103.30. Arctogaeic realm. thorny. 168. 69. Aniphipods. Astacus.Malayan region Axolotls. 13.38. 210. 8. 184. 172. water-buck. Balaena. 207. Auk. 123. 134. 149. 107. 11. 117. 80. Ass. 81. 128. 16. 106. dog-faced. 115. Baboons. Argentine. 106. 170. 107.3. African sable. 124. 18. 158.59. 89. 214. America. Abysfal fauna of sea. 112. 213. 19. 12<). 13. 52. 83. 188. Anthropoid apes. Africa. 224. 191. 61. 115. 132. 128. 12. 22. 165. 139. 107. 15. 177. liXi. 80-3. Aepvceros. Apes. 157. 101. Asia. 194. Arvicola. 61. Arabia.. Aquila. 34. 225. 55. Amu Daria. Amazon. R. Artemia. 82. 56. Central. 80. 128. 218. 59.i840.50. 182. Aniblyopsis. Adelops. 102. 90. Ant-eaters. 206. 169. 1. Atlantic. Azores. 133. 204. 129. Badger. Battln Land. Xi. African lakes. 129. 191. harnessed. Australian region. Lake. Alcyonarians. 215. American continent. 210. 126. 218. 116.?2. 89. 178. 215. 163. Central. Balaenoptera. 38.^. Asia. 109. 132. Apteryx. 195. 216. little. 32.3. 21. 201. 43. 200. 22. Argali. 88. Algae. 167. 100. 121. 98. 9. 129. 22. 126-40. 136. 129. Baikal. !«. 219. 54. 204. 90. Amphibians. Antelope. 132. 123. 84. Annelids. 99. wolf. 40.5. i:«. 74. 60.64. 160. Austro . Alca. . South. Alpine regions. 126. 142. 130. West. 163. 149. 16. Aphrodite.Africa. 20. 77. 217. South-eastern. 110. 106. 88. scaly. 42. 127. hare. 100-25. 163. 197. 208. 82. . 50. 153.5. 94. 62.5. 82. 145-52. hairy. 113. Bahamas. Arctomys. 195. Central. 112. 19. Babbling thrushes. 50. Asio. 61. Antechinoinvs. 215. waters. island of. 107. 131. Lake. i. African. 135. pala. 102. 58.34. 188. see Arctogaea. 110. 117. 209. 158. banded. 163. 62. 135. 49. 84. 87. 175. 214. Argyropclecus. two-toed. Arctictis. Alactaga. Eiistern. 130. 140. Alders. 130. North America. 12.25. North. 206. 210. 98. 147. 31. 51. 211. 72. steppes 218. 33. 131. 102. Babirusa.39. 68. 46. Angola.5. 132. black. Artemisia. 24. Balaton. Aral. Alashan. 58. Abyssinia. Acipenser. im. . 153. 21. 85. Northern. 22. 223. 182. Aniblyrhynchus. fox. 58. 32. 69. Atlas Mts. regions. 139.. 22.

184. 115. Chipmunks. 134. 223. Conurinac. 152. 106. 82. 84. Bear. Cercoleptes. 219. Bonasa. 222. 179. Cholydra. 148. 87. 41. 196. 218. 185. 116. 223. 222. 141. 152. Colies. 104. Birds. 193. 180. 79. brown. 94. Corals. 218. Ceylon. 200. frizzly. 1. 107. [alayan. 177. Congo basin. 118. 80. 113. Crinoids. Carnivores. 127. K). 100. Crabs. 221. . Crowberries. Chlamydophorus. 13. 146. 1. 220. 96. 106. 9. Crocidura. 159.^ave faunas. 224. 91. 133. water. Cockatoos. Bivalves. 45. 124. 44. 76. of. 200. Bull-frog. 106. INDEX L'accabis. 166. island Bustards. Crane. -221. 1-26.3. 147. 47. 193. 218. 45. 107. 195. 58. Central Europe. 169. 203. Bees. Carniola. Boa. 220. Belg^ium. 107. 110. Cre. 40. 216.!'!. 224. 154. Cetacea. 98. 85. 80.232 Barnaclets. 104. Centetes. 205. 81. Connochoetes. 147. Caspian Sea. 110. parti. horse-shoe. 19. Chili. 42. 191. 107. Bithynia. 47. 130. 68. island of. 19. 126. sloth. Climbing perch. 222. 69. 67. Cactus wren. Chamois. Corsica. 216. 219. Coryphaena. 89. 218. Bog moss. 224. 146. Benguela current. Upper. water. 90. 184. Chatterei-s. Caucasus. Convoluta. 207. 220. 203. 103. 205. 220. 89. 87. 132. 38. 116. Cavy. <'helonia. 219. 88. 42. 209. Chenopodiaceae. Brazil. Cape of Hope. 162. 216. 88. 168. Bar of Biscay. 224. 206. 83. 142. 182. 158. 222. 152. Cattle. i:». 205. Isles. 107. Chevrotains. 42. Cricetus. 134. Conolophus. 99. 193. 217. 99. Cariama. Bath ypelagic animal. 206. 149. Bulbuls. Cladonia. 85. 68. 44. 141. Cariacus. Cracidae. Cryptobranchus. Capra. 78. Cod. 141. Crayfish. 198. Clouded leopard. 169. 125. 153. 106. 72. 181. 113. 119. 219. 88. Capercaillie. California. Corvus. 170. 47. 49. 221.5. Chirogale. 221. 115. Copepods. 83. Coatis. Canis. 141. 116. 192. Burma. Canachites. Coelentera. 84. China. 182. 58. 140. R. 103. cat. Bathyscia. Blessbok.. Cemas. 161. 206. Chough-thrushes. Brahmaputra. 102. 61. 217. 87. 127. Ceratodus. Cercopithecus. 105. 79. 212. 180. Cervus. Bats. Alpine. 165. Capuchin monkevs. liermit. Calainoicthvs. 98. Colobus. Carinthia. 168. 106. Cervulus. 181. Chironomus. 67. 196. Canada. 207. Ctenomys. Crocodiles. 224. 147. 134. 28. Chough. Bonito. 134. 38. 193. C'himarrogale. 29. short-horned. 221. IM. 135. 167. 106. 85. 181. 218. 61. Cembra 49. Butterflies. 42. 77. Castor. 89. 219. 222. 201. 150. 162. 152. 126. Chimpanzee. 167. Chiru. 199. 220. Choloepus. 217. Beaver. 203. 151.coloured. Crossbill. 61. 105. 60. Cats. Chrvsomitris. 204. Bos. 207. 80. 78. Covpu. (.18. 179. 221. Cactus. 183. 47. 27. 180. 157. 167. Good Cichlidae. 206. Cephalophus. 116. 81. Bontcbok. 43. 114. of paradise. Beetles. 63. 65.52. Camel. 222. 75. British 154. Colombia. 222. fox. 63. tree. Civets. 170. 124. 49. Curas-sows. Black Sea. Canadian lakes. 193. Bobac. 130. 127. 155. 80. Carp. 218. ^aInpire. 110. Coenolestes. Chinchilla. 221. 44. 40. 217. 140. 165. 102. 193. 44. Condor. fruit. Centrocercus. Capreolus. 85. Borneo. 225. Chickaree. 119. 45. 117. 217. 31. 19. 81. 80. 222.205. 109. 38. 117. 42. Buru. Ccrthidea. Buffalo. Coliidae. Cryptoprocta. 81. Campylorhyncbus. 96. Cranberries. 89. 174. 38. Bird cherry.sted cockatoos. 28. 69. 80. palm. Bonv pike.?. black. Cape Colony. Cat-fish. Brittle-star. 113. Continental Slope. 47. Coelogenys. Bubalis. Bermudas. Bolivia. Brine-shrimps. 59. . 220. 123. 151. 115. 9. 106. Ctenophora. 218. 194. 196. Celebes. Continental Islands. 1. Chelone. 218.32. 38. Cassowary. 81. spectacled. 110. 163. 38. 165. 67. Bradypus. Crustacea. 160. 192. Bering Strait. 102. 66. 107. 11. 110. 206. 148. 134. Bufo. Cockles. 69. 168. 18. 177. insect-eating. Chameleon.30. Bison. 160. Centetidae. 40 see Deer. 23. 82. 68. Capivara. 167. pine. 45. 134. 16. 105. 102. rice. pygmies. 83. 120. Corixa. Bower-birds.

196. 146. 9:1. Dingo. i:iO. 175. 80. Gobies. Gecko. 185. 1:54. 122. 190. Cuttles. 177. 70. Cynopithecus. Delphinaptcrus. (58. 117. i. 37. Eel. snow. 70. 79. 192. 22. 24. Roedeer. Ganoids. Halolimnic fauna. 58. 43. 199. 147. Galeopithecu. Globigerina. 47. Goat. Darwin. 11(5. Cystophora. 78. 106. Deer. flying. 219. 26. 89. Finch. Demoiselle crane. or Ornitho- Flying lemur. 147. 72. 181. 201. 186. 23. Diatoms. 145-51. 220. 66. Halobates. 199. 217. 225. 167. Frog. 136. Gopher. Dabs. 69. 16. 216. flying. I )endrobatinae. 178. Falco. 200. 41. Duck. 44. 146. 42. Grinnell Land. Hangnests. 222. 192. Cape hunting. 37. 200. 181. 218. 206. 24.JO. 217. 222. Pallas's sand. (58. 88 also Polar hare. 184. 140. 9:5. Rocky Mts. :32.218. 40. 70. 16:5. Geese. 126. Geospiza. 98. deep-sea. 177. 116. 16.s. 44. 162. 181. 47. K. 224. Danube. Halicore. 88.. red. 71. long-eared. hawk. wild. Flycatchers. 49.s. Galago. 195. Dolichonyx. 23. 115. Eryx. 218. 170. 218. 185.. Gemsbok. 84.35. 129. 217. 216. 151. 8:3. 199. 22. Deinnark. 218. Grosbeak. 109. Cyuoiuys. 159. 16-29. Dolphin. ttsh-eating. Diuco.5. 219.w. Goniostoma. European Russia. 12(5.5. 118.219. Desert of Gobi. 224.30. Fox. Asiatic. 84. 24. 182. 115. 22. Duikcrboks. Gavialis. Elk. 68. Dalmatia. Virginian. 107. 21. 223. 202. 18. Rein218. 223. 39. hazel. 145. 214. Dzungarian Arctic. 131. 9. 44. 178. 76. Galapagos Islands.see . Canadian. 192. 207. fresh-water. 83. 166. 132. (ioral. 217. 47. Stream. 106. 210. Gannet. Gulf of Mexico. 42. 89. 115. i:J9. . 128. Fiber. Firth of Forth. 2(16. 131. England. 117. 118. 178. 42. 84. Gorilla. Felidao. 199. Grouse. Dugong. 164. Fish. 79. 233 Ganges. 117. 167. Echinoderms. 160. ISJ. 151. 110. 224. 128. 129. 219. Euneccs. Eremias. 200. 222. 89. 117. Euphorbia. Grasshopper. 47. 175. Grus. 220. 194. 178. 122. 216. Golden. 130. Arctic. 88. black. Cyclothone. 119. 211. 37. Gibbon. 26. 166. 18:5. 1. Diprotodont marsupials. Gazelle. fresh-water.. 149. American. . Dipus. 18:5. Eagle. 210. willow. Desman. 70. 174. Guillemots. Indian wild. Northern. 108. Friendly Islands. Great Salt Lake. Eland. '£i. nmd. Dodo. 102. Fruit-pigeon. 40. Elephant. Dog. ItM. UW. 47. 191. citril. 104. tree. 120. Dasyprocta. ruffed. 55. 219. Europe. Equus. Gudgeon. Dryopliis. 110. Gate. Germany. Glutton. 47. !«. (>4. 46. 83. 66. 46. Gulo. Hairmoss. 119. m. (59. Eastern. Exocoetus. Flies. 50. 59. 150. 40. 71. 225. Greenland. 204. 89. 151. Emvs. (59. Echidna. Ecuador. Arabian. 84. 8:5. 177. Graculus. Dinornis. Kskimo. Gerfalcon. Hare. Dipnoi. 85. p]rethizon. 199. 79. Gnu. 189. (Golden mole. 69. 200. . Greece. 176. 75. 218. cave. Guanaco. 2£i. Edentates. 152. 90. 125. Dormouse. 223. 69. deer. ttshing. 194. 16. Gypaetus. 59. 224. 166. 64. . 170. 162. 80. 217. 1(5:5. 40. l«(i. EljTuus. 178. corsac. 67. Guinea-fowl. 201. 205.218. 223. 219. Eurasiatic continent. 49. (Central. 1&5. see Eskimo. 27. 216. 13. 27. 113. 176. 70. IWi. 192. Dasypus. !«. 62. Fiji. Eider-duck. 98. 1(52. Didelphys. Ethiopian region. 157. (53. blind. Glass crab. 24. 176. 23. 104. 191. 47. R. 216. 147. Emu. 199. 19. Duck-mole rhynchus. 8. Hamster. 63. 40. l(i(j. 203. South. 88. Gecinus. :ii. 168. Dendrophis. 45. Ermine. 189. 117. 217. 167. 28. 129. 42. 58.INDEX CuiicwH. Fells. ISMi. 1. 214. F'ranz Josef Land. 20. 215. 128. Giraffe. Gastropods. 202. Haddock. pampa. 40. 67. 177. 159. 99. 217. rod. 75.

210. leaf. :Macaws. 6. Lobster. Honey Honey creepei-s. 20. tree. 58. Kangaroo. five-toed. Mackenzie K. Lithocranius. Tibetan. 115. 104. 117. 86. 82. 40. Insectivores. 83. 126. 129. 136. 102. 146. 165. 87. 158. 80. 88. of Ken- Jacamars. 204. 219. Jumping shrew. Arctic. Jaguar. Leopard. lynx. Larch. 78. Hydromedusae. 213. Kilimanjaro. Lepus. eaters. Indies. Lagidium. 129. five-toed. Llamas. Mackerel. Kalahari desert. 165. -224. 148. 11. 88. 149. -22. Lake. 220. 116. 22. Leven. 72. 214. steppe. 79. Khingan Kiang. 57.3. . 189. 108. 60. 220. 84. im. Manatee. fishes. Lagomys. 80. Isopods. 116. 88. 102. 90. 219. 218. 155. 160. Hill-tits.. 20. 162. 108. 142. Lophyrus. -222.5. 125. 8<t. Java. 56. 55. 174. 101. 191. 118. 216. 116.65. Kudu. 124. R.50. Hungary. 11. ring-tixiled. 215. tailless. 224. 22.5.5. 1. !Mts. VSi. 163. Mammoth. 136. 221. 135. West. Hjort. 170. :Macaques. 2l«. Lumpsucker. 11.sec also 212. 196. Indus. l. Ictonyx. 118. Linota. 215.. Jellyfish. 215.3. 162. 90. Labrador. 220. Iceland. Hydrophis. 49. •22(t. Malagasj. 44. Koala. 1. Humming-birds. 222. Leiotrichidae. 105. 101. 124. -223. 80.36. 78. 219. Sli. 72. 130. 209. Heterocephalus. Indian Ocean. 1*2. 2ii. Lobodon. 123. Ireland. Langur. 49. 134. 224.3. Lemming. 152. 222. 217. Tibetan. 25. 78. 207. 77. 165. i:J5-8. 21. Insects. 161. Iguana. 146. Hesperomys. 102. 20. Kirghiz. 196. 21. 78. 66. Lark. 218. 106. 176. VSi. 220. :Marmosets.65. 208. 162. 202. 213. 172. Macrorhinus. 153. 111. 88. 221. 193. Lyre-birds. 94. 117 142. 215. 217. 124. Lombok. 199. 78. 112. 217. 22. 222. 89. 148. King Oscar Land. 24. Limnocnifla. Jungle fowl. Macacus. Lion. 189. Limpet. Lung-fishes. Locusts. 59. fauna. 1. 16. 78. 60. 66. Lagopus. 16. 213. 83. 182. Hermit crabs.234 Hare.'J6. 123. 83. 221. Lemur. 76. Hipfjopotamus. Indo-Malayan region. 165. 147. Klipspringer.5. Hoatzin. 68. Hyrax. Ladak. 219. 6t. 62. Jumping mouse. 216. Icteridae. Lcptonyehotes. Koi'dofan desert. 102. Lutraria. 1S)0. 216. 77. 12. Leptocephalus. 164. Lyrurus.5. 115. Herring. Siberian. 94. 166.. 16ti. 10. 115-19. Magpie. 22. 201. 222. UK. 109. 168. 22:5. 1. 47. flying. 180. 112. Further.5. Lynx. Hartebeests. 16(5. birds.152. Holarctic region. Peninsula. 66. 65.5. mountain. 2:5. Lobelias. VM. 80-9. Laminarian zone. river. Semnopithecus. 128. 149. 62. 20. 215. 180. 85. L. 219. flying. Hypsiprymnodon. 220. 114. Kansu. 95. 128.61.scar. 218. 216. giant. 47. 71. 93. annelids. Littoral animals. m. wart. marine. Kiwi. 167 Himalayas. 213. Jerboa. 111. ilarkhor. Macroscelides. 109. Madaga. 176. 197. jumping. 217. Siberian. 104-7. . mouse. 37. 177. -221. 105. Kamchatka. 1. rat. 69. Hvla. 166. Kea. 167. 27. 2S.^3. 49. Lagostomus. 78. 216. 105. 100. 87. 219. 16. 155. 119. 127. 116. 188. region. 152. 130. 171. 219. tailless. India. 161. Macrurus. 49. Hedgehog. 1. Lacerta. Jackal.3. Juniper. Kinkajou. 79. 219. INDEX Japan. 69. 114. 78. 122. 111. Himalayan. 208. :Maral stag. Hylidae. Manunoth Cave 152. Lycaon. 77. 131. 169. Hydra. 12!).5. 77. 102. Hubara. Herpetodrvas.. 8. Hyaenas. 100. :j8. 153. 1-29. 126. Kuku Ibex. Lepidosiren. 47. 151. 221. 22. \oi-. 191. Hemitragus. 114. Horses. 210. 1-23. Loggerhead turtle. 224. Malay Archipelago. 99. 212. 117. 207. Norwegian. Hyperodon. 111. Jays. 101. 68. 167. 179. India. 5t\ Loxia. Hatteria. Limnaea. Hog. 219. 162. 124. 77. 222. 161. Kittiwake. 215. 1-20. 134. 155. 118. 224. 140.region.219. 199. 217. 153. 217. 219. Manis. 192. Laemmergeier. 125. 88. Hydrochoerus. 170. 218. fresh-water. Lerwa. 222. 2ti4. area. Lizard. 153. 104. 138. 105. tucky. 22l>-2. 166. 151. 64. 38. 66. 196.38.

161. 81. Oryx. 106. 209. 31. 178. 16. 176. Nyctea. Peccaries. lft3. 150. HI. Greek. 216. 88. 215. Mussel. 147. 128. 3. Otter. Periwinkles. 170. 11. 27. 170. Pangolins. 16. 41-51. 214. Ornithorhynchus. Montifringilla. 12. 221. Meliphagidae. Orang. 11. 64. Mediterranean. 110. 167. 220. Xeotropieal Palaearctic region. 179. 88. 223.INDEX Marmot. 58. Monkey. 215. 195. 113. Moloch. 221. 87. Patagonia. 98. 49. Medusoids. 67. Palms. 105. 10. 146. 199. 224. 215. Morar. 11. 83. 191. 218. 224. 140. Okapi. Xilgiri Hills. 223. 198. Ounce. 100. fish. 27. 87. Xectarinidae. 107. 205. 20. 135. HI. 77. Norway. Pacas. 140. 27. 90. 125. 58. Xansen. 151. 219. snow. . Muntjaes. 88. Monodon. wood. 217. Mountain shrimp. 114. 112. Prairie. Partridge. Periophthalmus. 196. Xyasa. 107.>().34.'IS. Marsupial mole. 46. 26. 222. 139. 184. 132. Xueifraga. 64. 165. 216. Moa. 86. 220. 107. 106. 9. 218. 85. 25. Mesopotamia.5. 75. 214. 87. 128. 180. 118. spider. Orca. 79. 45. 11. Marten. Mt. Persian gazelle. 85. 22. 89. Oreotragus.3. Marsupials. 102. Himalayan. 217. 1. Piehieiago. Parakeets. 167. 216. 49. Myrmecophaga. Notogaeic realm. 76. Opisthocomus. Xisaetus. XutcracKers. Mouse. 15. 168. \m. Xearctic . Mozambique. 190. 139. 145. 62. 72. 220.iS. 118. 191. 222. Phyllirhoe. 224. Xasua. region. 218. Petanrus. 170. 177.34. 42. 161. Mound Xotoryctes. Panda. 27. L. Xew Zealand. Mongolian gazelle. Pamirs. Otis. Xeoturus. 162. V. 158.176. 167. 12. 221. Meles. L. 139. N'arwhal. Pantholops. Nekton. 19. Perisoreus. 17. 221. 218. Mosquitoes. 177. Pig-footed bandicoot. Paraguay. 49. 110. 217. 214. 8. 223. 117. 134. 216. 21.. 13. 235 197 Musquash. 89. 219. 215. turkeys. Petrels. 134. Pelagic 214.. 56. 198. 100. 146. •209. larvae. 72. 61. Minx. 94. 57. 100. . broad-nosed. 19(5. 44. Palaeotropical region.3. 222. 217. X'^orth America. 224. 55. Xordenskiold. Pelias. 195. 218. howling. 20. 193. 223. 219. Mycetes. 214. 82. 167. 219. 225. Oxen. Orcyteropus. 122. 47. Melursus. Xoetiluca. Xew Siberian Islands.region. Papio. :\lolluscs. Xewfoiindland. 15:3. 46. 175. 18. 81. .. 174.57. IDS. 15. Mephitis. 45. 129.212. Paradoxurus.163. 223. Meerkat.194. 216. 59. Mountain ash". 90.5. Mudskippers. Palalo worm. Ovis. Ostrich. 44. 222. 94. Xorthern Russia.5. Parrots. Medusa. 225. 197. 224. 173-80. 16. Mirhacl Sars Expedition.s. 180. IKi. 224. 17. Musk ox.s. 216. 97-100. Owl parrot. 66. Mississippi. Pedioecetes. Mysis. 220. 83. Mole. Mice. 87. Picas. 100. region. 84. 208. 135. Xemorhoedus. Montieola. Xess. 217. 112. Oriental region. Xorth At]antic. 69. 78. 161. Ommatoiihoca. 212. 21. 66. 225. 119. 167. Myogale. 175. 134. Myodes. 223. Xestor. Owl. Nova iiembla. deer. 27. 181. Alpine. 115. Perch. 63. 24. Ovibos. 31. 214. Musophagidae. Polea. 216. Pacific Ocean. 217. 197. 133. 88. 90. Muscieapidae. 140. 89. 10. Xewts. 44. proboseis. 30. 38. Penguins. 225. 170. 161-7. 22. 93. 2U7. 224.. 123. Pampas. 100.j. 216. Myriapods. Opossum. 67.114. 95. 199. 87. Kockv 23. 69. 82. 93. Mesoiiekton. Pigeon. 198. Mongolia. Mongoose. 124. 207. 213. 138. i:i3. Xereis. 209. Xorth Sea. Pelagothuria. 79. L. 177. Otaria. Persia. thumble. N'asalis. 191. 224. Xorth Pacittc. 195. 191. 41. 219. Mustela. 184. 224. 159. 161. 41. 41. 84. 204. 106. Otocyon. 222. 220. Norwegian Sea. 28. Micropus. Microchiroptera. 23. 167. Oceanic islands. R. Mosehus. 1. Mosoplankton. 162. Mexico. 213. Myopotamus. 84. Mus. Mya. 34. . Motmots. 217. 220. 177. 102. Monotreme. 157. Petauroides. Pelagic animals. Xorthern Asia. 9(j. Melanocorvpha. 142. 194. 224.9. 46. 68. Xew Guinea. Pedetes. 118. 197. Kenia. 38. 59. 55. 43. 81. 224. Mountain. 100. shrew. 178. Myrmecobius.3.

184. Phacoehoerus. 90. 60. 61. 223. 127. 216. 218. 38. Sardinia. 33. 151. 59. 163. 133-6. 193. Phrynocephalus. -otter. 106. 90. 108. KB. 215. zoological. 20. 83. 114. 219. 118. -cow. -elephant. 42 . 176. Protopterus. 24. Potamogale. 19. 218. 79. Roach. 217. 184. 112. 169. Sechwan. 19. Rattlesnake. 162. 27. Russian Turkestan. 129. 168. 220 221. Seal. 126. 180. Sahara. 175. Ruwenzori. Salamander. Punjab. Pine grosbeak.l(»4. Pteromys. 33. Puma. 81. -leopard. 41. 200. 205. Queensland. 204. 94. Scorpion. 176. 147. Phvllostomatidae. 161. 182. 69. -snake. Philippine Islands. Running . 85. 20. 203. 166. 75. 166. 219. 94. 159. -hen. Prejevalski's horse. 192. 69. 216. 202. 113. 43. 199. 220. 125. cock. 124. 198. Sambar. 178. 193. 198. 222. Pyrosoma. 174. 112. 212. 20. Pond snails. Redpoll. 198. 132. Rhinolophidae. flying. 150. 96. 185. 107. 79. 175. 89. 78. 41. 90. Pipefish. 129. 26. 26. Realms. 170. River-hog. Planorbis. ringed. Prairie-chicken.217. 84.5. 19. 177. 82. 40. 217. 222. 206. Polvchaete worms. Rhinoceros. 82. 63. 32. Sandpiper. 33. 204. 117. 198. 72. bearded. 189. 189. -pen. 43. Secretary bird. 80 82 84 44. Pycnodontidae. . 33. Reitbok. 221. 108. 178. 107. 184. Saxicava. 221-3. 24. 125. 183. 219. Semnopithecus. 192. 182. 89. 73. 90. 152. 184. Porcupine. Phvtoplankton. INDEX Proteles. Podoces. 108. Scincus. 75. Phascolarctos. 49. 97. Pleetrophenax. Roedeer. 217. 184. 31. 76. 126. Plovers. 218. 175. 27. 173. 42. 142. 140. 151. 112. 41. Sand-rat. 78. 224. Plankton. Polynesia. 87. 76.236 Pigs. 69. 107. 120. 30. 131. 121. 199. 25. 184. 153. Pinicola. moss. negritos. 192. as. Primates. Pudua. tree. 214. Puflins. 142. Rice-bii-d. 161. Sandwich Islands. Prongbuck. 110. 80. 89. 69. 220. Rangifer. Pteropods. 218. 49. 79. Pisidium. crested. Greenland. 84. 223. 167. see also Hare. 139. 21. Planulae. ^^'eddelrs. fur.\ge-brush.153. 93. 136. Canadian. 151. Pholas. 147. 164. 11. 216. 117. 161. Pyrenees. Rodents. 107. 163. Selvas. Scotland. Pine saw-fly. 116. Sea-anemone. 167. Salmon. 142. 89. 69. 223. Sciurus. Rorqual. 31. 222. Pythons. Potamochoerus. 169. Pygmv hippopotamus. Seals. wUd. 68. Phoca. 166. S. 216. 113. Phvllodaetylus. 107. 33. Sebastes. 18. Serpentarius. 169. -butterfly. Red Sea. 39. 177. Rana. 109. 64-7. 115. Raccoons. -squirt. 70. 133. Potorous. Reptiles. Rupicapra. 118. 162. 58. Bighorn. 120. Polar bear. Rotifers. 142. 44. 197. Sai-gasso Sea. Arctic. Scandinavia. 174. Sheep. 127. 136. 118. Prawns. 110. 181. 137. 192. 201. Phrynosoma. 166. Ptarmigan. 72. 110. Polytrichum. crab-eating. 72. 59. 16. 196. 219. Rhebok. 70. Saiga antelope. Sayansk divide. hare. Proechidna. 67. 112. earless. 159. 82. 209. 164. 176. Phyllornithidae. 50. 170.' 202. 162. Porpoises. Platanista. 21 see Reindeer. 161. 91. 221. Quagga. -snake. \M. 158. Rhacophorus. 219. 134. 220. Rock thrush. birds. lakes of. 86. 47. Rat. 45. 88. 84. 140. 115. 124. Pike. bush. 82. 218. 60. 136. 32. Sarcorhampus. 86. -horse. 46. 203. 175. 220. 66. 162. 201. Plaice. Polecat. 219. Rosss. 94. 78. 216. 217. Phalangers. Barbary. 187. also Raven. Seriemas. 33. 82. 165. 58. 163. Pheasant. 191. Plateaux. Sciuropterus. Southern. Protozoa. -fir. 177. 218. 68. 80. -urchin. 191. 67. 224. 49. Plat j'rrhine monkeys. 30. 69. 205. 168. 223. 223. 78. Rhea. Rock lobster. 84. 108. 42. 105. 1*2. 90. Rocky Mountains. Plantain-eaters. Rock conev. 173. 23. 223. 149. 77. 17. 82. 78. Reindeer. 26. 106. 24. Salpa. Polvpterus. 189. 142. Russia. Physeter. Pine-marten. Sagitta. 190. 42. 161. Serpula. 55. 177. 93. Poecilogale. 146. 100. 141. 218. 128. Proteus. 141. Senecios. Serow.

Trypanosoines.3. Tai-sipes. 206. 1!W. 119. Squirrel. 20. 219. 62. TeratoscincuH.Siberian fir. Tucotucos. 105.56. 1. 27. 222. 85. 13. 93. 108. 31. 106. 110. 89. 1(52.50. Takin. Sponges. water. 104. 190.50. nmsk. . 1.33. 181. 168. 218. Stickleback.3. 1!M5. 147. 16. 214. 207. VU. 7. 207.5. 123. Texas. 125. 70. 112. 87. 87-9. 142. 4. 222. 224. 110. 21. Switzerland. 85. Vultures. Tyrant shrikes. 60. hemisphere.). Tasnifinian devil. Tinamus. 217. Syrrhaptes. 199. American snapping. Snipe. 88. 61. 8. 192. South America. 114. 78. 3. "Victoria. Terebella. 221. 4. 62. 108. wood.5ft. 224. whip. 57. 89. 222. 142. 72. Tarsier. Tropidonotus. Silurus. 222. giant. Sterlet. Sumatra. 86. 164. i:30. 223. 208.37. Shrews. 42. 82. 112. 198. 187.5. 131.3. 80. 78.5. Upper Egypt. Struthio. Tibet. 206. 174. Springbok. as. 95. 5. 62. 90. i:U. loggerhead. Tragelaphus. 1.3. 153. li:j-23. 46. 129. 116. Turkestan. Sind. 4fi. 26. 217. 213. Spotted cuscus. Syntheres. flvmg. hawksbill. 43. 213. 166. Stcinbok. Skate. 219. 76. Skua. 149.5. 127. 106. 220. 13. 208. 1-2. 44. Trigonocephalus. 108. 10.30. fresh-water. i:{. 115.5. 87. 37. -heath. 185. 21(5. Sturgeon.35. 142. Tree-creeper. 49. 80. Swift. 216. Titanethes. 44. 296. i:«). 192. 218. 70.34. 28. 77. 24. France. 113. Alpine. Siren. m. 104. Wallace. . 1. 125. lo'J. Skunk. Tichodroma. 149. Thynnus. 19:5. 130-4. Tupaia. -shrew. tree. 116. . Lake. 124. 124. Sunflsh. 105.5:i. 64. 174. 18. 148. 218. Wallabies. Viper. 16J1. 13&-47. 18. Sus. 110. Toucans. 127. 74. im. 218. fresh-water. Syria. 68. 8. im. Tympanuchus. 44. Southern England. 165. water.5. Turbot. Tardigrada. Unio. horned. 117. 117. 219. leathery. 194. 221. 6. 214.54. lai. 2. Wallace's line. 219. 112. 12S. 19. 148. Uropsilus. 194. 190. . 19. 69. 1. 237 Toad. 200-5. Trochospheres. i:». 191. - Tahr. Snail. Trigonia. 16. 90. 216. 69. Thalassochelys. -cock. 152.34. Alpine. 81. Strepsiptera. 2:1 Termites. 26. Tanagers. Tarsius. 104. 32. 12J).5. 175. 130. Taxidea. Ursus. 217. 222. 193.52.35. Syr Daria. 120. 165. Sonialiland. Starling. Spain. 42. 16. tree. 218. 107. 74. 152 154. Sloth. 123. 16. 56. 108. 104. 79. 120. 112. 111. 90. 209. 219. 11. 44. 89. 150. . 67. Ticks. l(i<). 136. flying. 76. 147. 47. 165. 193. 102. 222.5. sea. 22:}. Sun-birds.3. 137. 71. Voles. 204. 105. 1. Tench. 83. 22:}. Tetrao. 111. Tamias. 117. 194. 45.5. 197. 162. 1. 80-4. 121. 16. flying. 112. 89. •Spermophilus. 149. 86. Snake. 37. 202. wolf. . Tiger. 180. United States. Tasmania. Victoria Land. Talpa. 223. Siberia. 218. 7. 12. &i. Spur-rowl. swimming. 22. 218. . 2(M). . Tarim Ungulates. Tyrol.Sirex. Strepsiceros. basin. 43.{.Sphagnum. 177. 42. 117. Solitaire. Tenrec. 214. . Tortoise. 146. 142. 16.Snow-bunting. 217. ground. 47. Spi<lers. 1. 219. Suslik. Tunicates. 21. 112. 84. Viverridae. 1. 194. Tipula. 225. 184. Tropidurus. 147.5. Skink.5.). 117. Sverdrup Expedition. 108. Vicuna. 79. 98. 102. Himalavaii 78. Ifio. . Stringops. 220. 102. 58. . 218. 175. 221. 59. 21. 200. 110. -frog. Tsetse flies. 55. 112. 9. 1.5. 195.5. 128. Sudan. Tetrastes. 97. Tibetan mole. 75. 107. 89. Starfish. 182. 220. Tapir. Viscacha. 131. 65. 120. Turtle.INDEX Slu'll-fi. 1. 151. Sirenia. -22.jumping. Tanganyika. 24.'$8. .5. Sorex. green. 49. 175. Timelidac. Snowy owl. 220. Typhlomolge. 131. 199. Tctraogallus. 19. 109. 24. Terns. Spitsbergen. Stoat. 222. Stcnorhvnchus. pygmy sugar. Trichechus. (»). Trimeresurus. 85. 141. 220. grass. 101.3. 27 Solenodon. Suricata. 2(!. Tailless hedgehog. 58.tO. 39. tree. pond. marine. 1. Tomopteris. Tragulus.

Water Greenland.238 Wall-creeper. West 221. 193. . 197. 46. 39. 50. 24. Walrus. Zebra. 198. 34. 152. bottle-nose. 159. Wart-hogs. shrew. green. 219. 101. vole. Whortleberries. wasp. three-toed. 32. killer. 162. 60. 162. 191. 19. 176. 124.%3. 47. 198. Wolverene. Whales. 127. beetles. Whelks. 20. spotted. 184. Weasel. Willows. 33. dog. Wryneck. 162. 197. Wild cat. 47. fleas. 44. 23. 167. 47. boatmen. 30. 147. 107. sperm. Welwitschia. 167. 46. 45. l. chevrotain. 109. 31. Wolf. 44. 34. whalebone. 37. 214. 176. 40. spider. INDEX 89. 107. bears. 132. 48. 30. Wood Yak. 47. 81. 163. 206. 32. 184. 44. 113. Indies. 37. Woodpeckers. 82. 176. white. Wapiti. 38. 191. 39. 31. boar. mites. 42. 198. 44.




y .-.

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