FRONTISPIECE

ISLAND LIFE

OR

THE PHENOMENA AND CAUSES OF

INSULAR FAUNAS AND FLORAS

INCLUDING A REVISION AND ATTEMPTED SOLUTION OF
THE PROBLEM OF

GEOLOGICAL CLIMATES

BY

ALFRED RUSSEL WALLACE

AUTHOR OF "THE MALAY ARCHIPELAGO," "THE GEOGRAPHICAL DISTRIBUTION OF ANIMALS,"
"DARWINISM," ETC.

SECOND AND REVISED EDITION

London

MACMILLAN AND CO.

AND NEW YORK

1895

The Right of Translation and Reproduction is Reserved

Richard Clay and Sons, Limited,
LONDON AND BUNGAY.

First Edition printed 1880 (Med. 8vo).
Second Edition 1892 (Extra cr. 8vo). Reprinted 1895.

TO

SIR JOSEPH DALTON HOOKER,

K.C.S.I., C.B., F.R.S., ETC., ETC.

WHO, MORE THAN ANY OTHER WRITER,

HAS ADVANCED OUR KNOWLEDGE OF THE GEOGRAPHICAL

DISTRIBUTION OF PLANTS, AND ESPECIALLY

OF INSULAR FLORAS,

I Dedicate this Volume;

ON A KINDRED SUBJECT,

AS A TOKEN OF ADMIRATION AND REGARD.

CORRECTIONS IN PRESENT ISSUE.

The first issue of this Edition being exhausted, the opportunity is taken of making a few corrections, the most important of which are here stated:—

Page 163. Statement modified as to supposed glaciation of South Africa.

Pages 174 and 338. Many geologists now hold that there was no great submergence during the glacial epoch. The passages referring to it have therefore been re-written.

Page 182. Colonel Fielden's explanation of the occurrence of large trees on shores and in recent drift in high latitudes, is now added.

" 272. A species of Carex peculiar to Bermuda is now given.

" 356. Geomalacus maculosus, as a peculiar British species, is now omitted.

Verbal alterations have also been made at pages 41, 105, 356, and 360.

PREFACE TO THE SECOND EDITION

This edition has been carefully revised throughout, and owing to the great increase to our knowledge of the Natural History of some of the islands during the last twelve years considerable additions or alterations have been required. The more important of these changes are the following:—

Chapter VII. The account of the migrations of animals and plants during and since the Glacial Epoch, has been modified to accord with newer information.

Chapters VIII and IX. The discussion of the causes of Glacial Epochs and Mild Arctic Climates has been somewhat modified in view of the late Dr. Croll's remarks, and the argument rendered clearer.

Chapter XIII. Several additions to the Fauna of the Galapagos have been noted.

Chapter XV. Considerable additions have been made to this chapter embodying the recent discoveries of birds and insects new to the Sandwich Islands, while a much fuller account has been given of its highly peculiar and very interesting flora.

Chapter XVI. Important additions and corrections have been made in the lists of peculiar British animals and plants embodying the most recent information.

Chapter XVII. Very large additions have been made to the mammalia and birds of Borneo, and full lists of the peculiar species are given.

Chapter XVIII. A more accurate account is given of the birds of Japan.

Chapter XIX. The recent additions to the mammals and birds of Madagascar are embodied in this chapter, and a fuller sketch is given of the rich and peculiar flora of the island.

Chapter XXI. and XXII. Some important additions have been made to these chapters owing to more accurate information as to the depth of the sea around New Zealand, and to the discovery of abundant remains of fossil plants of the tertiary and cretaceous periods both in New Zealand and Australia.

In the body of the work I have in each case acknowledged the valuable information given me by naturalists of eminence in their various departments, and I return my best thanks to all who have so kindly assisted me. I am however indebted in a special manner to one gentleman—Mr. Theo. D. A. Cockerell, now Curator of the Museum of the Jamaica Institute—who supplied me with a large amount of information by searching the most recent works in the scientific libraries, by personal inquiries among naturalists, and also by giving me the benefit of his own copious notes and observations. Without his assistance it would have been difficult for me to have made the present edition so full and complete as I hope it now is. In a work of such wide range, and dealing with so large a body of facts some errors will doubtless be detected, though, I trust few of importance.

Parkstone, Dorset, December, 1891.

PREFACE TO THE FIRST EDITION

The present volume is the result of four years' additional thought and research on the lines laid down in my Geographical Distribution of Animals, and may be considered as a popular supplement to and completion of that work.

It is, however, at the same time a complete work in itself: and, from the mode of treatment adopted, it will, I hope, be well calculated to bring before the intelligent reader the wide scope and varied interest of this branch of natural history. Although some of the earlier chapters deal with the same questions as my former volumes, they are here treated from a different point of view; and as the discussion of them is more elementary and at the same time tolerably full, it is hoped that they will prove both instructive and interesting. The plan of my larger work required that genera only should be taken account of; in the present volume I often discuss the distribution of species, and this will help to render the work more intelligible to the unscientific reader.

The full statement of the scope and object of the present essay given in the "Introductory" chapter, together with the "Summary" of the whole work and the general view of the more important arguments given in the "Conclusion," render it unnecessary for me to offer any further remarks on these points. I may, however, state

generally that, so far as I am able to judge, a real advance has here been made in the mode of treating problems in Geographical Distribution, owing to the firm establishment of a number of preliminary doctrines or "principles," which in many cases lead to a far simpler and yet more complete solution of such problems than have been hitherto possible. The most important of these doctrines are those which establish and define—(1) The former wide extension of all groups now discontinuous, as being a necessary result of "evolution"; (2) The permanence of the great features of the distribution of land and water on the earth's surface; and, (3) The nature and frequency of climatal changes throughout geological time.

I have now only to thank the many friends and correspondents who have given me information or advice. Besides those whose assistance is acknowledged in the body of the work, I am especially indebted to four gentlemen who have been kind enough to read over the proofs of chapters dealing with questions on which they have special knowledge, giving me the benefit of valuable emendations and suggestions. Mr. Edward R. Alston has looked over those parts of the earlier chapters which relate to the mammals of Europe and the North Temperate zone; Mr. S. B. J. Skertchley, of the Geological Survey, has read the chapters which discuss the glacial epoch and other geological questions; Professor A. Newton has looked over the passages referring to the birds of the Madagascar group; while Sir Joseph D. Hooker has given me the invaluable benefit of his remarks on my two chapters dealing with the New Zealand flora.

Croydon, August, 1880.

CONTENTS

PART I

THE DISPERSAL OF ORGANISMS; ITS PHENOMENA, LAWS, AND CAUSES

CHAPTER I

INTRODUCTORY

Remarkable Contrasts in the Distribution of Animals—Britain and Japan—Australia and New Zealand—Bali and Lombok—Florida and Bahama Islands—Brazil and Africa—Borneo, Madagascar, and Celebes—Problems in Distribution to be found in every Country—Can be Solved only by the Combination of many distinct lines of inquiry, Biological and Physical—Islands offer the best Subjects for the Study of Distribution—Outline of the Subjects to be discussed in the Present Volume.

Pages [3]-[11]

CHAPTER II

THE ELEMENTARY FACTS OF DISTRIBUTION.

Importance of Locality as an Essential Character of Species—Areas of Distribution—Extent and Limitations of Specific Areas—Specific Range of Birds—Generic Areas—Separate and Overlapping Areas—The Species of Tits as illustrating Areas of Distribution—The Distribution of the Species of Jays—Discontinuous Generic Areas—Peculiarities of Generic and Family Distribution—General Features of Overlapping and Discontinuous Areas—Restricted Areas of Families—The Distribution of Orders

Pages [12]-[30]

CHAPTER III

CLASSIFICATION OF THE FACTS OF DISTRIBUTION.—ZOOLOGICAL REGIONS

The Geographical Divisions of the Globe do not Correspond to Zoological Divisions—The Range of British Mammals as Indicating a Zoological Region—Range of East Asian and North African Mammals—The Range of British Birds—Range of East Asian Birds—The Limits of the Palæarctic Region—Characteristic Features of the Palæarctic Region—Definition and Characteristic Groups of the Ethiopian Region—Of the Oriental Region—Of the Australian Region—Of the Nearctic Region—Of the Neotropical Region—Comparison of Zoological Regions with the Geographical Divisions of the Globe

Pages [31]-[54]

CHAPTER IV

EVOLUTION AS THE KEY TO DISTRIBUTION

Importance of the Doctrine of Evolution—The Origin of New Species—Variation in Animals—The amount of Variation in North American Birds—How New Species Arise from a Variable Species—Definition and Origin of Genera—Cause of the Extinction of Species—The Rise and Decay of Species and Genera—Discontinuous Specific Areas, why Rare—Discontinuity of the Area of Parus Palustris—Discontinuity of Emberiza Schœniclus—The European and Japanese Jays—Supposed examples of Discontinuity among North American Birds—Distribution and Antiquity of Families—Discontinuity a Proof of Antiquity—Concluding remarks

Pages [55]-[71]

CHAPTER V

THE POWERS OF DISPERSAL OF ANIMALS AND PLANTS

Statement of the General Question of Dispersal—The Ocean as a Barrier to the Dispersal of Mammals—The Dispersal of Birds—The Dispersal of Reptiles—The Dispersal of Insects—The Dispersal of Land Mollusca—Great Antiquity of Land-shells—Causes Favouring the Abundance of Land-shells—The Dispersal of Plants—Special Adaptability of Seeds for Dispersal—Birds as Agents in the Dispersal of Seeds—Ocean Currents as Agents in Plant Dispersal—Dispersal along Mountain Chains—Antiquity of Plants as Effecting their Distribution

Pages [72]-[82]

CHAPTER VI

GEOGRAPHICAL AND GEOLOGICAL CHANGES: THE PERMANENCE OF CONTINENTS

Changes of Land and Sea, their Nature and Extent—Shore-Deposits and Stratified Rocks—The Movements of Continents—Supposed Oceanic

Formations; the Origin of Chalk—Fresh-water and Shore-deposits as Proving the Permanence of Continents—Oceanic Islands as Indications of the Permanence of Continents and Oceans—General Stability of Continents with Constant Change of Form—Effect of Continental Changes on the Distribution of Animals—Changed Distribution Proved by the Extinct Animals of Different Epochs—Summary of Evidence for the General Permanence of Continents and Oceans.

Pages [83]-[105]

CHAPTER VII

CHANGES OF CLIMATE WHICH HAVE INFLUENCED THE DISPERSAL OF ORGANISMS: THE GLACIAL EPOCH

Proofs of the Recent Occurrence of a Glacial Epoch—Moraines—Travelled Blocks—Glacial Deposits of Scotland: the "Till"—Inferences from the Glacial Phenomena of Scotland—Glacial Phenomena of North America—Effects of the Glacial Epoch on Animal Life—Warm and Cold Periods—Palæontological Evidence of Alternate Cold and Warm Periods—Evidence of Interglacial Warm Periods on the Continent and in North America—Migrations and Extinctions of Organisms Caused by the Glacial Epoch

Pages [106]-[124]

CHAPTER VIII

THE CAUSES OF GLACIAL EPOCHS

Various Suggested Causes—Astronomical Causes of Changes of Climate—Difference of Temperature Caused by Varying Distances of the Sun—Properties of Air and Water, Snow and Ice, in Relation to Climate—Effects of Snow on Climate—High Land and Great Moisture Essential to the Initiation of a Glacial Epoch—Perpetual Snow nowhere Exists on Lowlands—Conditions Determining the Presence or Absence of Perpetual Snow—Efficiency of Astronomical causes in Producing Glaciation—Action of Meteorological Causes in Intensifying Glaciation—Summary of Causes of Glaciation—Effect of Clouds and Fog in Cutting off the Sun's Heat—South Temperate America as Illustrating the Influence of Astronomical Causes on Climate—Geographical Changes how far a Cause of Glaciation—Land Acting as a Barrier to Ocean-currents—The Theory of Interglacial Periods and their Probable Character—Probable Effect of Winter in aphelion on the Climate of Britain—The Essential Principle of Climatal Change Restated—Probable Date of the Last Glacial Epoch—Changes of the Sea-level Dependent on Glaciation—The Planet Mars as Bearing on the Theory of Excentricity as a Cause of Glacial Epochs

Pages [125]-[168]

CHAPTER IX

ANCIENT GLACIAL EPOCHS, AND MILD CLIMATES IN THE ARCTIC REGIONS

Mr. Croll's Views on Ancient Glacial Epochs—Effects of Denudation in Destroying the Evidence of Remote Glacial Epochs—Rise of Sea-level Connected with Glacial Epochs a Cause of Further Denudation—What Evidence of Early Glacial Epochs may be Expected—Evidences of Ice-action During the Tertiary Period—The Weight of the Negative Evidence—Temperate Climates in the Arctic Regions—The Miocene Arctic Flora—Mild Arctic Climates of the Cretaceous Period—Stratigraphical Evidence of Long-continued Mild Arctic Conditions—The Causes of Mild Arctic Climates—Geographical Conditions Favouring Mild Northern Climates in Tertiary Times—The Indian Ocean as a Source of Heat in Tertiary Times—Condition of North America During the Tertiary Period—Effect of High Excentricity on Warm Polar Climates—Evidences as to Climate in the Secondary and Palæozoic Epochs—Warm Arctic Climates in Early Secondary and Palæozoic Times—Conclusions as to the Climates of Secondary and Tertiary Periods—General View of Geological Climates as Dependent on the Physical Features of the Earth's Surface—Estimate of the Comparative Effects of Geographical and Physical Causes in Producing Changes of Climate.

Pages [169]-[209]

CHAPTER X

THE EARTH'S AGE, AND THE RATE OF DEVELOPMENT OF ANIMALS AND PLANTS

Various Estimates of Geological Time—Denudation and Deposition of Strata as a Measure of Time—How to Estimate the Thickness of the Sedimentary Rocks—How to Estimate the Average Rate of Deposition of the Sedimentary Rocks—The Rate of Geological Change Probably Greater in very Remote Times—Value of the Preceding Estimate of Geological Time—Organic Modification Dependent on Change of Conditions—Geographical Mutations as a Motive Power in Bringing about Organic Changes—Climatal Revolutions as an Agent in Producing Organic Changes—Present Condition of the Earth One of Exceptional Stability as Regards Climate—Date of Last Glacial Epoch and its Bearing on the Measurement of Geological Time—Concluding Remarks

Pages [210]-[237]

PART II

INSULAR FAUNAS AND FLORAS

CHAPTER XI

THE CLASSIFICATION OF ISLANDS

Importance of Islands in the Study of the Distribution of Organisms—Classification of Islands with Reference to Distribution—Continental Islands—Oceanic Islands

Pages [241]-[245]

CHAPTER XII

OCEANIC ISLANDS:—THE AZORES AND BERMUDA

The Azores, or Western Islands

Position and Physical Features—Chief Zoological Features of the Azores—Birds—Origin of the Azorean Bird-fauna—Insects of the Azores—Land-shells of the Azores—The Flora of the Azores—The Dispersal of Seeds—Birds as seed-carriers—Facilities for Dispersal of Azorean Plants—Important Deduction from the Peculiarities of the Azorean Fauna and Flora

Pages [246]-[262]

Bermuda

Position and Physical Features—The Red Clay of Bermuda—Zoology of Bermuda—Birds of Bermuda—Comparison of the Bird-faunas of Bermuda and the Azores—Insects of Bermuda—Land Mollusca—Flora of Bermuda—Concluding Remarks on the Azores and Bermuda

Pages [263]-[274]

CHAPTER XIII

THE GALAPAGOS ISLANDS

Position and Physical Features—Absence of Indigenous Mammalia and Amphibia—Reptiles—Birds—Insects and Land-shells—The Keeling Islands as Illustrating the Manner in which Oceanic Islands are Peopled—Flora of the Galapagos—Origin of the Flora of the Galapagos—Concluding remarks

Pages [273]-[291]

CHAPTER XIV

ST. HELENA

Position and Physical Features of St. Helena—Change Effected by European Occupation—The Insects of St. Helena—Coleoptera—Peculiarities and Origin of the Coleoptera of St. Helena—Land-shells of St. Helena—Absence of Fresh-water Organisms—Native Vegetation of St. Helena—The Relations of the St. Helena Compositæ—Concluding Remarks on St. Helena

Pages [292]-[309]

CHAPTER XV

THE SANDWICH ISLANDS

Position and Physical Features—Zoology of the Sandwich Islands—Birds—Reptiles—Land-shells—Insects—Vegetation of the Sandwich Islands—Peculiar Features of the Hawaiian Flora—Antiquity of the Hawaiian Fauna and Flora—Concluding Observations on the Fauna and Flora of the Sandwich Islands—General Remarks on Oceanic Islands

Pages [310]-[330]

CHAPTER XVI

CONTINENTAL ISLANDS OF RECENT ORIGIN: GREAT BRITAIN

Characteristic Features of Recent Continental Islands—Recent Physical Changes of the British Isles—Proofs of Former Elevation—Submerged Forests—Buried River Channels—Time of Last Union with the Continent—Why Britain is Poor in Species—Peculiar British Birds—-Fresh-water Fishes—Cause of Great Speciality in Fishes—Peculiar British Insects—Lepidoptera Confined to the British Isles—Peculiarities of the Isle of Man Lepidoptera—Coleoptera Confined to the British Isles—Trichoptera Peculiar to the British Isles—Land and Fresh-water Shells—Peculiarities of the British Flora—Peculiarities of the Irish Flora—Peculiar British Mosses and Hepaticæ—Concluding Remarks on the Peculiarities of the British Fauna and Flora

Pages [331]-[372]

CHAPTER XVII

BORNEO AND JAVA

Position and Physical Features of Borneo—Zoological Features of Borneo: Mammalia—Birds—The Affinities of the Borneo Fauna—Java, its Position and Physical Features—General Character of the Fauna of Java—Differences Between the Fauna of Java and that of the other Malay Islands—Special Relations of the Javan Fauna to that of the Asiatic Continent—Past Geographical Changes of Java and Borneo—The Philippine Islands—Concluding Remarks on the Malay Islands

Pages [373]-[390]

CHAPTER XVIII

JAPAN AND FORMOSA

Japan, its Position and Physical Features—Zoological Features of Japan—Mammalia—Birds—Birds Common to Great Britain and Japan—Birds Peculiar to Japan—Japan Birds Recurring in Distant Areas—Formosa—Physical Features of Formosa—Animal Life of Formosa—Mammalia—Land Birds Peculiar to Formosa—Formosan Birds Recurring in India or Malaya—Comparison of Faunas of Hainan, Formosa, and Japan—General Remarks on Recent Continental Islands

Pages [391]-[410]

CHAPTER XIX

ANCIENT CONTINENTAL ISLANDS: THE MADAGASCAR GROUP

Remarks on Ancient Continental Islands—Physical Features of Madagascar—Biological Features of Madagascar—Mammalia—Reptiles—Relation of Madagascar to Africa—Early History of Africa and Madagascar—Anomalies of Distribution and how to Explain Them—The Birds of Madagascar as Indicating a Supposed Lemurian Continent—Submerged Islands Between Madagascar and India—Concluding Remarks on "Lemuria"—The Mascarene Islands—The Comoro Islands—The Seychelles Archipelago—Birds of the Seychelles—Reptiles and Amphibia—Fresh-water Fishes—Land Shells—Mauritius, Bourbon, and Rodriguez—Birds—Extinct Birds and their Probable Origin—Reptiles—Flora of Madagascar and the Mascarene Islands—Curious Relations of Mascarene Plants—Endemic Genera of Mauritius and Seychelles—Fragmentary Character of the Mascarene Flora—Flora of Madagascar Allied to that of South Africa—Preponderance of Ferns in the Mascarene Flora—Concluding Remarks on the Madagascar Group

Pages [411]-[449]

CHAPTER XX

ANOMALOUS ISLANDS: CELEBES

Anomalous Relations of Celebes—Physical Features of the Island—Zoological Character of the Islands Around Celebes—The Malayan and Australian Banks—Zoology of Celebes: Mammalia—Probable Derivation of the Mammals of Celebes—Birds of Celebes—Bird-types Peculiar to Celebes—Celebes not Strictly a Continental Island—Peculiarities of the Insects of Celebes—Himalayan Types of Birds and Butterflies in Celebes—Peculiarities of Shape and Colour of Celebesian Butterflies—Concluding Remarks—Appendix on the Birds of Celebes

Pages [450]-[470]

CHAPTER XXI

ANOMALOUS ISLANDS: NEW ZEALAND

Position and Physical Features of New Zealand—Zoological Character of New Zealand—Mammalia—Wingless Birds Living and Extinct—Recent Existence of the Moa—Past Changes of New Zealand deduced from its Wingless Birds—Birds and Reptiles of New Zealand—Conclusions from the Peculiarities of the New Zealand Fauna

Pages [471]-[486]

CHAPTER XXII

THE FLORA OF NEW ZEALAND: ITS AFFINITIES AND PROBABLE ORIGIN

Relations of the New Zealand Flora to that of Australia—General Features of the Australian Flora—The Floras of South-eastern and South-western Australia—Geological Explanation of the Differences of these Two Floras—The Origin of the Australian Element in the New Zealand Flora—Tropical Character of the New Zealand Flora Explained—Species Common to New Zealand and Australia mostly Temperate Forms—Why Easily Dispersed Plants have often Restricted Ranges—Summary and Conclusion on the New Zealand Flora

Pages [487]-[508]

CHAPTER XXIII

ON THE ARCTIC ELEMENT IN SOUTH TEMPERATE FLORAS

European Species and Genera of Plants in the Southern Hemisphere—Aggressive Power of the Scandinavian Flora—Means by which Plants have Migrated from North to South—Newly Moved Soil as Affording Temporary Stations to Migrating Plants—Elevation and Depression of the Snow-line as Aiding the Migration of Plants—Changes of Climate Favourable to Migration—The Migration from North to South has been Long going on—Geological Changes as Aiding Migration—Proofs of Migration by way of the Andes—Proofs of Migration by way of the Himalayas and Southern Asia—Proofs of Migration by way of the African Highlands—Supposed Connection of South Africa and Australia—The Endemic Genera of Plants in New Zealand—The Absence of Southern Types from the Northern Hemisphere—Concluding Remarks on the New Zealand and South Temperate Floras

Pages [509]-[530]

CHAPTER XXIV

SUMMARY AND CONCLUSION

The Present Volume is the Development and Application of a Theory—Statement of the Biological and Physical Causes of Dispersal—Investigation of the Facts of Dispersal—Of the Means of Dispersal—Of Geographical Changes Affecting Dispersal—Of Climatal Changes Affecting Dispersal—The Glacial Epoch and its Causes—Alleged Ancient Glacial Epochs—Warm Polar Climates and their Causes—Conclusions as to Geological Climates—How Far Different from those of Mr. Croll—Supposed Limitations of Geological Time—Time Amply Sufficient both for Geological and Biological Development—Insular Faunas and Floras—The North Atlantic Islands—The Galapagos—St. Helena and the Sandwich Islands—Great Britain as a Recent Continental Island—Borneo and Java—Japan and Formosa—Madagascar as an Ancient Continental Island—Celebes and New Zealand as Anomalous Islands—The Flora of New Zealand and its Origin—The European Element in the South Temperate Floras—Concluding Remarks

Pages [531]-[545]

MAPS AND ILLUSTRATIONS

ISLAND LIFE

PART I

THE DISPERSAL OF ORGANISMS

ITS PHENOMENA, LAWS, AND CAUSES

CHAPTER I

INTRODUCTORY

Remarkable Contrasts in distribution of Animals—Britain and Japan—Australia and New Zealand—Bali and Lombok—Florida and Bahama Islands—Brazil and Africa—Borneo, Madagascar, and Celebes—Problems in distribution to be found in every country—Can be solved only by the combination of many distinct lines of inquiry, biological and physical—Islands offer the best subjects for the study of distribution—Outline of the subjects to be discussed in the present volume.

When an Englishman travels by the nearest sea-route from Great Britain to Northern Japan he passes by countries very unlike his own, both in aspect and natural productions. The sunny isles of the Mediterranean, the sands and date-palms of Egypt, the arid rocks of Aden, the cocoa groves of Ceylon, the tiger-haunted jungles of Malacca and Singapore, the fertile plains and volcanic peaks of Luzon, the forest-clad mountains of Formosa, and the bare hills of China, pass successively in review; till after a circuitous voyage of thirteen thousand miles he finds himself at Hakodadi in Japan. He is now separated from his starting-point by the whole width of Europe and Northern Asia, by an almost endless succession of plains and mountains, arid deserts or icy plateaux, yet when he visits the interior of the country he sees so many familiar natural objects that he can hardly help fancying he is close to his home. He finds the woods and fields tenanted by tits, hedge-sparrows, wrens, wagtails, larks, redbreasts,

thrushes, buntings, and house-sparrows, some absolutely identical with our own feathered friends, others so closely resembling them that it requires a practised ornithologist to tell the difference. If he is fond of insects he notices many butterflies and a host of beetles which, though on close examination they are found to be distinct from ours, are yet of the same general aspect, and seem just what might be expected in any part of Europe. There are also of course many birds and insects which are quite new and peculiar, but these are by no means so numerous or conspicuous as to remove the general impression of a wonderful resemblance between the productions of such remote islands as Britain and Yesso.

Now let an inhabitant of Australia sail to New Zealand, a distance of less than thirteen hundred miles, and he will find himself in a country whose productions are totally unlike those of his own. Kangaroos and wombats there are none, the birds are almost all entirely new, insects are very scarce and quite unlike the handsome or strange Australian forms, while even the vegetation is all changed, and no gum-tree, or wattle, or grass-tree meets the traveller's eye.

But there are some more striking cases even than this, of the diversity of the productions of countries not far apart. In the Malay Archipelago there are two islands, named Bali and Lombok, each about as large as Corsica, and separated by a strait only fifteen miles wide at its narrowest part. Yet these islands differ far more from each other in their birds and quadrupeds than do England and Japan. The birds of the one are extremely unlike those of the other, the difference being such as to strike even the most ordinary observer. Bali has red and green woodpeckers, barbets, weaver-birds, and black-and-white magpie-robins, none of which are found in Lombok, where, however, we find screaming cockatoos and friar-birds, and the strange mound-building megapodes, which are all equally unknown in Bali. Many of the kingfishers, crow-shrikes, and other birds, though of the same general form, are of very distinct species; and though a considerable number of birds are the same in both islands the difference

is none the less remarkable—as proving that mere distance is one of the least important of the causes which have determined the likeness or unlikeness in the animals of different countries.

In the western hemisphere we find equally striking examples. The Eastern United States possess very peculiar and interesting plants and animals, the vegetation becoming more luxuriant as we go south but not altering in essential character, so that when we reach Alabama or Florida we still find ourselves in the midst of pines, oaks, sumachs, magnolias, vines, and other characteristic forms of the temperate flora; while the birds, insects, and land-shells are of the same general character with those found further north.[^[1]] But if we now cross over the narrow strait, about fifty miles wide, which separates Florida from the Bahama Islands, we find ourselves in a totally different country, surrounded by a vegetation which is essentially tropical and generally identical with that of Cuba. The change is most striking, because there is little difference of climate, of soil, or apparently of position, to account for it; and when we find that the birds, the insects, and especially the land-shells of the Bahamas are almost all West Indian, while the North American types of plants and animals have almost all completely disappeared, we shall be convinced that such differences and resemblances cannot be due to existing conditions, but must depend upon laws and causes to which mere proximity of position offers no clue.

Hardly less uncertain and irregular are the effects of climate. Hot countries usually differ widely from cold ones in all their organic forms; but the difference is by no means constant, nor does it bear any proportion to difference of temperature. Between frigid Canada and sub-tropical Florida there are less marked differences in the animal productions than between Florida and Cuba or Yucatan, so much more alike in climate and so much nearer together. So the differences between the birds and quadrupeds of temperate Tasmania and tropical North

Australia are slight and unimportant as compared with the enormous differences we find when we pass from the latter country to equally tropical Java. If we compare corresponding portions of different continents, we find no indication that the almost perfect similarity of climate and general conditions has any tendency to produce similarity in the animal world. The equatorial parts of Brazil and of the West Coast of Africa are almost identical in climate and in luxuriance of vegetation, but their animal life is totally diverse. In the former we have tapirs, sloths, and prehensile-tailed monkeys; in the latter elephants, antelopes, and man-like apes; while among birds, the toucans, chatterers, and humming-birds of Brazil are replaced by the plantain-eaters, bee-eaters, and sun-birds of Africa. Parts of South-temperate America, South Africa, and South Australia, correspond closely in climate; yet the birds and quadrupeds of these three districts are as completely unlike each other as those of any parts of the world that can be named.

If we visit the great islands of the globe, we find that they present similar anomalies in their animal productions, for while some exactly resemble the nearest continents others are widely different. Thus the quadrupeds, birds and insects of Borneo correspond very closely to those of the Asiatic continent, while those of Madagascar are extremely unlike African forms, although the distance from the continent is less in the latter case than in the former. And if we compare the three great islands Sumatra, Borneo, and Celebes—lying as it were side by side in the same ocean—we find that the two former, although furthest apart, have almost identical productions, while the two latter, though closer together, are more unlike than Britain and Japan situated in different oceans and separated by the largest of the great continents.

These examples will illustrate the kind of questions it is the object of the present work to deal with. Every continent, every country, and every island on the globe, offers similar problems of greater or less complexity and interest, and the time has now arrived when their solution can be attempted with some prospect of success. Many

years study of this class of subjects has convinced me that there is no short and easy method of dealing with them; because they are, in their very nature, the visible outcome and residual product of the whole past history of the earth. If we take the organic productions of a small island, or of any very limited tract of country, such as a moderate-sized country parish, we have, in their relations and affinities—in the fact that they are there and others are not there, a problem which involves all the migrations of these species and their ancestral forms—all the vicissitudes of climate and all the changes of sea and land which have affected those migrations—the whole series of actions and reactions which have determined the preservation of some forms and the extinction of others,—in fact the whole history of the earth, inorganic and organic, throughout a large portion of geological time.

We shall perhaps better exhibit the scope and complexity of the subject, and show that any intelligent study of it was almost impossible till quite recently, if we concisely enumerate the great mass of facts and the number of scientific theories or principles which are necessary for its elucidation.

We require then in the first place an adequate knowledge of the fauna and flora of the whole world, and even a detailed knowledge of many parts of it, including the islands of more special interest and their adjacent continents. This kind of knowledge is of very slow growth, and is still very imperfect;[^[2]] and in many cases it can

never now be obtained owing to the reckless destruction of forests and with them of countless species of plants and animals. In the next place we require a true and natural classification of animals and plants, so that we may know their real affinities; and it is only now that this is being generally arrived at. We further have to make use of the theory of "descent with modification" as the only possible key to the interpretation of the facts of distribution, and this theory has only been generally accepted within the last twenty years. It is evident that, so long as the belief in "special creations" of each species prevailed, no explanation of the complex facts of distribution could be arrived at or even conceived; for if each species was created where it is now found no further inquiry can take us beyond that fact, and there is an end of the whole matter. Another important factor in our interpretation of the phenomena of distribution, is a knowledge of the extinct forms that have inhabited each country during the tertiary and secondary periods of geology. New facts of this kind are daily coming to light, but except as regards Europe, North America, and parts of India, they are extremely scanty; and even in the best-known countries the record itself is often very defective and fragmentary. Yet we have already obtained remarkable evidence of the migrations of many animals and plants in past ages, throwing an often unexpected light on the actual distribution of many groups.[^[3]] By this means alone can we obtain positive evidence of the past migrations of organisms; and when, as too frequently is the case, this is altogether wanting, we

have to trust to collateral evidence and more or less probable hypothetical explanations. Hardly less valuable is the evidence of stratigraphical geology; for this often shows us what parts of a country have been submerged at certain epochs, and thus enables us to prove that certain areas have been long isolated and the fauna and flora allowed time for special development. Here, too, our knowledge is exceedingly imperfect, though the blanks upon the geological map of the world are yearly diminishing in extent. Lastly, as a most valuable supplement to geology, we require to know approximately, the depth and contour of the ocean-bed, since this affords an important clue to the former existence of now-submerged lands, uniting islands to continents, or affording intermediate stations which have aided the migrations of many organisms. This kind of information has only been partially obtained during the last few years; and it will be seen in the latter part of this volume, that some of the most recent deep-sea soundings have afforded a basis for an explanation of one of the most difficult and interesting questions in geographical biology—the origin of the fauna and flora of New Zealand.

Such are the various classes of evidence that bear directly on the question of the distribution of organisms; but there are others of even a more fundamental character, and the importance of which is only now beginning to be recognised by students of nature. These are, firstly, the wonderful alterations of climate which have occurred in the temperate and polar zones, as proved by the evidences of glaciation in the one and of luxuriant vegetation in the other; and, secondly, the theory of the permanence of existing continents and oceans. If glacial epochs in temperate lands and mild climates near the poles have, as now believed by men of eminence, occurred several times over in the past history of the earth, the effects of such great and repeated changes, both on the migration, modification, and extinction, of species, must have been of overwhelming importance—of more importance perhaps than even the geological changes of sea and land. It is therefore necessary to consider the evidence for these climatal changes;

and then, by a critical examination of their possible causes, to ascertain whether they were isolated phenomena, were due to recurrent cosmical actions, or were the result of a great system of terrestrial development. The latter is the conclusion we arrive at; and this conclusion brings with it the conviction, that in the theory which accounts for both glacial epochs and warm polar climates, we have the key to explain and harmonize many of the most anomalous biological and geological phenomena, and one which is especially valuable for the light it throws on the dispersal and existing distribution of organisms. The other important theory, or rather corollary from the preceding theory—that of the permanence of oceans and the general stability of continents throughout all geological time, is as yet very imperfectly understood, and seems, in fact, to many persons in the nature of a paradox. The evidence for it, however, appears to me to be conclusive; and it is certainly the most fundamental question in regard to the subject we have to deal with: since, if we once admit that continents and oceans may have changed places over and over again (as many writers maintain), we lose all power of reasoning on the migrations of ancestral forms of life, and are at the mercy of every wild theorist who chooses to imagine the former existence of a now-submerged continent to explain the existing distribution of a group of frogs or a genus of beetles.

As already shown by the illustrative examples adduced in this chapter, some of the most remarkable and interesting facts in the distribution and affinities of organic forms are presented by islands in relation to each other and to the surrounding continents. The study of the productions of the Galapagos—so peculiar, and yet so decidedly related to the American continent—appears to have had a powerful influence in determining the direction of Mr. Darwin's researches into the origin of species; and every naturalist who studies them has always been struck by the unexpected relations or singular anomalies which are so often found to characterize the fauna and flora of islands. Yet their full importance in connection with the history of the earth and its inhabitants has hardly yet

been recognised; and it is in order to direct the attention of naturalists to this most promising field of research, that I restrict myself in this volume to an elucidation of some of the problems they present to us. By far the larger part of the islands of the globe are but portions of continents undergoing some of the various changes to which they are ever subject; and the correlative proposition, that every portion of our continents has again and again passed through insular conditions, has not been sufficiently considered, but is, I believe, the statement of a great and most suggestive truth, and one which lies at the foundation of all accurate conception of the physical and organic changes which have resulted in the present state of the earth.

The indications now given of the scope and purpose of the present volume renders it evident that, before we can proceed to the discussion of the remarkable phenomena presented by insular faunas and floras, and the complex causes which have produced them, we must go through a series of preliminary studies, adapted to give us a command of the more important facts and principles on which the solution of such problems depends. The succeeding eight chapters will therefore be devoted to the explanation of the mode of distribution, variation, modification, and dispersal, of species and groups, illustrated by facts and examples; of the true nature of geological change as affecting continents and islands; of changes of climate, their nature, causes, and effects; of the duration of geological time and the rate of organic development.

CHAPTER II

THE ELEMENTARY FACTS OF DISTRIBUTION

Importance of Locality as an essential character of Species—Areas of Distribution—Extent and Limitations of Specific Areas—Specific range of Birds—Generic Areas—Separate and overlapping areas—The species of Tits as illustrating Areas of Distribution—The distribution of the species of Jays—Discontinuous generic areas—Peculiarities of generic and family distribution—General features of overlapping and discontinuous areas—Restricted areas of Families—The distribution of Orders.

So long as it was believed that the several species of animals and plants were "special creations," and had been formed expressly to inhabit the countries in which they are now found, their habitat was an ultimate fact which required no explanation. It was assumed that every animal was exactly adapted to the climate and surroundings amid which it lived, and that the only, or, at all events, the chief reason why it did not inhabit another country was, that the climate or general conditions of that country were not suitable to it, but in what the unsuitability consisted we could rarely hope to discover. Hence the exact locality of any species was not thought of much importance from a scientific point of view, and the idea that anything could be learnt by a comparative study of different floras and faunas never entered the minds of the older naturalists.

But so soon as the theory of evolution came to be generally adopted, and it was seen that each animal could only have come into existence in some area where ancestral

forms closely allied to it already lived, a real and important relation was established between an animal and its native country, and a new set of problems at once sprang into existence. From the old point of view the diversities of animal life in the separate continents, even where physical conditions were almost identical, was the fact that excited astonishment; but seen by the light of the evolution theory, it is the resemblances rather than the diversities in these distant continents and islands that are most difficult to explain. It thus comes to be admitted that a knowledge of the exact area occupied by a species or a group is a real portion of its natural history, of as much importance as its habits, its structure, or its affinities; and that we can never arrive at any trustworthy conclusions as to how the present state of the organic world was brought about, until we have ascertained with some accuracy the general laws of the distribution of living things over the earth's surface.

Areas of Distribution.—Every species of animal has a certain area of distribution to which, as a rule, it is permanently confined, although, no doubt, the limits of its range fluctuate somewhat from year to year, and in some exceptional cases may be considerably altered in a few years or centuries. Each species is moreover usually limited to one continuous area, over the whole of which it is more or less frequently to be met with, but there are many apparent and some real exceptions to this rule. Some animals are so adapted to certain kinds of country—as to forests or marshes, mountains or deserts—that they cannot, permanently, live elsewhere. These may be found scattered over a wide area in suitable spots only, but can hardly on that account be said to have several distinct areas of distribution. As an example we may name the chamois, which lives only on high mountains, but is found in the Pyrenees, the Alps, the Carpathians, in some of the Greek mountains and the Caucasus. The variable hare is another and more remarkable case, being found all over Northern Europe and Asia beyond lat. 55°, and also in Scotland and Ireland. In central Europe it is unknown till we come to the Alps, the Pyrenees, and the Caucasus, where it again appears. This is one of the best cases known of the

discontinuous distribution of a species, there being a gap of about a thousand miles between its southern limits in Russia, and its reappearance in the Alps. There are of course numerous instances in which species occur in two or more islands, or in an island and continent, and are thus rendered discontinuous by the sea, but these involve questions of changes in sea and land which we shall have to consider further on. Other cases are believed to exist of still wider separation of a species, as with the marsh titmice and the reed buntings of Europe and Japan, where similar forms are found in the extreme localities, while distinct varieties or sub-species, inhabit the intervening districts.

Extent and Limitations of Specific Areas.—Leaving for the present these cases of want of continuity in a species, we find the most wide difference between the extent of country occupied, varying in fact from a few square miles to almost the entire land surface of the globe. Among the mammalia, however, the same species seldom inhabits both the old and new worlds, unless they are strictly arctic animals, as the reindeer, the elk, the arctic fox, the glutton, the ermine, and some others. The common wolf of Europe and Northern Asia is thought by many naturalists to be identical with the variously coloured wolves of North America extending from the Arctic Ocean to Mexico, in which case this will have perhaps the widest range of any species of mammal. Little doubt exists as to the identity of the brown bears and the beavers of Europe and North America; but all these species range up to the arctic circle, and there is no example of a mammal universally admitted to be identical yet confined to the temperate zones of the two hemispheres. Among the undisputed species of mammalia the leopard has an enormous range, extending all over Africa and South Asia to Borneo and the east of China, and thus having probably the widest range of any known mammal. The winged mammalia have not usually very wide ranges, there being only one bat common to the Old and New Worlds. This is a British species, Vesperugo serotinus, which is found over the larger part of North America, Europe and Asia, as far

as Pekin, and even extends into tropical Africa, thus rivalling the leopard and the wolf in the extent of country it occupies.

Of very restricted ranges there are many examples, but some of these are subject to doubts as to the distinctness of the species or as to its geographical limits being really known. In Europe we have a distinct species of ibex (Capra Pyrenaica) confined to the Pyrenean mountains, while the true marmot is restricted to the Alpine range. More remarkable is the Pyrenean water-mole (Mygale Pyrenaica), a curious small insectivorous animal found only in a few places in the northern valleys of the Pyrenees. In islands there are many cases of undoubted restriction of species to a small area, but these involve a different question from the range of species on continents where there is no apparent obstacle to their wider extension.

Specific range of Birds.—Among birds we find instances of much wider range of species, which is only what might be expected considering their powers of flight; but, what is very curious, we also find more striking (though perhaps not more frequent) examples of extreme limitation of range among birds than among mammals. Of the former phenomenon perhaps the most remarkable case is that afforded by the osprey or fishing-hawk, which ranges over the greater portion of all the continents, as far as Brazil, South Africa, the Malay Islands, and Tasmania. The barn owl (Strix flammea) has nearly as wide a range, but in this case there is more diversity of opinion as to the specific difference of many of the forms inhabiting remote countries, some of which seem undoubtedly to be distinct. Among passerine birds the raven has probably the widest range, extending from the arctic regions to Texas and New Mexico in America, and to North India and Lake Baikal in Asia; while the little northern willow-wren (Phylloscopus borealis) ranges from arctic Norway across Asia to Alaska, and southward to Ceylon, China, Borneo, and Timor.

Of very restricted continental ranges the best examples in Europe are, the little blue magpie (Cyanopica cooki) confined to the central portions of the Spanish peninsula; and the Italian sparrow found only in Italy and Corsica.

In Asia, Palestine affords some examples of birds of very restricted range—a beautiful sun-bird (Nectarinea osea) a peculiar starling (Amydrus tristramii) and some others, being almost or quite confined to the warmer portions of the valley of the Jordan. In the Himalayas there are numbers of birds which have very restricted ranges, but those of the Neilgherries are perhaps better known, several species of laughing thrushes and some other birds being found only on the summits of these mountains. The most wonderfully restricted ranges are, however, to be found among the humming-birds of tropical America. The great volcanic peaks of Chimborazo and Pichincha have each a peculiar species of humming-bird confined to a belt just below the limits of perpetual snow, while the extinct volcano of Chiriqui in Veragua has a species confined to its wooded crater. One of the most strange and beautiful of the humming-birds (Loddigesia mirabilis) was obtained once only, more than forty years ago, near Chachapoyas in the Andes of northern Peru; and though Mr. Gould sent many drawings of the bird to people visiting the district and for many years offered a high reward for a specimen, no other has ever been seen![^[4]]

The above details will sufficiently explain what is meant by the "specific area" or range of a species. The very wide and very narrow ranges are exceptional, the great majority of species both of mammals and birds ranging over moderately wide areas, which present no striking contrasts in climate and physical conditions. Thus a large proportion of European birds range over the whole continent in an east and west direction, but considerable numbers are restricted either to the northern or the southern half. In Africa some species range over all the continent south of the desert, while large numbers are restricted to the equatorial forests, or to the upland plains. In North America, if we exclude the tropical and the arctic portions, a considerable number of species range over all the temperate parts of the continent, while still

more are restricted to the east, the centre, or the west, respectively.

Generic Areas.—Having thus obtained a tolerably clear idea of the main facts as to the distribution of isolated species, let us now consider those collections of closely-allied species termed genera. What a genus is will be sufficiently understood by a few illustrations. All the different kinds of dogs, jackals, and wolves belong to the dog genus, Canis; the tiger, lion, leopard, jaguar, and the wild cats, to the cat genus, Felis; the blackbird, song-thrush, missel-thrush, fieldfare, and many others to the thrush genus, Turdus; the crow, rook, raven, and jackdaw, to the crow genus, Corvus; but the magpie belongs to another, though closely-allied genus, Pica, distinguished by the different form and proportions of its wings and tail from all the species of the crow genus. The number of species in a genus varies greatly, from one up to several hundreds. The giraffe, the glutton, the walrus, the bearded reedling, the secretary-bird, and many others, have no close allies, and each forms a genus by itself. The beaver genus, Castor, and the camel genus, Camelus, each consist of two species. On the other hand, the deer genus, Cervus has forty species; the mouse and rat genus, Mus more than a hundred species; and there is about the same number of the thrush genus; while among the lower classes of animals genera are often very extensive, the fine genus Papilio, or swallow-tailed butterflies, containing more than four hundred species; and Cicindela, which includes our native tiger beetles, has about the same number. Many genera of shells are very extensive, and one of them—the genus Helix, including the commonest snails, and ranging all over the world—is probably the most extensive in the animal kingdom, numbering about two thousand described species.[^[5]]

Separate and Overlapping Areas.—The species of a genus are distributed in two ways. Either they occupy distinct areas which do not touch each other and are sometimes widely separated, or they touch and occasionally overlap

each other, each species occupying an area of its own which rarely coincides exactly with that of any other species of the same genus. In some cases, when a river, a mountain-chain, or a change of conditions as from pasture to desert or forest, determines the range of species, the areas of two species of the same genus may just meet, one beginning where the other ends; but this is comparatively rare. It occurs, however, in the Amazon valley, where several species of monkeys, birds, and insects come up to the south bank of the river but do not pass it, while allied species come to the north bank, which in like manner forms their boundary. As examples we may mention that one of the Saki monkeys (Pithecia monachus?) comes up to the south bank of the Upper Amazon, while immediately we cross over to the north bank we find another species (Pithecia rufibarbata?). Among birds we have the green jacamar (Galbula viridis), abundant on the north bank of the Lower Amazon, while on the south bank we have two allied species (Galbula rufoviridis and G. cyaneicollis); and among insects we have at Santarem on the south bank of the Amazon, the beautiful blue butterfly, Callithea sapphira, while almost opposite to it, at Monte-alegre, an allied species, Callithea Leprieuri is alone found. Perhaps the most interesting and best known case of a series of allied species, whose ranges are separate but conterminous, is that of the beautiful South American wading birds, called trumpeters, and forming the genus Psophia. There are five species, all found in the Amazon valley, but each limited to a well-marked district bounded by great rivers. On the north bank of the Amazon there are two species, one in its lower valley extending up to the Rio Negro; and the other in the central part of the valley beyond that river; while to the south of the Amazon there are three, one above the Madeira, one below it, and a third near Para, probably separated from the last by the Tocantins river.

Overlapping areas among the species of a genus is a more common phenomenon, and is almost universal where these species are numerous in the same continent. It is, however, exceedingly irregular, so that we often find one

species extending over a considerable portion of the area occupied by the genus and including the entire areas of some of the other species. So little has been done to work out accurately the limits of species that it is very difficult to give examples. One of the best is to be found in the genus Dendrœca, a group of American wood-warblers. These little birds all migrate in the winter into the tropical regions, but in the summer they come north, each having its particular range. Thus, D. dominica comes as far as the middle Eastern States, D. cœrulea keeps west of the Alleghanies, D. discolor comes to Michigan and New England; four other species go farther north in Canada, while several extend to the borders of the Arctic zone.

The Species of Tits as Illustrating Areas of Distribution.—In our own hemisphere the overlapping of allied species may be well illustrated by the various kinds of titmice, constituting the genus Parus, several of which are among our best known English birds. The great titmouse (Parus major) has the widest range of all, extending from the Arctic circle to Algeria, Palestine, and Persia, and from Ireland right across Siberia to the Ochotsk sea, probably following the great northern forest belt. It does not extend into China and Japan, where distinct species are found. Next in extent of range is the coal tit (Parus ater) which inhabits all Europe from the Mediterranean to about 64° N. latitude, in Asia Minor to the Lebanon and Caucasus, and across Siberia to Amoorland and Japan. The marsh tit (Parus palustris) inhabits temperate and south Europe from 61° N. latitude in Norway to Poland and South-west Russia, and in the south from Spain to Asia Minor. Closely allied to this—of which it is probably only a variety or sub-species—is the northern marsh tit (Parus borealis), which overlaps the last in Norway and Sweden, and also in South Russia and the Alps, but extends further north into Lapland and North Russia, and thence probably in a south-easterly direction across Central Asia to North China. Yet another closely-allied species (Parus camtschatkensis) ranges from North-eastern Russia across Northern Siberia to Lake Baikal and to Hakodadi in Japan, thus overlapping Parus borealis in the

western portion of its area. Our little favourite, the blue tit (Parus cœruleus) ranges over all Europe from the Arctic circle to the Mediterranean, and on to Asia Minor and Persia, but does not seem to pass beyond the Ural mountains. Its lovely eastern ally the azure tit (Parus cyaneus) overlaps the range of P. cœruleus in Western Europe as far as St. Petersburg and Austria, rarely straggling to Denmark, while it stretches all across Central Asia between the latitudes 35° and 56° N. as far as the Amoor valley. Besides these wide-ranging species there are several others which are more restricted. Parus teneriffæ, a beautiful dark blue form of our blue tit, inhabits North-west Africa and the Canaries; Parus ledouci, closely allied to our coal tit, is found only in Algeria; Parus lugubris, allied to the marsh tit, is confined to South-east Europe and Asia Minor, from Hungary and South Russia to Palestine; and Parus cinctus, another allied form, is confined to the extreme north in Lapland, Finland, and perhaps Northern Russia and Siberia. Another beautiful little bird, the crested titmouse (Parus cristatus) is sometimes placed in a separate genus. It inhabits nearly all Central and South Europe, wherever there are pine forests, from 64° N. latitude to Austria and North Italy, and in the west to Spain and Gibraltar, while in the east it does not pass the Urals and the Caucasus range. Its nearest allies are in the high Himalayas.

These are all the European tits, but there are many others inhabiting Asia, Africa, and North America; so that the genus Parus has a very wide range, in Asia to Ceylon and the Malay Islands, in Africa to the Cape, and in North America to the highlands of Mexico.

The Distribution of the Species of Jays.—Owing to the very wide range of several of the tits, the uncertainty of the specific distinction of others, and the difficulty in many cases of ascertaining their actual distribution, it has not been found practicable to illustrate this genus by means of a map. For this purpose we have chosen the genus Garrulus or the jays, in which the species are less numerous, the specific areas less extensive, and the species generally better defined; while being large and handsome

birds they are sure to have been collected, or at least noticed, wherever they occur. There are, so far as yet known, twelve species of true jays, occupying an area extending from Western Europe to Eastern Asia and Japan, and nowhere passing the Arctic circle to the north, or the tropic of Cancer to the south, so that they constitute one of the most typical of the Palæarctic[^[6]] genera. The following are the species, beginning with the most westerly and proceeding towards the east. The numbers prefixed to each species correspond to those on the coloured map which forms the frontispiece to this volume.

1. Garrulus glandarius.—The common jay, inhabits the British Isles and all Europe except the extreme north, extending also into North Africa, where it has been observed in many parts of Algeria. It occurs near Constantinople, but apparently not in Asia Minor; and in Russia, up to, but not beyond, the Urals. The jays being woodland birds are not found in open plains or barren uplands, and their distribution is hence by no means uniform within the area they actually occupy.

2. Garrulus cervicalis.—The Algerian jay, is a very distinct species inhabiting a limited area in North Africa, and found in some places along with the common species.

3. Garrulus krynicki.—The black-headed jay, is closely allied to the common species, but quite distinct, inhabiting a comparatively small area in South-eastern Europe, and Western Asia.

4. Garrulus atricapillus.—The Syrian jay, is very closely allied to the last, and inhabits an adjoining area in Syria, Palestine, and Southern Persia.

5. Garrulus hyrcanus.—The Persian jay, is a small species allied to our jay and only known from the Elburz Mountains in the north of Persia.

6. Garrulus brandti.—Brandt's jay, is a very distinct species, having an extensive range across Asia from the Ural Mountains to North China, Mandchuria, and the northern island of Japan, and also crossing the Urals into

Russia where it has been found as far west as Kazan in districts where the common jay also occurs.

7. Garrulus lanceolatus.—The black-throated jay, is a very distinct form known only from the North-western Himalayas and Nepal, common about Simla, and extending into Cashmere beyond the range of the next species.

8. Garrulus bispecularis.—The Himalayan jay is also very distinct, having the head coloured like the back, and not striped as in all the western species. It inhabits the Himalayas east of Cashmere, but is more abundant in the western than the eastern division, though according to the Abbé David it reaches Moupin in East Thibet.

9. Garrulus sinensis.—The Chinese jay, is very closely allied to the Himalayan, of which it is sometimes classed as a sub-species. It seems to be found in all the southern mountains of China, from Foochow on the east to Sze-chuen and East Thibet on the west, as it is recorded from Moupin by the Abbé David as well as the Himalayan bird—a tolerable proof that it is a distinct form.

10. Garrulus taivanus.—The Formosan jay is a very close ally of the preceding, confined to the island of Formosa.

11. Garrulus japonicus.—The Japanese jay is nearly allied to our common British species, being somewhat smaller and less brightly coloured, and with black orbits; yet these are the most widely separated species of the genus. According to Mr. Seebohm this species is equally allied to the Chinese and Siberian jays.

In the accompanying map (see frontispiece) we have laid down the distribution of each species so far as it can be ascertained from the works of Sharpe and Dresser for Europe, Jerdon for India, Swinhoe for China, and Mr. Seebohm's recent work for Japan. There is, however, much uncertainty in many places, and gaps have to be filled up conjecturally, while such a large part of Asia is still very imperfectly explored, that considerable modifications may have to be made when the country becomes more accurately known. But though details may be modified we can hardly suppose that the great features of the several specific areas, or their relations to each other

will be much affected; and these are what we have chiefly to consider as bearing on the questions here discussed.

The first thing that strikes us on looking at the map, is, the small amount of overlapping of the several areas, and the isolation of many of the species; while the next most striking feature is the manner in which the Asiatic species almost surround a vast area in which no jays are found. The only species with large areas, are the European G. glandarius and the Asiatic G. Brandti. The former has three species overlapping it—in Algeria, in South-eastern and North-eastern Europe respectively. The Syrian jay (No. 4), is not known to occur anywhere with the black-headed jay (No. 3), and perhaps the two areas do not meet. The Persian jay (No. 5), is quite isolated. The Himalayan and Chinese jays (Nos. 7, 8, and 9) form a group which are isolated from the rest of the genus; while the Japanese jay (No. 11), is also completely isolated as regards the European jays to which it is nearly allied. These peculiarities of distribution are no doubt in part dependent on the habits of the jays, which live only in well-wooded districts, among deciduous trees, and are essentially non-migratory in their habits, though sometimes moving southwards in winter. This will explain their absence from the vast desert area of Central Asia, but it will not account for the gap between the North and South Chinese species, nor for the absence of jays from the wooded hills of Turkestan, where Mr. N. A. Severtzoff collected assiduously, obtaining 384 species of birds but no jay. These peculiarities, and the fact that jays are never very abundant anywhere, seem to indicate that the genus is now a decaying one, and that it has at no very distant epoch occupied a larger and more continuous area, such as that of the genus Parus at the present day.

Discontinuous generic Areas.—It is not very easy to find good examples of genera whose species occupy two or more quite disconnected areas, for though such cases may not be rare, we are seldom in a position to mark out the limits of the several species with sufficient accuracy. The best and most remarkable case among European birds is

that of the blue magpies, forming the genus Cyanopica. One species (C. cooki) is confined (as already stated) to the wooded and mountainous districts of Spain and Portugal, while the only other species of the genus (C. cyanus) is found far away in North-eastern Asia and Japan, so that the two species are separated by about 5,000 miles of continuous land. Another case is that of the curious little water-moles forming the genus Mygale, one species M. muscovitica, being found only on the banks of the Volga and Don in South-eastern Russia, while the other, M. pyrenaica, is confined to streams on the northern side of the Pyrenees. In tropical America there are four different kinds of bell-birds belonging to the genus Chasmorhynchus, each of which appears to inhabit a restricted area completely separated from the others. The most northerly is C. tricarunculatus of Costa Rica and Veragua, a brown bird with a white head and three long caruncles growing upwards at the base of the beak. Next comes C. variegatus, in Venezuela, a white bird with a brown head and numerous caruncles on the throat, perhaps conterminous with the last; in Guiana, extending to near the mouth of the Rio Negro, we have C. niveus, the bell-bird described by Waterton, which is pure white, with a single long fleshy caruncle at the base of the beak; the last species, C. nudicollis, inhabits South-east Brazil, and is also white, but with black stripes over the eyes, and with a naked throat. These birds are about the size of thrushes, and are all remarkable for their loud, ringing notes, like a bell or a blow on an anvil, as well as for their peculiar colours. They are therefore known to the native Indians wherever they exist, and we may be the more sure that they do not spread over the intervening areas where they have never been found, and where the natives know nothing of them.

A good example of isolated species of a group nearer home, is afforded by the snow-partridges of the genus Tetraogallus. One species inhabits the Caucasus range and nowhere else, keeping to the higher slopes from 6,000 to 11,000 feet above the sea, and accompanying the ibex in its wanderings, as both feed on the same plants. Another

has a wider range in Asia Minor and Persia, from the Taurus mountains to the South-east corner of the Caspian Sea; a third species inhabits the Western Himalayas, between the forests and perpetual snow, extending eastwards to Nepal; while a fourth is found on the north side of the mountains in Thibet, and the ranges of these two perhaps overlap; the last species inhabit the Altai mountains, and like the two first appears to be completely separated from all its allies.

There are some few still more extraordinary cases in which the species of one genus are separated in remote continents or islands. The most striking of these is that of the tapirs, forming the genus Tapirus, of which there are two or three species in South America, and one very distinct species in Malacca and Borneo, separated by nearly half the circumference of the globe. Another example among quadrupeds is a peculiar genus of moles named Urotrichus, of which one species inhabits Japan and the other British Columbia. The cuckoo-like honey-guides, forming the genus Indicator, are tolerably abundant in tropical Africa, but there are two outlying species, one in the Eastern Himalaya mountains, the other in Borneo, both very rare, and recently an allied species has been found in the Malay peninsula. The beautiful blue and green thrush-tits forming the genus Cochoa, have two species in the Eastern Himalayas and Eastern China, while the third is confined to Java; the curious genus Eupetes, supposed to be allied to the dippers, has one species in Sumatra and Malacca, while four other species are found two thousand miles distant in New Guinea; lastly, the lovely ground-thrushes of the genus Pitta, range from Hindostan to Australia, while a single species, far removed from all its near allies, inhabits West Africa.

Peculiarities of Generic, and Family Distribution.—The examples now given sufficiently illustrate the mode in which the several species of a genus are distributed. We have next to consider genera as the component parts of families, and families of orders, from the same point of view.

All the phenomena presented by the species of a genus are reproduced by the genera of a family, and often in a more marked degree. Owing, however, to the extreme restriction of genera by modern naturalists, there are not many among the higher animals that have a world-wide distribution. Among the mammalia there is no such thing as a truly cosmopolitan genus. This is owing to the absence of all the higher orders except the mice from Australia, while the genus Mus, which occurs there, is represented by a distinct group, Hesperomys, in America. If, however, we consider the Australian dingo as a native animal we might class the genus Canis as cosmopolite, but the wild dogs of South America are now formed into separate genera by some naturalists. Many genera, however, range over three or more continents, as Felis (the cat genus) absent only from Australia; Ursus (the bear genus) absent from Australia and tropical Africa; Cervus (the deer genus) with nearly the same range; and Sciurus (the squirrel genus) found in all the continents but Australia. Among birds Turdus, the thrush, and Hirundo, the swallow genus, are the only perching birds which are truly cosmopolites; but there are many genera of hawks, owls, wading and swimming birds, which have a world-wide range.

As a great many genera consist of single species there is no lack of cases of great restriction, such as the curious lemur called the "potto," which is found only at Sierra Leone, and forms the genus Perodicticus; the true chinchillas found only in the Andes of Peru and Chili south of 9° S. lat. and between 8,000 and 12,000 feet elevation; several genera of finches each confined to limited portions of the higher Himalayas, the blood-pheasants (Ithaginis) found only above 10,000 feet from Nepal to East Thibet; the bald-headed starling of the Philippine islands, the lyre-birds of East Australia, and a host of others.

It is among the different genera of the same family that we meet with the most striking examples of discontinuity, although these genera are often as unmistakably allied as are the species of a genus; and it is these cases that furnish the most interesting problems to the student of distribution.

We must therefore consider them somewhat more fully.

Among mammalia the most remarkable of these divided families is that of the camels, of which one genus Camelus, the true camels, comprising the camel and dromedary, is confined to Asia, while the other Auchenia, comprising the llamas and alpacas, is found only in the high Andes and in the plains of temperate South America. Not only are these two genera separated by the Atlantic and by the greater part of the land of two continents, but one is confined to the Northern and the other to the Southern hemisphere. The next case, though not so well known, is equally remarkable; it is that of the Centetidæ, a family of small insectivorous animals, which are wholly confined to Madagascar and the large West Indian islands Cuba and Hayti, the former containing five genera and the latter a single genus with a species in each island. Here again we have the whole continent of Africa as well as the Atlantic ocean separating allied genera. Two families (or subfamilies) of rat-like animals, Octodontidæ and Echimyidæ, are also divided by the Atlantic. Both are mainly South American, but the former has two genera in North and East Africa, and the latter also two in South and West Africa. Two other families of mammalia, though confined to the Eastern hemisphere, are yet markedly discontinuous. The Tragulidæ are small deer-like animals, known as chevrotains or mouse-deer, abundant in India and the larger Malay islands and forming the genus Tragulus; while another genus, Hyomoschus, is confined to West Africa. The other family is the Simiidæ or anthropoid apes, in which we have the gorilla and chimpanzee confined to West and Central Africa, while the allied orangs are found only in the islands of Sumatra and Borneo, the two groups being separated by a greater space than the Echimyidæ and other rodents of Africa and South America.

Among birds and reptiles we have several families, which, from being found only within the tropics of Asia, Africa, and America, have been termed tropicopolitan groups. The Megalæmidæ or barbets are gaily coloured

fruit-eating birds, almost equally abundant in tropical Asia and Africa, but less plentiful in America, where they probably suffer from the competition of the larger sized toucans. The genera of each country are distinct, but all are closely allied, the family being a very natural one. The trogons form a family of very gorgeously coloured and remarkable insect-eating birds very abundant in tropical America, less so in Asia, and with a single genus of two species in Africa.

Among reptiles we have two families of snakes—the Dendrophidæ or tree-snakes, and the Dryiophidæ or green whip-snakes—which are also found in the three tropical regions of Asia, Africa, and America, but in these cases even some of the genera are common to Asia and Africa, or to Africa and America. The lizards forming the family Amphisbænidæ are divided between tropical Africa and America, a few species only occurring in the southern portion of the adjacent temperate regions; while even the peculiarly American family of the iguanas is represented by two genera in Madagascar, and one in the Fiji and Friendly Islands. Passing on to the Amphibians the worm-like Cæciliadæ are tropicopolitan, as are also the toads of the family Engystomatidæ. Insects also furnish some analogous cases, three genera of Cicindelidæ, (Pogonostoma, Ctenostoma, and Peridexia) showing a decided connection between this family in South America and Madagascar; while the beautiful family of diurnal moths, Uraniidæ, is confined to the same two countries. A somewhat similar but better known illustration is afforded by the two genera of ostriches, one confined to Africa and Arabia, the other to the plains of temperate South America.

General features of Overlapping and Discontinuous Areas.—These numerous examples of discontinuous genera and families form an important section of the facts of animal dispersal which any true theory must satisfactorily account for. In greater or less prominence they are to be found all over the world, and in every group of animals, and they grade imperceptibly into those cases of conterminous and overlapping areas which we have seen to

prevail in most extensive groups of species, and which are perhaps even more common in those large families which consist of many closely allied genera. A sufficient proof of the overlapping of generic areas is the occurrence of a number of genera of the same family together. Thus in France or Italy about twenty genera of warblers (Sylviadæ) are found, and as each of the thirty-three genera of this family inhabiting temperate Europe and Asia has a different area, a great number must here overlap. So, in most parts of Africa, at least ten or twelve genera of antelopes may be found, and in South America a large proportion of the genera of monkeys of the family Cebidæ occur in many districts; and still more is this the case with the larger bird families, such as the tanagers, the tyrant shrikes, or the tree-creepers, so that there is in all these extensive families no genus whose area does not overlap that of many others. Then among the moderately extensive families we find a few instances of one or two genera isolated from the rest, as the spectacled bear, Tremarctos, found only in Chili, while the remainder of the family extends from Europe and Asia over North America to the Mountains of Mexico, but no further south; the Bovidæ, or hollow-horned ruminants, which have a few isolated genera in the Rocky Mountains and the islands of Sumatra and Celebes; and from these we pass on to the cases of wide separation already given.

Restricted Areas of Families.—As families sometimes consist of single genera and even single species, they often present examples of very restricted range; but what is perhaps more interesting are those cases in which a family contains numerous species and sometimes even several genera, and yet is confined to a narrow area. Such are the golden moles (Chrysochloridæ) consisting of two genera and three species, confined to extratropical South Africa; the hill-tits (Liotrichidæ), a family of numerous genera and species mainly confined to the Himalayas, but with a few straggling species in the Malay countries and the mountains of China; the Pteroptochidæ, large wren-like birds, consisting of eight genera and nineteen species, almost entirely confined to temperate South America and

the Andes; and the birds-of-paradise, consisting of nineteen or twenty genera and about thirty-five species, almost all inhabitants of New Guinea and the immediately surrounding islands, while a few, doubtfully belonging to the family, extend to East Australia. Among reptiles the most striking case of restriction is that of the rough-tailed burrowing snakes (Uropeltidæ), the five genera and eighteen species being strictly confined to Ceylon and the southern parts of the Indian Peninsula.

The Distribution of Orders.—When we pass to the larger groups, termed orders, comprising several families, we find comparatively few cases of restriction and many of worldwide distribution; and the families of which they are composed are strictly comparable to the genera of which families are composed, inasmuch as they present examples of overlapping, or conterminous, or isolated areas, though the latter are comparatively rare. Among mammalia the Insectivora offer the best example of an order, several of whose families inhabit areas more or less isolated from the rest; while the Marsupialia have six families in Australia, and one, the opossums, far off in America.

Perhaps, more important is the limitation of some entire orders to certain well-defined portions of the globe. Thus the Proboscidea, comprising the single family and genus of the elephants, and the Hyracoidea, that of the Hyrax or Syrian coney, are confined to parts of Africa and Asia; the Marsupials to Australia and America; and the Monotremata, the lowest of all mammals—comprising the duck-billed Platypus and the spiny Echidna, to Australia and New Guinea. Among birds the Struthiones or ostrich tribe are almost confined to the three Southern continents, South America, Africa and Australia; and among Amphibia the tailed Batrachia—the newts and salamanders—are similarly restricted to the northern hemisphere.

These various facts will receive their explanation in a future chapter.

CHAPTER III

CLASSIFICATION OF THE FACTS OF DISTRIBUTION.—ZOOLOGICAL REGIONS

The Geographical Divisions of the Globe do not correspond to Zoological divisions—The range of British Mammals as indicating a Zoological Region—Range of East Asian and North African Mammals—The Range of British Birds—Range of East Asian Birds—The limits of the Palæarctic Region—Characteristic features of the Palæarctic Region—Definition and characteristic groups of the Ethiopian Region—Of the Oriental Region—Of the Australian Region—Of the Nearctic Region—Of the Neotropical Region—Comparison of Zoological Regions with the Geographical Divisions of the Globe.

Having now obtained some notion of how animals are dispersed over the earth's surface, whether as single species or as collected in those groups termed genera, families, and orders, it will be well, before proceeding further, to understand something of the classification of the facts we have been considering, and some of the simpler conclusions these facts lead to.

We have hitherto described the distribution of species and groups of animals by means of the great geographical divisions of the globe in common use; but it will have been observed that in hardly any case do these define the limits of anything beyond species, and very seldom, or perhaps never, even those accurately. Thus the term "Europe" will not give, with any approach to accuracy, the range of any one genus of mammals or birds, and

perhaps not that of half-a-dozen species. Either they range into Siberia, or Asia Minor, or Palestine, or North Africa; and this seems to be always the case when their area of distribution occupies a large portion of Europe. There are, indeed, a few species limited to Central or Western or Southern Europe, and these are almost the only cases in which we can use the word for zoological purposes without having to add to it some portion of another continent. Still less useful is the term Asia for this purpose, since there is probably no single animal or group confined to Asia which is not also more or less nearly confined to the tropical or the temperate portion of it. The only exception is perhaps the tiger, which may really be called an Asiatic animal, as it occupies nearly two-thirds of the continent; but this is an unique example, while the cases in which Asiatic animals and groups are strictly limited to a portion of Asia, or extend also into Europe or into Africa or to the Malay Islands, are exceedingly numerous. So, in Africa, very few groups of animals range over the whole of it without going beyond either into Europe or Asia Minor or Arabia, while those which are purely African are generally confined to the portion south of the tropic of Cancer. Australia and America are terms which better serve the purpose of the zoologist. The former defines the limit of many important groups of animals; and the same may be said of the latter, but the division into North and South America introduces difficulties, for almost all the groups especially characteristic of South America are found also beyond the isthmus of Panama, in what is geographically part of the northern continent.

It being thus clear that the old and popular divisions of the globe are very inconvenient when used to describe the range of animals, we are naturally led to ask whether any other division can be made which will be more useful, and will serve to group together a considerable number of the facts we have to deal with. Such a division was made by Mr. P. L. Sclater more than twenty years ago, and it has, with some slight modifications, come into pretty general use in this country, and to some extent also

abroad; we shall therefore proceed to explain its nature and the principles on which it is established, as it will have to be often referred to in future chapters of this work, and will take the place of the old geographical divisions whose inconvenience has already been pointed out. The primary zoological divisions of the globe are called "regions," and we will begin by ascertaining the limits of the region of which our own country forms a part.

The Range of British Mammals as indicating a Zoological Region.—We will first take our commonest wild mammalia and see how far they extend, and especially whether they are confined to Europe or range over parts of other continents:

We thus see that out of thirteen of our commonest quadrupeds only one is confined to Europe, while seven are found also in Northern Africa, and eleven range into Siberia, most of them stretching quite across Asia to the valley of the Amoor on the extreme eastern side of that continent. Two of the above-named British species, the fox and weasel, are also inhabitants of the New World, being as common in the northern parts of North America as they are with us; but with these exceptions the entire range of our commoner species is given, and they clearly show that all Northern Asia and Northern Africa must be added to Europe in order to form the region which they collectively inhabit. If now we go into Central Europe and take, for example, the quadrupeds of Germany, we shall find that these too, although much more numerous, are confined to the same limits, except that some of the

more arctic kinds, as already stated, extend into the colder regions of North America.

Range of East Asian and North African Mammals.—Let us now pass to the other side of the great northern continent, and examine the list of the quadrupeds of Amoorland, in the same latitude as Germany. We find that there are forty-four terrestrial species (omitting the bats, the seals, and other marine animals), and of these no less than twenty-six are identical with European species, and twelve or thirteen more are closely allied representatives, leaving only five or six which are peculiarly Asiatic. We can hardly have a more convincing proof of the essential oneness of the mammalia of Europe and Northern Asia.

In Northern Africa we do not find so many European species (though even here they are very numerous) because a considerable number of West Asiatic and desert forms occur. Having, however, shown that Europe and Western Asia have almost identical animals, we may treat all these as really European, and we shall then be able to compare the quadrupeds of North Africa with those of Europe and West Asia. Taking those of Algeria as the best known, we find that there are thirty-three species identical with those of Europe and West Asia, while twenty-four more, though distinct, are closely allied, belonging to the same genera; thus making a total of fifty-seven of European type. On the other hand, we have seven species which are either identical with species of tropical Africa or allied to them, and six more which are especially characteristic of the African and Asiatic deserts which form a kind of neutral zone between the temperate and tropical regions. If now we consider that Algeria and the adjacent countries bordering the Mediterranean form part of Africa, while they are separated from Europe by a wide sea and are only connected with Asia by a narrow isthmus, we cannot but feel surprised at the wonderful preponderance of the European and West Asiatic elements in the mammalia which inhabit the district.

The Range of British Birds.—As it is very important that no doubt should exist as to the limits of the zoological

region of which Europe forms a part, we will now examine the birds, in order to see how far they agree in their distribution with the mammalia. Of late years great attention has been paid to the distribution of European and Asiatic birds, many ornithologists having travelled in North Africa, in Palestine, in Asia Minor, in Persia, in Siberia, in Mongolia, and in China; so that we are now able to determine the exact ranges of many species in a manner that would have been impossible a few years ago. These ranges are given for all British species in the new edition of Yarrell's History of British Birds edited by Professor Newton, while those of all European birds are given in still more detail in Mr. Dresser's beautiful work on the birds of Europe. In order to confine our examination within reasonable limits, and at the same time give it the interest attaching to familiar objects, we will take the whole series of British Passeres or perching birds given in Professor Newton's work (118 in number) and arrange them in series according to the extent of their range. These include not only the permanent residents and regular migrants to our country, but also those which occasionally straggle here, so that it really comprises a large proportion of all European birds.

We find, that out of a total of 118 British Passeres there are: