Island Life; Or, The Phenomena and Causes of Insular Faunas and Floras

Proofs of Migration by way of the Himalayas and Southern Asia.—But although we may thus explain the presence of a considerable portion of the European element in the floras of New Zealand and Australia, we cannot account for the whole of it by this means, because Australia itself contains a host of European and Asiatic genera of which we find no trace in New Zealand or South America, or any other Antarctic land. We find, in fact, in Australia two distinct sets of European plants. First we have a number of species identical with those of Northern Europe or Asia (of the most characteristic of which—thirty-eight in number—Sir Joseph Hooker gives a list); and in the second place a series of European genera usually of a somewhat more southern character, mostly represented by very distinct species, and all absent from New Zealand; such as Clematis, Papaver, Cleome, Polygala, Lavatera, Ajuga, &c. Now of the first set—the North European species—about three-fourths occur in some parts of America, and about half in South Temperate America or New Zealand; whence we may conclude that most of these, as well as some others, have reached Australia by the route already indicated. The second set of Australo-European genera, however, and many others characteristic of the South European or the Himalayan flora, have probably reached Australia by way of the mountains of Southern Asia, Borneo, the Moluccas, and New Guinea, at a somewhat remote period when loftier ranges and some intermediate peaks may have existed, sufficient to carry on the migration by the aid of the alternate climatal changes which are known to have occurred. The long belt of Secondary and Palæozoic formations in East Australia from Tasmania to Cape York continued by the lofty ranges of New Guinea, indicates the route of this immigration, and sufficiently explains how it is that these northern types are almost wholly confined to this part of the Australian continent. Some of the earlier immigrants of this class no doubt passed over to New Zealand and now form a portion of the peculiar genera confined to these two countries; but most of them are of later date, and have thus remained in Australia only.

Proofs of Migration by way of the African Highlands.—It is owing to this twofold current of vegetation flowing into Australia by widely different routes that we have in this distant land a better representation of the European flora, both as regards species and genera, than in any other part of the southern hemisphere; and, so far as I can judge of the facts, there is no general phenomenon—that is, nothing in the distribution of genera and other groups of plants as opposed to cases of individual species—that is not fairly accounted for by such an origin. It further receives support from the case of South Africa, which also contains a large and important representation of the northern flora. But here we see no indications (or very slight ones) of that southern influx which has given Australia such a community of vegetation with the Antarctic lands. There are no less than sixty genera of strictly north temperate plants in South Africa, none of which occur in Australia; while very few of the species, so characteristic of Australia, New Zealand, and Fuegia, are found there. It is clear, therefore, that South Africa has received its European plants by the direct route through the Abyssinian highlands and the lofty equatorial mountains, and mostly at a distant period when the conditions for migration were somewhat more favourable than they are now. The much greater directness of the route from Northern Europe to South Africa than to Australia; and the existence even now of lofty mountains and extensive highlands for a large portion of the distance, will explain (what Sir Joseph Hooker notes as "a very curious fact") why South Africa has more very northern European genera than Australia, while Australia has more identical species and a better representation on the whole of the European flora—this being clearly due to the large influx of species it has received from the Antarctic Islands, in addition to those which have entered it by way of Asia. The greater distance of South Africa even now from any of these islands, and the much deeper sea to the south of the African continent, than in the case of Tasmania and New Zealand, indicating a smaller recent extension southward, is all quite in harmony with the facts of distribution of the northern flora above referred to.

Supposed Connection of South Africa and Australia.—There remains, however, the small amount of direct affinity between the vegetation of South Africa and that of Australia, New Zealand, and Temperate South America, consisting in all of fifteen genera, five of which are confined to Australia and South Africa, while several natural orders are better represented in these two countries than in any other part of the world. This resemblance has been supposed to imply some former land-connection of all the great southern lands, but it appears to me that any such supposition is wholly unnecessary. The differences between the faunas and floras of these countries are too great and too radical to render it possible that any such connection should have existed except at a very remote period. But if we have to go back so far for an explanation, a much simpler one presents itself, and one more in accordance with what we have learnt of the general permanence of deep oceans and the great changes that have taken place in the distribution of all forms of life. Just as we explain the presence of marsupials in Australia and America and of Centetidæ in Madagascar and the Antilles, by the preservation in these localities of remnants of once wide-spread types, so we should prefer to consider the few genera common to Australia and South Africa as remnants of an ancient vegetation, once spread over the northern hemisphere, driven southward by the pressure of more specialised types, and now finding a refuge in these two widely separated southern lands. It is suggestive of such an explanation that these genera are either of very ancient groups—as Conifers and Cycads—or plants of low organisation as the Restiaceæ—or of world-wide distribution, as Melanthaceæ.

The Endemic Genera of Plants in New Zealand.—Returning now to the New Zealand flora, with which we are more especially concerned, there only remains to be considered the peculiar or endemic genera which characterise it. These are thirty-two in number, and are mostly very isolated. A few have affinities with Arctic groups, others with Himalayan, or Australian genera; several are tropical forms, but the majority appear to be altogether peculiar types of world-wide groups—as Leguminosæ, Saxifrageæ, Compositæ, Orchideæ, &c. We must evidently trace back these peculiar forms to the earliest immigrants, either from the north or from the south; and the great antiquity we are obliged to give to New Zealand—an antiquity supported by every feature in its fauna and flora, no less than by its geological structure, and its extinct forms of life[195 - Dr. Hector notes the occurrence of the genus Dammara in Triassic deposits, while in the Jurassic period New Zealand possessed the genera Palæozamia, Oleandrium, Alethopteris, Camptopteris, Cycadites, Echinostrobus, &c., all Indian forms of the same age. Neocomian beds contain a true dicotyledonous leaf with Dammara and Araucaria. The Cretaceous deposits have produced a rich flora of dicotyledonous plants, many of which are of the same genera as the existing flora; while the Miocene and other Tertiary deposits produce plants almost identical with those now inhabiting the country, together with many North Temperate genera which have since become extinct. (See p. 499, footnote (#x28_x_28_i99), and Trans. New Zealand Inst., Vol. XI. 1879, p. 536.)]—affords ample time for the changes in the general distribution of plants, and for those due to isolation and modification under the influence of changed conditions, which are manifested by the extreme peculiarity of many of these interesting endemic forms.

The Absence of Southern Types from the Northern Hemisphere.—We have now only to notice the singular want of reciprocity in the migrations of northern and southern types of vegetation. In return for the vast number of European plants which have reached Australia, not one single Australian plant has entered any part of the north temperate zone, and the same may be said of the typical southern vegetation in general, whether developed in the Antarctic lands, New Zealand, South America, or South Africa. The furthest northern outliers of the southern flora are a few genera of Antarctic type on the Bornean Alps; the genus Acæna which has a species in California; two representatives of the Australian flora—Casuarina and Stylidium, in the peninsula of India; while China and the Philippines have two strictly Australian genera of Orchideæ—Microtis and Thelymitra, as well as a Restiaceous genus. Several distinct causes appear to have combined to produce this curious inability of the southern flora to make its way into the northern hemisphere. The primary cause is, no doubt, the totally different distribution of land in the two hemispheres, so that in the south there is the minimum of land in the colder parts of the temperate zone and in the north the maximum. This is well shown by the fact that on the parallel of Lat. 50° N. we pass over 240° of land or shallow sea, while on the same parallel of south latitude we have only 4°, where we cross the southern part of Patagonia. Again the three most important south temperate land-areas—South Temperate America, South Africa, and Australia—are widely separated from each other, and have in all probability always been so; whereas the whole of the north temperate lands are practically continuous. It follows that, instead of the enormous northern area, in which highly organised and dominant groups of plants have been developed gifted with great colonising and aggressive powers, we have in the south three comparatively small and detached areas, in which rich floras have been developed with special adaptations to soil, climate, and organic environment, but comparatively impotent and inferior beyond their own domain.

Another circumstance which makes the contest between the northern and southern forms still more unequal, is the much greater hardiness of the former, from having been developed in a colder region, and one where alpine and arctic conditions extensively prevail; whereas the southern floras have been mainly developed in mild regions to which they have been altogether confined. While the northern plants have been driven north or south by each succeeding change of climate, the southern species have undergone comparatively slight changes of this nature, owing to the areas they occupy being unconnected with the ice-bearing Antarctic continent. It follows, that whereas the northern plants find in all these southern lands a milder and more equable climate than that to which they have been accustomed, and are thus often able to grow and flourish even more vigorously than in their native land, the southern plants would find in almost every part of Europe, North America or Northern Asia, a more severe and less equable climate, with winters that usually prove fatal to them even under cultivation. These causes, taken separately, are very powerful, but when combined they must, I think, be held to be amply sufficient to explain why examples of the typical southern vegetation are almost unknown in the north temperate zone, while a very few of them have extended so far as the northern tropic.[196 - The fact stated in the last edition of the Origin of Species (p. 340) on the authority of Sir Joseph Hooker, that Australian plants are rapidly sowing themselves and becoming naturalised on the Neilgherrie mountains in the southern part of the Indian Peninsula, though an exception to the rule of the inability of Australian plants to become naturalised in the Northern Hemisphere, is yet quite in harmony with the hypothesis here advocated. For not only is the climate of the Neilgherries more favourable to Australian plants than any part of the North Temperate zone, but the entire Indian Peninsula has existed for unknown ages as an island and thus possesses the "insular" characteristic of a comparatively poor and less developed flora and fauna as compared with the truly "continental" Malayan and Himalayan regions. Australian plants are thus enabled to compete with those of the Indian Peninsula highlands with a fair chance of success.]

Concluding Remarks on the Last Two Chapters.—Our inquiry into the external relations and probable origin of the fauna and flora of New Zealand, has thus led us on to a general theory as to the cause of the peculiar biological relations between the northern and the southern hemispheres; and no better or more typical example could be found of the wide range and great interest of the study of the geographical distribution of animals and plants.

The solution which has here been given of one of the most difficult of this class of problems, has been rendered possible solely by the knowledge very recently obtained of the form of the sea-bottom in the southern ocean, and of the geological structure of the great Australian continent. Without this knowledge we should have nothing but a series of guesses or probabilities on which to found our hypothetical explanation, which we have now been able to build up on a solid foundation of fact. The complete separation of East from West Australia during a portion of the Cretaceous and Tertiary periods, could never have been guessed till it was established by the laborious explorations of the Australian geologists; while the hypothesis of a comparatively shallow sea, uniting New Zealand by a long route with tropical Australia, while a profoundly deep ocean always separated it from temperate Australia, would have been rejected as too improbable a supposition for the foundation of even the most enticing theory. Yet it is mainly by means of these two facts, that we are enabled to give an adequate explanation of the strange anomalies in the flora of Australia and its relation to that of New Zealand.

In the more general explanation of the relations of the various northern and southern floras, I have shown what an important aid to any such explanation is the theory of repeated changes of climate, not necessarily of great amount, given in Chapters VIII. and IX.; while the whole discussion justifies the importance attached to the theory of the general permanence of continents and oceans, as demonstrated in Chapter VI., since any rational explanation based upon facts (as opposed to mere unsupported conjecture) must take such general permanence as a starting-point. The whole inquiry into the phenomena presented by islands, which forms the main subject of the present volume has, I think, shown that this theory does afford a firm foundation for the discussion of questions of distribution and dispersal; and that by its aid, combined with a clear perception of the wonderful powers of dispersion and modification in the organic world when long periods are considered, the most difficult problems connected with this subject cease to be insoluble.

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.

The present volume has gone over a very wide field both of facts and theories, and it will be well to recall these to the reader's attention and point out their connection with each other, in a concluding chapter. I hope to be able to show that, although at first sight somewhat fragmentary and disconnected, this work is really the development of a clear and definite theory, and its application to the solution of a number of biological problems. That theory is, briefly, that the distribution of the various species and groups of living things over the earth's surface, and their aggregation in definite assemblages in certain areas, is the direct result and outcome of a complex set of causes, which may be grouped as "biological" and "physical." The biological causes are mainly of two kinds—firstly, the constant tendency of all organisms to increase in numbers and to occupy a wider area, and their various powers of dispersion and migration through which, when unchecked, they are enabled to spread widely over the globe; and, secondly, those laws of evolution and extinction which determine the manner in which groups of organisms arise and grow, reach their maximum, and then dwindle away, often breaking up into separate portions which long survive in very remote regions. The physical causes are also mainly of two kinds. We have, first, the geographical changes which at one time isolate a whole fauna and flora, at another time lead to their dispersal and intermixture with adjacent faunas and floras—and it was here important to ascertain and define the exact nature and extent of these changes, and to determine the question of the general stability or instability of continents and oceans; in the second place, it was necessary to determine the exact nature, extent and frequency of the changes of climate which have occurred in various parts of the earth,—because such changes are among the most powerful agents in causing the dispersal and extinction of plants and animals. Hence the importance attached to the question of geological climates and their causes, which have been here investigated at some length with the aid of the most recent researches of geologists, physicists, and explorers. These various inquiries led on to an investigation of the mode of formation of stratified deposits, with a view to fix within some limits their probable age; and also to an estimate of the probable rate of development of the organic world; and both these processes are shown to involve, so far as we can judge, periods of time less vast than have generally been thought necessary.

The numerous facts and theories established in the First Part of the work are then applied to explain the phenomena presented by the floras and faunas of the chief islands of the globe, which are classified, in accordance with their physical origin, in three groups or classes, each of which are shown to exhibit certain well-marked biological features.

Having thus shown that the work is a connected whole, founded on the principle of tracing out the more recondite causes of the distribution of organisms, we will briefly indicate the scope and object of the several chapters, by means of which this general conception has been carried out.

Beginning with simple and familiar facts relating to British and European quadrupeds and birds, I have defined and shown the exact character of "areas of distribution," as applied to species, genera, and families, and have illustrated the subject by maps showing the peculiarities of distribution of some well-known groups of birds. Taking then our British mammals and land-birds, I follow them over the whole area they inhabit, and thus obtain a foundation for the establishment of "zoological regions," and a clear insight into their character as distinct from the usual geographical divisions of the globe.

The facts thus far established are then shown to be necessary results of the "law of evolution." The nature and amount of "variation" is exhibited by a number of curious examples; the origin, growth, and decay of species and genera are traced, and all the interesting phenomena of isolated groups and discontinuous generic and specific areas are shown to follow as logical consequences.

The next subject investigated is the means by which the various groups of animals are enabled to overcome the natural barriers which often seem to limit them to very restricted areas, how far those barriers are themselves liable to be altered or abolished, and what is the exact nature and amount of the changes of sea and land which our earth has undergone in past times. This latter part of the inquiry is shown to be the most important as it is the most fundamental; and as it is still a subject of controversy, and many erroneous views prevail in regard to it, it is discussed at some length. Several distinct classes of evidence are adduced to prove that the grand features of our globe—the position of the great oceans and the chief land-areas—have remained, on the whole, unchanged throughout geological time. Our continents are shown to be built up mainly of "shore-deposits"; and even the chalk, which is so often said to be the exact equivalent of the "globigerina ooze" now forming in mid-Atlantic, is shown to be a comparatively shallow-water deposit formed in inland seas, or in the immediate vicinity of land. The general stability of continents has, however, been accompanied by constant changes of form, and insular conditions have prevailed over every part in succession; and the effect of such changes on the distribution of organisms is pointed out.

We then approach the consideration of another set of changes—those of climate—which have probably been agents of the first importance in modifying the specific forms as well as the distribution of animals. Here again we find ourselves in the midst of fierce controversies. The occurrence of a recent glacial epoch of great severity in the northern hemisphere is now universally admitted, but the causes which brought it on are matter of dispute. But unless we can arrive at these causes, as well as at those which produced the equally well demonstrated mild climate in the Arctic regions, we shall be quite unable to determine the nature and amount of the changes of climate which have occurred throughout past ages, and shall thus be left without a most important clue to the explanation of many of the anomalies in the distribution of animals and plants.

I have therefore devoted three chapters to a full investigation of this question. I have first given such a sketch of the most salient facts as to render the phenomena of the glacial epoch clear and intelligible. I then review the various suggested explanations, and, taking up the two which alone seem tenable, I endeavour to determine the true principles of each. While adopting generally Mr. Croll's views as to the causes of the "glacial epoch," I have introduced certain limitations and modifications. I have pointed out, I believe, more clearly than has hitherto been done, the very different effects on climate of water in the liquid and in the solid state; and I have shown, by a variety of evidence, that without high land there can be no permanent snow and ice. From these facts and principles the very important conclusion is reached, that the alternate phases of precession—causing the winter of each hemisphere to be in aphelion and perihelion each 10,500 years—would produce a complete change of climate only where a country was partially snow-clad; while, whenever a large area became almost wholly buried in snow and ice—as was certainly the case with Northern Europe and America during the glacial epoch—then the glacial conditions would be continued and perhaps even intensified when the sun approached nearest to the earth in winter, instead of there being at that time, as Mr. Croll maintains, an almost perpetual spring. This important result is supported by reference to the existing differences between the climates of the northern and southern hemispheres, and by what is known to have occurred during the last glacial epoch; and it is shown to be in complete harmony with the geological evidence as to interglacial mild periods.

Discussing next the evidence for glacial epochs in earlier times, it is shown that Mr. Croll's views are opposed by a vast body of facts, and that the geological evidence leads irresistibly to the conclusion that during a large portion of the Secondary and Tertiary periods, uninterrupted warm climates prevailed in the north temperate zone, and so far ameliorated the climate of the Arctic regions as to admit of the growth of a luxuriant vegetation in the highest latitudes yet explored. The geographical condition of the northern hemisphere at these periods is then investigated, and it is shown to have been probably such as to admit the warm tropical waters freely to penetrate the land, and to reach the Arctic seas by several channels; and, adopting Mr. Croll's calculations as to the enormous quantity of heat that would thus be conveyed northwards, it is maintained that the mild Arctic climates are amply accounted for. With such favourable geographical conditions, it is shown, that changes of excentricity and of the phases of precession would have no other effect than to cause greater differences of temperature between summer and winter; but, wherever there was a considerable extent of very lofty mountains the snow-line would be lowered, and the snow-collecting area being thus largely increased a considerable amount of local glaciation might result. Thus may be explained the presence of enormous ice-borne rocks in Eocene and Miocene times in Central Europe, while at the very same period all the surrounding country enjoyed a tropical or sub-tropical climate.

The general conclusion is thus reached, that geographical conditions are the essential causes of great changes of climate, and that the radically different distribution of land and sea in the northern and southern hemispheres has generally led to great diversity of climate in the Arctic and Antarctic regions. The form and arrangement of the continents is shown to be such as to favour the transfer of warm oceanic currents to the north far in excess of those which move towards the south, and whenever these currents had free passage through the northern land-masses to the polar area, a mild climate must have prevailed over the whole northern hemisphere. It is only in very recent times that the great northern continents have become so completely consolidated as they now are, thus shutting out the warm water from their interiors, and rendering possible a wide-spread and intense glacial epoch. But this great climatal change was actually brought about by the high excentricity which occurred about 200,000 years ago; and it is doubtful if a similar glaciation in equally low latitudes could be produced by means of any such geographical combinations as actually occur, without the concurrence of a high excentricity.

A survey of the present condition of the earth supports this view, for though we have enormous mountain ranges in every latitude, there is no glaciated country south of Greenland in N. Lat. 61°. But directly we go back a very short period, we find the superficial evidences of glaciation to an enormous extent over three-fourths of the globe. In the Alps and Pyrenees, in the British Isles and Scandinavia, in Spain and the Atlas, in the Caucasus and the Himalayas, in Eastern North America and west of the Rocky Mountains, in the Andes of South Temperate America, in South Africa, and in New Zealand, huge moraines and other unmistakable ice-marks attest the universal descent of the snow-line for several thousand feet below its present level. If we reject the influence of high excentricity as the cause of this almost universal glaciation, we must postulate a general elevation of all these mountains about the same time, geologically speaking—for the general similarity in the state of preservation of the ice-marks and the known activity of denudation as a destroying agent, forbid the idea that they belong to widely separated epochs. It has, indeed, been suggested, that denudation alone has lowered these mountains so much during the post-tertiary epoch, that they were previously of sufficient height to account for the glaciation of all of them; but this hardly needs refutation, for it is clear that denudation could not at the same time have removed some thousands of feet of rock from many hundreds of square miles of lofty snow-collecting plateaus, and yet have left moraines, and blocks, and even glacial striæ, undisturbed and uneffaced on the slopes and in the valleys of these same mountains.

The theory of geological climates set forth in this volume, while founded on Mr. Croll's researches, differs from all that have yet been made public, in clearly tracing out the comparative influence of geographical and astronomical revolutions, showing that, while the former have been the chief, if not the exclusive, causes of the long-continued mild climates of the Arctic regions, the concurrence of the latter has been essential to the production of glacial epochs in the temperate zones, as well as of those local glaciations in low latitudes, of which there is such an abundance of evidence.

The next question discussed is that of geological time as bearing on the development of the organic world. The periods of time usually demanded by geologists have been very great, and it was often assumed that there was no occasion to limit them. But the theory of development demands far more; for the earliest fossiliferous rocks prove the existence of many and varied forms of life which require unrecorded ages for their development—ages probably far longer than those which have elapsed from that period to the present day. The physicists, however, deny that any such indefinitely long periods are available. The sun is ever losing heat far more rapidly than it can be renewed from any known or conceivable source. The earth is a cooling body, and must once have been too hot to support life; while the friction of the tides is checking the earth's rotation, and this cannot have gone on indefinitely without making our day much longer than it is. A limit is therefore placed to the age of the habitable earth, and it has been thought that the time so allowed is not sufficient for the long processes of geological change and organic development. It is therefore important to inquire whether these processes are either of them so excessively slow as has been supposed, and I devote a chapter to the inquiry.

Geologists have measured with some accuracy the maximum thickness of all the known sedimentary rocks. The rate of denudation has also been recently measured by a method which, if not precise, at all events gives results of the right order of magnitude and which err on the side of being too slow rather than too fast. If, then, the maximum thickness of the known sedimentary rocks is taken to represent the average thickness of all the sedimentary rocks, and we also know the amount of sediment carried to the sea or lakes, and the area over which that sediment is spread, we have a means of calculating the time required for the building up of all the sedimentary rocks of the geological system. I have here inquired how far the above suppositions are correct, or on which side they probably err; and the conclusion arrived at is, that the time required is very much less than has hitherto been supposed.

Another estimate is afforded by the date of the last glacial epoch if coincident with the last period of high excentricity, while the Alpine glaciation of the Miocene period is assumed to have been caused by the next earlier phase of very high excentricity. Taking these as data, the proportionate change of the species of mollusca affords a means of arriving at the whole lapse of time represented by the fossiliferous rocks; and these two estimates agree in the order of their magnitudes.

It is then argued that the changes of climate every 10,500 years during the numerous periods of high excentricity have acted as a motive power in hastening on both geological and biological change. By raising and lowering the snow-line in all mountain ranges it has caused increased denudation; while the same changes have caused much migration and disturbance in the organic world, and have thus tended to the more rapid modification of species. The present epoch being a period of very low excentricity, the earth is in a phase of exceptional stability both physical and organic; and it is from this period of exceptional stability that our notions of the very slow rate of change have been derived.

The conclusion is, on the whole, that the periods allowed by physicists are not only far in excess of such as are required for geological and organic change, but that they allow ample margin for a lapse of time anterior to the deposit of the earliest fossiliferous rocks several times longer than the time which has elapsed since their deposit to the present day.

Having thus laid the foundation for a scientific interpretation of the phenomena of distribution, we proceed to the Second Part of our work—the discussion of a series of typical Insular Faunas and Floras with a view to explain the interesting phenomena they present. Taking first two North Atlantic groups—the Azores and Bermuda—it is shown how important an agent in the dispersal of most animals and plants is a stormy atmosphere. Although 900 and 700 miles respectively from the nearest continents, their productions are very largely identical with those of Europe and America; and, what is more important, fresh arrivals of birds, insects, and plants, are now taking place almost annually. These islands afford, therefore, test examples of the great dispersive powers of certain groups of organisms, and thus serve as a basis on which to found our explanations of many anomalies of distribution. Passing on to the Galapagos we have a group less distant from a continent and of larger area, yet, owing to special conditions, of which the comparatively stormless equatorial atmosphere is the most important, exhibiting far more speciality in its productions than the more distant Azores. Still, however, its fauna and flora are as unmistakably derived from the American continent as those of the Azores are from the European.

We next take St. Helena and the Sandwich Islands, both wonderfully isolated in the midst of vast oceans, and no longer exhibiting in their productions an exclusive affinity to one continent. Here we have to recognise the results of immense antiquity, and of those changes of geography, of climate, and in the general distribution of organisms which we know have occurred in former geological epochs, and whose causes and consequences we have discussed in the first part of our volume. This concludes our review of the Oceanic Islands.

Coming now to Continental Islands we consider first those of most recent origin and offering the simplest phenomena; and begin with the British Isles as affording the best example of very recent and well known Continental Islands. Reviewing the interesting past history of Britain, we show why it is comparatively poor in species and why this poverty is still greater in Ireland. By a careful examination of its fauna and flora it is then shown that the British Isles are not so completely identical, biologically, with the continent as has been supposed. A considerable amount of speciality is shown to exist, and that this speciality is real and not apparent is supported by the fact, that small outlying islands, such as the Isle of Man, the Shetland Isles, Lundy Island, and the Isle of Wight, all possess certain species or varieties not found elsewhere.

Borneo and Java are next taken, as illustrations of tropical islands which may be not more ancient than Britain, but which, owing to their much larger area, greater distance from the continent, and the extreme richness of the equatorial fauna and flora, possess a large proportion of peculiar species, though these are in general very closely allied to those of the adjacent parts of Asia. The preliminary studies we have made enable us to afford a simpler and more definite interpretation of the peculiar relations of Java to the continent and its differences from Borneo and Sumatra, than was given in my former work (The Geographical Distribution of Animals).

Japan and Formosa are next taken, as examples of islands which are decidedly somewhat more ancient than those previously considered, and which present a number of very interesting phenomena, especially in their relations to each other, and to remote rather than to adjacent parts of the Asiatic continent.

We now pass to the group of Ancient Continental Islands, of which Madagascar is the most typical example. It is surrounded by a number of smaller islands which may be termed its satellites since they partake of many of its peculiarities; though some of these—as the Comoros and Seychelles—may be considered continental, while others—as Bourbon, Mauritius, and Rodriguez—are decidedly oceanic. In order to understand the peculiarities of the Madagascar fauna we have to consider the past history of the African and Asiatic continents, which it is shown are such as to account for all the main peculiarities of the fauna of these islands without having recourse to the hypothesis of a now-submerged Lemurian continent. Considerable evidence is further adduced to show that "Lemuria" is a myth, since not only is its existence unnecessary, but it can be proved that it would not explain the actual facts of distribution. The origin of the interesting Mascarene wingless birds is discussed, and the main peculiarities of the remarkable flora of Madagascar and the Mascarene islands pointed out; while it is shown that all these phenomena are to be explained on the general principles of the permanence of the great oceans and the comparatively slight fluctuations of the land area, and by taking account of established palæontological facts.

There remain two other islands—Celebes and New Zealand—which are classed as "anomalous," the one because it is almost impossible to place it in any of the six zoological regions, or determine whether it has ever been actually joined to a continent—the other because it combines the characteristics of continental and oceanic islands.

The peculiarities of the Celebesian fauna have already been dwelt upon in several previous works, but they are so remarkable and so unique that they cannot be omitted in a treatise on "Insular Faunas"; and here, as in the case of Borneo and Java, fuller consideration and the application of the general principles laid down in our First Part, lead to a solution of the problem at once more simple and more satisfactory than any which have been previously proposed. I now look upon Celebes as an outlying portion of the great Asiatic continent of Miocene times, which either by submergence or some other cause had lost the greater portion of its animal inhabitants, and since then has remained more or less completely isolated from every other land. It has thus preserved a fragment of a very ancient fauna along with a number of later types which have reached it from surrounding islands by the ordinary means of dispersal. This sufficiently explains all the peculiar affinities of its animals, though the peculiar and distinctive characters of some of them remain as mysterious as ever.

New Zealand is shown to be so completely continental in its geological structure, and its numerous wingless birds so clearly imply a former connection with some other land (as do its numerous lizards and its remarkable reptile, the Hatteria), that the total absence of indigenous land-mammalia was hardly to be expected. Some attention is therefore given to the curious animal which has been seen but never captured, and this is shown to be probably identical with an animal referred to by Captain Cook. The more accurate knowledge which has recently been obtained of the sea bottom around New Zealand enables us to determine that the former connection of that island with Australia was towards the north, and this is found to agree well with many of the peculiarities of its fauna.

The flora of New Zealand and that of Australia are now both so well known, and they present so many peculiarities, and relations of so anomalous a character, as to present in Sir Joseph Hooker's opinion an almost insoluble problem. Much additional information on the physical and geological history of these two countries has, however, been obtained since the appearance of Sir Joseph Hooker's works, and I therefore determined to apply to them the same method of discussion and treatment which has been usually successful with similar problems in the case of animals. The fact above noted, that New Zealand was connected with Australia in its northern and tropical portion only, of itself affords a clue to one portion of the specialities of the New Zealand flora—the presence of an unusual number of tropical families and genera, while the temperate forms consist mainly of species either identical with those found in Australia or closely allied to them. But a still more important clue is obtained in the geological structure of Australia itself, which is shown to have been for long periods divided into an eastern and a western island, in the latter of which the highly peculiar flora of temperate Australia was developed. This is found to explain with great exactness the remarkable absence from New Zealand of all the most abundant and characteristic Australian genera, both of plants and of animals, since these existed at that time only in the western island, while New Zealand was in connection with the eastern island alone and with the tropical portion of it. From these geological and physical facts, and the known powers of dispersal of plants, all the main features, and many of the detailed peculiarities of the New Zealand flora are shown necessarily to result.

Our last chapter is devoted to a wider, and if possible more interesting subject—the origin of the European element in the floras of New Zealand and Australia, and also in those of South America and South Africa. This is so especially a botanical question, that it was with some diffidence I entered upon it, yet it arose so naturally from the study of the New Zealand and Australian floras, and seemed to have so much light thrown upon it by our preliminary studies as to changes of climate and the causes which have favoured the distribution of plants, that I felt my work would be incomplete without a consideration of it. The subject will be so fresh in the reader's mind that a complete summary of it is unnecessary. I venture to think, however, that I have shown, not only the several routes by which the northern plants have reached the various southern lands, but have pointed out the special aids to their migration, and the motive power which has urged them on.

In this discussion, if nowhere else, will be found a complete justification of that lengthy investigation of the exact nature of past changes of climate, which to some readers may have seemed unnecessary and unsuited to such a work as the present. Without the clear and definite conclusions arrived at by that discussion, and those equally important views as to the permanence of the great features of the earth's surface, and the wonderful dispersive powers of plants which have been so frequently brought before us in our studies of insular floras, I should not have ventured to attack the wide and difficult problem of the northern element in southern floras.

In concluding a work dealing with subjects which have occupied my attention for many years, I trust that the reader who has followed me throughout will be imbued with the conviction that ever presses upon myself, of the complete interdependence of organic and inorganic nature. Not only does the marvellous structure of each organised being involve the whole past history of the earth, but such apparently unimportant facts as the presence of certain types of plants or animals in one island rather than in another, are now shown to be dependent on the long series of past geological changes—on those marvellous astronomical revolutions which cause a periodic variation of terrestrial climates—on the apparently fortuitous action of storms and currents in the conveyance of germs—and on the endlessly varied actions and reactions of organised beings on each other. And although these various causes are far too complex in their combined action to enable us to follow them out in the case of any one species, yet their broad results are clearly recognisable; and we are thus encouraged to study more completely every detail and every anomaly in the distribution of living things, in the firm conviction that by so doing we shall obtain a fuller and clearer insight into the course of nature, and with increased confidence that the "mighty maze" of Being we see everywhere around us is "not without a plan."

notes

1

A small number of species belonging to the West Indies are found in the extreme southern portion of the Florida Peninsula.

2

I cannot avoid here referring to the enormous waste of labour and money with comparatively scanty and unimportant results to natural history of most of the great scientific voyages of the various civilized governments during the present century. All these expeditions combined have done far less than private collectors in making known the products of remote lands and islands. They have brought home fragmentary collections, made in widely scattered localities, and these have been usually described in huge folios or quartos, whose value is often in inverse proportion to their bulk and cost. The same species have been collected again and again, often described several times over under new names, and not unfrequently stated to be from places they never inhabited. The result of this wretched system is that the productions of some of the most frequently visited and most interesting islands on the globe are still very imperfectly known, while their native plants and animals are being yearly exterminated, and this is the case even with countries under the rule or protection of European governments. Such are the Sandwich Islands, Tahiti, the Marquesas, the Philippine Islands, and a host of smaller ones; while Bourbon and Mauritius, St. Helena, and several others, have only been adequately explored after an important portion of their productions has been destroyed by cultivation or the reckless introduction of goats and pigs. The employment in each of our possessions, and those of other European powers, of a resident naturalist at a very small annual expense, would have done more for the advancement of knowledge in this direction than all the expensive expeditions that have again and again circumnavigated the globe.

3

The general facts of Palæontology, as bearing on the migrations of animal groups, are summarised in my Geographical Distribution of Animals, Vol. I. Chapters VI., VII., and VIII.

4