Chapter XIX: Introduction (3)

I have dwelt on the importance of what I may call short-distance dispersal as a necessary condition of plant life, because I think it suggests the solution of a difficulty which leads Guppy to a conclusion with which I am unable to agree. But the work which he has done taken as a whole appears to me so admirable that I do so with the utmost respect. He points out, as Bentham had already done, that long-distance dispersal is fortuitous. And being so it cannot have been provided for by previous adaptation. He says (Guppy, op. cit. II. page 99.): "It is not conceivable that an organism can be adapted to conditions outside its environment." To this we must agree; but, it may be asked, do the general means of plant dispersal violate so obvious a principle? He proceeds: "The great variety of the modes of dispersal of seeds is in itself an indication that the dispersing agencies avail themselves in a hap-hazard fashion of characters and capacities that have been developed in other connections." (Loc. cit. page 102.) "Their utility in these respects is an accident in the plant's life." (Loc. cit. page 100.) He attributes this utility to a "determining agency," an influence which constantly reappears in various shapes in the literature of Evolution and is ultra-scientific in the sense that it bars the way to the search for material causes. He goes so far as to doubt whether fleshy fruits are an adaptation for the dispersal of their contained seeds. (Loc. cit. page 102.) Writing as I am from a hillside which is covered by hawthorn bushes sown by birds, I confess I can feel little doubt on the subject myself. The essential fact which Guppy brings out is that long-distance unlike short-distance dispersal is not universal and purposeful, but selective and in that sense accidental. But it is not difficult to see how under favouring conditions one must merge into the other.

Guppy has raised one novel point which can only be briefly referred to but which is of extreme interest. There are grounds for thinking that flowers and insects have mutually reacted upon one another in their evolution. Guppy suggests that something of the same kind may be true of birds. I must content myself with the quotation of a single sentence. "With the secular drying of the globe and the consequent differentiation of climate is to be connected the suspension to a great extent of the agency of birds as plant dispersers in later ages, not only in the Pacific Islands but all over the tropics. The changes of climate, birds and plants have gone on together, the range of the bird being controlled by the climate, and the distribution of the plant being largely dependent on the bird." (Loc.cit. II. page 221.)

Darwin was clearly prepared to go further than Hooker in accounting for the southern flora by dispersion from the north. Thus he says: "We must, I suppose, admit that every yard of land has been successively covered with a beech-forest between the Caucasus and Japan." ("More Letters", II. page 9.) Hooker accounted for the dissevered condition of the southern flora by geographical change, but this Darwin could not admit. He suggested to Hooker that the Australian and Cape floras might have had a point of connection through Abyssinia (Ibid. I. page 447.), an idea which was promptly snuffed out. Similarly he remarked to Bentham (1869): "I suppose you think that the Restiaceae, Proteaceae, etc., etc. once extended over the whole world, leaving fragments in the south." (Ibid. I. page 380.) Eventually he conjectured "that there must have been a Tertiary Antarctic continent, from which various forms radiated to the southern extremities of our present continents." ("Life and Letters", III. page 231.) But characteristically he could not admit any land connections and trusted to "floating ice for transporting seed." ("More Letters", I. page 116.) I am far from saying that this theory is not deserving of serious attention, though there seems to be no positive evidence to support it, and it immediately raises the difficulty how did such a continent come to be stocked?

We must, however, agree with Hooker that the common origin of the northern and southern floras must be referred to a remote past. That Darwin had this in his mind at the time of the publication of the "Origin" is clear from a letter to Hooker. "The view which I should have looked at as perhaps most probable (though it hardly differs from yours) is that the whole world during the Secondary ages was inhabited by marsupials, araucarias (Mem.--Fossil wood of this nature in South America), Banksia, etc.; and that these were supplanted and exterminated in the greater area of the north, but were left alive in the south." (Ibid. I. page 453.) Remembering that Araucaria, unlike Banksia, belongs to the earlier Jurassic not to the angiospermous flora, this view is a germinal idea of the widest generality.

The extraordinary congestion in species of the peninsulas of the Old World points to the long-continued action of a migration southwards. Each is in fact a cul-de-sac into which they have poured and from which there is no escape. On the other hand the high degree of specialisation in the southern floras and the little power the species possess of holding their own in competition or in adaptation to new conditions point to long-continued isolation. "An island... will prevent free immigration and competition, hence a greater number of ancient forms will survive." (Ibid. I. page 481.) But variability is itself subject to variation. The nemesis of a high degree of protected specialisation is the loss of adaptability. (See Lyell, "The Geological Evidences of the Antiquity of Man", London, 1863, page 446.) It is probable that many elements of the southern flora are doomed: there is, for example, reason to think that the singular Stapelieae of S. Africa are a disappearing group. The tree Lobelias which linger in the mountains of Central Africa, in Tropical America and in the Sandwich Islands have the aspect of extreme antiquity. I may add a further striking illustration from Professor Seward: "The tall, graceful fronds of Matonia pectinata, forming miniature forests on the slopes of Mount Ophir and other districts in the Malay Peninsula in association with Dipteris conjugata and Dipteris lobbiana, represent a phase of Mesozoic life which survives 'Like a dim picture of the drowned past.'" ("Report of the 73rd Meeting of the British Assoc." (Southport, 1903), London, 1904, page 844.)

The Matonineae are ferns with an unusually complex vascular system and were abundant "in the northern hemisphere during the earlier part of the Mesozoic era."

It was fortunate for science that Wallace took up the task which his colleague had abandoned. Writing to him on the publication of his "Geographical Distribution of Animals" Darwin said: "I feel sure that you have laid a broad and safe foundation for all future work on Distribution. How interesting it will be to see hereafter plants treated in strict relation to your views." ("More Letters", II. page 12.) This hope was fulfilled in "Island Life". I may quote a passage from it which admirably summarises the contrast between the northern and the southern floras.

"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." (Wallace, "Island Life", pages 527, 528.)

It will be noticed that in the summary I have attempted to give of the history of the subject, efforts have been concentrated on bringing into relation the temperate floras of the northern and southern hemispheres, but no account has been taken of the rich tropical vegetation which belts the world and little to account for the original starting-point of existing vegetation generally. It must be remembered on the one hand that our detailed knowledge of the floras of the tropics is still very incomplete and far inferior to that of temperate regions; on the other hand palaeontological discoveries have put the problem in an entirely new light. Well might Darwin, writing to Heer in 1875, say: "Many as have been the wonderful discoveries in Geology during the last half-century, I think none have exceeded in interest your results with respect to the plants which formerly existed in the arctic regions." ("More Letters", II. page 240.)

As early as 1848 Debey had described from the Upper Cretaceous rocks of Aix-la-Chapelle Flowering plants of as high a degree of development as those now existing. The fact was commented upon by Hooker ("Introd. Essay to the Flora of Tasmania", page xx.), but its full significance seems to have been scarcely appreciated. For it implied not merely that their evolution must have taken place but the foundations of existing distribution must have been laid in a preceding age. We now know from the discoveries of the last fifty years that the remains of the Neocomian flora occur over an area extending through 30 deg of latitude. The conclusion is irresistible that within this was its centre of distribution and probably of origin.

Darwin was immensely impressed with the outburst on the world of a fully fledged angiospermous vegetation. He warmly approved the brilliant theory of Saporta that this happened "as soon (as) flower-frequenting insects were developed and favoured intercrossing." ("More Letters", II. page 21.) Writing to him in 1877 he says: "Your idea that dicotyledonous plants were not developed in force until sucking insects had been evolved seems to me a splendid one. I am surprised that the idea never occurred to me, but this is always the case when one first hears a new and simple explanation of some mysterious phenomenon." ("Life and Letters", III. page 285. Substantially the same idea had occurred earlier to F.W.A. Miquel. Remarking that "sucking insects (Haustellata)... perform in nature the important duty of maintaining the existence of the vegetable kingdom, at least as far as the higher orders are concerned," he points our that "the appearance in great numbers of haustellate insects occurs at and after the Cretaceous epoch, when the plants with pollen and closed carpels (Angiosperms) are found, and acquire little by little the preponderance in the vegetable kingdom." "Archives Neerlandaises", III. (1868). English translation in "Journ. of Bot." 1869, page 101.)

Even with this help the abruptness still remains an almost insoluble problem, though a forecast of floral structure is now recognised in some Jurassic and Lower Cretaceous plants. But the gap between this and the structural complexity and diversity of angiosperms is enormous. Darwin thought that the evolution might have been accomplished during a period of prolonged isolation. Writing to Hooker (1881) he says: "Nothing is more extraordinary in the history of the Vegetable Kingdom, as it seems to me, than the APPARENTLY very sudden or abrupt development of the higher plants. I have sometimes speculated whether there did not exist somewhere during long ages an extremely isolated continent, perhaps near the South Pole." ("Life and Letters", III. page 248.)

The present trend of evidence is, however, all in favour of a northern origin for flowering plants, and we can only appeal to the imperfection of the geological record as a last resource to extricate us from the difficulty of tracing the process. But Darwin's instinct that at some time or other the southern hemisphere had played an important part in the evolution of the vegetable kingdom did not mislead him. Nothing probably would have given him greater satisfaction than the masterly summary in which Seward has brought together the evidence for the origin of the Glossopteris flora in Gondwana land.

"A vast continental area, of which remnants are preserved in Australia, South Africa and South America... A tract of enormous extent occupying an area, part of which has since given place to a southern ocean, while detached masses persist as portions of more modern continents, which have enabled us to read in their fossil plants and ice-scratched boulders the records of a lost continent, in which the Mesozoic vegetation of the northern continent had its birth." ("Encycl. Brit." (10th edition 1902), Vol. XXXI. ("Palaeobotany; Mesozoic"), page 422.) Darwin would probably have demurred on physical grounds to the extent of the continent, and preferred to account for the transoceanic distribution of its flora by the same means which must have accomplished it on land.

It must in fairness be added that Guppy's later views give some support to the conjectural existence of the "lost continent." "The distribution of the genus Dammara" (Agathis) led him to modify his earlier conclusions. He tells us:--"In my volume on the geology of Vanua Levu it was shown that the Tertiary period was an age of submergence in the Western Pacific, and a disbelief in any previous continental condition was expressed. My later view is more in accordance with that of Wichmann, who, on geological grounds, contended that the islands of the Western Pacific were in a continental condition during the Palaeozoic and Mesozoic periods, and that their submergence and subsequent emergence took place in Tertiary times." (Guppy, op. cit. II. page 304.)

The weight of the geological evidence I am unable to scrutinise. But though I must admit the possibility of some unconscious bias in my own mind on the subject, I am impressed with the fact that the known distribution of the Glossopteris flora in the southern hemisphere is precisely paralleled by that of Proteaceae and Restiaceae in it at the present time. It is not unreasonable to suppose that both phenomena, so similar, may admit of the same explanation. I confess it would not surprise me if fresh discoveries in the distribution of the Glossopteris flora were to point to the possibility of its also having migrated southwards from a centre of origin in the northern hemisphere.

Darwin, however, remained sceptical "about the travelling of plants from the north EXCEPT DURING THE TERTIARY PERIOD." But he added, "such speculations seem to me hardly scientific, seeing how little we know of the old floras." ("Life and Letters", III. page 247.) That in later geological times the south has been the grave of the weakened offspring of the aggressive north can hardly be doubted. But if we look to the Glossopteris flora for the ancestry of Angiosperms during the Secondary period, Darwin's prevision might be justified, though he has given us no clue as to how he arrived at it.

It may be true that technically Darwin was not a botanist. But in two pages of the "Origin" he has given us a masterly explanation of "the relationship, with very little identity, between the productions of North America and Europe." (Pages 333, 334.) He showed that this could be accounted for by their migration southwards from a common area, and he told Wallace that he "doubted much whether the now called Palaearctic and Neartic regions ought to be separated." ("Life and Letters", III. page 230.) Catkin-bearing deciduous trees had long been seen to justify Darwin's doubt: oaks, chestnuts, beeches, hazels, hornbeams, birches, alders, willows and poplars are common both to the Old and New World. Newton found that the separate regions could not be sustained for birds, and he is now usually followed in uniting them as the Holartic. One feels inclined to say in reading the two pages, as Lord Kelvin did to a correspondent who asked for some further development of one of his papers, It is all there. We have only to apply the principle to previous geological ages to understand why the flora of the Southern United States preserves a Cretaceous facies. Applying it still further we can understand why, when the northern hemisphere gradually cooled through the Tertiary period, the plants of the Eocene "suggest a comparison of the climate and forests with those of the Malay Archipelago and Tropical America." (Clement Reid, "Encycl. Brit." (10th edition), Vol. XXXI. ("Palaeobotany; Tertiary"), page 435.) Writing to Asa Gray in 1856 with respect to the United States flora, Darwin said that "nothing has surprised me more than the greater generic and specific affinity with East Asia than with West America." ("More Letters", I. page 434.) The recent discoveries of a Tulip tree and a Sassafras in China afford fresh illustrations. A few years later Asa Gray found the explanation in both areas being centres of preservation of the Cretaceous flora from a common origin. It is interesting to note that the paper in which this was enunciated at once established his reputation.

In Europe the latitudinal range of the great mountain chains gave the Miocene flora no chance of escape during the Glacial period, and the Mediterranean appears to have equally intercepted the flow of alpine plants to the Atlas. (John Ball in Appendix G, page 438, in "Journal of a Tour in Morocco and the Great Atlas", J.D. Hooker and J. Ball, London, 1878.) In Southern Europe the myrtle, the laurel, the fig and the dwarf-palm are the sole representatives of as many great tropical families. Another great tropical family, the Gesneraceae has left single representatives from the Pyrenees to the Balkans; and in the former a diminutive yam still lingers. These are only illustrations of the evidence which constantly accumulates and which finds no rational explanation except that which Darwin has given to it.

The theory of southward migration is the key to the interpretation of the geographical distribution of plants. It derived enormous support from the researches of Heer and has now become an accepted commonplace. Saporta in 1888 described the vegetable kingdom as "emigrant pour suivre une direction determinee et marcher du nord au sud, a la recherche de regions et de stations plus favorables, mieux appropriees aux adaptations acquises, a meme que la temperature terrestre perd ses conditions premieres." ("Origine Paleontologique des arbres", Paris, 1888, page 28.) If, as is so often the case, the theory now seems to be a priori inevitable, the historian of science will not omit to record that the first germ sprang from the brain of Darwin.

In attempting this sketch of Darwin's influence on Geographical Distribution, I have found it impossible to treat it from an external point of view. His interest in it was unflagging; all I could say became necessarily a record of that interest and could not be detached from it. He was in more or less intimate touch with everyone who was working at it. In reading the letters we move amongst great names. With an extraordinary charm of persuasive correspondence he was constantly suggesting, criticising and stimulating. It is hardly an exaggeration to say that from the quiet of his study at Down he was founding and directing a wide-world school.

POSTSCRIPTUM.

Since this essay was put in type Dr Ernst's striking account of the "New Flora of the Volcanic Island of Krakatau" (Cambridge, 1909.) has reached me. All botanists must feel a debt of gratitude to Prof. Seward for his admirable translation of a memoir which in its original form is practically unprocurable and to the liberality of the Cambridge University Press for its publication. In the preceding pages I have traced the laborious research by which the methods of Plant Dispersal were established by Darwin. In the island of Krakatau nature has supplied a crucial experiment which, if it had occurred earlier, would have at once secured conviction of their efficiency. A quarter of a century ago every trace of organic life in the island was "destroyed and buried under a thick covering of glowing stones." Now, it is "again covered with a mantle of green, the growth being in places so luxuriant that it is necessary to cut one's way laboriously through the vegetation." (Op. cit. page 4.) Ernst traces minutely how this has been brought about by the combined action of wind, birds and sea currents, as means of transport. The process will continue, and he concludes:--"At last after a long interval the vegetation on the desolated island will again acquire that wealth of variety and luxuriance which we see in the fullest development which Nature has reached in the primaeval forest in the tropics." (Op. cit. page 72.) The possibility of such a result revealed itself to the insight of Darwin with little encouragement or support from contemporary opinion.

One of the most remarkable facts established by Ernst is that this has not been accomplished by the transport of seeds alone. "Tree stems and branches played an important part in the colonisation of Krakatau by plants and animals. Large piles of floating trees, stems, branches and bamboos are met with everywhere on the beach above high-water mark and often carried a considerable distance inland. Some of the animals on the island, such as the fat Iguana (Varanus salvator) which suns itself in the beds of streams, may have travelled on floating wood, possibly also the ancestors of the numerous ants, but certainly plants." (Op. cit. page 56.) Darwin actually had a prevision of this. Writing to Hooker he says:--"Would it not be a prodigy if an unstocked island did not in the course of ages receive colonists from coasts whence the currents flow, trees are drifted and birds are driven by gales?" ("More Letters", I. page 483.) And ten years earlier:--"I must believe in the... whole plant or branch being washed into the sea; with floods and slips and earthquakes; this must continually be happening." ("Life and Letters", II. pages 56, 57.) If we give to "continually" a cosmic measure, can the fact be doubted? All this, in the light of our present knowledge, is too obvious to us to admit of discussion. But it seems to me nothing less than pathetic to see how in the teeth of the obsession as to continental extension, Darwin fought single-handed for what we now know to be the truth.

Guppy's heart failed him when he had to deal with the isolated case of Agathis which alone seemed inexplicable by known means of transport. But when we remember that it is a relic of the pre-Angiospermous flora, and is of Araucarian ancestry, it cannot be said that the impossibility, in so prolonged a history, of the bodily transference of cone-bearing branches or even of trees, compels us as a last resort to fall back on continental extension to account for its existing distribution.

When Darwin was in the Galapagos Archipelago, he tells us that he fancied himself "brought near to the very act of creation." He saw how new species might arise from a common stock. Krakatau shows us an earlier stage and how by simple agencies, continually at work, that stock might be supplied. It also shows us how the mixed and casual elements of a new colony enter into competition for the ground and become mutually adjusted. The study of Plant Distribution from a Darwinian standpoint has opened up a new field of research in Ecology. The means of transport supply the materials for a flora, but their ultimate fate depends on their equipment for the "struggle for existence." The whole subject can no longer be regarded as a mere statistical inquiry which has seemed doubtless to many of somewhat arid interest. The fate of every element of the earth's vegetation has sooner or later depended on its ability to travel and to hold its own under new conditions. And the means by which it has secured success is an each case a biological problem which demands and will reward the most attentive study. This is the lesson which Darwin has bequeathed to us. It is summed up in the concluding paragraph of the "Origin" ("Origin of Species" (6th edition), page 429.):--"It is interesting to contemplate a tangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent upon each other in so complex a manner, have all been produced by laws acting around us."

XVII. GEOGRAPHICAL DISTRIBUTION OF ANIMALS. By Hans Gadow, M.A., Ph.D., F.R.S.

Strickland Curator and Lecturer on Zoology in the University of Cambridge.

The first general ideas about geographical distribution may be found in some of the brilliant speculations contained in Buffon's "Histoire Naturelle". The first special treatise on the subject was however written in 1777 by E.A.W. Zimmermann, Professor of Natural Science at Brunswick, whose large volume, "Specimen Zoologiae Geographicae Quadrupedum"..., deals in a statistical way with the mammals; important features of the large accompanying map of the world are the ranges of mountains and the names of hundreds of genera indicating their geographical range. In a second work he laid special stress on domesticated animals with reference to the spreading of the various races of Mankind.

In the following year appeared the "Philosophia Entomologica" by J.C. Fabricius, who was the first to divide the world into eight regions. In 1803 G.R. Treviranus ("Biologie oder Philosophie der lebenden Natur", Vol. II. Gottingen, 1803.) devoted a long chapter of his great work on "Biologie" to a philosophical and coherent treatment of the distribution of the whole animal kingdom. Remarkable progress was made in 1810 by F. Tiedemann ("Anatomie und Naturgeschichte der Vogel". Heidelberg, 1810.) of Heidelberg. Few, if any, of the many subsequent Ornithologists seem to have appreciated, or known of, the ingenious way in which Tiedemann marshalled his statistics in order to arrive at general conclusions. There are, for instance, long lists of birds arranged in accordance with their occurrence in one or more continents: by correlating the distribution of the birds with their food he concludes "that the countries of the East Indian flora have no vegetable feeders in common with America," and "that it is probably due to the great peculiarity of the African flora that Africa has few phytophagous kinds in common with other countries, whilst zoophagous birds have a far more independent, often cosmopolitan, distribution." There are also remarkable chapters on the influence of environment, distribution, and migration, upon the structure of the Birds! In short, this anatomist dealt with some of the fundamental causes of distribution.

Whilst Tiedemann restricted himself to Birds, A. Desmoulins in 1822 wrote a short but most suggestive paper on the Vertebrata, omitting the birds; he combated the view recently proposed by the entomologist Latreille that temperature was the main factor in distribution. Some of his ten main conclusions show a peculiar mixture of evolutionary ideas coupled with the conception of the stability of species: whilst each species must have started from but one creative centre, there may be several "analogous centres of creation" so far as genera and families are concerned. Countries with different faunas, but lying within the same climatic zones, are proof of the effective and permanent existence of barriers preventing an exchange between the original creative centres.

The first book dealing with the "geography and classification" of the whole animal kingdom was written by W. Swainson ("A Treatise on the Geography and Classification of Animals", Lardner's "Cabinet Cyclopaedia" London, 1835.) in 1835. He saw in the five races of Man the clue to the mapping of the world into as many "true zoological divisions," and he reconciled the five continents with his mystical quinary circles.

Lyell's "Principles of Geology" should have marked a new epoch, since in his "Elements" he treats of the past history of the globe and the distribution of animals in time, and in his "Principles" of their distribution in space in connection with the actual changes undergone by the surface of the world. But as the sub-title of his great work "Modern changes of the Earth and its inhabitants" indicates, he restricted himself to comparatively minor changes, and, emphatically believing in the permanency of the great oceans, his numerous and careful interpretations of the effect of the geological changes upon the dispersal of animals did after all advance the problem but little.

Hitherto the marine faunas had been neglected. This was remedied by E. Forbes, who established nine homozoic zones, based mainly on the study of the mollusca, the determining factors being to a great extent the isotherms of the sea, whilst the 25 provinces were given by the configuration of the land. He was followed by J.D. Dana, who, taking principally the Crustacea as a basis, and as leading factors the mean temperatures of the coldest and of the warmest months, established five latitudinal zones. By using these as divisors into an American, Afro-European, Oriental, Arctic and Antarctic realm, most of which were limited by an eastern and western land-boundary, he arrived at about threescore provinces.

In 1853 appeared L.K. Schmarda's ("Die geographische Verbreitung der Thiere", Wien, 1853.) two volumes, embracing the whole subject. Various centres of creation being, according to him, still traceable, he formed the hypothesis that these centres were originally islands, which later became enlarged and joined together to form the great continents, so that the original faunas could overlap and mix whilst still remaining pure at their respective centres. After devoting many chapters to the possible physical causes and modes of dispersal, he divided the land into 21 realms which he shortly characterises, e.g. Australia as the only country inhabited by marsupials, monotremes and meliphagous birds. Ten main marine divisions were diagnosed in a similar way. Although some of these realms were not badly selected from the point of view of being applicable to more than one class of animals, they were obviously too numerous for general purposes, and this drawback was overcome, in 1857, by P.L. Sclater. ("On the general Geographical Distribution of the members of the class Aves", "Proc. Linn. Soc." (Zoology II. 1858, pages 130-145.)) Starting with the idea, that "each species must have been created within and over the geographical area, which it now occupies," he concluded "that the most natural primary ontological divisions of the Earth's surface" were those six regions, which since their adoption by Wallace in his epoch-making work, have become classical. Broadly speaking, these six regions are equivalent to the great masses of land; they are convenient terms for geographical facts, especially since the Palaearctic region expresses the unity of Europe with the bulk of Asia. Sclater further brigaded the regions of the Old World as Palaeogaea and the two Americas as Neogaea, a fundamental mistake, justifiable to a certain extent only since he based his regions mainly upon the present distribution of the Passerine birds.

Unfortunately these six regions are not of equal value. The Indian countries and the Ethiopian region (Africa south of the Sahara) are obviously nothing but the tropical, southern continuations or appendages of one greater complex. Further, the great eastern mass of land is so intimately connected with North America that this continent has much more in common with Europe and Asia than with South America. Therefore, instead of dividing the world longitudinally as Sclater had done, Huxley, in 1868 ("On the classification and distribution of the Alectoromorphae and Heteromorphae", "Proc. Zool. Soc." 1868, page 294.), gave weighty reasons for dividing it transversely. Accordingly he established two primary divisions, Arctogaea or the North world in a wider sense, comprising Sclater's Indian, African, Palaearctic and Neartic regions; and Notogaea, the Southern world, which he divided into (1) Austro-Columbia (an unfortunate substitute for the neotropical region), (2) Australasia, and (3) New Zealand, the number of big regions thus being reduced to three but for the separation of New Zealand upon rather negative characters. Sclater was the first to accept these four great regions and showed, in 1874 ("The geographical distribution of Mammals", "Manchester Science Lectures", 1874.), that they were well borne out by the present distribution of the Mammals.

Although applicable to various other groups of animals, for instance to the tailless Amphibia and to Birds (Huxley himself had been led to found his two fundamental divisions on the distribution of the Gallinaceous birds), the combination of South America with Australia was gradually found to be too sweeping a measure. The obvious and satisfactory solution was provided by W.T. Blanford (Anniversary address (Geological Society, 1889), "Proc. Geol. Soc." 1889-90, page 67; "Quart. Journ." XLVI 1890.), who in 1890 recognised three main divisions, namely Australian, South American, and the rest, for which the already existing terms (although used partly in a new sense, as proposed by an anonymous writer in "Natural Science", III. page 289) "Notogaea," "Neogaea" and "Arctogaea" have been gladly accepted by a number of English writers.

After this historical survey of the search for larger and largest or fundamental centres of animal creation, which resulted in the mapping of the world into zoological regions and realms of after all doubtful value, we have to return to the year 1858. The eleventh and twelfth chapters of "The Origin of Species" (1859), dealing with "Geographical Distribution," are based upon a great amount of observation, experiment and reading. As Darwin's main problem was the origin of species, nature's way of making species by gradual changes from others previously existing, he had to dispose of the view, held universally, of the independent creation of each species and at the same time to insist upon a single centre of creation for each species; and in order to emphasise his main point, the theory of descent, he had to disallow convergent, or as they were then called, analogous forms. To appreciate the difficulty of his position we have to take the standpoint of fifty years ago, when the immutability of the species was an axiom and each was supposed to have been created within or over the geographical area which it now occupies. If he once admitted that a species could arise from many individuals instead of from one pair, there was no way of shutting the door against the possibility that these individuals may have been so numerous that they occupied a very large district, even so large that it had become as discontinuous as the distribution of many a species actually is. Such a concession would at once be taken as an admission of multiple, independent, origin instead of descent in Darwin's sense.

For the so-called multiple, independently repeated creation of species as an explanation of their very wide and often quite discontinuous distribution, he substituted colonisation from the nearest and readiest source together with subsequent modification and better adaptation to their new home.

He was the first seriously to call attention to the many accidental means, "which more properly should be called occasional means of distribution," especially to oceanic islands. His specific, even individual, centres of creation made migrations all the more necessary, but their extent was sadly baulked by the prevailing dogma of the permanency of the oceans. Any number of small changes ("many islands having existed as halting places, of which not a wreck now remains" ("The Origin of Species" (1st edition), page 396.).) were conceded freely, but few, if any, great enough to permit migration of truly terrestrial creatures. The only means of getting across the gaps was by the principle of the "flotsam and jetsam," a theory which Darwin took over from Lyell and further elaborated so as to make it applicable to many kinds of plants and animals, but sadly deficient, often grotesque, in the case of most terrestrial creatures.

Another very fertile source was Darwin's strong insistence upon the great influence which the last glacial epoch must have had upon the distribution of animals and plants. Why was the migration of northern creatures southwards of far-reaching and most significant importance? More northerners have established themselves in southern lands than vice versa, because there is such a great mass of land in the north and greater continents imply greater intensity of selection. "The productions of real islands have everywhere largely yielded to continental forms." (Ibid. page 380.)... "The Alpine forms have almost everywhere largely yielded to the more dominant forms generated in the larger areas and more efficient workshops of the North."

Let us now pass in rapid survey the influence of the publication of "The Origin of Species" upon the study of Geographical Distribution in its wider sense.

Hitherto the following thought ran through the minds of most writers: Wherever we examine two or more widely separated countries their respective faunas are very different, but where two faunas can come into contact with each other, they intermingle. Consequently these faunas represent centres of creation, whence the component creatures have spread peripherally so far as existing boundaries allowed them to do so. This is of course the fundamental idea of "regions." There is not one of the numerous writers who considered the possibility that these intermediate belts might represent not a mixture of species but transitional forms, the result of changes undergone by the most peripheral migrants in adaptation to their new surroundings. The usual standpoint was also that of Pucheran ("Note sur l'equateur zoologique", "Rev. et Mag. de Zoologie", 1855; also several other papers, ibid. 1865, 1866, and 1867.) in 1855. But what a change within the next ten years! Pucheran explains the agreement in coloration between the desert and its fauna as "une harmonie post-etablie"; the Sahara, formerly a marine basin, was peopled by immigrants from the neighbouring countries, and these new animals adapted themselves to the new environment. He also discusses, among other similar questions, the Isthmus of Panama with regard to its having once been a strait. From the same author may be quoted the following passage as a strong proof of the new influence: "By the radiation of the contemporaneous faunas, each from one centre, whence as the various parts of the world successively were formed and became habitable, they spread and became modified according to the local physical conditions."

The "multiple" origin of each species as advocated by Sclater and Murray, although giving the species a broader basis, suffered from the same difficulties. There was only one alternative to the old orthodox view of independent creation, namely the bold acceptance of land-connections to an extent for which geological and palaeontological science was not yet ripe. Those who shrank from either view, gave up the problem as mysterious and beyond the human intellect. This was the expressed opinion of men like Swainson, Lyell and Humboldt. Only Darwin had the courage to say that the problem was not insoluble. If we admit "that in the long course of time the individuals of the same species, and likewise of allied species, have proceeded from some one source; then I think all the grand leading facts of geographical distribution are explicable on the theory of migration... together with subsequent modification and the multiplication of new forms." We can thus understand how it is that in some countries the inhabitants "are linked to the extinct beings which formerly inhabited the same continent." We can see why two areas, having nearly the same physical conditions, should often be inhabited by very different forms of life,... and "we can see why in two areas, however distant from each other, there should be a correlation, in the presence of identical species... and of distinct but representative species." ("The Origin of Species" (1st edition), pages 408, 409.)

Darwin's reluctance to assume great geological changes, such as a land-connection of Europe with North America, is easily explained by the fact that he restricted himself to the distribution of the present and comparatively recent species. "I do not believe that it will ever be proved that within the recent period continents which are now quite separate, have been continuously, or almost continuously, united with each other, and with the many existing oceanic islands." (Ibid. page 357.) Again, "believing... that our continents have long remained in nearly the same relative position, though subjected to large, but partial oscillations of level," that means to say within the period of existing species, or "within the recent period." (Ibid. page. 370.) The difficulty was to a great extent one of his own making. Whilst almost everybody else believed in the immutability of the species, which implies an enormous age, logically since the dawn of creation, to him the actually existing species as the latest results of evolution, were necessarily something very new, so young that only the very latest of the geological epochs could have affected them. It has since come to our knowledge that a great number of terrestrial "recent" species, even those of the higher classes of Vertebrates, date much farther back than had been thought possible. Many of them reach well into the Miocene, a time since which the world seems to have assumed the main outlines of the present continents.

In the year 1866 appeared A. Murray's work on the "Geographical Distribution of Mammals", a book which has perhaps received less recognition than it deserves. His treatment of the general introductory questions marks a considerable advance of our problem, although, and partly because, he did not entirely agree with Darwin's views as laid down in the first edition of "The Origin of Species", which after all was the great impulse given to Murray's work. Like Forbes he did not shrink from assuming enormous changes in the configuration of the continents and oceans because the theory of descent, with its necessary postulate of great migrations, required them. He stated, for instance, "that a Miocene Atlantis sufficiently explains the common distribution of animals and plants in Europe and America up to the glacial epoch." And next he considers how, and by what changes, the rehabilitation and distribution of these lands themselves were effected subsequent to that period. Further, he deserves credit for having cleared up a misunderstanding of the idea of specific centres of creation. Whilst for instance Schmarda assumed without hesitation that the same species, if occurring at places separated by great distances, or apparently insurmountable barriers, had been there created independently (multiple centres), Lyell and Darwin held that each species had only one single centre, and with this view most of us agree, but their starting point was to them represented by one individual, or rather one single pair. According to Murray, on the other hand, this centre of a species is formed by all the individuals of a species, all of which equally undergo those changes which new conditions may impose upon them. In this respect a new species has a multiple origin, but this in a sense very different from that which was upheld by L. Agassiz. As Murray himself puts it: "To my multiple origin, communication and direct derivation is essential. The species is compounded of many influences brought together through many individuals, and distilled by Nature into one species; and, being once established it may roam and spread wherever it finds the conditions of life not materially different from those of its original centre." (Murray, "The Geographical Distribution of Mammals", page 14. London, 1866.) This declaration fairly agrees with more modern views, and it must be borne in mind that the application of the single-centre principle to the genera, families and larger groups in the search for descent inevitably leads to one creative centre for the whole animal kingdom, a condition as unwarrantable as the myth of Adam and Eve being the first representatives of Mankind.

It looks as if it had required almost ten years for "The Origin of Species" to show its full effect, since the year 1868 marks the publication of Haeckel's "Naturliche Schoepfungsgeschichte" in addition to other great works. The terms "Oecology" (the relation of organisms to their environment) and "Chorology" (their distribution in space) had been given us in his "Generelle Morphologie" in 1866. The fourteenth chapter of the "History of Creation" is devoted to the distribution of organisms, their chorology, with the emphatic assertion that "not until Darwin can chorology be spoken of as a separate science, since he supplied the acting causes for the elucidation of the hitherto accumulated mass of facts." A map (a "hypothetical sketch") shows the monophyletic origin and the routes of distribution of Man.

Natural Selection may be all-mighty, all-sufficient, but it requires time, so much that the countless aeons required for the evolution of the present fauna were soon felt to be one of the most serious drawbacks of the theory. Therefore every help to ease and shorten this process should have been welcomed. In 1868 M. Wagner (The first to formulate clearly the fundamental idea of a theory of migration and its importance in the origin of new species was L. von Buch, who in his "Physikalische Beschreibung der Canarischen Inseln", written in 1825, wrote as follows: "Upon the continents the individuals of the genera by spreading far, form, through differences of the locality, food and soil, varieties which finally become constant as new species, since owing to the distances they could never be crossed with other varieties and thus be brought back to the main type. Next they may again, perhaps upon different roads, return to the old home where they find the old type likewise changed, both having become so different that they can interbreed no longer. Not so upon islands, where the individuals shut up in narrow valleys or within narrow districts, can always meet one another and thereby destroy every new attempt towards the fixing of a new variety." Clearly von Buch explains here why island types remain fixed, and why these types themselves have become so different from their continental congeners.--Actually von Buch is aware of a most important point, the difference in the process of development which exists between a new species b, which is the result of an ancestral species a having itself changed into b and thereby vanished itself, and a new species c which arose through separation out of the same ancestral a, which itself persists as such unaltered. Von Buch's prophetic view seems to have escaped Lyell's and even Wagner's notice.) came to the rescue with his "Darwin'sche Theorie und das Migrations-Gesetz der Organismen". (Leipzig, 1868.) He shows that migration, i.e. change of locality, implies new environmental conditions (never mind whether these be new stimuli to variation, or only acting as their selectors or censors), and moreover secures separation from the original stock and thus eliminates or lessens the reactionary dangers of panmixia. Darwin accepted Wagner's theory as "advantageous." Through the heated polemics of the more ardent selectionists Wagner's theory came to grow into an alternative instead of a help to the theory of selectional evolution. Separation is now rightly considered a most important factor by modern students of geographical distribution.

For the same year, 1868, we have to mention Huxley, whose Arctogaea and Notogaea are nothing less than the reconstructed main masses of land of the Mesozoic period. Beyond doubt the configuration of land at that remote period has left recognisable traces in the present continents, but whether they can account for the distribution of such a much later group as the Gallinaceous birds is more than questionable. In any case he took for his text a large natural group of birds, cosmopolitan as a whole, but with a striking distribution. The Peristeropodes, or pigeon-footed division, are restricted to the Australian and Neotropical regions, in distinction to the Alectoropodes (with the hallux inserted at a level above the front toes) which inhabit the whole of the Arctogaea, only a few members having spread into the South World. Further, as Asia alone has its Pheasants and allies, so is Africa characterised by its Guinea-fowls and relations, America has the Turkey as an endemic genus, and the Grouse tribe in a wider sense has its centre in the holarctic region: a splendid object lesson of descent, world-wide spreading and subsequent differentiation. Huxley, by the way, was the first--at least in private talk--to state that it will be for the morphologist, the well-trained anatomist, to give the casting vote in questions of geographical distribution, since he alone can determine whether we have to deal with homologous, or analogous, convergent, representative forms.

It seems late to introduce Wallace's name in 1876, the year of the publication of his standard work. ("The Geographical Distribution of Animals", 2 vols. London, 1876.) We cannot do better than quote the author's own words, expressing the hope that his "book should bear a similar relation to the eleventh and twelfth chapters of the "Origin of Species" as Darwin's "Animals and Plants under Domestication" does to the first chapter of that work," and to add that he has amply succeeded. Pleading for a few primary centres he accepts Sclater's six regions and does not follow Huxley's courageous changes which Sclater himself had accepted in 1874. Holding the view of the permanence of the oceans he accounts for the colonisation of outlying islands by further elaborating the views of Lyell and Darwin, especially in his fascinating "Island Life", with remarkable chapters on the Ice Age, Climate and Time and other fundamental factors. His method of arriving at the degree of relationship of the faunas of the various regions is eminently statistical. Long lists of genera determine by their numbers the affinity and hence the source of colonisation. In order to make sure of his material he performed the laborious task of evolving a new classification of the host of Passerine birds. This statistical method has been followed by many authors, who, relying more upon quantity than quality, have obscured the fact that the key to the present distribution lies in the past changes of the earth's surface. However, with Wallace begins the modern study of the geographical distribution of animals and the sudden interest taken in this subject by an ever widening circle of enthusiasts far beyond the professional brotherhood.

A considerable literature has since grown up, almost bewildering in its range, diversity of aims and style of procedure. It is a chaos, with many paths leading into the maze, but as yet very few take us to a position commanding a view of the whole intricate terrain with its impenetrable tangle and pitfalls.

One line of research, not initiated but greatly influenced by Wallace's works, became so prominent as to almost constitute a period which may be characterised as that of the search by specialists for either the justification or the amending of his regions. As class after class of animals was brought up to reveal the secret of the true regions, some authors saw in their different results nothing but the faultiness of previously established regions; others looked upon eventual agreements as their final corroboration, especially when for instance such diverse groups as mammals and scorpions could, with some ingenuity, be made to harmonise. But the obvious result of all these efforts was the growing knowledge that almost every class seemed to follow principles of its own. The regions tallied neither in extent nor in numbers, although most of them gravitated more and more towards three centres, namely Australia, South America and the rest of the world. Still zoologists persisted in the search, and the various modes and capabilities of dispersal of the respective groups were thought sufficient explanation of the divergent results in trying to bring the mapping of the world under one scheme.

Contemporary literature is full of devices for the mechanical dispersal of animals. Marine currents, warm and cold, were favoured all the more since they showed the probable original homes of the creatures in question. If these could not stand sea-water, they floated upon logs or icebergs, or they were blown across by storms; fishes were lifted over barriers by waterspouts, and there is on record even an hypothetical land tortoise, full of eggs, which colonised an oceanic island after a perilous sea voyage upon a tree trunk. Accidents will happen, and beyond doubt many freaks of discontinuous distribution have to be accounted for by some such means. But whilst sufficient for the scanty settlers of true oceanic islands, they cannot be held seriously to account for the rich fauna of a large continent, over which palaeontology shows us that the immigrants have passed like waves. It should also be borne in mind that there is a great difference between flotsam and jetsam. A current is an extension of the same medium and the animals in it may suffer no change during even a long voyage, since they may be brought from one litoral to another where they will still be in the same or but slightly altered environment. But the jetsam is in the position of a passenger who has been carried off by the wrong train. Almost every year some American land birds arrive at our western coasts and none of them have gained a permanent footing although such visits must have taken place since prehistoric times. It was therefore argued that only those groups of animals should be used for locating and defining regions which were absolutely bound to the soil. This method likewise gave results not reconcilable with each other, even when the distribution of fossils was taken into account, but it pointed to the absolute necessity of searching for former land-connections regardless of their extent and the present depths to which they may have sunk.