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Чарльз Дарвин
The Foundations of the Origin of Species

Alpine Floras

We will now turn to the floras of mountain-summits which are well known to differ from the floras of the neighbouring lowlands. In certain characters, such as dwarfness of stature, hairiness, &c., the species from the most distant mountains frequently resemble each other, – a kind of analogy like that for instance of the succulency of most desert plants. Besides this analogy, Alpine plants present some eminently curious facts in their distribution. In some cases the summits of mountains, although immensely distant from each other, are clothed by the same identical species³⁶² which are likewise the same with those growing on the likewise very distant Arctic shores. In other cases, although few or none of the species may be actually identical, they are closely related; whilst the plants of the lowland districts surrounding the two mountains in question will be wholly dissimilar. As mountain-summits, as far as their plants are concerned, are islands rising out of an ocean of land in which the Alpine species cannot live, nor across which is there any known means of transport, this fact appears directly opposed to the conclusion which we have come to from considering the general distribution of organisms both on continents and on islands – namely, that the degree of relationship between the inhabitants of two points depends on the completeness and nature of the barriers between those points³⁶³. I believe, however, this anomalous case admits, as we shall presently see, of some explanation. We might have expected that the flora of a mountain summit would have presented the same relation to the flora of the surrounding lowland country, which any isolated part of a continent does to the whole, or an island does to the mainland, from which it is separated by a rather wide space of sea. This in fact is the case with the plants clothing the summits of some mountains, which mountains it may be observed are particularly isolated; for instance, all the species are peculiar, but they belong to the forms characteristic of the surrounding continent, on the mountains of Caraccas, of Van Dieman's Land and of the Cape of Good Hope³⁶⁴. On some other mountains, for instance «in» Tierra del Fuego and in Brazil, some of the plants though distinct species are S. American forms; whilst others are allied to or are identical with the Alpine species of Europe. In islands of which the lowland flora is distinct «from» but allied to that of the nearest continent, the Alpine plants are sometimes (or perhaps mostly) eminently peculiar and distinct³⁶⁵; this is the case on Teneriffe, and in a lesser degree even on some of the Mediterranean islands.

If all Alpine floras had been characterised like that of the mountain of Caraccas, or of Van Dieman’s Land, &c., whatever explanation is possible of the general laws of geographical distribution would have applied to them. But the apparently anomalous case just given, namely of the mountains of Europe, of some mountains in the United States (Dr Boott) and of the summits of the Himalaya (Royle), having many identical species in common conjointly with the Arctic regions, and many species, though not identical, closely allied, require a separate explanation. The fact likewise of several of the species on the mountains of Tierra del Fuego (and in a lesser degree on the mountains of Brazil) not belonging to American forms, but to those of Europe, though so immensely remote, requires also a separate explanation.

Cause of the similarity in the floras of some distant mountains

Now we may with confidence affirm, from the number of the then floating icebergs and low descent of the glaciers, that within a period so near that species of shells have remained the same, the whole of Central Europe and of North America (and perhaps of Eastern Asia) possessed a very cold climate; and therefore it is probable that the floras of these districts were the same as the present Arctic one, – as is known to have been to some degree the case with then existing sea-shells, and those now living on the Arctic shores. At this period the mountains must have been covered with ice of which we have evidence in the surfaces polished and scored by glaciers. What then would be the natural and almost inevitable effects of the gradual change into the present more temperate climate³⁶⁶? The ice and snow would disappear from the mountains, and as new plants from the more temperate regions of the south migrated northward, replacing the Arctic plants, these latter would crawl³⁶⁷ up the now uncovered mountains, and likewise be driven northward to the present Arctic shores. If the Arctic flora of that period was a nearly uniform one, as the present one is, then we should have the same plants on these mountain-summits and on the present Arctic shores. On this view the Arctic flora of that period must have been a widely extended one, more so than even the present one; but considering how similar the physical conditions must always be of land bordering on perpetual frost, this does not appear a great difficulty; and may we not venture to suppose that the almost infinitely numerous icebergs, charged with great masses of rocks, soil and brushwood³⁶⁸ and often driven high up on distant beaches, might have been the means of widely distributing the seeds of the same species?

I will only hazard one other observation, namely that during the change from an extremely cold climate to a more temperate one the conditions, both on lowland and mountain, would be singularly favourable for the diffusion of any existing plants, which could live on land, just freed from the rigour of eternal winter; for it would possess no inhabitants; and we cannot doubt that preoccupation³⁶⁹ is the chief bar to the diffusion of plants. For amongst many other facts, how otherwise can we explain the circumstance that the plants on the opposite, though similarly constituted sides of a wide river in Eastern Europe (as I was informed by Humboldt) should be widely different; across which river birds, swimming quadrupeds and the wind must often transport seeds; we can only suppose that plants already occupying the soil and freely seeding check the germination of occasionally transported seeds.

At about the same period when icebergs were transporting boulders in N. America as far as 36° south, where the cotton tree now grows in South America, in latitude 42° (where the land is now clothed with forests having an almost tropical aspect with the trees bearing epiphytes and intertwined with canes), the same ice action was going on; is it not then in some degree probable that at this period the whole tropical parts of the two Americas possessed³⁷⁰ (as Falconer asserts that India did) a more temperate climate? In this case the Alpine plants of the long chain of the Cordillera would have descended much lower and there would have been a broad high-road³⁷¹ connecting those parts of North and South America which were then frigid. As the present climate supervened, the plants occupying the districts which now are become in both hemispheres temperate and even semi-tropical must have been driven to the Arctic and Antarctic³⁷² regions; and only a few of the loftiest points of the Cordillera can have retained their former connecting flora. The transverse chain of Chiquitos might perhaps in a similar manner during the ice-action period have served as a connecting road (though a broken one) for Alpine plants to become dispersed from the Cordillera to the highlands of Brazil. It may be observed that some (though not strong) reasons can be assigned for believing that at about this same period the two Americas were not so thoroughly divided as they now are by the West Indies and tableland of Mexico. I will only further remark that the present most singularly close similarity in the vegetation of the lowlands of Kerguelen’s Land³⁷³ and of Tierra del Fuego (Hooker), though so far apart, may perhaps be explained by the dissemination of seeds during this same cold period, by means of icebergs, as before alluded to³⁷⁴.

Finally, I think we may safely grant from the foregoing facts and reasoning that the anomalous similarity in the vegetation of certain very distant mountain-summits is not in truth opposed to the conclusion of the intimate relation subsisting between proximity in space (in accordance with the means of transport in each class) and the degree of affinity of the inhabitants of any two countries. In the case of several quite isolated mountains, we have seen that the general law holds good.

Whether the same species has been created more than once

As the fact of the same species of plants having been found on mountain-summits immensely remote has been one chief cause of the belief of some species having been contemporaneously produced or created at two different points³⁷⁵, I will here briefly discuss this subject. On the ordinary theory of creation, we can see no reason why on two similar mountain-summits two similar species may not have been created; but the opposite view, independently of its simplicity, has been generally received from the analogy of the general distribution of all organisms, in which (as shown in this chapter) we almost always find that great and continuous barriers separate distinct series; and we are naturally led to suppose that the two series have been separately created. When taking a more limited view we see a river, with a quite similar country on both sides, with one side well stocked with a certain animal and on the other side not one (as is the case with the Bizcacha³⁷⁶ on the opposite sides of the Plata), we are at once led to conclude that the Bizcacha was produced on some one point or area on the western side of the river. Considering our ignorance of the many strange chances of diffusion by birds (which occasionally wander to immense distances) and quadrupeds swallowing seeds and ova (as in the case of the flying water-beetle which disgorged the eggs of a fish), and of whirlwinds carrying seeds and animals into strong upper currents (as in the case of volcanic ashes and showers of hay, grain and fish³⁷⁷), and of the possibility of species having survived for short periods at intermediate spots and afterwards becoming extinct there³⁷⁸; and considering our knowledge of the great changes which have taken place from subsidence and elevation in the surface of the earth, and of our ignorance of the greater changes which may have taken place, we ought to be very slow in admitting the probability of double creations. In the case of plants on mountain-summits, I think I have shown how almost necessarily they would, under the past conditions of the northern hemisphere, be as similar as are the plants on the present Arctic shores; and this ought to teach us a lesson of caution.

But the strongest argument against double creations may be drawn from considering the case of mammifers³⁷⁹ in which, from their nature and from the size of their offspring, the means of distribution are more in view. There are no cases where the same species is found in very remote localities, except where there is a continuous belt of land: the Arctic region perhaps offers the strongest exception, and here we know that animals are transported on icebergs³⁸⁰. The cases of lesser difficulty may all receive a more or less simple explanation; I will give only one instance; the nutria³⁸¹, I believe, on the eastern coast of S. America live exclusively in fresh-water rivers, and I was much surprised how they could have got into rivulets, widely apart, on the coast of Patagonia; but on the opposite coast I found these quadrupeds living exclusively in the sea, and hence their migration along the Patagonian coast is not surprising. There is no case of the same mammifer being found on an island far from the coast, and on the mainland, as happens with plants³⁸². On the idea of double creations it would be strange if the same species of several plants should have been created in Australia and Europe; and no one instance of the same species of mammifer having been created, or aboriginally existing, in two as nearly remote and equally isolated points. It is more philosophical, in such cases, as that of some plants being found in Australia and Europe, to admit that we are ignorant of the means of transport. I will allude only to one other case, namely, that of the Mydas³⁸³, an Alpine animal, found only on the distant peaks of the mountains of Java: who will pretend to deny that during the ice period of the northern and southern hemispheres, and when India is believed to have been colder, the climate might not have permitted this animal to haunt a lower country, and thus to have passed along the ridges from summit to summit? Mr Lyell has further observed that, as in space, so in time, there is no reason to believe that after the extinction of a species, the self-same form has ever reappeared³⁸⁴. I think, then, we may, notwithstanding the many cases of difficulty, conclude with some confidence that every species has been created or produced on a single point or area.

On the number of species, and of the classes to which they belong in different regions

The last fact in geographical distribution, which, as far as I can see, in any way concerns the origin of species, relates to the absolute number and nature of the organic beings inhabiting different tracts of land. Although every species is admirably adapted (but not necessarily better adapted than every other species, as we have seen in the great increase of introduced species) to the country and station it frequents; yet it has been shown that the entire difference between the species in distant countries cannot possibly be explained by the difference of the physical conditions of these countries. In the same manner, I believe, neither the number of the species, nor the nature of the great classes to which they belong, can possibly in all cases be explained by the conditions of their country. New Zealand³⁸⁵, a linear island stretching over about 700 miles of latitude, with forests, marshes, plains and mountains reaching to the limits of eternal snow, has far more diversified habitats than an equal area at the Cape of Good Hope; and yet, I believe, at the Cape of Good Hope there are, of phanerogamic plants, from five to ten times the number of species as in all New Zealand. Why on the theory of absolute creations should this large and diversified island only have from 400 to 500 (? Dieffenbach) phanerogamic plants? and why should the Cape of Good Hope, characterised by the uniformity of its scenery, swarm with more species of plants than probably any other quarter of the world? Why on the ordinary theory should the Galapagos Islands abound with terrestrial reptiles? and why should many equal-sized islands in the Pacific be without a single one³⁸⁶ or with only one or two species? Why should the great island of New Zealand be without one mammiferous quadruped except the mouse, and that was probably introduced with the aborigines? Why should not one island (it can be shown, I think, that the mammifers of Mauritius and St Iago have all been introduced) in the open ocean possess a mammiferous quadruped? Let it not be said that quadrupeds cannot live in islands, for we know that cattle, horses and pigs during a long period have run wild in the West Indian and Falkland Islands; pigs at St Helena; goats at Tahiti; asses in the Canary Islands; dogs in Cuba; cats at Ascension; rabbits at Madeira and the Falklands; monkeys at St Iago and the Mauritius; even elephants during a long time in one of the very small Sooloo Islands; and European mice on very many of the smallest islands far from the habitations of man³⁸⁷. Nor let it be assumed that quadrupeds are more slowly created and hence that the oceanic islands, which generally are of volcanic formation, are of too recent origin to possess them; for we know (Lyell) that new forms of quadrupeds succeed each other quicker than Mollusca or Reptilia. Nor let it be assumed (though such an assumption would be no explanation) that quadrupeds cannot be created on small islands; for islands not lying in mid-ocean do possess their peculiar quadrupeds; thus many of the smaller islands of the East Indian Archipelago possess quadrupeds; as does Fernando Po on the West Coast of Africa; as the Falkland Islands possess a peculiar wolf-like fox³⁸⁸; so do the Galapagos Islands a peculiar mouse of the S. American type. These two last are the most remarkable cases with which I am acquainted; inasmuch as the islands lie further from other land. It is possible that the Galapagos mouse may have been introduced in some ship from the S. American coast (though the species is at present unknown there), for the aboriginal species soon haunts the goods of man, as I noticed in the roof of a newly erected shed in a desert country south of the Plata. The Falkland Islands, though between 200 and 300 miles from the S. American coast, may in one sense be considered as intimately connected with it; for it is certain that formerly many icebergs loaded with boulders were stranded on its southern coast, and the old canoes which are occasionally now stranded, show that the currents still set from Tierra del Fuego. This fact, however, does not explain the presence of the Canis antarcticus on the Falkland Islands, unless we suppose that it formerly lived on the mainland and became extinct there, whilst it survived on these islands, to which it was borne (as happens with its northern congener, the common wolf) on an iceberg, but this fact removes the anomaly of an island, in appearance effectually separated from other land, having its own species of quadruped, and makes the case like that of Java and Sumatra, each having their own rhinoceros.

Before summing up all the facts given in this section on the present condition of organic beings, and endeavouring to see how far they admit of explanation, it will be convenient to state all such facts in the past geographical distribution of extinct beings as seem anyway to concern the theory of descent.

Section Second

Geographical distribution of extinct organisms

I have stated that if the land of the entire world be divided into (we will say) three sections, according to the amount of difference of the terrestrial mammifers inhabiting them, we shall have three unequal divisions of (1st) Australia and its dependent islands, (2nd) South America, (3rd) Europe, Asia and Africa. If we now look to the mammifers which inhabited these three divisions during the later Tertiary periods, we shall find them almost as distinct as at the present day, and intimately related in each division to the existing forms in that division³⁸⁹. This is wonderfully the case with the several fossil Marsupial genera in the caverns of New South Wales and even more wonderfully so in South America, where we have the same peculiar group of monkeys, of a guanaco-like animal, of many rodents, of the Marsupial Didelphys, of Armadilloes and other Edentata. This last family is at present very characteristic of S. America, and in a late Tertiary epoch it was even more so, as is shown by the numerous enormous animals of the Megatheroid family, some of which were protected by an osseous armour like that, but on a gigantic scale, of the recent Armadillo. Lastly, over Europe the remains of the several deer, oxen, bears, foxes, beavers, field-mice, show a relation to the present inhabitants of this region; and the contemporaneous remains of the elephant, rhinoceros, hippopotamus, hyæna, show a relation with the grand Africo-Asiatic division of the world. In Asia the fossil mammifers of the Himalaya (though mingled with forms long extinct in Europe) are equally related to the existing forms of the Africo-Asiatic division; but especially to those of India itself. As the gigantic and now extinct quadrupeds of Europe have naturally excited more attention than the other and smaller remains, the relation between the past and the present mammiferous inhabitants of Europe has not been sufficiently attended to. But in fact the mammifers of Europe are at present nearly as much Africo-Asiatic as they were formerly when Europe had its elephants and rhinoceroses, etc.; Europe neither now nor then possessed peculiar groups as does Australia and S. America. The extinction of certain peculiar forms in one quarter does not make the remaining mammifers of that quarter less related to its own great division of the world: though Tierra del Fuego possesses only a fox, three rodents, and the guanaco, no one (as these all belong to S. American types, but not to the most characteristic forms) would doubt for one minute «as to» classifying this district with S. America; and if fossil Edentata, Marsupials and monkeys were to be found in Tierra del Fuego, it would not make this district more truly S. American than it now is. So it is with Europe³⁹⁰, and so far as is known with Asia, for the lately past and present mammifers all belong to the Africo-Asiatic division of the world. In every case, I may add, the forms which a country has is of more importance in geographical arrangement than what it has not.

We find some evidence of the same general fact in a relation between the recent and the Tertiary sea-shells, in the different main divisions of the marine world.

This general and most remarkable relation between the lately past and present mammiferous inhabitants of the three main divisions of the world is precisely the same kind of fact as the relation between the different species of the several sub-regions of any one of the main divisions. As we usually associate great physical changes with the total extinction of one series of beings, and its succession by another series, this identity of relation between the past and the present races of beings in the same quarters of the globe is more striking than the same relation between existing beings in different sub-regions: but in truth we have no reason for supposing that a change in the conditions has in any of these cases supervened, greater than that now existing between the temperate and tropical, or between the highlands and lowlands of the same main divisions, now tenanted by related beings. Finally, then, we clearly see that in each main division of the world the same relation holds good between its inhabitants in time as over space³⁹¹.