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Чарльз Дарвин
On the Origin of Species By Means of Natural Selection

It is a remarkable fact, strongly insisted on by Hooker in regard to America, and by Alph. de Candolle in regard to Australia, that many more identical plants and allied forms have apparently migrated from the north to the south, than in a reversed direction. We see, however, a few southern vegetable forms on the mountains of Borneo and Abyssinia. I suspect that this preponderant migration from north to south is due to the greater extent of land in the north, and to the northern forms having existed in their own homes in greater numbers, and having consequently been advanced through natural selection and competition to a higher stage of perfection or dominating power, than the southern forms. And thus, when they became commingled during the Glacial period, the northern forms were enabled to beat the less powerful southern forms. Just in the same manner as we see at the present day, that very many European productions cover the ground in La Plata, and in a lesser degree in Australia, and have to a certain extent beaten the natives; whereas extremely few southern forms have become naturalised in any part of Europe, though hides, wool, and other objects likely to carry seeds have been largely imported into Europe during the last two or three centuries from La Plata, and during the last thirty or forty years from Australia. Something of the same kind must have occurred on the intertropical mountains: no doubt before the Glacial period they were stocked with endemic Alpine forms; but these have almost everywhere largely yielded to the more dominant forms, generated in the larger areas and more efficient workshops of the north. In many islands the native productions are nearly equalled or even outnumbered by the naturalised; and if the natives have not been actually exterminated, their numbers have been greatly reduced, and this is the first stage towards extinction. A mountain is an island on the land; and the intertropical mountains before the Glacial period must have been completely isolated; and I believe that the productions of these islands on the land yielded to those produced within the larger areas of the north, just in the same way as the productions of real islands have everywhere lately yielded to continental forms, naturalised by man's agency.

I am far from supposing that all difficulties are removed on the view here given in regard to the range and affinities of the allied species which live in the northern and southern temperate zones and on the mountains of the intertropical regions. Very many difficulties remain to be solved. I do not pretend to indicate the exact lines and means of migration, or the reason why certain species and not others have migrated; why certain species have been modified and have given rise to new groups of forms, and others have remained unaltered. We cannot hope to explain such facts, until we can say why one species and not another becomes naturalised by man's agency in a foreign land; why one ranges twice or thrice as far, and is twice or thrice as common, as another species within their own homes.

I have said that many difficulties remain to be solved: some of the most remarkable are stated with admirable clearness by Dr. Hooker in his botanical works on the antarctic regions. These cannot be here discussed. I will only say that as far as regards the occurrence of identical species at points so enormously remote as Kerguelen Land, New Zealand, and Fuegia, I believe that towards the close of the Glacial period, icebergs, as suggested by Lyell, have been largely concerned in their dispersal. But the existence of several quite distinct species, belonging to genera exclusively confined to the south, at these and other distant points of the southern hemisphere, is, on my theory of descent with modification, a far more remarkable case of difficulty. For some of these species are so distinct, that we cannot suppose that there has been time since the commencement of the Glacial period for their migration, and for their subsequent modification to the necessary degree. The facts seem to me to indicate that peculiar and very distinct species have migrated in radiating lines from some common centre; and I am inclined to look in the southern, as in the northern hemisphere, to a former and warmer period, before the commencement of the Glacial period, when the antarctic lands, now covered with ice, supported a highly peculiar and isolated flora. I suspect that before this flora was exterminated by the Glacial epoch, a few forms were widely dispersed to various points of the southern hemisphere by occasional means of transport, and by the aid, as halting-places, of existing and now sunken islands, and perhaps at the commencement of the Glacial period, by icebergs. By these means, as I believe, the southern shores of America, Australia, New Zealand have become slightly tinted by the same peculiar forms of vegetable life.

Sir C. Lyell in a striking passage has speculated, in language almost identical with mine, on the effects of great alternations of climate on geographical distribution. I believe that the world has recently felt one of his great cycles of change; and that on this view, combined with modification through natural selection, a multitude of facts in the present distribution both of the same and of allied forms of life can be explained. The living waters may be said to have flowed during one short period from the north and from the south, and to have crossed at the equator; but to have flowed with greater force from the north so as to have freely inundated the south. As the tide leaves its drift in horizontal lines, though rising higher on the shores where the tide rises highest, so have the living waters left their living drift on our mountain-summits, in a line gently rising from the arctic lowlands to a great height under the equator. The various beings thus left stranded may be compared with savage races of man, driven up and surviving in the mountain-fastnesses of almost every land, which serve as a record, full of interest to us, of the former inhabitants of the surrounding lowlands.

12. GEOGRAPHICAL DISTRIBUTION – continued

Distribution of fresh-water productions. On the inhabitants of oceanic islands. Absence of Batrachians and of terrestrial Mammals. On the relation of the inhabitants of islands to those of the nearest mainland. On colonisation from the nearest source with subsequent modification. Summary of the last and present chapters.

As lakes and river-systems are separated from each other by barriers of land, it might have been thought that fresh-water productions would not have ranged widely within the same country, and as the sea is apparently a still more impassable barrier, that they never would have extended to distant countries. But the case is exactly the reverse. Not only have many fresh-water species, belonging to quite different classes, an enormous range, but allied species prevail in a remarkable manner throughout the world. I well remember, when first collecting in the fresh waters of Brazil, feeling much surprise at the similarity of the fresh-water insects, shells, etc., and at the dissimilarity of the surrounding terrestrial beings, compared with those of Britain.

But this power in fresh-water productions of ranging widely, though so unexpected, can, I think, in most cases be explained by their having become fitted, in a manner highly useful to them, for short and frequent migrations from pond to pond, or from stream to stream; and liability to wide dispersal would follow from this capacity as an almost necessary consequence. We can here consider only a few cases. In regard to fish, I believe that the same species never occur in the fresh waters of distant continents. But on the same continent the species often range widely and almost capriciously; for two river-systems will have some fish in common and some different. A few facts seem to favour the possibility of their occasional transport by accidental means; like that of the live fish not rarely dropped by whirlwinds in India, and the vitality of their ova when removed from the water. But I am inclined to attribute the dispersal of fresh-water fish mainly to slight changes within the recent period in the level of the land, having caused rivers to flow into each other. Instances, also, could be given of this having occurred during floods, without any change of level. We have evidence in the loess of the Rhine of considerable changes of level in the land within a very recent geological period, and when the surface was peopled by existing land and fresh-water shells. The wide difference of the fish on opposite sides of continuous mountain-ranges, which from an early period must have parted river-systems and completely prevented their inosculation, seems to lead to this same conclusion. With respect to allied fresh-water fish occurring at very distant points of the world, no doubt there are many cases which cannot at present be explained: but some fresh-water fish belong to very ancient forms, and in such cases there will have been ample time for great geographical changes, and consequently time and means for much migration. In the second place, salt-water fish can with care be slowly accustomed to live in fresh water; and, according to Valenciennes, there is hardly a single group of fishes confined exclusively to fresh water, so that we may imagine that a marine member of a fresh-water group might travel far along the shores of the sea, and subsequently become modified and adapted to the fresh waters of a distant land.

Some species of fresh-water shells have a very wide range, and allied species, which, on my theory, are descended from a common parent and must have proceeded from a single source, prevail throughout the world. Their distribution at first perplexed me much, as their ova are not likely to be transported by birds, and they are immediately killed by sea water, as are the adults. I could not even understand how some naturalised species have rapidly spread throughout the same country. But two facts, which I have observed – and no doubt many others remain to be observed – throw some light on this subject. When a duck suddenly emerges from a pond covered with duck-weed, I have twice seen these little plants adhering to its back; and it has happened to me, in removing a little duck-weed from one aquarium to another, that I have quite unintentionally stocked the one with fresh-water shells from the other. But another agency is perhaps more effectual: I suspended a duck's feet, which might represent those of a bird sleeping in a natural pond, in an aquarium, where many ova of fresh-water shells were hatching; and I found that numbers of the extremely minute and just hatched shells crawled on the feet, and clung to them so firmly that when taken out of the water they could not be jarred off, though at a somewhat more advanced age they would voluntarily drop off. These just hatched molluscs, though aquatic in their nature, survived on the duck's feet, in damp air, from twelve to twenty hours; and in this length of time a duck or heron might fly at least six or seven hundred miles, and would be sure to alight on a pool or rivulet, if blown across sea to an oceanic island or to any other distant point. Sir Charles Lyell also informs me that a Dyticus has been caught with an Ancylus (a fresh-water shell like a limpet) firmly adhering to it; and a water-beetle of the same family, a Colymbetes, once flew on board the 'Beagle,' when forty-five miles distant from the nearest land: how much farther it might have flown with a favouring gale no one can tell.

With respect to plants, it has long been known what enormous ranges many fresh-water and even marsh-species have, both over continents and to the most remote oceanic islands. This is strikingly shown, as remarked by Alph. de Candolle, in large groups of terrestrial plants, which have only a very few aquatic members; for these latter seem immediately to acquire, as if in consequence, a very wide range. I think favourable means of dispersal explain this fact. I have before mentioned that earth occasionally, though rarely, adheres in some quantity to the feet and beaks of birds. Wading birds, which frequent the muddy edges of ponds, if suddenly flushed, would be the most likely to have muddy feet. Birds of this order I can show are the greatest wanderers, and are occasionally found on the most remote and barren islands in the open ocean; they would not be likely to alight on the surface of the sea, so that the dirt would not be washed off their feet; when making land, they would be sure to fly to their natural fresh-water haunts. I do not believe that botanists are aware how charged the mud of ponds is with seeds: I have tried several little experiments, but will here give only the most striking case: I took in February three table-spoonfuls of mud from three different points, beneath water, on the edge of a little pond; this mud when dry weighed only 6 3/4 ounces; I kept it covered up in my study for six months, pulling up and counting each plant as it grew; the plants were of many kinds, and were altogether 537 in number; and yet the viscid mud was all contained in a breakfast cup! Considering these facts, I think it would be an inexplicable circumstance if water-birds did not transport the seeds of fresh-water plants to vast distances, and if consequently the range of these plants was not very great. The same agency may have come into play with the eggs of some of the smaller fresh-water animals.

Other and unknown agencies probably have also played a part. I have stated that fresh-water fish eat some kinds of seeds, though they reject many other kinds after having swallowed them; even small fish swallow seeds of moderate size, as of the yellow water-lily and Potamogeton. Herons and other birds, century after century, have gone on daily devouring fish; they then take flight and go to other waters, or are blown across the sea; and we have seen that seeds retain their power of germination, when rejected in pellets or in excrement, many hours afterwards. When I saw the great size of the seeds of that fine water-lily, the Nelumbium, and remembered Alph. de Candolle's remarks on this plant, I thought that its distribution must remain quite inexplicable; but Audubon states that he found the seeds of the great southern water-lily (probably, according to Dr. Hooker, the Nelumbium luteum) in a heron's stomach; although I do not know the fact, yet analogy makes me believe that a heron flying to another pond and getting a hearty meal of fish, would probably reject from its stomach a pellet containing the seeds of the Nelumbium undigested; or the seeds might be dropped by the bird whilst feeding its young, in the same way as fish are known sometimes to be dropped.

In considering these several means of distribution, it should be remembered that when a pond or stream is first formed, for instance, on a rising islet, it will be unoccupied; and a single seed or egg will have a good chance of succeeding. Although there will always be a struggle for life between the individuals of the species, however few, already occupying any pond, yet as the number of kinds is small, compared with those on the land, the competition will probably be less severe between aquatic than between terrestrial species; consequently an intruder from the waters of a foreign country, would have a better chance of seizing on a place, than in the case of terrestrial colonists. We should, also, remember that some, perhaps many, fresh-water productions are low in the scale of nature, and that we have reason to believe that such low beings change or become modified less quickly than the high; and this will give longer time than the average for the migration of the same aquatic species. We should not forget the probability of many species having formerly ranged as continuously as fresh-water productions ever can range, over immense areas, and having subsequently become extinct in intermediate regions. But the wide distribution of fresh-water plants and of the lower animals, whether retaining the same identical form or in some degree modified, I believe mainly depends on the wide dispersal of their seeds and eggs by animals, more especially by fresh-water birds, which have large powers of flight, and naturally travel from one to another and often distant piece of water. Nature, like a careful gardener, thus takes her seeds from a bed of a particular nature, and drops them in another equally well fitted for them.

ON THE INHABITANTS OF OCEANIC ISLANDS.

We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe. I have already stated that I cannot honestly admit Forbes's view on continental extensions, which, if legitimately followed out, would lead to the belief that within the recent period all existing islands have been nearly or quite joined to some continent. This view would remove many difficulties, but it would not, I think, explain all the facts in regard to insular productions. In the following remarks I shall not confine myself to the mere question of dispersal; but shall consider some other facts, which bear on the truth of the two theories of independent creation and of descent with modification.

The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. de Candolle admits this for plants, and Wollaston for insects. If we look to the large size and varied stations of New Zealand, extending over 780 miles of latitude, and compare its flowering plants, only 750 in number, with those on an equal area at the Cape of Good Hope or in Australia, we must, I think, admit that something quite independently of any difference in physical conditions has caused so great a difference in number. Even the uniform county of Cambridge has 847 plants, and the little island of Anglesea 764, but a few ferns and a few introduced plants are included in these numbers, and the comparison in some other respects is not quite fair. We have evidence that the barren island of Ascension aboriginally possessed under half-a-dozen flowering plants; yet many have become naturalised on it, as they have on New Zealand and on every other oceanic island which can be named. In St. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions. He who admits the doctrine of the creation of each separate species, will have to admit, that a sufficient number of the best adapted plants and animals have not been created on oceanic islands; for man has unintentionally stocked them from various sources far more fully and perfectly than has nature.

Although in oceanic islands the number of kinds of inhabitants is scanty, the proportion of endemic species (i.e. those found nowhere else in the world) is often extremely large. If we compare, for instance, the number of the endemic land-shells in Madeira, or of the endemic birds in the Galapagos Archipelago, with the number found on any continent, and then compare the area of the islands with that of the continent, we shall see that this is true. This fact might have been expected on my theory, for, as already explained, species occasionally arriving after long intervals in a new and isolated district, and having to compete with new associates, will be eminently liable to modification, and will often produce groups of modified descendants. But it by no means follows, that, because in an island nearly all the species of one class are peculiar, those of another class, or of another section of the same class, are peculiar; and this difference seems to depend on the species which do not become modified having immigrated with facility and in a body, so that their mutual relations have not been much disturbed. Thus in the Galapagos Islands nearly every land-bird, but only two out of the eleven marine birds, are peculiar; and it is obvious that marine birds could arrive at these islands more easily than land-birds. Bermuda, on the other hand, which lies at about the same distance from North America as the Galapagos Islands do from South America, and which has a very peculiar soil, does not possess one endemic land bird; and we know from Mr. J. M. Jones's admirable account of Bermuda, that very many North American birds, during their great annual migrations, visit either periodically or occasionally this island. Madeira does not possess one peculiar bird, and many European and African birds are almost every year blown there, as I am informed by Mr. E. V. Harcourt. So that these two islands of Bermuda and Madeira have been stocked by birds, which for long ages have struggled together in their former homes, and have become mutually adapted to each other; and when settled in their new homes, each kind will have been kept by the others to their proper places and habits, and will consequently have been little liable to modification. Madeira, again, is inhabited by a wonderful number of peculiar land-shells, whereas not one species of sea-shell is confined to its shores: now, though we do not know how seashells are dispersed, yet we can see that their eggs or larvae, perhaps attached to seaweed or floating timber, or to the feet of wading-birds, might be transported far more easily than land-shells, across three or four hundred miles of open sea. The different orders of insects in Madeira apparently present analogous facts.

Oceanic islands are sometimes deficient in certain classes, and their places are apparently occupied by the other inhabitants; in the Galapagos Islands reptiles, and in New Zealand gigantic wingless birds, take the place of mammals. In the plants of the Galapagos Islands, Dr. Hooker has shown that the proportional numbers of the different orders are very different from what they are elsewhere. Such cases are generally accounted for by the physical conditions of the islands; but this explanation seems to me not a little doubtful. Facility of immigration, I believe, has been at least as important as the nature of the conditions.

Many remarkable little facts could be given with respect to the inhabitants of remote islands. For instance, in certain islands not tenanted by mammals, some of the endemic plants have beautifully hooked seeds; yet few relations are more striking than the adaptation of hooked seeds for transportal by the wool and fur of quadrupeds. This case presents no difficulty on my view, for a hooked seed might be transported to an island by some other means; and the plant then becoming slightly modified, but still retaining its hooked seeds, would form an endemic species, having as useless an appendage as any rudimentary organ, – for instance, as the shrivelled wings under the soldered elytra of many insular beetles. Again, islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species; now trees, as Alph. de Candolle has shown, generally have, whatever the cause may be, confined ranges. Hence trees would be little likely to reach distant oceanic islands; and an herbaceous plant, though it would have no chance of successfully competing in stature with a fully developed tree, when established on an island and having to compete with herbaceous plants alone, might readily gain an advantage by growing taller and taller and overtopping the other plants. If so, natural selection would often tend to add to the stature of herbaceous plants when growing on an island, to whatever order they belonged, and thus convert them first into bushes and ultimately into trees.

With respect to the absence of whole orders on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts) have never been found on any of the many islands with which the great oceans are studded. I have taken pains to verify this assertion, and I have found it strictly true. I have, however, been assured that a frog exists on the mountains of the great island of New Zealand; but I suspect that this exception (if the information be correct) may be explained through glacial agency. This general absence of frogs, toads, and newts on so many oceanic islands cannot be accounted for by their physical conditions; indeed it seems that islands are peculiarly well fitted for these animals; for frogs have been introduced into Madeira, the Azores, and Mauritius, and have multiplied so as to become a nuisance. But as these animals and their spawn are known to be immediately killed by sea-water, on my view we can see that there would be great difficulty in their transportal across the sea, and therefore why they do not exist on any oceanic island. But why, on the theory of creation, they should not have been created there, it would be very difficult to explain.

Mammals offer another and similar case. I have carefully searched the oldest voyages, but have not finished my search; as yet I have not found a single instance, free from doubt, of a terrestrial mammal (excluding domesticated animals kept by the natives) inhabiting an island situated above 300 miles from a continent or great continental island; and many islands situated at a much less distance are equally barren. The Falkland Islands, which are inhabited by a wolf-like fox, come nearest to an exception; but this group cannot be considered as oceanic, as it lies on a bank connected with the mainland; moreover, icebergs formerly brought boulders to its western shores, and they may have formerly transported foxes, as so frequently now happens in the arctic regions. Yet it cannot be said that small islands will not support small mammals, for they occur in many parts of the world on very small islands, if close to a continent; and hardly an island can be named on which our smaller quadrupeds have not become naturalised and greatly multiplied. It cannot be said, on the ordinary view of creation, that there has not been time for the creation of mammals; many volcanic islands are sufficiently ancient, as shown by the stupendous degradation which they have suffered and by their tertiary strata: there has also been time for the production of endemic species belonging to other classes; and on continents it is thought that mammals appear and disappear at a quicker rate than other and lower animals. Though terrestrial mammals do not occur on oceanic islands, aerial mammals do occur on almost every island. New Zealand possesses two bats found nowhere else in the world: Norfolk Island, the Viti Archipelago, the Bonin Islands, the Caroline and Marianne Archipelagoes, and Mauritius, all possess their peculiar bats. Why, it may be asked, has the supposed creative force produced bats and no other mammals on remote islands? On my view this question can easily be answered; for no terrestrial mammal can be transported across a wide space of sea, but bats can fly across. Bats have been seen wandering by day far over the Atlantic Ocean; and two North American species either regularly or occasionally visit Bermuda, at the distance of 600 miles from the mainland. I hear from Mr. Tomes, who has specially studied this family, that many of the same species have enormous ranges, and are found on continents and on far distant islands. Hence we have only to suppose that such wandering species have been modified through natural selection in their new homes in relation to their new position, and we can understand the presence of endemic bats on islands, with the absence of all terrestrial mammals.

Besides the absence of terrestrial mammals in relation to the remoteness of islands from continents, there is also a relation, to a certain extent independent of distance, between the depth of the sea separating an island from the neighbouring mainland, and the presence in both of the same mammiferous species or of allied species in a more or less modified condition. Mr. Windsor Earl has made some striking observations on this head in regard to the great Malay Archipelago, which is traversed near Celebes by a space of deep ocean; and this space separates two widely distinct mammalian faunas. On either side the islands are situated on moderately deep submarine banks, and they are inhabited by closely allied or identical quadrupeds. No doubt some few anomalies occur in this great archipelago, and there is much difficulty in forming a judgment in some cases owing to the probable naturalisation of certain mammals through man's agency; but we shall soon have much light thrown on the natural history of this archipelago by the admirable zeal and researches of Mr. Wallace. I have not as yet had time to follow up this subject in all other quarters of the world; but as far as I have gone, the relation generally holds good. We see Britain separated by a shallow channel from Europe, and the mammals are the same on both sides; we meet with analogous facts on many islands separated by similar channels from Australia. The West Indian Islands stand on a deeply submerged bank, nearly 1000 fathoms in depth, and here we find American forms, but the species and even the genera are distinct. As the amount of modification in all cases depends to a certain degree on the lapse of time, and as during changes of level it is obvious that islands separated by shallow channels are more likely to have been continuously united within a recent period to the mainland than islands separated by deeper channels, we can understand the frequent relation between the depth of the sea and the degree of affinity of the mammalian inhabitants of islands with those of a neighbouring continent, – an inexplicable relation on the view of independent acts of creation.