Geographical Distribution Continued

On The Origin of Species

By Charles Robert Darwin

Chapter XIII: Geographical Distribution Continued

Fresh-water Productions

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 formidable barrier, that they would never have extended to distant countries. But the case is exactly the reverse. Not only have many fresh-water species, belonging to different classes, an enormous range, but allied species prevail in a remarkable manner throughout the world. When first collecting in the fresh waters of Brazil, I well remember feeling much surprise at the similarity of the fresh-water insects, shells , and at the dissimilarity of the surrounding terrestrial beings, compared with those of Britain.

But the wide ranging power of fresh-water productions 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, within their own countries; and liability to wide dispersal would follow from this capacity as an almost necessary consequence. We can here consider only a few cases; of these, some of the most difficult to explain are presented by fish. It was formerly believed that the same fresh-water species never existed on two continents distant from each other. But Dr. Gunther has lately shown that the Galaxias attenuatus inhabits Tasmania, New Zealand, the Falkland Islands, and the mainland of South America. This is a wonderful case, and probably indicates dispersal from an Antarctic centre during a former warm period. This case, however, is rendered in some degree less surprising by the species of this genus having the power of crossing by some unknown means considerable spaces of open ocean: thus there is one species common to New Zealand and to the Auckland Islands, though separated by a distance of about 230 miles. On the same continent fresh-water fish often range widely, and as if capriciously; for in two adjoining river-systems some of the species may be the same, and some wholly different.

It is probable that they are occasionally transported by what may be called accidental means. Thus fishes still alive are not very rarely dropped at distant points by whirlwinds; and it is known that the ova retain their vitality for a considerable time after removal from the water. Their dispersal may, however, be mainly attributed to changes in the level of the land within the recent period, causing rivers to flow into each other. Instances, also, could be given of this having occurred during floods, without any change of level. The wide difference of the fish on the opposite sides of most mountain-ranges, which are continuous, and which consequently must from an early period have completely prevented the inosculation of the river-systems on the two sides, leads to the same conclusion. 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. Moreover, Dr. Gunther has recently been led by several considerations to infer that with fishes the same forms have a long endurance. 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 which an the members are confined to fresh water, so that a marine species belonging to a fresh-water group might travel far along the shores of the sea, and could, it is probable, become adapted without much difficulty to the fresh waters of a distant land.

Some species of fresh-water shells have very wide ranges, and allied species which, on our 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 the ova, as well as the adults, are immediately killed by sea-water. I could not even understand how some naturalised species have spread rapidly throughout the same country. But two facts, which I have observed- and many others no doubt will be discovered- throw some light on this subject. When ducks suddenly emerge from a pond covered with duck-weed, I have twice seen these little plants adhering to their backs; and it has happened to me, in removing a little duck-weed from one aquarium to another, that I have unintentionally stocked the one with fresh-water shells from the other. But another agency is perhaps more effectual: I suspended the feet of a duck 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 if blown across the sea to an oceanic island, or to any other distant point, would be sure to alight on a pool or rivulet. Sir Charles Lyell informs me that a Dytiscus 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 been blown by 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 illustrated, according to Alph. de Candolle, in those large groups of terrestrial plants, which have very few aquatic members; for the latter seem immediately to acquire, as if in consequence, a wide range. I think favourable means of dispersal explain this fact. I have before mentioned that earth occasionally 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 wander more than those of any other; and they are occasionally found on the most remote and barren islands of the open ocean; they would not be likely to alight on the surface of the sea, so that any dirt on their feet would not be washed off; and when gaining the 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 tablespoonfuls of mud from three different points, beneath water, on the edge of a little pond: this mud when dried weighed only 63/4 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 unstocked ponds and streams, situated at very distant points. 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 many hours afterwards in pellets or in the excrement. When I saw the great size of the seeds of that fine water-lily, the Nelumbium, and remembered Alph. de Candolle's remarks on the distribution of this plant, I thought that the means of its dispersal must remain 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. Now this bird must often have flown with its stomach thus well stocked to distant ponds, and then getting a hearty meal of fish, analogy makes me believe that it would have rejected the seeds in a pellet in a fit state for germination.

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 inhabitants of the same pond, however few in kind, yet as the number even in a well-stocked pond is small in comparison with the number of species inhabiting an equal area of land, the competition between them will probably be less severe than between terrestrial species; consequently an intruder from the waters of a foreign country would have a better chance of seizing on new place, than in the case of terrestrial colonists. We should also remember that many fresh-water productions are low in the scale of nature, and we have reason to believe that such beings become modified more slowly than the high; and this will give time for the migration of aquatic species. We should not forget the probability of many fresh-water forms laving formerly ranged continuously over immense areas, and then having become extinct at intermediate points. But the wide distribution of fresh-water plants and of the lower animals, whether retaining the same identical form or in some degree modified, apparently depends in main part on the wide dispersal of their seeds and eggs by animals, more especially by fresh-water birds, which have great powers of flight, and naturally travel from one piece of water to another.

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 with respect to distribution, on the view that not only all the individuals of the same species have migrated from some one area, but that allied species, although now inhabiting the most distant points, have proceeded from a single area,- the birthplace of their early progenitors. I have already given my reasons for disbelieving in continental extensions within the period of existing species, on so enormous a scale that all the many islands of the several oceans were thus stocked with their present terrestrial inhabitants. This view removes many difficulties, but it does not accord with all the facts in regard to the productions of islands. In the following remarks I shall not confine myself to the mere question of dispersal, but shall consider some other cases bearing 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. New Zealand, for instance, with its lofty mountains and diversified stations, extending over 780 miles of latitude, together with the outlying islands of Auckland, Campbell and Chatham, contain altogether only 960 kinds of flowering plants; if we compare this moderate number with the species which swarm over equal areas in South-Western Australia or at the Cape of Good Hope, we must admit that some cause, independently of different physical conditions, has given rise to 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 less than half-a-dozen flowering plants; yet many species have now become naturalised on it, as they have in 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 were not created for oceanic islands; for man has unintentionally stocked them far more fully and perfectly than did nature.

Although in oceanic islands the species are few in number, the proportion of endemic kinds (i.e., those found nowhere else in the world) is often extremely large. If we compare, for instance, the number of endemic landshells in Madeira, or of endemic birds in the Galapagos Archipelago, with the number found on any continent, and then compare the area of the island with that of the continent, we shall see that this is true. This fact might have been theoretically expected, for, as already explained, species occasionally arriving after long intervals of time in the new and isolated district, and having to compete with new associates, would be eminently liable to modification, and would 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 partly on the species which are not modified having immigrated in a body, so that their mutual relations have not been much disturbed; and partly on the frequent arrival of unmodified immigrants from the mother-country, with which the insular forms have intercrossed. It should be borne in mind that the offspring of such crosses would certainly gain in vigour, so that even an occasional cross would produce more effect than might have been anticipated. I will give a few illustrations of the foregoing remarks: in the Galapagos Islands there are 9.6 land-birds; of these 21 (or perhaps 93) are peculiar, whereas of the 11 marine birds only 2 are peculiar; and it is obvious that marine birds could arrive at these islands much more easily and frequently 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 a single endemic landbird, and we know from Mr. J. M. Jones's admirable account of Bermuda, that very many North American birds occasionally or even frequently visit this, island. Almost every year, as I am informed by Mr. E. V. Harcourt, many European and African birds are blown to Madeira; this island is inhabited by 99 kinds of which one alone is peculiar, though very closely related to a European form; and three or four other species are confined to this island and to the Canaries. So that the islands of Bermuda and Madeira have been stocked from the neighbouring continents with birds, which for long ages have there struggled together, and have become mutually co-adapted. Hence when settled in their new homes, each kind will have been kept by the others to its proper place and habits, and will consequently have been but little liable to modification. Any tendency to modification will also have been checked by intercrossing with the unmodified immigrants, often arriving from the mother-country. Madeira again is inhabited by a wonderful number of peculiar land-shells, whereas not one species of sea-shell is peculiar 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 across three or four hundred miles of open sea far more easily than land-shells. The different orders of insects inhabiting Madeira present nearly parallel cases.

Oceanic islands are sometimes deficient in animals of certain whole classes, and their places are occupied by other classes; thus in the Galapagos Islands reptiles, and in New Zealand gigantic wingless birds, take, or recently took, the place of mammals. Although New Zealand is here spoken of as an oceanic island, it is in some degree doubtful whether it should be so ranked; it is of large size, and is not separated from Australia by a profoundly deep sea; from its geological character and the direction of its mountain-ranges, the Rev. W. B. Clarke has lately maintained that this island, as well as New Caledonia, should be considered as appurtenances of Australia. Turning to plants, Dr. Hooker has shown that in the Galapagos Islands the proportional numbers of the different orders are very different from what they are elsewhere. All such differences in number, and the absence of certain whole groups of animals and plants, are generally accounted for by supposed differences in the physical conditions of the islands; but this explanation is not a little doubtful. Facility of immigration seems to have been fully as important as the nature of the conditions.

Many remarkable little facts could be given with respect to the inhabitants of oceanic islands. For instance, in certain islands not tenanted by a single mammal, some of the endemic plants have beautifully hooked seeds; yet few relations are more manifest than that hooks serve for the transportal of seeds in the wool or fur of quadrupeds. But a hooked seed might be carried to an island by other means; and the plant then becoming modified would form an endemic species, still retaining its hooks, which would form a useless appendage like the shrivelled wings under the soldered wing-covers 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, which had no chance of successfully competing with the many fully developed trees growing on a continent, might, when established on an island, gain an advantage over other herbaceous plants by growing taller and taller and overtopping them. In this case, natural selection would tend to add to the stature of the plant, to whatever order it belonged, and thus first convert it into a bush and then into a tree.

Absence of Batrachians and Terrestrial Mammals on Oceanic Islands

With respect to the absence of whole orders of animals on oceanic islands, Bory St. Vincent long ago remarked that batrachians (frogs, toads, newts) are never found on any of the many islands with which the great oceans are studded. I have taken pains to verify this assertion, and have found it true, with the exception of New Zealand, New Caledonia, the Andaman Islands, and perhaps the Solomon Islands and the Seychelles. But I have already remarked that it is doubtful whether New Zealand and New Caledonia ought to be classed as oceanic islands; and this is still more doubtful with respect to the Andaman and Solomon groups and the Seychelles. This general absence of frogs, toads, and newts on so many true oceanic islands cannot be accounted for by their physical conditions: indeed it seems that islands are peculiarly 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 immediately killed (with the exception, as far as known, of one Indian species) by sea-water, there would be great difficulty in their transportal across the sea, and therefore we can see why they do not exist on strictly oceanic islands. 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, and 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 in connection with the mainland at the distance of about 280 miles; moreover, icebergs formerly brought boulders to its western shores, and they may have formerly transported foxes, as now frequently happens in the arctic regions. Yet it cannot be said that small islands will not support at least small mammals, for they occur in many parts of the world on very small islands, when lying 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 known that new species of mammals appear and disappear at a quicker rate than other and lower animals. Although 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 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 in their new homes in relation to their new position, and we can understand the presence of endemic bats on oceanic islands, with the absence of all other terrestrial mammals.

Another interesting relation exists, namely, between the depth of the sea separating islands from each other or from the nearest continent, and the degree of affinity of their mammalian inhabitants. Mr. Windsor Earl has made some striking observations on this head, since greatly extended by Mr. Wallace's admirable researches, in regard to the great Malay Archipelago, which is traversed near Celebes by a space of deep ocean, and this separates two widely distinct mammalian faunas. On either side the islands stand on a moderately shallow submarine bank, and these islands are inhabited by the same or by closely allied quadrupeds. I have not as yet had time to follow up this subject in all quarters of the world; but as far as I have gone, the relation holds good. For instance, Britain is separated by a shallow channel from Europe, and the mammals are the same on both sides; and so it is with all the islands near the shores of Australia. The West Indian Islands, on the other hand, 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 quite distinct. As the amount of modification which animals of all kinds undergo partly depends on the lapse of time, and as the islands which are separated from each other or from the mainland by shallow channels, are more likely to have been continuously united within a recent period than the islands separated by deeper channels, we can understand how it is that a relation exists between the depth of the sea separating two mammalian faunas, and the degree of their affinity,- a relation which is quite inexplicable on the theory of independent acts of creation.

The foregoing statements in regard to the inhabitants of oceanic islands,- namely, the fewness of the species, with a large proportion consisting of endemic forms,- the members of certain groups, but not those of other groups in the same class, having been modified,- the absence of certain whole orders, as of batrachians and of terrestrial mammals, notwithstanding the presence of aerial bats,- the singular proportions of certain orders of plants,- herbaceous forms having been developed into trees,- seem to me to accord better with the belief in the efficiency of occasional means of transport, carried on during a long course of time, than with the belief in the former connection of all oceanic islands with the nearest continent; for on this latter view it is probable that the various classes would have immigrated more uniformly, and from the species having entered in a body their mutual relations would not have been much disturbed, and consequently they would either have not been modified, or all the species in a more equable manner.

I do not deny that there are many and serious difficulties in understanding how many Of the inhabitants of the inhabitants of the more remote islands, whether still retaining the same specific form or subsequently modified, have reached their present homes. But the probability of other islands having once existed as halting-places, of which not a wreck now remains, must not be overlooked. I will specify one difficult case. Almost all oceanic islands, even the most isolated and smallest, are inhabited by landshells, generally by endemic species, but sometimes by species found elsewhere,- striking instances of which have been given by Dr. A. A. Gould in relation to the Pacific. Now it is notorious that land-shells are easily killed by sea-water; their eggs, at least such as I have tried, sink in it and are killed. Yet there must be some unknown, but occasionally efficient means for their transportal. Would the just-hatched young sometimes adhere to the feet of birds roosting on the ground, and thus get transported? It occurred to me that landshells, when hibernating and having a membranous diaphragm over the mouth of the shell, might be floated in chinks of drifted timber across moderately wide arms of the sea. And I find that several species in this state withstand uninjured an immersion in sea-water during seven days: one shell, the Helix pomatia, after having been thus treated and again hibernating was put into sea-water for twenty days, and perfectly recovered. During this length of time the shell might have been carried by a marine current of average swiftness, to a distance of 660 geographical miles. As this Helix has a thick calcareous operculum, I removed it, and

when it had formed a new membranous one, I again immersed it for fourteen days in sea-water, and again it recovered and crawled away. Baron Aucapitaine has since tried similar experiments: he placed 100 landshells, belonging to ten species, in a box pierced with holes, and immersed it for a fortnight in the sea. Out of the hundred shells, twenty-seven recovered. The presence of an operculum seems to have been of importance, as out of twelve specimens of Cyclostoma elegans, which is thus furnished, eleven revived. It is remarkable, seeing how well the Helix pomatia resisted with me the salt-water, that not one of fifty-four specimens belonging to four other species of Helix tried by Aucapitaine, recovered. It is, however, not at all probable that land-shells have often been thus transported; the feet of birds offer a more probable method.

On the Relations of the Inhabitants of Islands to those of the nearest Mainland

The most striking and important fact for us is the affinity of the species which inhabit islands to those of the nearest mainland, without being actually the same. Numerous instances could be given. The Galapagos Archipelago, situated under the equator, lies at the distance of between 500 and 600 miles from the shores of South America. Here almost every product of the land and of the water bears the unmistakable stamp of the American continent. There are twenty-six land-birds; of these, twenty-one, or perhaps twenty-three are ranked as distinct species, and would commonly be assumed to have been here created; yet the close affinity of most of these birds to American species is manifest in every character, in their habits, gestures, and tones of voice. So it is with the other animals, and with a large proportion of the plants, as shown by Dr. Hooker in his admirable Flora of this archipelago. The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, feels that he is standing on American land. Why should this be so? Why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plainly the stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which closely resemble; the conditions of the South American coast: in fact, there is a considerable dissimilarity in all these respects. On the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in the climate, height, and size of the islands, between the Galapagos and Cape Verde Archipelagoes: but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape Verde Islands are related to those of Africa, like those of the Galapagos to America. Facts such as these admit of no sort of explanation on the ordinary view of independent creation; whereas on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists from America, whether by occasional means of transport or (though I do not believe in this doctrine) by formerly continuous land, and the Cape Verde Islands from Africa; such colonists would be liable to modification,- the principle of inheritance still betraying their original birthplace.

Many analogous facts could be given: indeed it is an almost universal rule that the endemic productions of islands are related to those of the nearest continent, or of the nearest large island. The exceptions are few, and most of them can be explained. Thus although Kerguelen Land stands nearer to Africa than to America, the plants are related, and that very closely, as we know from Dr. Hooker's account, to those of America: but on the view that this island has been mainly stocked by seeds brought with earth and stones on icebergs, drifted by the prevailing currents, this anomaly disappears. New Zealand in its endemic planes is much more closely related to Australia, the nearest mainland, than to any other region: and this is what might have been expected; but it is also plainly related to South America, which, although the next nearest continent, is so enormously remote, that the fact becomes an anomaly. But this difficulty partially disappears on the view that New Zealand, South America, and the other southern lands have been stocked in part from a nearly intermediate though distant point, namely from the antarctic islands, when they were clothed with vegetation, during a warmer tertiary period, before the commencement of the last Glacial period. The affinity, which though feeble, I am assured by Dr. Hooker is real, between the flora of the south-western corner of Australia and of the Cape of Good Hope, is a far more remarkable case; but this affinity is confined to the plants, and will, no doubt, some day be explained.

The same law which has determined the relationship between the inhabitants of islands and the nearest mainland, is sometimes displayed on a small scale, but in a most interesting manner, within the limits of the same archipelago. Thus each separate island of the Galapagos Archipelago is tenanted, and the fact is a marvellous one, by many different species; but these species are related to each other in a very much closer manner than to the inhabitants of the American continent, or of any other quarter of the world. This is what might have been expected, for islands situated so near to each other would almost necessarily receive immigrants from the same original source, and from each other. But how is it that many of the immigrants have been differently modified, though only in a small degree, in islands situated within sight of each other, having the same geological nature, the same height, climate? This long appeared to me a great difficulty: but it arises in chief part from the deeply-seated error of considering the physical conditions of a country as the most important; whereas it cannot be disputed that the nature of the other species with which each has to compete, is at least as important, and generally a far more important element of success. Now if we look to the species which inhabit the Galapagos Archipelago, and are likewise found in other parts of the world, we find that they differ considerably in the several islands. This difference might indeed have been expected if the islands had been stocked by occasional means of transport- a seed, for instance, of one plant having been brought to one island, and that of another plant to another island, though all proceeding from the same general source. Hence, when in former times an immigrant first settled on one of the islands, or when it subsequently spread from one to another, it would undoubtedly be exposed to different conditions in the different islands, for it would have to compete with a different set of organisms; a plant, for instance, would find the ground best fitted for it occupied by somewhat different species in the different islands, and would be exposed to the attacks of somewhat different enemies. If then it varied, natural selection would probably favour different varieties in the different islands. Some species, however, might spread and yet retain the same character throughout the group, just as we see some species spreading widely throughout a continent and remaining the same.

The really surprising fact in this case of the Galapagos Archipelago, and in a lesser degree in some analogous cases, is that each new species after being formed in any one island, did not spread quickly to the other islands. But the islands, though in sight of each other, are separated by deep arms of the sea, in most cases wider than the British Channel, and there is no reason to suppose that they have at any former period been continuously united. The currents of the sea are rapid and sweep between the islands, and gales of wind are extraordinarily rare; so that the islands are far more effectually separated from each other than they appear on a map. Nevertheless some of the species, both of those found in other parts of the world and of those confined to the archipelago, are common to the several islands; and we may infer from their present manner of distribution, that they have spread from one island to the others. But we often take, I think, an erroneous view of the probability of closely-allied species invading each other's territory, when put into free intercommunication. Undoubtedly, if one species has any advantage over another, it will in a very brief time wholly or in part supplant it; but if both are equally well fitted for their own places, both will probably hold their separate places for almost any length of time. Being familiar with the fact that many species, naturalised through man's agency, have spread with astonishing rapidity over wide areas, we are apt to infer that most species would thus spread; but we should remember that the species which become naturalised in new countries are not generally closely allied to the aboriginal inhabitants, but are very distinct forms, belonging in a large proportion of cases, as shown by Alph. de Candolle, to distinct genera. In the Galapagos Archipelago, many even of the birds, though so well adapted for flying from island to island, differ on the different islands; thus there are three closely-allied species of mocking-thrush, each confined to its own island. Now let us suppose the mocking-thrush of Chatham Island to be blown to Charles Island, which has its own mocking-thrush, why should it succeed in establishing itself there? We may safely infer that Charles Island is well stocked with its own species, for annually more eggs are laid and young birds hatched, than can possibly be reared; and we may infer that the mocking-thrush peculiar to Charles's Island is at least as well fitted for its home as is the species peculiar to Chatham Island. Sir C. Lyell and Mr. Wollaston have communicated to me a remarkable fact bearing on this subject; namely, that Madeira and the adjoining islet of Porto Santo possess many distinct but representative species of land-shells, some of which live in crevices of stone; and although large quantities of stone are annually transported from Porto Santo to Madeira, yet this latter island has not become colonised by the Porto Santo species; nevertheless both islands have been colonised by European land-shells, which no doubt had some advantage over the indigenous species. From these considerations I think we need not greatly marvel at the endemic species which inhabit the several islands of the Galapagos Archipelago, not having all spread from island to island. On the same continent, also, preoccupation has probably played an important part in checking the commingling of the species which inhabit different districts with nearly the same physical conditions. Thus, the south-east and south-west corners of Australia have nearly the same physical conditions, and are united by continuous land, yet they are inhabited by a vast number of distinct mammals, birds, and plants; so it is, according to Mr. Bates, with the butterflies and other animals inhabiting the great, open, and continuous valley of the Amazons.

The same principle which governs the general character of the inhabitants of oceanic islands, namely, the relation to the source whence colonists could have been most easily derived, together with their subsequent modification, is of the widest application throughout nature. We see this on every mountain summit, in every lake and marsh. For Alpine species, excepting in as far as the same species have become widely spread during the Glacial epoch, are related to those of the surrounding lowlands; thus we have in South America, Alpine humming-birds, Alpine rodents, Alpine plants, all strictly belonging to American forms; and it is obvious that a mountain, as it became slowly unheaved, would be colonised from the surrounding lowlands. So it is with the inhabitants of lakes and marshes, excepting in so far as great facility of transport has allowed the same forms to prevail throughout large portions of the world. We see this same principle in the character of most of the blind animals inhabiting the caves of America and of Europe. Other analogous facts could be given. It will, I believe, be found universally true, that wherever in two regions, let them be ever so distant, many closely allied or representative species occur, there will likewise be found some identical species; and wherever many closely-allied species occur, there will be found many forms which some naturalists rank as distinct species, and others as mere varieties; these doubtful forms showing us the steps in the progress of modification.

The relation between the power and extent of migration in certain species, either at the present or at some former period, and the existence at remote points of the world of closely-allied species, is shown in another and more general way. Mr. Gould remarked to me long ago, that in those genera of birds which range over the world, many of the species have very wide ranges. I can hardly doubt that this rule is generally true, though difficult of proof. Amongst mammals, we see it strikingly displayed in bats, and in a lesser degree in the Felidae and Canidae. We see the same rule in the distribution of butterflies and beetles. So it is with most of the inhabitants of fresh water, for many of the genera in the most distinct classes range over the world, and many of the species have enormous ranges. It is not meant that all, but that some of the species have very wide ranges in the genera which range very widely. Nor is it meant that the species in such genera have on an average a very wide range; for this will largely depend on how far the process of modification has gone; for instance, two varieties of the same species inhabit America and Europe, and thus the species has an immense range; but, if variation were to be carried a little further, the two varieties would be ranked as distinct species, and their range would be greatly reduced. Still less is it meant, that species which have the capacity of crossing barriers and ranging widely, as in the case of certain powerfully-winged birds, will necessarily range widely; for we should never forget that to range widely implies not only the power of crossing barriers, but the more important power of being victorious in distant lands in the struggle for life with foreign associates. But according to the view that all the species of a genus, though distributed to the most remote points of the world, are descended from a single progenitor, we ought to find, and I believe as a general rule we do find, that some at least of the species range very widely.

We should bear in mind that many genera in all classes are of ancient origin, and the species in this case will have had ample time for dispersal and subsequent modification. There is also reason to believe from geological evidence, that within each great class the lower organisms change at a slower rate than the higher; consequently they will have had a better chance of ranging widely and of still retaining the same specific character. This fact, together with that of the seeds and eggs of most lowly organised forms being very minute and better fitted for distant transportal, probably accounts for a law which has long been observed, and which has lately been discussed by Alph. de Candolle in regard to plants, namely, that the lower any group of organisms stands the more widely it ranges.

The relations just discussed,- namely, lower organisms ranging more widely than the higher,- some of the species of widely-ranging genera themselves ranging widely,- such facts, as Alpine, lacustrine, and marsh productions being generally related to those which live on the surrounding low lands and dry lands,- the striking relationship between the inhabitants of islands and those of the nearest mainland, the still closer relationship of the distinct inhabitants of the islands in the same archipelago,- are inexplicable on the ordinary view of the independent creation of each species, but are explicable if we admit colonisation from the nearest or readiest source, together with the subsequent adaptation of the colonists to their new homes.

Summary of the last and present Chapters

In these chapters I have endeavoured to show, that if we make due allowance for our ignorance of the full effects of changes of climate and of the level of the land, which have certainly occurred within the recent period, and of other changes which have probably occurred,- if we remember how ignorant we are with respect to the many curious means of occasional transport,- if we bear in mind, and this is a very important consideration, how often a species may have ranged continuously over a wide area, and then have become extinct in the intermediate tracts,- the difficulty is not insuperable in believing that all the individuals of the same species, wherever found, are descended from common parents. And we are led to this conclusion, which has been arrived at by many naturalists under the designation of single centres of creation, by various general considerations, more especially from the importance of barriers of all kinds, and from the analogical distribution of subgenera, genera, and families.

With respect to distinct species belonging to the same genus, which on our theory have spread from one parent-source; if we make the same allowances as before for our ignorance, and remember that some forms of life have changed very slowly, enormous periods of time having been thus granted for their migration, the difficulties are far from insuperable; though in this case, as in that of the individuals of the same species, they are often great.

As exemplifying the effects of climatal changes on distribution, I have attempted to show how important a part the last Glacial period has played, which affected even the equatorial regions, and which, during the alternations of the cold in the north and south, allowed the productions of opposite hemispheres to mingle, and left some of them stranded on the mountain-summits in all parts of the world. As showing how diversified are the means of occasional transport, I have discussed at some little length the means of dispersal of fresh-water productions.

If the difficulties be not insuperable in admitting that in the long course of time all the individuals of the same species, and likewise of the several species belonging to the same genus, have proceeded from some one source; then 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 the high importance of barriers, whether of land or water, in not only separating, but in apparently forming the several zoological and botanical provinces. We can thus understand the concentration of related species within the same areas; and how it is that under different latitudes, for instance in South America, the inhabitants of the plains and mountains, of the forests, marshes, and deserts, are linked together in so mysterious a manner, and are likewise linked to the extinct beings which formerly inhabited the same continent. Bearing in mind that the mutual relation of organism to organism is of the highest importance, we can see why two areas having nearly the same physical conditions should often be inhabited by very different forms of life; for according to the length of time which has elapsed since the colonists entered one of the regions, or both; according to the nature of the communication which allowed certain forms and not others to enter, either in greater or lesser numbers; according or not, as those which entered happened to come into more or less direct competition with each other and with the aborigines; and according as the immigrants were capable of varying more or less rapidly, there would ensue in the two or more regions, independently of their physical conditions, infinitely diversified conditions of life,- there would be an almost endless amount of organic action and reaction,- and we should find some groups of beings greatly, and some only slightly modified,- some developed in great force, some existing in scanty numbers- and this we do find in the several great geographical provinces of the world.

On these same principles we can understand, as I have endeavoured to show, why oceanic islands should have few inhabitants, but that of these, a large proportion should be endemic or peculiar; and why, in relation to the means of migration, one group of beings should have all its species peculiar, and another group, even within the same class, should have all its species the same with those in an adjoining quarter of the world. We can see why whole groups of organisms, as batrachians and terrestrial mammals, should be absent from oceanic islands, whilst the most isolated islands should possess their own peculiar species of aerial mammals or bats. We can see why, in islands, there should be some relation between the presence of mammals, in a more or less modified condition, and the depth of the sea between such islands and the mainland. We can clearly see why all the inhabitants of an archipelago, though specifically distinct on the several islets, should be closely related to each other; and should likewise be related, but less closely, to those of the nearest continent, or other source whence immigrants might have been derived. We can see why, if there exists very closely allied or representative species in two areas, however distant from each other, some identical species will almost always there be found.

As the late Edward Forbes often insisted, there is a striking parallelism in the laws of life throughout time and space; the laws governing the succession of forms in past times being nearly the same with those governing at the present time the differences in different areas. We see this in many facts. The endurance of each species and group of species is continuous in time; for the apparent exceptions to the rule are so few, that they may fairly be attributed to our not having as yet discovered in an intermediate deposit certain forms which are absent in it, but which occur both above and below: so in space, it certainly is the general rule that the area inhabited by a single species, or by a group of species, is continuous, and the exceptions, which are not rare, may, as I have attempted to show, be accounted for by former migrations under different circumstances, or through occasional means of transport, or by the species having become extinct in the intermediate tracts. Both in time and space species and groups of species have their points of maximum development. Groups of species, living during the same period of time, or living within the same area, are often characterised by trifling features in common, as of sculpture or colour. In looking to the long succession of past ages, as in looking to distant provinces throughout the world, we find that species in certain classes differ little from each other, whilst those in another class, or only in a different section of the same order, differ greatly from each other. In both time and space the lowly organised members of each class generally change less than the highly organised; but there are in both cases marked exceptions to the rule. According to our theory, these several relations throughout time and space are intelligible; for whether we look to the allied forms of life which have changed during successive ages, or to those which have changed after having migrated into distant quarters, in both cases they are connected by the same bond of ordinary generation; in both cases the laws of variation have been the same, and modifications have been accumulated by the same means of natural selection.