The Origin of Species by Means of Natural Selection

In South America, Dr. Hooker has shown that besides many closely allied species, between forty and fifty of the flowering plants of Tierra del Fuego, forming no inconsiderable part of its scanty flora, are common to North America and Europe, enormously remote as these areas in opposite hemispheres are from each other. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur. On the Organ Mountains of Brazil some few temperate European, some Antarctic and some Andean genera were found by Gardner which do not exist in the low intervening hot countries. On the Silla of Caraccas the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera.

In Africa, several forms characteristic of Europe, and some few representatives of the flora of the Cape of Good Hope, occur on the mountains of Abyssinia. At the Cape of Good Hope a very few European species, believed not to have been introduced by man, and on the mountains several representative European forms are found which have not been discovered in the intertropical parts of Africa. Dr. Hooker has also lately shown that several of the plants living on the upper parts of the lofty island of Fernando Po, and on the neighbouring Cameroon Mountains, in the Gulf of Guinea, are closely related to those on the mountains of Abyssinia, and likewise to those of temperate Europe. It now also appears, as I hear from Dr. Hooker, that some of these same temperate plants have been discovered by the Rev. R.T. Lowe on the mountains of the Cape Verde Islands. This extension of the same temperate forms, almost under the equator, across the whole continent of Africa and to the mountains of the Cape Verde archipelago, is one of the most astonishing facts ever recorded in the distribution of plants.

On the Himalaya, and on the isolated mountain ranges of the peninsula of India, on the heights of Ceylon, and on the volcanic cones of Java, many plants occur either identically the same or representing each other, and at the same time representing plants of Europe not found in the intervening hot lowlands. A list of the genera of plants collected on the loftier peaks of Java, raises a picture of a collection made on a hillock in Europe. Still more striking is the fact that peculiar Australian forms are represented by certain plants growing on the summits of the mountains of Borneo. Some of these Australian forms, as I hear from Dr. Hooker, extend along the heights of the peninsula of Malacca, and are thinly scattered on the one hand over India, and on the other hand as far north as Japan.

On the southern mountains of Australia, Dr. F. Muller has discovered several European species; other species, not introduced by man, occur on the lowlands; and a long list can be given, as I am informed by Dr. Hooker, of European genera, found in Australia, but not in the intermediate torrid regions. In the admirable "Introduction to the Flora of New Zealand," by Dr. Hooker, analogous and striking facts are given in regard to the plants of that large island. Hence, we see that certain plants growing on the more lofty mountains of the tropics in all parts of the world, and on the temperate plains of the north and south, are either the same species or varieties of the same species. It should, however, be observed that these plants are not strictly arctic forms; for, as Mr. H.C. Watson has remarked, "in receding from polar toward equatorial latitudes, the Alpine or mountain flora really become less and less Arctic." Besides these identical and closely allied forms, many species inhabiting the same widely sundered areas, belong to genera not now found in the intermediate tropical lowlands.

These brief remarks apply to plants alone; but some few analogous facts could be given in regard to terrestrial animals. In marine productions, similar cases likewise occur; as an example, I may quote a statement by the highest authority, Prof. Dana, that "it is certainly a wonderful fact that New Zealand should have a closer resemblance in its crustacea to Great Britain, its antipode, than to any other part of the world." Sir J. Richardson, also, speaks of the reappearance on the shores of New Zealand, Tasmania, etc., of northern forms of fish. Dr. Hooker informs me that twenty-five species of Algae are common to New Zealand and to Europe, but have not been found in the intermediate tropical seas.

From the foregoing facts, namely, the presence of temperate forms on the highlands across the whole of equatorial Africa, and along the Peninsula of India, to Ceylon and the Malay Archipelago, and in a less well-marked manner across the wide expanse of tropical South America, it appears almost certain that at some former period, no doubt during the most severe part of a Glacial period, the lowlands of these great continents were everywhere tenanted under the equator by a considerable number of temperate forms. At this period the equatorial climate at the level of the sea was probably about the same with that now experienced at the height of from five to six thousand feet under the same latitude, or perhaps even rather cooler. During this, the coldest period, the lowlands under the equator must have been clothed with a mingled tropical and temperate vegetation, like that described by Hooker as growing luxuriantly at the height of from four to five thousand feet on the lower slopes of the Himalaya, but with perhaps a still greater preponderance of temperate forms. So again in the mountainous island of Fernando Po, in the Gulf of Guinea, Mr. Mann found temperate European forms beginning to appear at the height of about five thousand feet. On the mountains of Panama, at the height of only two thousand feet, Dr. Seemann found the vegetation like that of Mexico, "with forms of the torrid zone harmoniously blended with those of the temperate."

Now let us see whether Mr. Croll's conclusion that when the northern hemisphere suffered from the extreme cold of the great Glacial period, the southern hemisphere was actually warmer, throws any clear light on the present apparently inexplicable distribution of various organisms in the temperate parts of both hemispheres, and on the mountains of the tropics. The Glacial period, as measured by years, must have been very long; and when we remember over what vast spaces some naturalised plants and animals have spread within a few centuries, this period will have been ample for any amount of migration. As the cold became more and more intense, we know that Arctic forms invaded the temperate regions; and from the facts just given, there can hardly be a doubt that some of the more vigorous, dominant and widest-spreading temperate forms invaded the equatorial lowlands. The inhabitants of these hot lowlands would at the same time have migrated to the tropical and subtropical regions of the south, for the southern hemisphere was at this period warmer. On the decline of the Glacial period, as both hemispheres gradually recovered their former temperature, the northern temperate forms living on the lowlands under the equator, would have been driven to their former homes or have been destroyed, being replaced by the equatorial forms returning from the south. Some, however, of the northern temperate forms would almost certainly have ascended any adjoining high land, where, if sufficiently lofty, they would have long survived like the Arctic forms on the mountains of Europe. They might have survived, even if the climate was not perfectly fitted for them, for the change of temperature must have been very slow, and plants undoubtedly possess a certain capacity for acclimatisation, as shown by their transmitting to their offspring different constitutional powers of resisting heat and cold.

In the regular course of events the southern hemisphere would in its turn be subjected to a severe Glacial period, with the northern hemisphere rendered warmer; and then the southern temperate forms would invade the equatorial lowlands. The northern forms which had before been left on the mountains would now descend and mingle with the southern forms. These latter, when the warmth returned, would return to their former homes, leaving some few species on the mountains, and carrying southward with them some of the northern temperate forms which had descended from their mountain fastnesses. Thus, we should have some few species identically the same in the northern and southern temperate zones and on the mountains of the intermediate tropical regions. But the species left during a long time on these mountains, or in opposite hemispheres, would have to compete with many new forms and would be exposed to somewhat different physical conditions; hence, they would be eminently liable to modification, and would generally now exist as varieties or as representative species; and this is the case. We must, also, bear in mind the occurrence in both hemispheres of former Glacial periods; for these will account, in accordance with the same principles, for the many quite distinct species inhabiting the same widely separated areas, and belonging to genera not now found in the intermediate torrid zones.

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 or slightly modified species have migrated from the north to the south, than in a reversed direction. We see, however, a few southern forms on the mountains of Borneo and Abyssinia. I suspect that this preponderant migration from the north to the 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 the two sets became commingled in the equatorial regions, during the alternations of the Glacial periods, the northern forms were the more powerful and were able to hold their places on the mountains, and afterwards migrate southward with the southern forms; but not so the southern in regard to the northern forms. In the same manner, at the present day, we see that very many European productions cover the ground in La Plata, New Zealand, and to a lesser degree in Australia, and have beaten the natives; whereas extremely few southern forms have become naturalised in any part of the northern hemisphere, 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 forty or fifty years from Australia. The Neilgherrie Mountains in India, however, offer a partial exception; for here, as I hear from Dr. Hooker, Australian forms are rapidly sowing themselves and becoming naturalised. Before the last great Glacial period, no doubt the intertropical mountains were stocked with endemic Alpine forms; but these have almost everywhere 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 those which have become naturalised; and this is the first stage towards their extinction. Mountains are islands on the land; and their inhabitants have yielded to those produced within the larger areas of the north, just in the same way as the inhabitants of real islands have everywhere yielded and are still yielding to continental forms naturalised through man's agency.

The same principles apply to the distribution of terrestrial animals and of marine productions, in the northern and southern temperate zones, and on the intertropical mountains. When, during the height of the Glacial period, the ocean-currents were widely different to what they now are, some of the inhabitants of the temperate seas might have reached the equator; of these a few would perhaps at once be able to migrate southwards, by keeping to the cooler currents, while others might remain and survive in the colder depths until the southern hemisphere was in its turn subjected to a glacial climate and permitted their further progress; in nearly the same manner as, according to Forbes, isolated spaces inhabited by Arctic productions exist to the present day in the deeper parts of the northern temperate seas.

I am far from supposing that all the difficulties in regard to the distribution and affinities of the identical and allied species, which now live so widely separated in the north and south, and sometimes on the intermediate mountain ranges, are removed on the views above given. The exact lines of migration cannot be indicated. We cannot say why certain species and not others have migrated; why certain species have been modified and have given rise to new forms, while 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 species ranges twice or thrice as far, and is twice or thrice as common, as another species within their own homes.

Various special difficulties also remain to be solved; for instance, the occurrence, as shown by Dr. Hooker, of the same plants at points so enormously remote as Kerguelen Land, New Zealand, and Fuegia; but icebergs, as suggested by Lyell, may have been concerned in their dispersal. The existence at these and other distant points of the southern hemisphere, of species, which, though distinct, belong to genera exclusively confined to the south, is a more remarkable case. Some of these species are so distinct, that we cannot suppose that there has been time since the commencement of the last Glacial period for their migration and subsequent modification to the necessary degree. The facts seem to indicate that distinct species belonging to the same genera have migrated in radiating lines from a 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 last Glacial period, when the Antarctic lands, now covered with ice, supported a highly peculiar and isolated flora. It may be suspected that before this flora was exterminated during the last Glacial epoch, a few forms had been already widely dispersed to various points of the southern hemisphere by occasional means of transport, and by the aid, as halting-places, of now sunken islands. Thus the southern shores of America, Australia, and New Zealand may have become slightly tinted by the same peculiar forms of life.

Sir C. Lyell in a striking passage has speculated, in language almost identical with mine, on the effects of great alternations of climate throughout the world on geographical distribution. And we have now seen that Mr. Croll's conclusion that successive Glacial periods in the one hemisphere coincide with warmer periods in the opposite hemisphere, together with the admission of the slow modification of species, explains a multitude of facts in the distribution of the same and of the allied forms of life in all parts of the globe. The living waters have flowed during one period from the north and during another from the south, and in both cases have reached the equator; but the stream of life has flowed with greater force from the north than in the opposite direction, and has consequently more freely inundated the south. As the tide leaves its drift in horizontal lines, 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 latitude 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 serves as a record, full of interest to us, of the former inhabitants of the surrounding lowlands.

CHAPTER XIII. 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.

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, etc., 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 differences of the fish on the opposite sides of most mountain-ranges, which are continuous and 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 all 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 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 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 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 dry weighed only 6 and 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 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 the 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 a 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 having 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 Southwestern 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 land-shells 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 twenty-six land birds; of these twenty-one (or perhaps twenty-three) are peculiar; whereas of the eleven marine birds only two 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 land bird; 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 ninety-nine 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 sea-shells 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.