The Origin of Species by Means of Natural Selection

The inhabitants of the world at each successive period in its history have beaten their predecessors in the race for life, and are, in so far, higher in the scale, and their structure has generally become more specialised; and this may account for the common belief held by so many palaeontologists, that organisation on the whole has progressed. Extinct and ancient animals resemble to a certain extent the embryos of the more recent animals belonging to the same classes, and this wonderful fact receives a simple explanation according to our views. The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious, and is intelligible on the principle of inheritance.

If, then, the geological record be as imperfect as many believe, and it may at least be asserted that the record cannot be proved to be much more perfect, the main objections to the theory of natural selection are greatly diminished or disappear. On the other hand, all the chief laws of palaeontology plainly proclaim, as it seems to me, that species have been produced by ordinary generation: old forms having been supplanted by new and improved forms of life, the products of variation and the survival of the fittest.

CHAPTER XII. GEOGRAPHICAL DISTRIBUTION

Present distribution cannot be accounted for by differences in physical conditions – Importance of barriers – Affinity of the productions of the same continent – Centres of creation – Means of dispersal by changes of climate and of the level of the land, and by occasional means – Dispersal during the Glacial period – Alternate Glacial periods in the North and South.

In considering the distribution of organic beings over the face of the globe, the first great fact which strikes us is, that neither the similarity nor the dissimilarity of the inhabitants of various regions can be wholly accounted for by climatal and other physical conditions. Of late, almost every author who has studied the subject has come to this conclusion. The case of America alone would almost suffice to prove its truth; for if we exclude the arctic and northern temperate parts, all authors agree that one of the most fundamental divisions in geographical distribution is that between the New and Old Worlds; yet if we travel over the vast American continent, from the central parts of the United States to its extreme southern point, we meet with the most diversified conditions; humid districts, arid deserts, lofty mountains, grassy plains, forests, marshes, lakes and great rivers, under almost every temperature. There is hardly a climate or condition in the Old World which cannot be paralleled in the New – at least so closely as the same species generally require. No doubt small areas can be pointed out in the Old World hotter than any in the New World; but these are not inhabited by a fauna different from that of the surrounding districts; for it is rare to find a group of organisms confined to a small area, of which the conditions are peculiar in only a slight degree. Notwithstanding this general parallelism in the conditions of Old and New Worlds, how widely different are their living productions!

In the southern hemisphere, if we compare large tracts of land in Australia, South Africa, and western South America, between latitudes 25 and 35 degrees, we shall find parts extremely similar in all their conditions, yet it would not be possible to point out three faunas and floras more utterly dissimilar. Or, again, we may compare the productions of South America south of latitude 35 degrees with those north of 25 degrees, which consequently are separated by a space of ten degrees of latitude, and are exposed to considerably different conditions; yet they are incomparably more closely related to each other than they are to the productions of Australia or Africa under nearly the same climate. Analogous facts could be given with respect to the inhabitants of the sea.

A second great fact which strikes us in our general review is, that barriers of any kind, or obstacles to free migration, are related in a close and important manner to the differences between the productions of various regions. We see this in the great difference in nearly all the terrestrial productions of the New and Old Worlds, excepting in the northern parts, where the land almost joins, and where, under a slightly different climate, there might have been free migration for the northern temperate forms, as there now is for the strictly arctic productions. We see the same fact in the great difference between the inhabitants of Australia, Africa, and South America under the same latitude; for these countries are almost as much isolated from each other as is possible. On each continent, also, we see the same fact; for on the opposite sides of lofty and continuous mountain-ranges, and of great deserts and even of large rivers, we find different productions; though as mountain chains, deserts, etc., are not as impassable, or likely to have endured so long, as the oceans separating continents, the differences are very inferior in degree to those characteristic of distinct continents.

Turning to the sea, we find the same law. The marine inhabitants of the eastern and western shores of South America are very distinct, with extremely few shells, crustacea, or echinodermata in common; but Dr. Gunther has recently shown that about thirty per cent of the fishes are the same on the opposite sides of the isthmus of Panama; and this fact has led naturalists to believe that the isthmus was formerly open. Westward of the shores of America, a wide space of open ocean extends, with not an island as a halting-place for emigrants; here we have a barrier of another kind, and as soon as this is passed we meet in the eastern islands of the Pacific with another and totally distinct fauna. So that three marine faunas range northward and southward in parallel lines not far from each other, under corresponding climate; but from being separated from each other by impassable barriers, either of land or open sea, they are almost wholly distinct. On the other hand, proceeding still further westward from the eastern islands of the tropical parts of the Pacific, we encounter no impassable barriers, and we have innumerable islands as halting-places, or continuous coasts, until, after travelling over a hemisphere, we come to the shores of Africa; and over this vast space we meet with no well-defined and distinct marine faunas. Although so few marine animals are common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands, yet many fishes range from the Pacific into the Indian Ocean, and many shells are common to the eastern islands of the Pacific and the eastern shores of Africa on almost exactly opposite meridians of longitude.

A third great fact, partly included in the foregoing statement, is the affinity of the productions of the same continent or of the same sea, though the species themselves are distinct at different points and stations. It is a law of the widest generality, and every continent offers innumerable instances. Nevertheless, the naturalist, in travelling, for instance, from north to south, never fails to be struck by the manner in which successive groups of beings, specifically distinct, though nearly related, replace each other. He hears from closely allied, yet distinct kinds of birds, notes nearly similar, and sees their nests similarly constructed, but not quite alike, with eggs coloured in nearly the same manner. The plains near the Straits of Magellan are inhabited by one species of Rhea (American ostrich), and northward the plains of La Plata by another species of the same genus; and not by a true ostrich or emu, like those inhabiting Africa and Australia under the same latitude. On these same plains of La Plata we see the agouti and bizcacha, animals having nearly the same habits as our hares and rabbits, and belonging to the same order of Rodents, but they plainly display an American type of structure. We ascend the lofty peaks of the Cordillera, and we find an alpine species of bizcacha; we look to the waters, and we do not find the beaver or muskrat, but the coypu and capybara, rodents of the South American type. Innumerable other instances could be given. If we look to the islands off the American shore, however much they may differ in geological structure, the inhabitants are essentially American, though they may be all peculiar species. We may look back to past ages, as shown in the last chapter, and we find American types then prevailing on the American continent and in the American seas. We see in these facts some deep organic bond, throughout space and time, over the same areas of land and water, independently of physical conditions. The naturalist must be dull who is not led to inquire what this bond is.

The bond is simply inheritance, that cause which alone, as far as we positively know, produces organisms quite like each other, or, as we see in the case of varieties, nearly alike. The dissimilarity of the inhabitants of different regions may be attributed to modification through variation and natural selection, and probably in a subordinate degree to the definite influence of different physical conditions. The degrees of dissimilarity will depend on the migration of the more dominant forms of life from one region into another having been more or less effectually prevented, at periods more or less remote – on the nature and number of the former immigrants – and on the action of the inhabitants on each other in leading to the preservation of different modifications; the relation of organism to organism in the struggle for life being, as I have already often remarked, the most important of all relations. Thus the high importance of barriers comes into play by checking migration; as does time for the slow process of modification through natural selection. Widely-ranging species, abounding in individuals, which have already triumphed over many competitors in their own widely-extended homes, will have the best chance of seizing on new places, when they spread out into new countries. In their new homes they will be exposed to new conditions, and will frequently undergo further modification and improvement; and thus they will become still further victorious, and will produce groups of modified descendants. On this principle of inheritance with modification we can understand how it is that sections of genera, whole genera, and even families, are confined to the same areas, as is so commonly and notoriously the case.

There is no evidence, as was remarked in the last chapter, of the existence of any law of necessary development. As the variability of each species is an independent property, and will be taken advantage of by natural selection, only so far as it profits each individual in its complex struggle for life, so the amount of modification in different species will be no uniform quantity. If a number of species, after having long competed with each other in their old home, were to migrate in a body into a new and afterwards isolated country, they would be little liable to modification; for neither migration nor isolation in themselves effect anything. These principles come into play only by bringing organisms into new relations with each other and in a lesser degree with the surrounding physical conditions. As we have seen in the last chapter that some forms have retained nearly the same character from an enormously remote geological period, so certain species have migrated over vast spaces, and have not become greatly or at all modified.

According to these views, it is obvious that the several species of the same genus, though inhabiting the most distant quarters of the world, must originally have proceeded from the same source, as they are descended from the same progenitor. In the case of those species which have undergone, during whole geological periods, little modification, there is not much difficulty in believing that they have migrated from the same region; for during the vast geographical and climatical changes which have supervened since ancient times, almost any amount of migration is possible. But in many other cases, in which we have reason to believe that the species of a genus have been produced within comparatively recent times, there is great difficulty on this head. It is also obvious that the individuals of the same species, though now inhabiting distant and isolated regions, must have proceeded from one spot, where their parents were first produced: for, as has been explained, it is incredible that individuals identically the same should have been produced from parents specifically distinct.

SINGLE CENTRES OF SUPPOSED CREATION.

We are thus brought to the question which has been largely discussed by naturalists, namely, whether species have been created at one or more points of the earth's surface. Undoubtedly there are many cases of extreme difficulty in understanding how the same species could possibly have migrated from some one point to the several distant and isolated points, where now found. Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. He who rejects it, rejects the vera causa of ordinary generation with subsequent migration, and calls in the agency of a miracle. It is universally admitted, that in most cases the area inhabited by a species is continuous; and that when a plant or animal inhabits two points so distant from each other, or with an interval of such a nature, that the space could not have been easily passed over by migration, the fact is given as something remarkable and exceptional. The incapacity of migrating across a wide sea is more clear in the case of terrestrial mammals than perhaps with any other organic beings; and, accordingly, we find no inexplicable instances of the same mammals inhabiting distant points of the world. No geologist feels any difficulty in Great Britain possessing the same quadrupeds with the rest of Europe, for they were no doubt once united. But if the same species can be produced at two separate points, why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same, so that a multitude of European animals and plants have become naturalised in America and Australia; and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres? The answer, as I believe, is, that mammals have not been able to migrate, whereas some plants, from their varied means of dispersal, have migrated across the wide and broken interspaces. The great and striking influence of barriers of all kinds, is intelligible only on the view that the great majority of species have been produced on one side, and have not been able to migrate to the opposite side. Some few families, many subfamilies, very many genera, a still greater number of sections of genera, are confined to a single region; and it has been observed by several naturalists that the most natural genera, or those genera in which the species are most closely related to each other, are generally confined to the same country, or if they have a wide range that their range is continuous. What a strange anomaly it would be if a directly opposite rule were to prevail when we go down one step lower in the series, namely to the individuals of the same species, and these had not been, at least at first, confined to some one region!

Hence, it seems to me, as it has to many other naturalists, that the view of each species having been produced in one area alone, and having subsequently migrated from that area as far as its powers of migration and subsistence under past and present conditions permitted, is the most probable. Undoubtedly many cases occur in which we cannot explain how the same species could have passed from one point to the other. But the geographical and climatical changes which have certainly occurred within recent geological times, must have rendered discontinuous the formerly continuous range of many species. So that we are reduced to consider whether the exceptions to continuity of range are so numerous, and of so grave a nature, that we ought to give up the belief, rendered probable by general considerations, that each species has been produced within one area, and has migrated thence as far as it could. It would be hopelessly tedious to discuss all the exceptional cases of the same species, now living at distant and separated points; nor do I for a moment pretend that any explanation could be offered of many instances. But, after some preliminary remarks, I will discuss a few of the most striking classes of facts, namely, the existence of the same species on the summits of distant mountain ranges, and at distant points in the Arctic and Antarctic regions; and secondly (in the following chapter), the wide distribution of fresh water productions; and thirdly, the occurrence of the same terrestrial species on islands and on the nearest mainland, though separated by hundreds of miles of open sea. If the existence of the same species at distant and isolated points of the earth's surface can in many instances be explained on the view of each species having migrated from a single birthplace; then, considering our ignorance with respect to former climatical and geographical changes, and to the various occasional means of transport, the belief that a single birthplace is the law seems to me incomparably the safest.

In discussing this subject we shall be enabled at the same time to consider a point equally important for us, namely, whether the several species of a genus which must on our theory all be descended from a common progenitor, can have migrated, undergoing modification during their migration from some one area. If, when most of the species inhabiting one region are different from those of another region, though closely allied to them, it can be shown that migration from the one region to the other has probably occurred at some former period, our general view will be much strengthened; for the explanation is obvious on the principle of descent with modification. A volcanic island, for instance, upheaved and formed at the distance of a few hundreds of miles from a continent, would probably receive from it in the course of time a few colonists, and their descendants, though modified, would still be related by inheritance to the inhabitants of that continent. Cases of this nature are common, and are, as we shall hereafter see, inexplicable on the theory of independent creation. This view of the relation of the species of one region to those of another, does not differ much from that advanced by Mr. Wallace, who concludes that "every species has come into existence coincident both in space and time with a pre-existing closely allied species." And it is now well known that he attributes this coincidence to descent with modification.

The question of single or multiple centres of creation differs from another though allied question, namely, whether all the individuals of the same species are descended from a single pair, or single hermaphrodite, or whether, as some authors suppose, from many individuals simultaneously created. With organic beings which never intercross, if such exist, each species, must be descended from a succession of modified varieties, that have supplanted each other, but have never blended with other individuals or varieties of the same species, so that, at each successive stage of modification, all the individuals of the same form will be descended from a single parent. But in the great majority of cases, namely, with all organisms which habitually unite for each birth, or which occasionally intercross, the individuals of the same species inhabiting the same area will be kept nearly uniform by intercrossing; so that many individuals will go on simultaneously changing, and the whole amount of modification at each stage will not be due to descent from a single parent. To illustrate what I mean: our English race-horses differ from the horses of every other breed; but they do not owe their difference and superiority to descent from any single pair, but to continued care in the selecting and training of many individuals during each generation.

Before discussing the three classes of facts, which I have selected as presenting the greatest amount of difficulty on the theory of "single centres of creation," I must say a few words on the means of dispersal.

MEANS OF DISPERSAL.

Sir C. Lyell and other authors have ably treated this subject. I can give here only the briefest abstract of the more important facts. Change of climate must have had a powerful influence on migration. A region now impassable to certain organisms from the nature of its climate, might have been a high road for migration, when the climate was different. I shall, however, presently have to discuss this branch of the subject in some detail. Changes of level in the land must also have been highly influential: a narrow isthmus now separates two marine faunas; submerge it, or let it formerly have been submerged, and the two faunas will now blend together, or may formerly have blended. Where the sea now extends, land may at a former period have connected islands or possibly even continents together, and thus have allowed terrestrial productions to pass from one to the other. No geologist disputes that great mutations of level have occurred within the period of existing organisms. Edward Forbes insisted that all the islands in the Atlantic must have been recently connected with Europe or Africa, and Europe likewise with America. Other authors have thus hypothetically bridged over every ocean, and united almost every island with some mainland. If, indeed, the arguments used by Forbes are to be trusted, it must be admitted that scarcely a single island exists which has not recently been united to some continent. This view cuts the Gordian knot of the dispersal of the same species to the most distant points, and removes many a difficulty; but to the best of my judgment we are not authorized in admitting such enormous geographical changes within the period of existing species. It seems to me that we have abundant evidence of great oscillations in the level of the land or sea; but not of such vast changes in the position and extension of our continents, as to have united them within the recent period to each other and to the several intervening oceanic islands. I freely admit the former existence of many islands, now buried beneath the sea, which may have served as halting places for plants and for many animals during their migration. In the coral-producing oceans such sunken islands are now marked by rings of coral or atolls standing over them. Whenever it is fully admitted, as it will some day be, that each species has proceeded from a single birthplace, and when in the course of time we know something definite about the means of distribution, we shall be enabled to speculate with security on the former extension of the land. But I do not believe that it will ever be proved that within the recent period most of our continents which now stand quite separate, have been continuously, or almost continuously united with each other, and with the many existing oceanic islands. Several facts in distribution – such as the great difference in the marine faunas on the opposite sides of almost every continent – the close relation of the tertiary inhabitants of several lands and even seas to their present inhabitants – the degree of affinity between the mammals inhabiting islands with those of the nearest continent, being in part determined (as we shall hereafter see) by the depth of the intervening ocean – these and other such facts are opposed to the admission of such prodigious geographical revolutions within the recent period, as are necessary on the view advanced by Forbes and admitted by his followers. The nature and relative proportions of the inhabitants of oceanic islands are likewise opposed to the belief of their former continuity of continents. Nor does the almost universally volcanic composition of such islands favour the admission that they are the wrecks of sunken continents; if they had originally existed as continental mountain ranges, some at least of the islands would have been formed, like other mountain summits, of granite, metamorphic schists, old fossiliferous and other rocks, instead of consisting of mere piles of volcanic matter.

I must now say a few words on what are called accidental means, but which more properly should be called occasional means of distribution. I shall here confine myself to plants. In botanical works, this or that plant is often stated to be ill adapted for wide dissemination; but the greater or less facilities for transport across the sea may be said to be almost wholly unknown. Until I tried, with Mr. Berkeley's aid, a few experiments, it was not even known how far seeds could resist the injurious action of sea-water. To my surprise I found that out of eighty-seven kinds, sixty-four germinated after an immersion of twenty-eight days, and a few survived an immersion of 137 days. It deserves notice that certain orders were far more injured than others: nine Leguminosae were tried, and, with one exception, they resisted the salt-water badly; seven species of the allied orders, Hydrophyllaceae and Polemoniaceae, were all killed by a month's immersion. For convenience sake I chiefly tried small seeds without the capsules or fruit; and as all of these sank in a few days, they could not have been floated across wide spaces of the sea, whether or not they were injured by salt water. Afterwards I tried some larger fruits, capsules, etc., and some of these floated for a long time. It is well known what a difference there is in the buoyancy of green and seasoned timber; and it occurred to me that floods would often wash into the sea dried plants or branches with seed-capsules or fruit attached to them. Hence I was led to dry the stems and branches of ninety-four plants with ripe fruit, and to place them on sea-water. The majority sank quickly, but some which, whilst green, floated for a very short time, when dried floated much longer; for instance, ripe hazel-nuts sank immediately, but when dried they floated for ninety days, and afterwards when planted germinated; an asparagus plant with ripe berries floated for twenty-three days, when dried it floated for eighty-five days, and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days, when dried they floated for above ninety days, and afterwards germinated. Altogether, out of the ninety-four dried plants, eighteen floated for above twenty-eight days; and some of the eighteen floated for a very much longer period. So that as 64/87 kinds of seeds germinated after an immersion of twenty-eight days; and as 18/94 distinct species with ripe fruit (but not all the same species as in the foregoing experiment) floated, after being dried, for above twenty-eight days, we may conclude, as far as anything can be inferred from these scanty facts, that the seeds of 14/100 kinds of plants of any country might be floated by sea-currents during twenty-eight days, and would retain their power of germination. In Johnston's Physical Atlas, the average rate of the several Atlantic currents is thirty-three miles per diem (some currents running at the rate of sixty miles per diem); on this average, the seeds of 14/100 plants belonging to one country might be floated across 924 miles of sea to another country; and when stranded, if blown by an inland gale to a favourable spot, would germinate.

Subsequently to my experiments, M. Martens tried similar ones, but in a much better manner, for he placed the seeds in a box in the actual sea, so that they were alternately wet and exposed to the air like really floating plants. He tried ninety-eight seeds, mostly different from mine, but he chose many large fruits, and likewise seeds, from plants which live near the sea; and this would have favoured both the average length of their flotation and their resistance to the injurious action of the salt-water. On the other hand, he did not previously dry the plants or branches with the fruit; and this, as we have seen, would have caused some of them to have floated much longer. The result was that 18/98 of his seeds of different kinds floated for forty-two days, and were then capable of germination. But I do not doubt that plants exposed to the waves would float for a less time than those protected from violent movement as in our experiments. Therefore, it would perhaps be safer to assume that the seeds of about 10/100 plants of a flora, after having been dried, could be floated across a space of sea 900 miles in width, and would then germinate. The fact of the larger fruits often floating longer than the small, is interesting; as plants with large seeds or fruit which, as Alph. de Candolle has shown, generally have restricted ranges, could hardly be transported by any other means.

Seeds may be occasionally transported in another manner. Drift timber is thrown up on most islands, even on those in the midst of the widest oceans; and the natives of the coral islands in the Pacific procure stones for their tools, solely from the roots of drifted trees, these stones being a valuable royal tax. I find that when irregularly shaped stones are embedded in the roots of trees, small parcels of earth are very frequently enclosed in their interstices and behind them, so perfectly that not a particle could be washed away during the longest transport: out of one small portion of earth thus COMPLETELY enclosed by the roots of an oak about fifty years old, three dicotyledonous plants germinated: I am certain of the accuracy of this observation. Again, I can show that the carcasses of birds, when floating on the sea, sometimes escape being immediately devoured; and many kinds of seeds in the crops of floating birds long retain their vitality: peas and vetches, for instance, are killed by even a few days' immersion in sea-water; but some taken out of the crop of a pigeon, which had floated on artificial sea-water for thirty days, to my surprise nearly all germinated.

Living birds can hardly fail to be highly effective agents in the transportation of seeds. I could give many facts showing how frequently birds of many kinds are blown by gales to vast distances across the ocean. We may safely assume that under such circumstances their rate of flight would often be thirty-five miles an hour; and some authors have given a far higher estimate. I have never seen an instance of nutritious seeds passing through the intestines of a bird; but hard seeds of fruit pass uninjured through even the digestive organs of a turkey. In the course of two months, I picked up in my garden twelve kinds of seeds, out of the excrement of small birds, and these seemed perfect, and some of them, which were tried, germinated. But the following fact is more important: the crops of birds do not secrete gastric juice, and do not, as I know by trial, injure in the least the germination of seeds; now, after a bird has found and devoured a large supply of food, it is positively asserted that all the grains do not pass into the gizzard for twelve or even eighteen hours. A bird in this interval might easily be blown to the distance of five hundred miles, and hawks are known to look out for tired birds, and the contents of their torn crops might thus readily get scattered. Some hawks and owls bolt their prey whole, and after an interval of from twelve to twenty hours, disgorge pellets, which, as I know from experiments made in the Zoological Gardens, include seeds capable of germination. Some seeds of the oat, wheat, millet, canary, hemp, clover, and beet germinated after having been from twelve to twenty-one hours in the stomachs of different birds of prey; and two seeds of beet grew after having been thus retained for two days and fourteen hours. Fresh-water fish, I find, eat seeds of many land and water plants; fish are frequently devoured by birds, and thus the seeds might be transported from place to place. I forced many kinds of seeds into the stomachs of dead fish, and then gave their bodies to fishing-eagles, storks, and pelicans; these birds, after an interval of many hours, either rejected the seeds in pellets or passed them in their excrement; and several of these seeds retained the power of germination. Certain seeds, however, were always killed by this process.

Locusts are sometimes blown to great distances from the land. I myself caught one 370 miles from the coast of Africa, and have heard of others caught at greater distances. The Rev. R.T. Lowe informed Sir C. Lyell that in November, 1844, swarms of locusts visited the island of Madeira. They were in countless numbers, as thick as the flakes of snow in the heaviest snowstorm, and extended upward as far as could be seen with a telescope. During two or three days they slowly careered round and round in an immense ellipse, at least five or six miles in diameter, and at night alighted on the taller trees, which were completely coated with them. They then disappeared over the sea, as suddenly as they had appeared, and have not since visited the island. Now, in parts of Natal it is believed by some farmers, though on insufficient evidence, that injurious seeds are introduced into their grass-land in the dung left by the great flights of locusts which often visit that country. In consequence of this belief Mr. Weale sent me in a letter a small packet of the dried pellets, out of which I extracted under the microscope several seeds, and raised from them seven grass plants, belonging to two species, of two genera. Hence a swarm of locusts, such as that which visited Madeira, might readily be the means of introducing several kinds of plants into an island lying far from the mainland.

Although the beaks and feet of birds are generally clean, earth sometimes adheres to them: in one case I removed sixty-one grains, and in another case twenty-two grains of dry argillaceous earth from the foot of a partridge, and in the earth there was a pebble as large as the seed of a vetch. Here is a better case: the leg of a woodcock was sent to me by a friend, with a little cake of dry earth attached to the shank, weighing only nine grains; and this contained a seed of the toad-rush (Juncus bufonius) which germinated and flowered. Mr. Swaysland, of Brighton, who during the last forty years has paid close attention to our migratory birds, informs me that he has often shot wagtails (Motacillae), wheatears, and whinchats (Saxicolae), on their first arrival on our shores, before they had alighted; and he has several times noticed little cakes of earth attached to their feet. Many facts could be given showing how generally soil is charged with seeds. For instance, Professor Newton sent me the leg of a red-legged partridge (Caccabis rufa) which had been wounded and could not fly, with a ball of hard earth adhering to it, and weighing six and a half ounces. The earth had been kept for three years, but when broken, watered and placed under a bell glass, no less than eighty-two plants sprung from it: these consisted of twelve monocotyledons, including the common oat, and at least one kind of grass, and of seventy dicotyledons, which consisted, judging from the young leaves, of at least three distinct species. With such facts before us, can we doubt that the many birds which are annually blown by gales across great spaces of ocean, and which annually migrate – for instance, the millions of quails across the Mediterranean – must occasionally transport a few seeds embedded in dirt adhering to their feet or beaks? But I shall have to recur to this subject.

As icebergs are known to be sometimes loaded with earth and stones, and have even carried brushwood, bones, and the nest of a land-bird, it can hardly be doubted that they must occasionally, as suggested by Lyell, have transported seeds from one part to another of the arctic and antarctic regions; and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of plants common to Europe, in comparison with the species on the other islands of the Atlantic, which stand nearer to the mainland, and (as remarked by Mr. H.C. Watson) from their somewhat northern character, in comparison with the latitude, I suspected that these islands had been partly stocked by ice-borne seeds during the Glacial epoch. At my request Sir C. Lyell wrote to M. Hartung to inquire whether he had observed erratic boulders on these islands, and he answered that he had found large fragments of granite and other rocks, which do not occur in the archipelago. Hence we may safely infer that icebergs formerly landed their rocky burdens on the shores of these mid-ocean islands, and it is at least possible that they may have brought thither the seeds of northern plants.

Considering that these several means of transport, and that other means, which without doubt remain to be discovered, have been in action year after year for tens of thousands of years, it would, I think, be a marvellous fact if many plants had not thus become widely transported. These means of transport are sometimes called accidental, but this is not strictly correct: the currents of the sea are not accidental, nor is the direction of prevalent gales of wind. It should be observed that scarcely any means of transport would carry seeds for very great distances; for seeds do not retain their vitality when exposed for a great length of time to the action of sea water; nor could they be long carried in the crops or intestines of birds. These means, however, would suffice for occasional transport across tracts of sea some hundred miles in breadth, or from island to island, or from a continent to a neighbouring island, but not from one distant continent to another. The floras of distant continents would not by such means become mingled; but would remain as distinct as they now are. The currents, from their course, would never bring seeds from North America to Britain, though they might and do bring seeds from the West Indies to our western shores, where, if not killed by their very long immersion in salt water, they could not endure our climate. Almost every year, one or two land-birds are blown across the whole Atlantic Ocean, from North America to the western shores of Ireland and England; but seeds could be transported by these rare wanderers only by one means, namely, by dirt adhering to their feet or beaks, which is in itself a rare accident. Even in this case, how small would be the chance of a seed falling on favourable soil, and coming to maturity! But it would be a great error to argue that because a well-stocked island, like Great Britain, has not, as far as is known (and it would be very difficult to prove this), received within the last few centuries, through occasional means of transport, immigrants from Europe or any other continent, that a poorly-stocked island, though standing more remote from the mainland, would not receive colonists by similar means. Out of a hundred kinds of seeds or animals transported to an island, even if far less well-stocked than Britain, perhaps not more than one would be so well fitted to its new home, as to become naturalised. But this is no valid argument against what would be effected by occasional means of transport, during the long lapse of geological time, whilst the island was being upheaved, and before it had become fully stocked with inhabitants. On almost bare land, with few or no destructive insects or birds living there, nearly every seed which chanced to arrive, if fitted for the climate, would germinate and survive.

DISPERSAL DURING THE GLACIAL PERIOD.

The identity of many plants and animals, on mountain-summits, separated from each other by hundreds of miles of lowlands, where Alpine species could not possibly exist, is one of the most striking cases known of the same species living at distant points, without the apparent possibility of their having migrated from one point to the other. It is indeed a remarkable fact to see so many plants of the same species living on the snowy regions of the Alps or Pyrenees, and in the extreme northern parts of Europe; but it is far more remarkable, that the plants on the White Mountains, in the United States of America, are all the same with those of Labrador, and nearly all the same, as we hear from Asa Gray, with those on the loftiest mountains of Europe. Even as long ago as 1747, such facts led Gmelin to conclude that the same species must have been independently created at many distinct points; and we might have remained in this same belief, had not Agassiz and others called vivid attention to the Glacial period, which, as we shall immediately see, affords a simple explanation of these facts. We have evidence of almost every conceivable kind, organic and inorganic, that, within a very recent geological period, central Europe and North America suffered under an Arctic climate. The ruins of a house burnt by fire do not tell their tale more plainly than do the mountains of Scotland and Wales, with their scored flanks, polished surfaces, and perched boulders, of the icy streams with which their valleys were lately filled. So greatly has the climate of Europe changed, that in Northern Italy, gigantic moraines, left by old glaciers, are now clothed by the vine and maize. Throughout a large part of the United States, erratic boulders and scored rocks plainly reveal a former cold period.

The former influence of the glacial climate on the distribution of the inhabitants of Europe, as explained by Edward Forbes, is substantially as follows. But we shall follow the changes more readily, by supposing a new glacial period slowly to come on, and then pass away, as formerly occurred. As the cold came on, and as each more southern zone became fitted for the inhabitants of the north, these would take the places of the former inhabitants of the temperate regions. The latter, at the same time would travel further and further southward, unless they were stopped by barriers, in which case they would perish. The mountains would become covered with snow and ice, and their former Alpine inhabitants would descend to the plains. By the time that the cold had reached its maximum, we should have an arctic fauna and flora, covering the central parts of Europe, as far south as the Alps and Pyrenees, and even stretching into Spain. The now temperate regions of the United States would likewise be covered by arctic plants and animals and these would be nearly the same with those of Europe; for the present circumpolar inhabitants, which we suppose to have everywhere travelled southward, are remarkably uniform round the world.

As the warmth returned, the arctic forms would retreat northward, closely followed up in their retreat by the productions of the more temperate regions. And as the snow melted from the bases of the mountains, the arctic forms would seize on the cleared and thawed ground, always ascending, as the warmth increased and the snow still further disappeared, higher and higher, whilst their brethren were pursuing their northern journey. Hence, when the warmth had fully returned, the same species, which had lately lived together on the European and North American lowlands, would again be found in the arctic regions of the Old and New Worlds, and on many isolated mountain-summits far distant from each other.

Thus we can understand the identity of many plants at points so immensely remote as the mountains of the United States and those of Europe. We can thus also understand the fact that the Alpine plants of each mountain-range are more especially related to the arctic forms living due north or nearly due north of them: for the first migration when the cold came on, and the re-migration on the returning warmth, would generally have been due south and north. The Alpine plants, for example, of Scotland, as remarked by Mr. H.C. Watson, and those of the Pyrenees, as remarked by Ramond, are more especially allied to the plants of northern Scandinavia; those of the United States to Labrador; those of the mountains of Siberia to the arctic regions of that country. These views, grounded as they are on the perfectly well-ascertained occurrence of a former Glacial period, seem to me to explain in so satisfactory a manner the present distribution of the Alpine and Arctic productions of Europe and America, that when in other regions we find the same species on distant mountain-summits, we may almost conclude, without other evidence, that a colder climate formerly permitted their migration across the intervening lowlands, now become too warm for their existence.

As the arctic forms moved first southward and afterwards backward to the north, in unison with the changing climate, they will not have been exposed during their long migrations to any great diversity of temperature; and as they all migrated in a body together, their mutual relations will not have been much disturbed. Hence, in accordance with the principles inculcated in this volume, these forms will not have been liable to much modification. But with the Alpine productions, left isolated from the moment of the returning warmth, first at the bases and ultimately on the summits of the mountains, the case will have been somewhat different; for it is not likely that all the same arctic species will have been left on mountain ranges far distant from each other, and have survived there ever since; they will also, in all probability, have become mingled with ancient Alpine species, which must have existed on the mountains before the commencement of the Glacial epoch, and which during the coldest period will have been temporarily driven down to the plains; they will, also, have been subsequently exposed to somewhat different climatical influences. Their mutual relations will thus have been in some degree disturbed; consequently they will have been liable to modification; and they have been modified; for if we compare the present Alpine plants and animals of the several great European mountain ranges, one with another, though many of the species remain identically the same, some exist as varieties, some as doubtful forms or sub-species and some as distinct yet closely allied species representing each other on the several ranges.

In the foregoing illustration, I have assumed that at the commencement of our imaginary Glacial period, the arctic productions were as uniform round the polar regions as they are at the present day. But it is also necessary to assume that many sub-arctic and some few temperate forms were the same round the world, for some of the species which now exist on the lower mountain slopes and on the plains of North America and Europe are the same; and it may be asked how I account for this degree of uniformity of the sub-arctic and temperate forms round the world, at the commencement of the real Glacial period. At the present day, the sub-arctic and northern temperate productions of the Old and New Worlds are separated from each other by the whole Atlantic Ocean and by the northern part of the Pacific. During the Glacial period, when the inhabitants of the Old and New Worlds lived further southwards than they do at present, they must have been still more completely separated from each other by wider spaces of ocean; so that it may well be asked how the same species could then or previously have entered the two continents. The explanation, I believe, lies in the nature of the climate before the commencement of the Glacial period. At this, the newer Pliocene period, the majority of the inhabitants of the world were specifically the same as now, and we have good reason to believe that the climate was warmer than at the present day. Hence, we may suppose that the organisms which now live under latitude 60 degrees, lived during the Pliocene period further north, under the Polar Circle, in latitude 66-67 degrees; and that the present arctic productions then lived on the broken land still nearer to the pole. Now, if we look at a terrestrial globe, we see under the Polar Circle that there is almost continuous land from western Europe through Siberia, to eastern America. And this continuity of the circumpolar land, with the consequent freedom under a more favourable climate for intermigration, will account for the supposed uniformity of the sub-arctic and temperate productions of the Old and New Worlds, at a period anterior to the Glacial epoch.

Believing, from reasons before alluded to, that our continents have long remained in nearly the same relative position, though subjected to great oscillations of level, I am strongly inclined to extend the above view, and to infer that during some earlier and still warmer period, such as the older Pliocene period, a large number of the same plants and animals inhabited the almost continuous circumpolar land; and that these plants and animals, both in the Old and New Worlds, began slowly to migrate southwards as the climate became less warm, long before the commencement of the Glacial period. We now see, as I believe, their descendants, mostly in a modified condition, in the central parts of Europe and the United States. On this view we can understand the relationship with very little identity, between the productions of North America and Europe – a relationship which is highly remarkable, considering the distance of the two areas, and their separation by the whole Atlantic Ocean. We can further understand the singular fact remarked on by several observers that the productions of Europe and America during the later tertiary stages were more closely related to each other than they are at the present time; for during these warmer periods the northern parts of the Old and New Worlds will have been almost continuously united by land, serving as a bridge, since rendered impassable by cold, for the intermigration of their inhabitants.

During the slowly decreasing warmth of the Pliocene period, as soon as the species in common, which inhabited the New and Old Worlds, migrated south of the Polar Circle, they will have been completely cut off from each other. This separation, as far as the more temperate productions are concerned, must have taken place long ages ago. As the plants and animals migrated southward, they will have become mingled in the one great region with the native American productions, and would have had to compete with them; and in the other great region, with those of the Old World. Consequently we have here everything favourable for much modification – for far more modification than with the Alpine productions, left isolated, within a much more recent period, on the several mountain ranges and on the arctic lands of Europe and North America. Hence, it has come, that when we compare the now living productions of the temperate regions of the New and Old Worlds, we find very few identical species (though Asa Gray has lately shown that more plants are identical than was formerly supposed), but we find in every great class many forms, which some naturalists rank as geographical races, and others as distinct species; and a host of closely allied or representative forms which are ranked by all naturalists as specifically distinct.

As on the land, so in the waters of the sea, a slow southern migration of a marine fauna, which, during the Pliocene or even a somewhat earlier period, was nearly uniform along the continuous shores of the Polar Circle, will account, on the theory of modification, for many closely allied forms now living in marine areas completely sundered. Thus, I think, we can understand the presence of some closely allied, still existing and extinct tertiary forms, on the eastern and western shores of temperate North America; and the still more striking fact of many closely allied crustaceans (as described in Dana's admirable work), some fish and other marine animals, inhabiting the Mediterranean and the seas of Japan – these two areas being now completely separated by the breadth of a whole continent and by wide spaces of ocean.

These cases of close relationship in species either now or formerly inhabiting the seas on the eastern and western shores of North America, the Mediterranean and Japan, and the temperate lands of North America and Europe, are inexplicable on the theory of creation. We cannot maintain that such species have been created alike, in correspondence with the nearly similar physical conditions of the areas; for if we compare, for instance, certain parts of South America with parts of South Africa or Australia, we see countries closely similar in all their physical conditions, with their inhabitants utterly dissimilar.

ALTERNATE GLACIAL PERIODS IN THE NORTH AND SOUTH.

But we must return to our more immediate subject. I am convinced that Forbes's view may be largely extended. In Europe we meet with the plainest evidence of the Glacial period, from the western shores of Britain to the Ural range, and southward to the Pyrenees. We may infer from the frozen mammals and nature of the mountain vegetation, that Siberia was similarly affected. In the Lebanon, according to Dr. Hooker, perpetual snow formerly covered the central axis, and fed glaciers which rolled 4,000 feet down the valleys. The same observer has recently found great moraines at a low level on the Atlas range in North Africa. Along the Himalaya, at points 900 miles apart, glaciers have left the marks of their former low descent; and in Sikkim, Dr. Hooker saw maize growing on ancient and gigantic moraines. Southward of the Asiatic continent, on the opposite side of the equator, we know, from the excellent researches of Dr. J. Haast and Dr. Hector, that in New Zealand immense glaciers formerly descended to a low level; and the same plants, found by Dr. Hooker on widely separated mountains in this island tell the same story of a former cold period. From facts communicated to me by the Rev. W.B. Clarke, it appears also that there are traces of former glacial action on the mountains of the south-eastern corner of Australia.

Looking to America: in the northern half, ice-borne fragments of rock have been observed on the eastern side of the continent, as far south as latitude 36 and 37 degrees, and on the shores of the Pacific, where the climate is now so different, as far south as latitude 46 degrees. Erratic boulders have, also, been noticed on the Rocky Mountains. In the Cordillera of South America, nearly under the equator, glaciers once extended far below their present level. In central Chile I examined a vast mound of detritus with great boulders, crossing the Portillo valley, which, there can hardly be a doubt, once formed a huge moraine; and Mr. D. Forbes informs me that he found in various parts of the Cordillera, from latitude 13 to 30 degrees south, at about the height of 12,000 feet, deeply-furrowed rocks, resembling those with which he was familiar in Norway, and likewise great masses of detritus, including grooved pebbles. Along this whole space of the Cordillera true glaciers do not now exist even at much more considerable heights. Further south, on both sides of the continent, from latitude 41 degrees to the southernmost extremity, we have the clearest evidence of former glacial action, in numerous immense boulders transported far from their parent source.

From these several facts, namely, from the glacial action having extended all round the northern and southern hemispheres – from the period having been in a geological sense recent in both hemispheres – from its having lasted in both during a great length of time, as may be inferred from the amount of work effected – and lastly, from glaciers having recently descended to a low level along the whole line of the Cordillera, it at one time appeared to me that we could not avoid the conclusion that the temperature of the whole world had been simultaneously lowered during the Glacial period. But now, Mr. Croll, in a series of admirable memoirs, has attempted to show that a glacial condition of climate is the result of various physical causes, brought into operation by an increase in the eccentricity of the earth's orbit. All these causes tend towards the same end; but the most powerful appears to be the indirect influence of the eccentricity of the orbit upon oceanic currents. According to Mr. Croll, cold periods regularly recur every ten or fifteen thousand years; and these at long intervals are extremely severe, owing to certain contingencies, of which the most important, as Sir C. Lyell has shown, is the relative position of the land and water. Mr. Croll believes that the last great glacial period occurred about 240,000 years ago, and endured, with slight alterations of climate, for about 160,000 years. With respect to more ancient glacial periods, several geologists are convinced, from direct evidence, that such occurred during the miocene and eocene formations, not to mention still more ancient formations. But the most important result for us, arrived at by Mr. Croll, is that whenever the northern hemisphere passes through a cold period the temperature of the southern hemisphere is actually raised, with the winters rendered much milder, chiefly through changes in the direction of the ocean currents. So conversely it will be with the northern hemisphere, while the southern passes through a glacial period. This conclusion throws so much light on geographical distribution that I am strongly inclined to trust in it; but I will first give the facts which demand an explanation.

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.