The Origin of Species by Means of Natural Selection

He who rejects this view of the imperfection of the geological record, will rightly reject the whole theory. For he may ask in vain where are the numberless transitional links which must formerly have connected the closely allied or representative species, found in the successive stages of the same great formation? He may disbelieve in the immense intervals of time which must have elapsed between our consecutive formations; he may overlook how important a part migration has played, when the formations of any one great region, as those of Europe, are considered; he may urge the apparent, but often falsely apparent, sudden coming in of whole groups of species. He may ask where are the remains of those infinitely numerous organisms which must have existed long before the Cambrian system was deposited? We now know that at least one animal did then exist; but I can answer this last question only by supposing that where our oceans now extend they have extended for an enormous period, and where our oscillating continents now stand they have stood since the commencement of the Cambrian system; but that, long before that epoch, the world presented a widely different aspect; and that the older continents, formed of formations older than any known to us, exist now only as remnants in a metamorphosed condition, or lie still buried under the ocean.

Passing from these difficulties, the other great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and natural selection. We can thus understand how it is that new species come in slowly and successively; how species of different classes do not necessarily change together, or at the same rate, or in the same degree; yet in the long run that all undergo modification to some extent. The extinction of old forms is the almost inevitable consequence of the production of new forms. We can understand why, when a species has once disappeared, it never reappears. Groups of species increase in numbers slowly, and endure for unequal periods of time; for the process of modification is necessarily slow, and depends on many complex contingencies. The dominant species belonging to large and dominant groups tend to leave many modified descendants, which form new sub-groups and groups. As these are formed, the species of the less vigorous groups, from their inferiority inherited from a common progenitor, tend to become extinct together, and to leave no modified offspring on the face of the earth. But the utter extinction of a whole group of species has sometimes been a slow process, from the survival of a few descendants, lingering in protected and isolated situations. When a group has once wholly disappeared, it does not reappear; for the link of generation has been broken.

We can understand how it is that dominant forms which spread widely and yield the greatest number of varieties tend to people the world with allied, but modified, descendants; and these will generally succeed in displacing the groups which are their inferiors in the struggle for existence. Hence, after long intervals of time, the productions of the world appear to have changed simultaneously.

We can understand how it is that all the forms of life, ancient and recent, make together a few grand classes. We can understand, from the continued tendency to divergence of character, why the more ancient a form is, the more it generally differs from those now living. Why ancient and extinct forms often tend to fill up gaps between existing forms, sometimes blending two groups, previously classed as distinct into one; but more commonly bringing them only a little closer together. The more ancient a form is, the more often it stands in some degree intermediate between groups now distinct; for the more ancient a form is, the more nearly it will be related to, and consequently resemble, the common progenitor of groups, since become widely divergent. Extinct forms are seldom directly intermediate between existing forms; but are intermediate only by a long and circuitous course through other extinct and different forms. We can clearly see why the organic remains of closely consecutive formations are closely allied; for they are closely linked together by generation. We can clearly see why the remains of an intermediate formation are intermediate in character.

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.