species being given, but this is unimportant for us. The horizontal lines may represent successive geological formations, and all the forms beneath the uppermost line may be considered as extinct. The three existing genera, a14, q14, p14, will form a small family; b14 and f14 a closely allied family or sub-family; and o14, e14, m14, a third family. These three families, together with the many extinct genera on the several lines of descent diverging from the parent-form A, will form an order; for all will have inherited something in common from their ancient and common progenitor. On the principle of the continued tendency to divergence of character, which was formerly illustrated by this diagram, the more recent any form is, the more it will generally differ from its ancient progenitor. Hence we can understand the rule that the most ancient fossils differ most from existing forms. We must not, however, assume that divergence of character is a necessary contingency; it depends solely on the descendants from a species being thus enabled to seize on many and different places in the economy of nature. Therefore it is quite possible, as we have seen in the case of some Silurian forms, that a species might go on being slightly modified in relation to its slightly altered conditions of life, and yet retain throughout a vast period the same general characteristics. This is represented in the diagram by the letter F14.
All the many forms, extinct and recent, descended from A, make, as before remarked, one order; and this order, from the continued effects of extinction and divergence of character, has become divided into several sub-families and families, some of which are supposed to have perished at different periods, and some to have endured to the present day.
By looking at the diagram we can see that if many of the extinct forms, supposed to be embedded in the successive formations, were discovered at several points low down in the series, the three existing families on the uppermost line would be rendered less distinct from each other. If, for instance, the genera a1, a5, a10, f8, m3, m6, m9 were disinterred, these three families would be so closely linked together that they probably would have to be united into one great family, in nearly the same manner as has occurred with ruminants and pachyderms. Yet he who objected to call the extinct genera, which thus linked the living genera of three families together, intermediate in character, would be justified, as they are intermediate, not directly, but only by a long and circuitous course through many widely different forms. If many extinct forms were to be discovered above one of the middle horizontal lines or geological formations–for instance, above Number VI.–but none from beneath this line, then only the two families on the left hand (namely, a14, etc., and b14, etc.) would have to be united into one family; and the two other families (namely, a14 to f14 now including five genera, and o14 to m14) would yet remain distinct. These two families, however, would be less distinct from each other than they were before the discovery of the fossils. If, for instance, we suppose the existing genera of the two families to differ from each other by a dozen characters, in this case the genera, at the early period marked VI., would differ by a lesser number of characters; for at this early stage of descent they have not diverged in character from the common progenitor of the order, nearly so much as they subsequently diverged. Thus it comes that ancient and extinct genera are often in some slight degree intermediate in character between their modified descendants, or between their collateral relations.
In nature the case will be far more complicated than is represented in the diagram; for the groups will have been more numerous, they will have endured for extremely unequal lengths of time, and will have been modified in various degrees. As we possess only the last volume of the geological record, and that in a very broken condition, we have no right to expect, except in very rare cases, to fill up wide intervals in the natural system, and thus unite distinct families or orders. All that we have a right to expect, is that those groups, which have within known geological periods undergone much modification, should in the older formations make some slight approach to each other; so that the older members should differ less from each other in some of their characters than do the existing members of the same groups; and this by the concurrent evidence of our best palaeontologists seems frequently to be the case.
Thus, on the theory of descent with modification, the main facts with respect to the mutual affinities of the extinct forms of life to each other and to living forms, seem to me explained in a satisfactory manner. And they are wholly inexplicable on any other view.
On this same theory, it is evident that the fauna of any great period in the earth’s history will be intermediate in general character between that which preceded and that which succeeded it. Thus, the species which lived at the sixth great stage of descent in the diagram are the modified offspring of those which lived at the fifth stage, and are the parents of those which became still more modified at the seventh stage; hence they could hardly fail to be nearly intermediate in character between the forms of life above and below. We must, however, allow for the entire extinction of some preceding forms, and for the coming in of quite new forms by immigration, and for a large amount of modification, during the long and blank intervals between the successive formations. Subject to these allowances, the fauna of each geological period undoubtedly is intermediate in character, between the preceding and succeeding faunas. I need give only one instance, namely, the manner in which the fossils of the Devonian system, when this system was first discovered, were at once recognised by palaeontologists as intermediate in character between those of the overlying carboniferous, and underlying Silurian system. But each fauna is not necessarily exactly intermediate, as unequal intervals of time have elapsed between consecutive formations.
It is no real objection to the truth of the statement, that the fauna of each period as a whole is nearly intermediate in character between the preceding and succeeding faunas, that certain genera offer exceptions to the rule. For instance, mastodons and elephants, when arranged by Dr. Falconer in two series, first according to their mutual affinities and then according to their periods of existence, do not accord in arrangement. The species extreme in character are not the oldest, or the most recent; nor are those which are intermediate in character, intermediate in age. But supposing for an instant, in this and other such cases, that the record of the first appearance and disappearance of the species was perfect, we have no reason to believe that forms successively produced necessarily endure for corresponding lengths of time: a very ancient form might occasionally last much longer than a form elsewhere subsequently produced, especially in the case of terrestrial productions inhabiting separated districts. To compare small things with great: if the principal living and extinct races of the domestic pigeon were arranged as well as they could be in serial affinity, this arrangement would not closely accord with the order in time of their production, and still less with the order of their disappearance; for the parent rock-pigeon now lives; and many varieties between the rock-pigeon and the carrier have become extinct; and carriers which are extreme in the important character of length of beak originated earlier than short-beaked tumblers, which are at the opposite end of the series in this same respect.
Closely connected with the statement, that the organic remains from an intermediate formation are in some degree intermediate in character, is the fact, insisted on by all palaeontologists, that fossils from two consecutive formations are far more closely related to each other, than are the fossils from two remote formations. Pictet gives as a well-known instance, the general resemblance of the organic remains from the several stages of the chalk formation, though the species are distinct in each stage. This fact alone, from its generality, seems to have shaken Professor Pictet in his firm belief in the immutability of species. He who is acquainted with the distribution of existing species over the globe, will not attempt to account for the close resemblance of the distinct species in closely consecutive formations, by the physical conditions of the ancient areas having remained nearly the same. Let it be remembered that the forms of life, at least those inhabiting the sea, have changed almost simultaneously throughout the world, and therefore under the most different climates and conditions. Consider the prodigious vicissitudes of climate during the pleistocene period, which includes the whole glacial period, and note how little the specific forms of the inhabitants of the sea have been affected.
On the theory of descent, the full meaning of the fact of fossil remains from closely consecutive formations, though ranked as distinct species, being closely related, is obvious. As the accumulation of each formation has often been interrupted, and as long blank intervals have intervened between successive formations, we ought not to expect to find, as I attempted to show in the last chapter, in any one or two formations all the intermediate varieties between the species which appeared at the commencement and close of these periods; but we ought to find after intervals, very long as measured by years, but only moderately long as measured geologically, closely allied forms, or, as they have been called by some authors, representative species; and these we assuredly do find. We find, in short, such evidence of the slow and scarcely sensible mutation of specific forms, as we have a just right to expect to find.
ON THE STATE OF DEVELOPMENT OF ANCIENT FORMS.
There has been much discussion whether recent forms are more highly developed than ancient. I will not here enter on this subject, for naturalists have not as yet defined to each other’s satisfaction what is meant by high and low forms. But in one particular sense the more recent forms must, on my theory, be higher than the more ancient; for each new species is formed by having had some advantage in the struggle for life over other and preceding forms. If under a nearly similar climate, the eocene inhabitants of one quarter of the world were put into competition with the existing inhabitants of the same or some other quarter, the eocene fauna or flora would certainly be beaten and exterminated; as would a secondary fauna by an eocene, and a palaeozoic fauna by a secondary fauna. I do not doubt that this process of improvement has affected in a marked and sensible manner the organisation of the more recent and victorious forms of life, in comparison with the ancient and beaten forms; but I can see no way of testing this sort of progress. Crustaceans, for instance, not the highest in their own class, may have beaten the highest molluscs. From the extraordinary manner in which European productions have recently spread over New Zealand, and have seized on places which must have been previously occupied, we may believe, if all the animals and plants of Great Britain were set free in New Zealand, that in the course of time a multitude of British forms would become thoroughly naturalized there, and would exterminate many of the natives. On the other hand, from what we see now occurring in New Zealand, and from hardly a single inhabitant of the southern hemisphere having become wild in any part of Europe, we may doubt, if all the productions of New Zealand were set free in Great Britain, whether any considerable number would be enabled to seize on places now occupied by our native plants and animals. Under this point of view, the productions of Great Britain may be said to be higher than those of New Zealand. Yet the most skilful naturalist from an examination of the species of the two countries could not have foreseen this result.
Agassiz insists that ancient animals resemble to a certain extent the embryos of recent animals of the same classes; or that the geological succession of extinct forms is in some degree parallel to the embryological development of recent forms. I must follow Pictet and Huxley in thinking that the truth of this doctrine is very far from proved. Yet I fully expect to see it hereafter confirmed, at least in regard to subordinate groups, which have branched off from each other within comparatively recent times. For this doctrine of Agassiz accords well with the theory of natural selection. In a future chapter I shall attempt to show that the adult differs from its embryo, owing to variations supervening at a not early age, and being inherited at a corresponding age. This process, whilst it leaves the embryo almost unaltered, continually adds, in the course of successive generations, more and more difference to the adult.
Thus the embryo comes to be left as a sort of picture, preserved by nature, of the ancient and less modified condition of each animal. This view may be true, and yet it may never be capable of full proof. Seeing, for instance, that the oldest known mammals, reptiles, and fish strictly belong to their own proper classes, though some of these old forms are in a slight degree less distinct from each other than are the typical members of the same groups at the present day, it would be vain to look for animals having the common embryological character of the Vertebrata, until beds far beneath the lowest Silurian strata are discovered–a discovery of which the chance is very small.
ON THE SUCCESSION OF THE SAME TYPES WITHIN THE SAME AREAS, DURING THE LATER TERTIARY PERIODS.
Mr. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent. In South America, a similar relationship is manifest, even to an uneducated eye, in the gigantic pieces of armour like those of the armadillo, found in several parts of La Plata; and Professor Owen has shown in the most striking manner that most of the fossil mammals, buried there in such numbers, are related to South American types. This relationship is even more clearly seen in the wonderful collection of fossil bones made by MM. Lund and Clausen in the caves of Brazil. I was so much impressed with these facts that I strongly insisted, in 1839 and 1845, on this “law of the succession of types,”–on “this wonderful relationship in the same continent between the dead and the living.” Professor Owen has subsequently extended the same generalisation to the mammals of the Old World. We see the same law in this author’s restorations of the extinct and gigantic birds of New Zealand. We see it also in the birds of the caves of Brazil. Mr. Woodward has shown that the same law holds good with sea-shells, but from the wide distribution of most genera of molluscs, it is not well displayed by them. Other cases could be added, as the relation between the extinct and living land-shells of Madeira; and between the extinct and living brackish-water shells of the Aralo-Caspian Sea.
Now what does this remarkable law of the succession of the same types within the same areas mean? He would be a bold man, who after comparing the present climate of Australia and of parts of South America under the same latitude, would attempt to account, on the one hand, by dissimilar physical conditions for the dissimilarity of the inhabitants of these two continents, and, on the other hand, by similarity of conditions, for the uniformity of the same types in each during the later tertiary periods. Nor can it be pretended that it is an immutable law that marsupials should have been chiefly or solely produced in Australia; or that Edentata and other American types should have been solely produced in South America. For we know that Europe in ancient times was peopled by numerous marsupials; and I have shown in the publications above alluded to, that in America the law of distribution of terrestrial mammals was formerly different from what it now is. North America formerly partook strongly of the present character of the southern half of the continent; and the southern half was formerly more closely allied, than it is at present, to the northern half. In a similar manner we know from Falconer and Cautley’s discoveries, that northern India was formerly more closely related in its mammals to Africa than it is at the present time. Analogous facts could be given in relation to the distribution of marine animals.
On the theory of descent with modification, the great law of the long enduring, but not immutable, succession of the same types within the same areas, is at once explained; for the inhabitants of each quarter of the world will obviously tend to leave in that quarter, during the next succeeding period of time, closely allied though in some degree modified descendants. If the inhabitants of one continent formerly differed greatly from those of another continent, so will their modified descendants still differ in nearly the same manner and degree. But after very long intervals of time and after great geographical changes, permitting much inter-migration, the feebler will yield to the more dominant forms, and there will be nothing immutable in the laws of past and present distribution.
It may be asked in ridicule, whether I suppose that the megatherium and other allied huge monsters have left behind them in South America the sloth, armadillo, and anteater, as their degenerate descendants. This cannot for an instant be admitted. These huge animals have become wholly extinct, and have left no progeny. But in the caves of Brazil, there are many extinct species which are closely allied in size and in other characters to the species still living in South America; and some of these fossils may be the actual progenitors of living species. It must not be forgotten that, on my theory, all the species of the same genus have descended from some one species; so that if six genera, each having eight species, be found in one geological formation, and in the next succeeding formation there be six other allied or representative genera with the same number of species, then we may conclude that only one species of each of the six older genera has left modified descendants, constituting the six new genera. The other seven species of the old genera have all died out and have left no progeny. Or, which would probably be a far commoner case, two or three species of two or three alone of the six older genera will have been the parents of the six new genera; the other old species and the other whole genera having become utterly extinct. In failing orders, with the genera and species decreasing in numbers, as apparently is the case of the Edentata of South America, still fewer genera and species will have left modified blood-descendants.
SUMMARY OF THE PRECEDING AND PRESENT CHAPTERS.
I have attempted to show that the geological record is extremely imperfect; that only a small portion of the globe has been geologically explored with care; that only certain classes of organic beings have been largely preserved in a fossil state; that the number both of specimens and of species, preserved in our museums, is absolutely as nothing compared with the incalculable number of generations which must have passed away even during a single formation; that, owing to subsidence being necessary for the accumulation of fossiliferous deposits thick enough to resist future degradation, enormous intervals of time have elapsed between the successive formations; that there has probably been more extinction during the periods of subsidence, and more variation during the periods of elevation, and during the latter the record will have been least perfectly kept; that each single formation has not been continuously deposited; that the duration of each formation is, perhaps, short compared with the average duration of specific forms; that migration has played an important part in the first appearance of new forms in any one area and formation; that widely ranging species are those which have varied most, and have oftenest given rise to new species; and that varieties have at first often been local. All these causes taken conjointly, must have tended to make the geological record extremely imperfect, and will to a large extent explain why we do not find interminable varieties, connecting together all the extinct and existing forms of life by the finest graduated steps.
He who rejects these views on the nature of the geological record, will rightly reject my 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 several stages of the same great formation. He may disbelieve in the enormous intervals of time which have elapsed between our consecutive formations; he may overlook how important a part migration must have played, when the formations of any one great region alone, as that 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 first bed of the Silurian system was deposited: I can answer this latter question only hypothetically, by saying that as far as we can see, where our oceans now extend they have for an enormous period extended, and where our oscillating continents now stand they have stood ever since the Silurian epoch; but that long before that period, the world may have presented a wholly different aspect; and that the older continents, formed of formations older than any known to us, may now all be in a metamorphosed condition, or may lie buried under the ocean.
Passing from these difficulties, all the other great leading facts in palaeontology seem to me simply to follow on the theory of descent with modification through 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 of the larger dominant groups tend to leave many modified descendants, and thus new sub-groups and groups are formed. 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 may often be a very 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 the spreading of the dominant forms of life, which are those that oftenest vary, will in the long run tend to people the world with allied, but modified, descendants; and these will generally succeed in taking the places of those groups of species which are their inferiors in the struggle for existence. Hence, after long intervals of time, the productions of the world will appear to have changed simultaneously.
We can understand how it is that all the forms of life, ancient and recent, make together one grand system; for all are connected by generation. 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 only bringing them a little closer together. The more ancient a form is, the more often, apparently, it displays characters 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 many extinct and very different forms. We can clearly see why the organic remains of closely consecutive formations are more closely allied to each other, than are those of remote formations; for the forms are more closely linked together by generation: we can clearly see why the remains of an intermediate formation are intermediate in character.
The inhabitants of each successive period in the world’s history have beaten their predecessors in the race for life, and are, in so far, higher in the scale of nature; and this may account for that vague yet ill-defined sentiment, felt by many palaeontologists, that organisation on the whole has progressed. If it should hereafter be proved that ancient animals resemble to a certain extent the embryos of more recent animals of the same class, the fact will be intelligible. The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious, and is simply explained by inheritance.
If then the geological record be as imperfect as I believe it to be, 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, produced by the laws of variation still acting round us, and preserved by Natural Selection.
CHAPTER 11. 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 co-extensive with the world.
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 accounted for by their 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 northern parts where the circumpolar land is almost continuous, 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; the most 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 as closely as the same species generally require; for it is a most rare case to find a group of organisms confined to any small spot, having conditions peculiar in only a slight degree; for instance, small areas in the Old World could be pointed out hotter than any in the New World, yet these are not inhabited by a peculiar fauna or flora. Notwithstanding this parallelism in the conditions of the 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 deg and 35 deg, 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 lat. 35 deg with those north of 25 deg, which consequently inhabit a considerably different climate, and they will be found 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 of 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 sometimes 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. No two marine faunas are more distinct, with hardly a fish, shell, or crab in common, than those of the eastern and western shores of South and Central America; yet these great faunas are separated only by the narrow, but impassable, isthmus of Panama. 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 here three marine faunas range far northward and southward, in parallel lines not far from each other, under corresponding climates; but from being separated from each other by impassable barriers, either of land or open sea, they are 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, 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 hardly one shell, crab or fish is common to the above-named three approximate faunas of Eastern and Western America and the eastern Pacific islands, yet many fish 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 statements, is the affinity of the productions of the same continent or 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, yet clearly 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 emeu, like those found in 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 musk-rat, but the coypu and capybara, rodents of the 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, though they may be all peculiar species, are essentially American. We may look back to past ages, as shown in the last chapter, and we find American types then prevalent on the American continent and in the American seas. We see in these facts some deep organic bond, prevailing throughout space and time, over the same areas of land and water, and independent of their physical conditions. The naturalist must feel little curiosity, who is not led to inquire what this bond is.
This bond, on my theory, is simply inheritance, that cause which alone, as far as we positively know, produces organisms quite like, or, as we see in the case of varieties nearly like each other. The dissimilarity of the inhabitants of different regions may be attributed to modification through natural selection, and in a quite subordinate degree to the direct influence of different physical conditions. The degree of dissimilarity will depend on the migration of the more dominant forms of life from one region into another having been effected with more or less ease, at periods more or less remote;–on the nature and number of the former immigrants;–and on their action and reaction, in their mutual struggles for life;–the relation of organism to organism 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 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.
I believe, as was remarked in the last chapter, in no 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 the individual in its complex struggle for life, so the degree of modification in different species will be no uniform quantity. If, for instance, a number of species, which stand in direct competition with each other, migrate in a body into a new and afterwards isolated country, they will be little liable to modification; for neither migration nor isolation in themselves can do 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 modified.
On 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 have descended from the same progenitor. In the case of those species, which have undergone during whole geological periods but little modification, there is not much difficulty in believing that they may have migrated from the same region; for during the vast geographical and climatal changes which will 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 explained in the last chapter, it is incredible that individuals identically the same should ever have been produced through natural selection from parents specifically distinct.
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 very 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 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 be easily passed over by migration, the fact is given as something remarkable and exceptional. The capacity of migrating across the sea is more distinctly limited in terrestrial mammals, than perhaps in any other organic beings; and, accordingly, we find no inexplicable cases of the same mammal inhabiting distant points of the world. No geologist will feel any difficulty in such cases as Great Britain having been formerly united to Europe, and consequently possessing the same quadrupeds. 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 vast and broken interspace. The great and striking influence which barriers of every kind have had on distribution, is intelligible only on the view that the great majority of species have been produced on one side alone, and have not been able to migrate to the other side. Some few families, many sub-families, very many genera, and 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 local, or confined to one area. What a strange anomaly it would be, if, when coming one step lower in the series, to the individuals of the same species, a directly opposite rule prevailed; and species were not local, but had been produced in two or more distinct areas!
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 climatal changes, which have certainly occurred within recent geological times, must have interrupted or 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 such cases. 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 freshwater productions; and thirdly, the occurrence of the same terrestrial species on islands and on the 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 climatal and geographical changes and various occasional means of transport, the belief that this has been the universal 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 distinct species of a genus, which on my theory have all descended from a common progenitor, can have migrated (undergoing modification during some part of their migration) from the area inhabited by their progenitor. If it can be shown to be almost invariably the case, that a region, of which most of its inhabitants are closely related to, or belong to the same genera with the species of a second region, has probably received at some former period immigrants from this other region, my theory will be strengthened; for we can clearly understand, on the principle of modification, why the inhabitants of a region should be related to those of another region, whence it has been stocked. 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 plainly related by inheritance to the inhabitants of the continent. Cases of this nature are common, and are, as we shall hereafter more fully see, inexplicable on the theory of independent creation. This view of the relation of species in one region to those in another, does not differ much (by substituting the word variety for species) from that lately advanced in an ingenious paper by Mr. Wallace, in which he concludes, that “every species has come into existence coincident both in space and time with a pre-existing closely allied species.” And I now know from correspondence, that this coincidence he attributes to generation with modification.
The previous remarks on “single and multiple centres of creation” do not directly bear on another allied question,–namely whether all the individuals of the same species have descended from a single pair, or single hermaphrodite, or whether, as some authors suppose, from many individuals simultaneously created. With those organic beings which never intercross (if such exist), the species, on my theory, must have descended from a succession of improved varieties, which will never have blended with other individuals or varieties, but will have supplanted each other; so that, at each successive stage of modification and improvement, all the individuals of each variety will have descended from a single parent. But in the majority of cases, namely, with all organisms which habitually unite for each birth, or which often intercross, I believe that during the slow process of modification the individuals of the species will have been kept nearly uniform by intercrossing; so that many individuals will have gone on simultaneously changing, and the whole amount of modification will not have been due, at each stage, to descent from a single parent. To illustrate what I mean: our English racehorses differ slightly 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 selecting and training many individuals during many generations.
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 when its climate was different may have been a high road for migration, but now be impassable; 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 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 will dispute that great mutations of level have occurred within the period of existing organisms. Edward Forbes insisted that all the islands in the Atlantic must recently have been connected with Europe or Africa, and Europe likewise with America. Other authors have thus hypothetically bridged over every ocean, and have united almost every island to 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 of level in our continents; but not of such vast changes in their position and extension, 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, as I believe, by rings of coral or atolls standing over them. Whenever it is fully admitted, as I believe 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 continents which are now 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,–a certain degree of relation (as we shall hereafter see) between the distribution of mammals and the depth of the sea,–these and other such facts seem to me opposed to the admission of such prodigious geographical revolutions within the recent period, as are necessitated on the view advanced by Forbes and admitted by his many followers. The nature and relative proportions of the inhabitants of oceanic islands likewise seem to me opposed to the belief of their former continuity with continents. Nor does their almost universally volcanic composition favour the admission that they are the wrecks of sunken continents;–if they had originally existed as mountain-ranges on the land, some at least of the islands would have been formed, like other mountain-summits, of granite, metamorphic schists, old fossiliferous or other such 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 might be called occasional means of distribution. I shall here confine myself to plants. In botanical works, this or that plant is stated to be ill adapted for wide dissemination; but for transport across the sea, the greater or less facilities 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 87 kinds, 64 germinated after an immersion of 28 days, and a few survived an immersion of 137 days. For convenience sake I chiefly tried small seeds, without the capsule or fruit; and as all of these sank in a few days, they could not be floated across wide spaces of the sea, whether or not they were injured by the 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 might wash down plants or branches, and that these might be dried on the banks, and then by a fresh rise in the stream be washed into the sea. Hence I was led to dry stems and branches of 94 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 90 days and afterwards when planted they germinated; an asparagus plant with ripe berries floated for 23 days, when dried it floated for 85 days, and the seeds afterwards germinated: the ripe seeds of Helosciadium sank in two days, when dried they floated for above 90 days, and afterwards germinated. Altogether out of the 94 dried plants, 18 floated for above 28 days, and some of the 18 floated for a very much longer period. So that as 64/87 seeds germinated after an immersion of 28 days; and as 18/94 plants with ripe fruit (but not all the same species as in the foregoing experiment) floated, after being dried, for above 28 days, as far as we may infer anything from these scanty facts, we may conclude that the seeds of 14/100 plants of any country might be floated by sea-currents during 28 days, and would retain their power of germination. In Johnston’s Physical Atlas, the average rate of the several Atlantic currents is 33 miles per diem (some currents running at the rate of 60 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 to a favourable spot by an inland gale, they 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 98 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 the average length of their flotation and of 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 floated for 42 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 could hardly be transported by any other means; and Alph. de Candolle has shown that such plants generally have restricted ranges.
But 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 on examination, 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 in the longest transport: out of one small portion of earth thus COMPLETELY enclosed by wood in an oak about 50 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 seeds of many kinds 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 salt-water for 30 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 I think safely assume that under such circumstances their rate of flight would often be 35 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 will pass uninjured through even the digestive organs of a turkey. In the course of two months, I picked up in my garden 12 kinds of seeds, out of the excrement of small birds, and these seemed perfect, and some of them, which I tried, germinated. But the following fact is more important: the crops of birds do not secrete gastric juice, and do not in the least injure, as I know by trial, 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 12 or even 18 hours. A bird in this interval might easily be blown to the distance of 500 miles, and hawks are known to look out for tired birds, and the contents of their torn crops might thus readily get scattered. Mr. Brent informs me that a friend of his had to give up flying carrier-pigeons from France to England, as the hawks on the English coast destroyed so many on their arrival. 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. Freshwater 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 their power of germination. Certain seeds, however, were always killed by this process.
Although the beaks and feet of birds are generally quite clean, I can show that earth sometimes adheres to them: in one instance I removed twenty-two grains of dry argillaceous earth from one foot of a partridge, and in this earth there was a pebble quite as large as the seed of a vetch. Thus seeds might occasionally be transported to great distances; for many facts could be given showing that soil almost everywhere is charged with seeds. Reflect for a moment on the millions of quails which annually cross the Mediterranean; and can we doubt that the earth adhering to their feet would sometimes include a few minute seeds? But I shall presently 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, I can hardly doubt that they must occasionally have transported seeds from one part to another of the arctic and antarctic regions, as suggested by Lyell; and during the Glacial period from one part of the now temperate regions to another. In the Azores, from the large number of the species of plants common to Europe, in comparison with the plants of other oceanic islands nearer to the mainland, and (as remarked by Mr. H. C. Watson) from the somewhat northern character of the flora 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 burthens 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 the several above means of transport, and that several other means, which without doubt remain to be discovered, have been in action year after year, for centuries and 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 seawater; 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 in any great degree; but would remain as distinct as we now see them to be. 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 so long an 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 wanderers only by one means, namely, in dirt sticking to their feet, which is in itself a rare accident. Even in this case, how small would the chance be 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. I do not doubt that out of twenty seeds or animals transported to an island, even if far less well-stocked than Britain, scarcely more than one would be so well fitted to its new home, as to become naturalised. But this, as it seems to me, is no valid argument against what would be effected by occasional means of transport, during the long lapse of geological time, whilst an island was being upheaved and formed, 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, would be sure to 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 the 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 to the other. It is indeed a remarkable fact to see so many of the same plants 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 several 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 rocks scored by drifted icebergs and coast-ice, plainly reveal a former cold period.
The former influence of the glacial climate on the distribution of the inhabitants of Europe, as explained with remarkable clearness by Edward Forbes, is substantially as follows. But we shall follow the changes more readily, by supposing a new glacial period to come slowly on, and then pass away, as formerly occurred. As the cold came on, and as each more southern zone became fitted for arctic beings and ill-fitted for their former more temperate inhabitants, the latter would be supplanted and arctic productions would take their places. The inhabitants of the more temperate regions would at the same time travel 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 a uniform 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. We may suppose that the Glacial period came on a little earlier or later in North America than in Europe, so will the southern migration there have been a little earlier or later; but this will make no difference in the final result.
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 higher and higher, as the warmth increased, whilst their brethren were pursuing their northern journey. Hence, when the warmth had fully returned, the same arctic species, which had lately lived in a body together on the lowlands of the Old and New Worlds, would be left isolated on distant mountain-summits (having been exterminated on all lesser heights) and in the arctic regions of both hemispheres.
Thus we can understand the identity of many plants at points so immensely remote as on the mountains of the United States and 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 migration as the cold came on, and the re-migration on the returning warmth, will 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 permitted their former migration across the low intervening tracts, since become too warm for their existence.
If the climate, since the Glacial period, has ever been in any degree warmer than at present (as some geologists in the United States believe to have been the case, chiefly from the distribution of the fossil Gnathodon), then the arctic and temperate productions will at a very late period have marched a little further north, and subsequently have retreated to their present homes; but I have met with no satisfactory evidence with respect to this intercalated slightly warmer period, since the Glacial period.
The arctic forms, during their long southern migration and re-migration northward, will have been exposed to nearly the same climate, and, as is especially to be noticed, they will have kept in a body together; consequently their mutual relations will not have been much disturbed, and, in accordance with the principles inculcated in this volume, they will not have been liable to much modification. But with our 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 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 its coldest period will have been temporarily driven down to the plains; they will, also, have been exposed to somewhat different climatal influences. Their mutual relations will thus have been in some degree disturbed; consequently they will have been liable to modification; and this we find has been the case; for if we compare the present Alpine plants and animals of the several great European mountain-ranges, though very many of the species are identically the same, some present varieties, some are ranked as doubtful forms, and some few are distinct yet closely allied or representative species.
In illustrating what, as I believe, actually took place during the Glacial period, I assumed that at its commencement the arctic productions were as uniform round the polar regions as they are at the present day. But the foregoing remarks on distribution apply not only to strictly arctic forms, but also to many sub-arctic and to some few northern temperate forms, for some of these are the same on the lower mountains and on the plains of North America and Europe; and it may be reasonably asked how I account for the necessary degree of uniformity of the sub-arctic and northern temperate forms round the world, at the commencement of the 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 Atlantic Ocean and by the extreme northern part of the Pacific. During the Glacial period, when the inhabitants of the Old and New Worlds lived further southwards than at present, they must have been still more completely separated by wider spaces of ocean. I believe the above difficulty may be surmounted by looking to still earlier changes of climate of an opposite nature. We have good reason to believe that during the newer Pliocene period, before the Glacial epoch, and whilst the majority of the inhabitants of the world were specifically the same as now, the climate was warmer than at the present day. Hence we may suppose that the organisms now living under the climate of latitude 60 deg, during the Pliocene period lived further north under the Polar Circle, in latitude 66 deg-67 deg; and that the strictly arctic productions then lived on the broken land still nearer to the pole. Now if we look at a globe, we shall see that under the Polar Circle there is almost continuous land from western Europe, through Siberia, to eastern America. And to this continuity of the circumpolar land, and to the consequent freedom for intermigration under a more favourable climate, I attribute the necessary amount of uniformity in the sub-arctic and northern 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 large, but partial 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 most remarkable, considering the distance of the two areas, and their separation by the 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 inter-migration 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 must have been completely cut off from each other. This separation, as far as the more temperate productions are concerned, took place long ages ago. And as the plants and animals migrated southward, they will have become mingled in the one great region with the native American productions, and 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 the two Worlds. 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 areas completely sundered. Thus, I think, we can understand the presence of many existing and tertiary representative forms on the eastern and western shores of temperate North America; and the still more striking case of many closely allied crustaceans (as described in Dana’s admirable work), of some fish and other marine animals, in the Mediterranean and in the seas of Japan,–areas now separated by a continent and by nearly a hemisphere of equatorial ocean.
These cases of relationship, without identity, of the inhabitants of seas now disjoined, and likewise of the past and present inhabitants of the temperate lands of North America and Europe, are inexplicable on the theory of creation. We cannot say that they 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 the southern continents of the Old World, we see countries closely corresponding in all their physical conditions, but with their inhabitants utterly dissimilar.
But we must return to our more immediate subject, the Glacial period. I am convinced that Forbes’s view may be largely extended. In Europe we have the plainest evidence of the cold period, from the western shores of Britain to the Oural range, and southward to the Pyrenees. We may infer, from the frozen mammals and nature of the mountain vegetation, that Siberia was similarly affected. 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 gigantic ancient moraines. South of the equator, we have some direct evidence of former glacial action in New Zealand; and the same plants, found on widely separated mountains in this island, tell the same story. If one account which has been published can be trusted, we have direct evidence of glacial action in 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 as far south as lat. 36 deg-37 deg, and on the shores of the Pacific, where the climate is now so different, as far south as lat. 46 deg; erratic boulders have, also, been noticed on the Rocky Mountains. In the Cordillera of Equatorial South America, glaciers once extended far below their present level. In central Chile I was astonished at the structure of a vast mound of detritus, about 800 feet in height, crossing a valley of the Andes; and this I now feel convinced was a gigantic moraine, left far below any existing glacier. Further south on both sides of the continent, from lat. 41 deg to the southernmost extremity, we have the clearest evidence of former glacial action, in huge boulders transported far from their parent source.
We do not know that the Glacial epoch was strictly simultaneous at these several far distant points on opposite sides of the world. But we have good evidence in almost every case, that the epoch was included within the latest geological period. We have, also, excellent evidence, that it endured for an enormous time, as measured by years, at each point. The cold may have come on, or have ceased, earlier at one point of the globe than at another, but seeing that it endured for long at each, and that it was contemporaneous in a geological sense, it seems to me probable that it was, during a part at least of the period, actually simultaneous throughout the world. Without some distinct evidence to the contrary, we may at least admit as probable that the glacial action was simultaneous on the eastern and western sides of North America, in the Cordillera under the equator and under the warmer temperate zones, and on both sides of the southern extremity of the continent. If this be admitted, it is difficult to avoid believing that the temperature of the whole world was at this period simultaneously cooler. But it would suffice for my purpose, if the temperature was at the same time lower along certain broad belts of longitude.
On this view of the whole world, or at least of broad longitudinal belts, having been simultaneously colder from pole to pole, much light can be thrown on the present distribution of identical and allied species. In America, Dr. Hooker has shown that between forty and fifty of the flowering plants of Tierra del Fuego, forming no inconsiderable part of its scanty flora, are common to Europe, enormously remote as these two points are; and there are many closely allied species. On the lofty mountains of equatorial America a host of peculiar species belonging to European genera occur. On the highest mountains of Brazil, some few European genera were found by Gardner, which do not exist in the wide intervening hot countries. So on the Silla of Caraccas the illustrious Humboldt long ago found species belonging to genera characteristic of the Cordillera. On the mountains of Abyssinia, several European forms and some few representatives of the peculiar flora of the Cape of Good Hope occur. At the Cape of Good Hope a very few European species, believed not to have been introduced by man, and on the mountains, some few representative European forms are found, which have not been discovered in the intertropical parts of Africa. 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 collected on the loftier peaks of Java raises a picture of a collection made on a hill in Europe! Still more striking is the fact that southern Australian forms are clearly represented by 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 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 throughout the world, the plants growing on the more lofty mountains, and on the temperate lowlands of the northern and southern hemispheres, are sometimes identically the same; but they are much oftener specifically distinct, though related to each other in a most remarkable manner.
This brief abstract applies to plants alone: some strictly analogous facts could be given on the distribution of terrestrial animals. In marine productions, similar cases occur; as an example, I may quote a remark by the highest authority, Professor 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.
It should be observed that the northern species and forms found in the southern parts of the southern hemisphere, and on the mountain-ranges of the intertropical regions, are not arctic, but belong to the northern temperate zones. As Mr. H. C. Watson has recently remarked, “In receding from polar towards equatorial latitudes, the Alpine or mountain floras really become less and less arctic.” Many of the forms living on the mountains of the warmer regions of the earth and in the southern hemisphere are of doubtful value, being ranked by some naturalists as specifically distinct, by others as varieties; but some are certainly identical, and many, though closely related to northern forms, must be ranked as distinct species.
Now let us see what light can be thrown on the foregoing facts, on the belief, supported as it is by a large body of geological evidence, that the whole world, or a large part of it, was during the Glacial period simultaneously much colder than at present. 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 came slowly on, all the tropical plants and other productions will have retreated from both sides towards the equator, followed in the rear by the temperate productions, and these by the arctic; but with the latter we are not now concerned. The tropical plants probably suffered much extinction; how much no one can say; perhaps formerly the tropics supported as many species as we see at the present day crowded together at the Cape of Good Hope, and in parts of temperate Australia. As we know that many tropical plants and animals can withstand a considerable amount of cold, many might have escaped extermination during a moderate fall of temperature, more especially by escaping into the warmest spots. But the great fact to bear in mind is, that all tropical productions will have suffered to a certain extent. On the other hand, the temperate productions, after migrating nearer to the equator, though they will have been placed under somewhat new conditions, will have suffered less. And it is certain that many temperate plants, if protected from the inroads of competitors, can withstand a much warmer climate than their own. Hence, it seems to me possible, bearing in mind that the tropical productions were in a suffering state and could not have presented a firm front against intruders, that a certain number of the more vigorous and dominant temperate forms might have penetrated the native ranks and have reached or even crossed the equator. The invasion would, of course, have been greatly favoured by high land, and perhaps by a dry climate; for Dr. Falconer informs me that it is the damp with the heat of the tropics which is so destructive to perennial plants from a temperate climate. On the other hand, the most humid and hottest districts will have afforded an asylum to the tropical natives. The mountain-ranges north-west of the Himalaya, and the long line of the Cordillera, seem to have afforded two great lines of invasion: and it is a striking fact, lately communicated to me by Dr. Hooker, that all the flowering plants, about forty-six in number, common to Tierra del Fuego and to Europe still exist in North America, which must have lain on the line of march. But I do not doubt that some temperate productions entered and crossed even the LOWLANDS of the tropics at the period when the cold was most intense,–when arctic forms had migrated some twenty-five degrees of latitude from their native country and covered the land at the foot of the Pyrenees. At this period of extreme cold, I believe that the climate under the equator at the level of the sea was about the same with that now felt there at the height of six or seven thousand feet. During this the coldest period, I suppose that large spaces of the tropical lowlands were clothed with a mingled tropical and temperate vegetation, like that now growing with strange luxuriance at the base of the Himalaya, as graphically described by Hooker.
Thus, as I believe, a considerable number of plants, a few terrestrial animals, and some marine productions, migrated during the Glacial period from the northern and southern temperate zones into the intertropical regions, and some even crossed the equator. As the warmth returned, these temperate forms would naturally ascend the higher mountains, being exterminated on the lowlands; those which had not reached the equator, would re-migrate northward or southward towards their former homes; but the forms, chiefly northern, which had crossed the equator, would travel still further from their homes into the more temperate latitudes of the opposite hemisphere. Although we have reason to believe from geological evidence that the whole body of arctic shells underwent scarcely any modification during their long southern migration and re-migration northward, the case may have been wholly different with those intruding forms which settled themselves on the intertropical mountains, and in the southern hemisphere. These being surrounded by strangers will have had to compete with many new forms of life; and it is probable that selected modifications in their structure, habits, and constitutions will have profited them. Thus many of these wanderers, though still plainly related by inheritance to their brethren of the northern or southern hemispheres, now exist in their new homes as well-marked varieties or as distinct species.
It is a remarkable fact, strongly insisted on by Hooker in regard to America, and by Alph. de Candolle in regard to Australia, that many more identical plants and allied forms have apparently migrated from the north to the south, than in a reversed direction. We see, however, a few southern vegetable forms on the mountains of Borneo and Abyssinia. I suspect that this preponderant migration from north to south is due to the greater extent of land in the north, and to the northern forms having existed in their own homes in greater numbers, and having consequently been advanced through natural selection and competition to a higher stage of perfection or dominating power, than the southern forms. And thus, when they became commingled during the Glacial period, the northern forms were enabled to beat the less powerful southern forms. Just in the same manner as we see at the present day, that very many European productions cover the ground in La Plata, and in a lesser degree in Australia, and have to a certain extent beaten the natives; whereas extremely few southern forms have become naturalised in any part of Europe, though hides, wool, and other objects likely to carry seeds have been largely imported into Europe during the last two or three centuries from La Plata, and during the last thirty or forty years from Australia. Something of the same kind must have occurred on the intertropical mountains: no doubt before the Glacial period they were stocked with endemic Alpine forms; but these have almost everywhere largely yielded to the more dominant forms, generated in the larger areas and more efficient workshops of the north. In many islands the native productions are nearly equalled or even outnumbered by the naturalised; and if the natives have not been actually exterminated, their numbers have been greatly reduced, and this is the first stage towards extinction. A mountain is an island on the land; and the intertropical mountains before the Glacial period must have been completely isolated; and I believe that the productions of these islands on the land yielded to those produced within the larger areas of the north, just in the same way as the productions of real islands have everywhere lately yielded to continental forms, naturalised by man’s agency.
I am far from supposing that all difficulties are removed on the view here given in regard to the range and affinities of the allied species which live in the northern and southern temperate zones and on the mountains of the intertropical regions. Very many difficulties remain to be solved. I do not pretend to indicate the exact lines and means of migration, or the reason why certain species and not others have migrated; why certain species have been modified and have given rise to new groups of forms, and others have remained unaltered. We cannot hope to explain such facts, until we can say why one species and not another becomes naturalised by man’s agency in a foreign land; why one ranges twice or thrice as far, and is twice or thrice as common, as another species within their own homes.
I have said that many difficulties remain to be solved: some of the most remarkable are stated with admirable clearness by Dr. Hooker in his botanical works on the antarctic regions. These cannot be here discussed. I will only say that as far as regards the occurrence of identical species at points so enormously remote as Kerguelen Land, New Zealand, and Fuegia, I believe that towards the close of the Glacial period, icebergs, as suggested by Lyell, have been largely concerned in their dispersal. But the existence of several quite distinct species, belonging to genera exclusively confined to the south, at these and other distant points of the southern hemisphere, is, on my theory of descent with modification, a far more remarkable case of difficulty. For some of these species are so distinct, that we cannot suppose that there has been time since the commencement of the Glacial period for their migration, and for their subsequent modification to the necessary degree. The facts seem to me to indicate that peculiar and very distinct species have migrated in radiating lines from some common centre; and I am inclined to look in the southern, as in the northern hemisphere, to a former and warmer period, before the commencement of the Glacial period, when the antarctic lands, now covered with ice, supported a highly peculiar and isolated flora. I suspect that before this flora was exterminated by the Glacial epoch, a few forms were widely dispersed to various points of the southern hemisphere by occasional means of transport, and by the aid, as halting-places, of existing and now sunken islands, and perhaps at the commencement of the Glacial period, by icebergs. By these means, as I believe, the southern shores of America, Australia, New Zealand have become slightly tinted by the same peculiar forms of vegetable life.
Sir C. Lyell in a striking passage has speculated, in language almost identical with mine, on the effects of great alternations of climate on geographical distribution. I believe that the world has recently felt one of his great cycles of change; and that on this view, combined with modification through natural selection, a multitude of facts in the present distribution both of the same and of allied forms of life can be explained. The living waters may be said to have flowed during one short period from the north and from the south, and to have crossed at the equator; but to have flowed with greater force from the north so as to have freely inundated the south. As the tide leaves its drift in horizontal lines, though rising higher on the shores where the tide rises highest, so have the living waters left their living drift on our mountain-summits, in a line gently rising from the arctic lowlands to a great height under the equator. The various beings thus left stranded may be compared with savage races of man, driven up and surviving in the mountain-fastnesses of almost every land, which serve as a record, full of interest to us, of the former inhabitants of the surrounding lowlands.
CHAPTER 12. GEOGRAPHICAL DISTRIBUTION–continued.
Distribution of fresh-water productions. On the inhabitants of oceanic islands.
Absence of Batrachians and of terrestrial Mammals. On the relation of the inhabitants of islands to those of the nearest mainland.
On colonisation from the nearest source with subsequent modification. Summary of the last and present chapters.
As lakes and river-systems are separated from each other by barriers of land, it might have been thought that fresh-water productions would not have ranged widely within the same country, and as the sea is apparently a still more impassable barrier, that they never would have extended to distant countries. But the case is exactly the reverse. Not only have many fresh-water species, belonging to quite different classes, an enormous range, but allied species prevail in a remarkable manner throughout the world. I well remember, when first collecting in the fresh waters of Brazil, feeling much surprise at the similarity of the fresh-water insects, shells, etc., and at the dissimilarity of the surrounding terrestrial beings, compared with those of Britain.
But this power in fresh-water productions of ranging widely, though so unexpected, can, I think, in most cases be explained by their having become fitted, in a manner highly useful to them, for short and frequent migrations from pond to pond, or from stream to stream; and liability to wide dispersal would follow from this capacity as an almost necessary consequence. We can here consider only a few cases. In regard to fish, I believe that the same species never occur in the fresh waters of distant continents. But on the same continent the species often range widely and almost capriciously; for two river-systems will have some fish in common and some different. A few facts seem to favour the possibility of their occasional transport by accidental means; like that of the live fish not rarely dropped by whirlwinds in India, and the vitality of their ova when removed from the water. But I am inclined to attribute the dispersal of fresh-water fish mainly to slight changes within the recent period in the level of the land, having caused rivers to flow into each other. Instances, also, could be given of this having occurred during floods, without any change of level. We have evidence in the loess of the Rhine of considerable changes of level in the land within a very recent geological period, and when the surface was peopled by existing land and fresh-water shells. The wide difference of the fish on opposite sides of continuous mountain-ranges, which from an early period must have parted river-systems and completely prevented their inosculation, seems to lead to this same conclusion. With respect to allied fresh-water fish occurring at very distant points of the world, no doubt there are many cases which cannot at present be explained: but some fresh-water fish belong to very ancient forms, and in such cases there will have been ample time for great geographical changes, and consequently time and means for much migration. In the second place, salt-water fish can with care be slowly accustomed to live in fresh water; and, according to Valenciennes, there is hardly a single group of fishes confined exclusively to fresh water, so that we may imagine that a marine member of a fresh-water group might travel far along the shores of the sea, and subsequently become modified and adapted to the fresh waters of a distant land.
Some species of fresh-water shells have a very wide range, and allied species, which, on my theory, are descended from a common parent and must have proceeded from a single source, prevail throughout the world. Their distribution at first perplexed me much, as their ova are not likely to be transported by birds, and they are immediately killed by sea water, as are the adults. I could not even understand how some naturalised species have rapidly spread throughout the same country. But two facts, which I have observed–and no doubt many others remain to be observed–throw some light on this subject. When a duck suddenly emerges from a pond covered with duck-weed, I have twice seen these little plants adhering to its back; and it has happened to me, in removing a little duck-weed from one aquarium to another, that I have quite unintentionally stocked the one with fresh-water shells from the other. But another agency is perhaps more effectual: I suspended a duck’s feet, which might represent those of a bird sleeping in a natural pond, in an aquarium, where many ova of fresh-water shells were hatching; and I found that numbers of the extremely minute and just hatched shells crawled on the feet, and clung to them so firmly that when taken out of the water they could not be jarred off, though at a somewhat more advanced age they would voluntarily drop off. These just hatched molluscs, though aquatic in their nature, survived on the duck’s feet, in damp air, from twelve to twenty hours; and in this length of time a duck or heron might fly at least six or seven hundred miles, and would be sure to alight on a pool or rivulet, if blown across sea to an oceanic island or to any other distant point. Sir Charles Lyell also informs me that a Dyticus has been caught with an Ancylus (a fresh-water shell like a limpet) firmly adhering to it; and a water-beetle of the same family, a Colymbetes, once flew on board the ‘Beagle,’ when forty-five miles distant from the nearest land: how much farther it might have flown with a favouring gale no one can tell.
With respect to plants, it has long been known what enormous ranges many fresh-water and even marsh-species have, both over continents and to the most remote oceanic islands. This is strikingly shown, as remarked by Alph. de Candolle, in large groups of terrestrial plants, which have only a very few aquatic members; for these latter seem immediately to acquire, as if in consequence, a very wide range. I think favourable means of dispersal explain this fact. I have before mentioned that earth occasionally, though rarely, adheres in some quantity to the feet and beaks of birds. Wading birds, which frequent the muddy edges of ponds, if suddenly flushed, would be the most likely to have muddy feet. Birds of this order I can show are the greatest wanderers, and are occasionally found on the most remote and barren islands in the open ocean; they would not be likely to alight on the surface of the sea, so that the dirt would not be washed off their feet; when making land, they would be sure to fly to their natural fresh-water haunts. I do not believe that botanists are aware how charged the mud of ponds is with seeds: I have tried several little experiments, but will here give only the most striking case: I took in February three table-spoonfuls of mud from three different points, beneath water, on the edge of a little pond; this mud when dry weighed only 6 3/4 ounces; I kept it covered up in my study for six months, pulling up and counting each plant as it grew; the plants were of many kinds, and were altogether 537 in number; and yet the viscid mud was all contained in a breakfast cup! Considering these facts, I think it would be an inexplicable circumstance if water-birds did not transport the seeds of fresh-water plants to vast distances, and if consequently the range of these plants was not very great. The same agency may have come into play with the eggs of some of the smaller fresh-water animals.
Other and unknown agencies probably have also played a part. I have stated that fresh-water fish eat some kinds of seeds, though they reject many other kinds after having swallowed them; even small fish swallow seeds of moderate size, as of the yellow water-lily and Potamogeton. Herons and other birds, century after century, have gone on daily devouring fish; they then take flight and go to other waters, or are blown across the sea; and we have seen that seeds retain their power of germination, when rejected in pellets or in excrement, many hours afterwards. When I saw the great size of the seeds of that fine water-lily, the Nelumbium, and remembered Alph. de Candolle’s remarks on this plant, I thought that its distribution must remain quite inexplicable; but Audubon states that he found the seeds of the great southern water-lily (probably, according to Dr. Hooker, the Nelumbium luteum) in a heron’s stomach; although I do not know the fact, yet analogy makes me believe that a heron flying to another pond and getting a hearty meal of fish, would probably reject from its stomach a pellet containing the seeds of the Nelumbium undigested; or the seeds might be dropped by the bird whilst feeding its young, in the same way as fish are known sometimes to be dropped.
In considering these several means of distribution, it should be remembered that when a pond or stream is first formed, for instance, on a rising islet, it will be unoccupied; and a single seed or egg will have a good chance of succeeding. Although there will always be a struggle for life between the individuals of the species, however few, already occupying any pond, yet as the number of kinds is small, compared with those on the land, the competition will probably be less severe between aquatic than between terrestrial species; consequently an intruder from the waters of a foreign country, would have a better chance of seizing on a place, than in the case of terrestrial colonists. We should, also, remember that some, perhaps many, fresh-water productions are low in the scale of nature, and that we have reason to believe that such low beings change or become modified less quickly than the high; and this will give longer time than the average for the migration of the same aquatic species. We should not forget the probability of many species having formerly ranged as continuously as fresh-water productions ever can range, over immense areas, and having subsequently become extinct in intermediate regions. But the wide distribution of fresh-water plants and of the lower animals, whether retaining the same identical form or in some degree modified, I believe mainly depends on the wide dispersal of their seeds and eggs by animals, more especially by fresh-water birds, which have large powers of flight, and naturally travel from one to another and often distant piece of water. Nature, like a careful gardener, thus takes her seeds from a bed of a particular nature, and drops them in another equally well fitted for them.
ON THE INHABITANTS OF OCEANIC ISLANDS.
We now come to the last of the three classes of facts, which I have selected as presenting the greatest amount of difficulty, on the view that all the individuals both of the same and of allied species have descended from a single parent; and therefore have all proceeded from a common birthplace, notwithstanding that in the course of time they have come to inhabit distant points of the globe. I have already stated that I cannot honestly admit Forbes’s view on continental extensions, which, if legitimately followed out, would lead to the belief that within the recent period all existing islands have been nearly or quite joined to some continent. This view would remove many difficulties, but it would not, I think, explain all the facts in regard to insular productions. In the following remarks I shall not confine myself to the mere question of dispersal; but shall consider some other facts, which bear on the truth of the two theories of independent creation and of descent with modification.
The species of all kinds which inhabit oceanic islands are few in number compared with those on equal continental areas: Alph. de Candolle admits this for plants, and Wollaston for insects. If we look to the large size and varied stations of New Zealand, extending over 780 miles of latitude, and compare its flowering plants, only 750 in number, with those on an equal area at the Cape of Good Hope or in Australia, we must, I think, admit that something quite independently of any difference in physical conditions has caused so great a difference in number. Even the uniform county of Cambridge has 847 plants, and the little island of Anglesea 764, but a few ferns and a few introduced plants are included in these numbers, and the comparison in some other respects is not quite fair. We have evidence that the barren island of Ascension aboriginally possessed under half-a-dozen flowering plants; yet many have become naturalised on it, as they have on New Zealand and on every other oceanic island which can be named. In St. Helena there is reason to believe that the naturalised plants and animals have nearly or quite exterminated many native productions. He who admits the doctrine of the creation of each separate species, will have to admit, that a sufficient number of the best adapted plants and animals have not been created on oceanic islands; for man has unintentionally stocked them from various sources far more fully and perfectly than has nature.
Although in oceanic islands the number of kinds of inhabitants is scanty, the proportion of endemic species (i.e. those found nowhere else in the world) is often extremely large. If we compare, for instance, the number of the endemic land-shells in Madeira, or of the endemic birds in the Galapagos Archipelago, with the number found on any continent, and then compare the area of the islands with that of the continent, we shall see that this is true. This fact might have been expected on my theory, for, as already explained, species occasionally arriving after long intervals in a new and isolated district, and having to compete with new associates, will be eminently liable to modification, and will often produce groups of modified descendants. But it by no means follows, that, because in an island nearly all the species of one class are peculiar, those of another class, or of another section of the same class, are peculiar; and this difference seems to depend on the species which do not become modified having immigrated with facility and in a body, so that their mutual relations have not been much disturbed. Thus in the Galapagos Islands nearly every land-bird, but only two out of the eleven marine birds, are peculiar; and it is obvious that marine birds could arrive at these islands more easily than land-birds. Bermuda, on the other hand, which lies at about the same distance from North America as the Galapagos Islands do from South America, and which has a very peculiar soil, does not possess one endemic land bird; and we know from Mr. J. M. Jones’s admirable account of Bermuda, that very many North American birds, during their great annual migrations, visit either periodically or occasionally this island. Madeira does not possess one peculiar bird, and many European and African birds are almost every year blown there, as I am informed by Mr. E. V. Harcourt. So that these two islands of Bermuda and Madeira have been stocked by birds, which for long ages have struggled together in their former homes, and have become mutually adapted to each other; and when settled in their new homes, each kind will have been kept by the others to their proper places and habits, and will consequently have been little liable to modification. Madeira, again, is inhabited by a wonderful number of peculiar land-shells, whereas not one species of sea-shell is confined to its shores: now, though we do not know how seashells are dispersed, yet we can see that their eggs or larvae, perhaps attached to seaweed or floating timber, or to the feet of wading-birds, might be transported far more easily than land-shells, across three or four hundred miles of open sea. The different orders of insects in Madeira apparently present analogous facts.
Oceanic islands are sometimes deficient in certain classes, and their places are apparently occupied by the other inhabitants; in the Galapagos Islands reptiles, and in New Zealand gigantic wingless birds, take the place of mammals. In the plants of the Galapagos Islands, Dr. Hooker has shown that the proportional numbers of the different orders are very different from what they are elsewhere. Such cases are generally accounted for by the physical conditions of the islands; but this explanation seems to me not a little doubtful. Facility of immigration, I believe, has been at least as important as the nature of the conditions.
Many remarkable little facts could be given with respect to the inhabitants of remote islands. For instance, in certain islands not tenanted by mammals, some of the endemic plants have beautifully hooked seeds; yet few relations are more striking than the adaptation of hooked seeds for transportal by the wool and fur of quadrupeds. This case presents no difficulty on my view, for a hooked seed might be transported to an island by some other means; and the plant then becoming slightly modified, but still retaining its hooked seeds, would form an endemic species, having as useless an appendage as any rudimentary organ,–for instance, as the shrivelled wings under the soldered elytra of many insular beetles. Again, islands often possess trees or bushes belonging to orders which elsewhere include only herbaceous species; now trees, as Alph. de Candolle has shown, generally have, whatever the cause may be, confined ranges. Hence trees would be little likely to reach distant oceanic islands; and an herbaceous plant, though it would have no chance of successfully competing in stature with a fully developed tree, when established on an island and having to compete with herbaceous plants alone, might readily gain an advantage by growing taller and taller and overtopping the other plants. If so, natural selection would often tend to add to the stature of herbaceous plants when growing on an island, to whatever order they belonged, and thus convert them first into bushes and ultimately into trees.
With respect to the absence of whole orders on oceanic islands, Bory St. Vincent long ago remarked that Batrachians (frogs, toads, newts) have never been found on any of the many islands with which the great oceans are studded. I have taken pains to verify this assertion, and I have found it strictly true. I have, however, been assured that a frog exists on the mountains of the great island of New Zealand; but I suspect that this exception (if the information be correct) may be explained through glacial agency. This general absence of frogs, toads, and newts on so many oceanic islands cannot be accounted for by their physical conditions; indeed it seems that islands are peculiarly well fitted for these animals; for frogs have been introduced into Madeira, the Azores, and Mauritius, and have multiplied so as to become a nuisance. But as these animals and their spawn are known to be immediately killed by sea-water, on my view we can see that there would be great difficulty in their transportal across the sea, and therefore why they do not exist on any oceanic island. But why, on the theory of creation, they should not have been created there, it would be very difficult to explain.
Mammals offer another and similar case. I have carefully searched the oldest voyages, but have not finished my search; as yet I have not found a single instance, free from doubt, of a terrestrial mammal (excluding domesticated animals kept by the natives) inhabiting an island situated above 300 miles from a continent or great continental island; and many islands situated at a much less distance are equally barren. The Falkland Islands, which are inhabited by a wolf-like fox, come nearest to an exception; but this group cannot be considered as oceanic, as it lies on a bank connected with the mainland; moreover, icebergs formerly brought boulders to its western shores, and they may have formerly transported foxes, as so frequently now happens in the arctic regions. Yet it cannot be said that small islands will not support small mammals, for they occur in many parts of the world on very small islands, if close to a continent; and hardly an island can be named on which our smaller quadrupeds have not become naturalised and greatly multiplied. It cannot be said, on the ordinary view of creation, that there has not been time for the creation of mammals; many volcanic islands are sufficiently ancient, as shown by the stupendous degradation which they have suffered and by their tertiary strata: there has also been time for the production of endemic species belonging to other classes; and on continents it is thought that mammals appear and disappear at a quicker rate than other and lower animals. Though terrestrial mammals do not occur on oceanic islands, aerial mammals do occur on almost every island. New Zealand possesses two bats found nowhere else in the world: Norfolk Island, the Viti Archipelago, the Bonin Islands, the Caroline and Marianne Archipelagoes, and Mauritius, all possess their peculiar bats. Why, it may be asked, has the supposed creative force produced bats and no other mammals on remote islands? On my view this question can easily be answered; for no terrestrial mammal can be transported across a wide space of sea, but bats can fly across. Bats have been seen wandering by day far over the Atlantic Ocean; and two North American species either regularly or occasionally visit Bermuda, at the distance of 600 miles from the mainland. I hear from Mr. Tomes, who has specially studied this family, that many of the same species have enormous ranges, and are found on continents and on far distant islands. Hence we have only to suppose that such wandering species have been modified through natural selection in their new homes in relation to their new position, and we can understand the presence of endemic bats on islands, with the absence of all terrestrial mammals.
Besides the absence of terrestrial mammals in relation to the remoteness of islands from continents, there is also a relation, to a certain extent independent of distance, between the depth of the sea separating an island from the neighbouring mainland, and the presence in both of the same mammiferous species or of allied species in a more or less modified condition. Mr. Windsor Earl has made some striking observations on this head in regard to the great Malay Archipelago, which is traversed near Celebes by a space of deep ocean; and this space separates two widely distinct mammalian faunas. On either side the islands are situated on moderately deep submarine banks, and they are inhabited by closely allied or identical quadrupeds. No doubt some few anomalies occur in this great archipelago, and there is much difficulty in forming a judgment in some cases owing to the probable naturalisation of certain mammals through man’s agency; but we shall soon have much light thrown on the natural history of this archipelago by the admirable zeal and researches of Mr. Wallace. I have not as yet had time to follow up this subject in all other quarters of the world; but as far as I have gone, the relation generally holds good. We see Britain separated by a shallow channel from Europe, and the mammals are the same on both sides; we meet with analogous facts on many islands separated by similar channels from Australia. The West Indian Islands stand on a deeply submerged bank, nearly 1000 fathoms in depth, and here we find American forms, but the species and even the genera are distinct. As the amount of modification in all cases depends to a certain degree on the lapse of time, and as during changes of level it is obvious that islands separated by shallow channels are more likely to have been continuously united within a recent period to the mainland than islands separated by deeper channels, we can understand the frequent relation between the depth of the sea and the degree of affinity of the mammalian inhabitants of islands with those of a neighbouring continent,–an inexplicable relation on the view of independent acts of creation.
All the foregoing remarks on the inhabitants of oceanic islands,–namely, the scarcity of kinds–the richness in endemic forms in particular classes or sections of classes,–the absence of whole groups, as of batrachians, and of terrestrial mammals notwithstanding the presence of aerial bats,–the singular proportions of certain orders of plants,–herbaceous forms having been developed into trees, etc.,–seem to me to accord better with the view of occasional means of transport having been largely efficient in the long course of time, than with the view of all our oceanic islands having been formerly connected by continuous land with the nearest continent; for on this latter view the migration would probably have been more complete; and if modification be admitted, all the forms of life would have been more equally modified, in accordance with the paramount importance of the relation of organism to organism.
I do not deny that there are many and grave difficulties in understanding how several of the inhabitants of the more remote islands, whether still retaining the same specific form or modified since their arrival, could have reached their present homes. But the probability of many islands having existed as halting-places, of which not a wreck now remains, must not be overlooked. I will here give a single instance of one of the cases of difficulty. Almost all oceanic islands, even the most isolated and smallest, are inhabited by land-shells, generally by endemic species, but sometimes by species found elsewhere. Dr. Aug. A. Gould has given several interesting cases in regard to the land-shells of the islands of the Pacific. Now it is notorious that land-shells are very easily killed by salt; their eggs, at least such as I have tried, sink in sea-water and are killed by it. Yet there must be, on my view, some unknown, but highly efficient means for their transportal. Would the just-hatched young occasionally crawl on and adhere to the feet of birds roosting on the ground, and thus get transported? It occurred to me that land-shells, when hybernating and having a membranous diaphragm over the mouth of the shell, might be floated in chinks of drifted timber across moderately wide arms of the sea. And I found that several species did in this state withstand uninjured an immersion in sea-water during seven days: one of these shells was the Helix pomatia, and after it had again hybernated I put it in sea-water for twenty days, and it perfectly recovered. As this species has a thick calcareous operculum, I removed it, and when it had formed a new membranous one, I immersed it for fourteen days in sea-water, and it recovered and crawled away: but more experiments are wanted on this head. The most striking and important fact for us in regard to the inhabitants of islands, is their affinity to those of the nearest mainland, without being actually the same species. Numerous instances could be given of this fact. I will give only one, that of the Galapagos Archipelago, situated under the equator, between 500 and 600 miles from the shores of South America. Here almost every product of the land and water bears the unmistakeable stamp of the American continent. There are twenty-six land birds, and twenty-five of these are ranked by Mr. Gould as distinct species, supposed to have been created here; yet the close affinity of most of these birds to American species in every character, in their habits, gestures, and tones of voice, was manifest. So it is with the other animals, and with nearly all the plants, as shown by Dr. Hooker in his admirable memoir on the Flora of this archipelago. The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, yet feels that he is standing on American land. Why should this be so? why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plain a stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which resembles closely the conditions of the South American coast: in fact there is a considerable dissimilarity in all these respects. On the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in climate, height, and size of the islands, between the Galapagos and Cape de Verde Archipelagos: but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape de Verde Islands are related to those of Africa, like those of the Galapagos to America. I believe this grand fact can receive no sort of explanation on the ordinary view of independent creation; whereas on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists, whether by occasional means of transport or by formerly continuous land, from America; and the Cape de Verde Islands from Africa; and that such colonists would be liable to modification;–the principle of inheritance still betraying their original birthplace.
Many analogous facts could be given: indeed it is an almost universal rule that the endemic productions of islands are related to those of the nearest continent, or of other near islands. The exceptions are few, and most of them can be explained. Thus the plants of Kerguelen Land, though standing nearer to Africa than to America, are related, and that very closely, as we know from Dr. Hooker’s account, to those of America: but on the view that this island has been mainly stocked by seeds brought with earth and stones on icebergs, drifted by the prevailing currents, this anomaly disappears. New Zealand in its endemic plants is much more closely related to Australia, the nearest mainland, than to any other region: and this is what might have been expected; but it is also plainly related to South America, which, although the next nearest continent, is so enormously remote, that the fact becomes an anomaly. But this difficulty almost disappears on the view that both New Zealand, South America, and other southern lands were long ago partially stocked from a nearly intermediate though distant point, namely from the antarctic islands, when they were clothed with vegetation, before the commencement of the Glacial period. The affinity, which, though feeble, I am assured by Dr. Hooker is real, between the flora of the south-western corner of Australia and of the Cape of Good Hope, is a far more remarkable case, and is at present inexplicable: but this affinity is confined to the plants, and will, I do not doubt, be some day explained.
The law which causes the inhabitants of an archipelago, though specifically distinct, to be closely allied to those of the nearest continent, we sometimes see displayed on a small scale, yet in a most interesting manner, within the limits of the same archipelago. Thus the several islands of the Galapagos Archipelago are tenanted, as I have elsewhere shown, in a quite marvellous manner, by very closely related species; so that the inhabitants of each separate island, though mostly distinct, are related in an incomparably closer degree to each other than to the inhabitants of any other part of the world. And this is just what might have been expected on my view, for the islands are situated so near each other that they would almost certainly receive immigrants from the same original source, or from each other. But this dissimilarity between the endemic inhabitants of the islands may be used as an argument against my views; for it may be asked, how has it happened in the several islands situated within sight of each other, having the same geological nature, the same