The Foundations of the Origin of Species

What evidence311 is there of a number of intermediate forms having existed, making a passage in the above sense, between the species of the same groups? Some naturalists have supposed that if every fossil which now lies entombed, together with all existing species, were collected together, a perfect series in every great class would be formed. Considering the enormous number of species requisite to effect this, especially in the above sense of the forms not being directly intermediate between the existing species and genera, but only intermediate by being linked through a common but often widely different ancestor, I think this supposition highly improbable. I am however far from underrating the probable number of fossilised species: no one who has attended to the wonderful progress of palæontology during the last few years will doubt that we as yet have found only an exceedingly small fraction of the species buried in the crust of the earth. Although the almost infinitely numerous intermediate forms in no one class may have been preserved, it does not follow that they have not existed. The fossils which have been discovered, it is important to remark, do tend, the little way they go, to make good the series; for as observed by Buckland they all fall into or between existing groups312. Moreover, those that fall between our existing groups, fall in, according to the manner required by our theory, for they do not directly connect two existing species of different groups, but they connect the groups themselves: thus the Pachydermata and Ruminantia are now separated by several characters, «for instance» the Pachydermata313 have both a tibia and fibula, whilst Ruminantia have only a tibia; now the fossil Macrauchenia has a leg bone exactly intermediate in this respect, and likewise has some other intermediate characters. But the Macrauchenia does not connect any one species of Pachydermata with some one other of Ruminantia but it shows that these two groups have at one time been less widely divided. So have fish and reptiles been at one time more closely connected in some points than they now are. Generally in those groups in which there has been most change, the more ancient the fossil, if not identical with recent, the more often it falls between existing groups, or into small existing groups which now lie between other large existing groups. Cases like the foregoing, of which there are many, form steps, though few and far between, in a series of the kind required by my theory.

As I have admitted the high improbability, that if every fossil were disinterred, they would compose in each of the Divisions of Nature a perfect series of the kind required; consequently I freely admit, that if those geologists are in the right who consider the lowest known formation as contemporaneous with the first appearances of life314; or the several formations as at all closely consecutive; or any one formation as containing a nearly perfect record of the organisms which existed during the whole period of its deposition in that quarter of the globe; – if such propositions are to be accepted, my theory must be abandoned.

If the Palæozoic system is really contemporaneous with the first appearance of life, my theory must be abandoned, both inasmuch as it limits from shortness of time the total number of forms which can have existed on this world, and because the organisms, as fish, mollusca315 and star-fish found in its lower beds, cannot be considered as the parent forms of all the successive species in these classes. But no one has yet overturned the arguments of Hutton and Lyell, that the lowest formations known to us are only those which have escaped being metamorphosed «illegible»; if we argued from some considerable districts, we might have supposed that even the Cretaceous system was that in which life first appeared. From the number of distant points, however, in which the Silurian system has been found to be the lowest, and not always metamorphosed, there are some objections to Hutton’s and Lyell’s view; but we must not forget that the now existing land forms only 1/5 part of the superficies of the globe, and that this fraction is only imperfectly known. With respect to the fewness of the organisms found in the Silurian and other Palæozoic formations, there is less difficulty, inasmuch as (besides their gradual obliteration) we can expect formations of this vast antiquity to escape entire denudation, only when they have been accumulated over a wide area, and have been subsequently protected by vast superimposed deposits: now this could generally only hold good with deposits accumulating in a wide and deep ocean, and therefore unfavourable to the presence of many living things. A mere narrow and not very thick strip of matter, deposited along a coast where organisms most abound, would have no chance of escaping denudation and being preserved to the present time from such immensely distant ages316.

If the several known formations are at all nearly consecutive in time, and preserve a fair record of the organisms which have existed, my theory must be abandoned. But when we consider the great changes in mineralogical nature and texture between successive formations, what vast and entire changes in the geography of the surrounding countries must generally have been effected, thus wholly to have changed the nature of the deposits on the same area. What time such changes must have required! Moreover how often has it not been found, that between two conformable and apparently immediately successive deposits a vast pile of water-worn matter is interpolated in an adjoining district. We have no means of conjecturing in many cases how long a period317 has elapsed between successive formations, for the species are often wholly different: as remarked by Lyell, in some cases probably as long a period has elapsed between two formations as the whole Tertiary system, itself broken by wide gaps.

Consult the writings of any one who has particularly attended to any one stage in the Tertiary system (and indeed of every system) and see how deeply impressed he is with the time required for its accumulation318. Reflect on the years elapsed in many cases, since the latest beds containing only living species have been formed; – see what Jordan Smith says of the 20,000 years since the last bed, which is above the boulder formation in Scotland, has been upraised; or of the far longer period since the recent beds of Sweden have been upraised 400 feet, what an enormous period the boulder formation must have required, and yet how insignificant are the records (although there has been plenty of elevation to bring up submarine deposits) of the shells, which we know existed at that time. Think, then, over the entire length of the Tertiary epoch, and think over the probable length of the intervals, separating the Secondary deposits. Of these deposits, moreover, those consisting of sand and pebbles have seldom been favourable, either to the embedment or to the preservation of fossils319.

Nor can it be admitted as probable that any one Secondary formation contains a fair record even of those organisms which are most easily preserved, namely hard marine bodies. In how many cases have we not certain evidence that between the deposition of apparently closely consecutive beds, the lower one existed for an unknown time as land, covered with trees. Some of the Secondary formations which contain most marine remains appear to have been formed in a wide and not deep sea, and therefore only those marine animals which live in such situations would be preserved320. In all cases, on indented rocky coasts, or any other coast, where sediment is not accumulating, although often highly favourable to marine animals, none can be embedded: where pure sand and pebbles are accumulating few or none will be preserved. I may here instance the great western line of the S. American coast321, tenanted by many peculiar animals, of which none probably will be preserved to a distant epoch. From these causes, and especially from such deposits as are formed along a line of coast, steep above and below water, being necessarily of little width, and therefore more likely to be subsequently denuded and worn away, we can see why it is improbable that our Secondary deposits contain a fair record of the Marine Fauna of any one period. The East Indian Archipelago offers an area, as large as most of our Secondary deposits, in which there are wide and shallow seas, teeming with marine animals, and in which sediment is accumulating; now supposing that all the hard marine animals, or rather those having hard parts to preserve, were preserved to a future age, excepting those which lived on rocky shores where no sediment or only sand and gravel were accumulating, and excepting those embedded along the steeper coasts, where only a narrow fringe of sediment was accumulating, supposing all this, how poor a notion would a person at a future age have of the Marine Fauna of the present day. Lyell322 has compared the geological series to a work of which only the few latter but not consecutive chapters have been preserved; and out of which, it may be added, very many leaves have been torn, the remaining ones only illustrating a scanty portion of the Fauna of each period. On this view, the records of anteceding ages confirm my theory; on any other they destroy it.

Finally, if we narrow the question into, why do we not find in some instances every intermediate form between any two species? the answer may well be that the average duration of each specific form (as we have good reason to believe) is immense in years, and that the transition could, according to my theory, be effected only by numberless small gradations; and therefore that we should require for this end a most perfect record, which the foregoing reasoning teaches us not to expect. It might be thought that in a vertical section of great thickness in the same formation some of the species ought to be found to vary in the upper and lower parts323, but it may be doubted whether any formation has gone on accumulating without any break for a period as long as the duration of a species; and if it had done so, we should require a series of specimens from every part. How rare must be the chance of sediment accumulating for some 20 or 30 thousand years on the same spot324, with the bottom subsiding, so that a proper depth might be preserved for any one species to continue living: what an amount of subsidence would be thus required, and this subsidence must not destroy the source whence the sediment continued to be derived. In the case of terrestrial animals, what chance is there when the present time is become a pleistocene formation (at an earlier period than this, sufficient elevation to expose marine beds could not be expected), what chance is there that future geologists will make out the innumerable transitional sub-varieties, through which the short-horned and long-horned cattle (so different in shape of body) have been derived from the same parent stock325? Yet this transition has been effected in the same country, and in a far shorter time, than would be probable in a wild state, both contingencies highly favourable for the future hypothetical geologists being enabled to trace the variation.

CHAPTER V

GRADUAL APPEARANCE AND DISAPPEARANCE OF SPECIES 326

In the Tertiary system, in the last uplifted beds, we find all the species recent and living in the immediate vicinity; in rather older beds we find only recent species, but some not living in the immediate vicinity327; we then find beds with two or three or a few more extinct or very rare species; then considerably more extinct species, but with gaps in the regular increase; and finally we have beds with only two or three or not one living species. Most geologists believe that the gaps in the percentage, that is the sudden increments, in the number of the extinct species in the stages of the Tertiary system are due to the imperfection of the geological record. Hence we are led to believe that the species in the Tertiary system have been gradually introduced; and from analogy to carry on the same view to the Secondary formations. In these latter, however, entire groups of species generally come in abruptly; but this would naturally result, if, as argued in the foregoing chapter, these Secondary deposits are separated by wide epochs. Moreover it is important to observe that, with our increase of knowledge, the gaps between the older formations become fewer and smaller; geologists of a few years standing remember how beautifully has the Devonian system328 come in between the Carboniferous and Silurian formations. I need hardly observe that the slow and gradual appearance of new forms follows from our theory, for to form a new species, an old one must not only be plastic in its organization, becoming so probably from changes in the conditions of its existence, but a place in the natural economy of the district must [be made,] come to exist, for the selection of some new modification of its structure, better fitted to the surrounding conditions than are the other individuals of the same or other species329.

In the Tertiary system the same facts, which make us admit as probable that new species have slowly appeared, lead to the admission that old ones have slowly disappeared, not several together, but one after another; and by analogy one is induced to extend this belief to the Secondary and Palæozoic epochs. In some cases, as the subsidence of a flat country, or the breaking or the joining of an isthmus, and the sudden inroad of many new and destructive species, extinction might be locally sudden. The view entertained by many geologists, that each fauna of each Secondary epoch has been suddenly destroyed over the whole world, so that no succession could be left for the production of new forms, is subversive of my theory, but I see no grounds whatever to admit such a view. On the contrary, the law, which has been made out, with reference to distinct epochs, by independent observers, namely, that the wider the geographical range of a species the longer is its duration in time, seems entirely opposed to any universal extermination330. The fact of species of mammiferous animals and fish being renewed at a quicker rate than mollusca, though both aquatic; and of these the terrestrial genera being renewed quicker than the marine; and the marine mollusca being again renewed quicker than the Infusorial animalcula, all seem to show that the extinction and renewal of species does not depend on general catastrophes, but on the particular relations of the several classes to the conditions to which they are exposed331.

Some authors seem to consider the fact of a few species having survived332 amidst a number of extinct forms (as is the case with a tortoise and a crocodile out of the vast number of extinct sub-Himalayan fossils) as strongly opposed to the view of species being mutable. No doubt this would be the case, if it were presupposed with Lamarck that there was some inherent tendency to change and development in all species, for which supposition I see no evidence. As we see some species at present adapted to a wide range of conditions, so we may suppose that such species would survive unchanged and unexterminated for a long time; time generally being from geological causes a correlative of changing conditions. How at present one species becomes adapted to a wide range, and another species to a restricted range of conditions, is of difficult explanation.

Extinction of species

The extinction of the larger quadrupeds, of which we imagine we better know the conditions of existence, has been thought little less wonderful than the appearance of new species; and has, I think, chiefly led to the belief of universal catastrophes. When considering the wonderful disappearance within a late period, whilst recent shells were living, of the numerous great and small mammifers of S. America, one is strongly induced to join with the catastrophists. I believe, however, that very erroneous views are held on this subject. As far as is historically known, the disappearance of species from any one country has been slow – the species becoming rarer and rarer, locally extinct, and finally lost333. It may be objected that this has been effected by man’s direct agency, or by his indirect agency in altering the state of the country; in this latter case, however, it would be difficult to draw any just distinction between his agency and natural agencies. But we now know in the later Tertiary deposits, that shells become rarer and rarer in the successive beds, and finally disappear: it has happened, also, that shells common in a fossil state, and thought to have been extinct, have been found to be still living species, but very rare ones334. If the rule is that organisms become extinct by becoming rarer and rarer, we ought not to view their extinction, even in the case of the larger quadrupeds, as anything wonderful and out of the common course of events. For no naturalist thinks it wonderful that one species of a genus should be rare and another abundant, notwithstanding he be quite incapable of explaining the causes of the comparative rareness335. Why is one species of willow-wren or hawk or woodpecker common in England, and another extremely rare: why at the Cape of Good Hope is one species of rhinoceros or antelope far more abundant than other species? Why again is the same species much more abundant in one district of a country than in another district? No doubt there are in each case good causes: but they are unknown and unperceived by us. May we not then safely infer that as certain causes are acting unperceived around us, and are making one species to be common and another exceedingly rare, that they might equally well cause the final extinction of some species without being perceived by us? We should always bear in mind that there is a recurrent struggle for life in every organism, and that in every country a destroying agency is always counteracting the geometrical tendency to increase in every species; and yet without our being able to tell with certainty at what period of life, or at what period of the year, the destruction falls the heaviest. Ought we then to expect to trace the steps by which this destroying power, always at work and scarcely perceived by us, becomes increased, and yet if it continues to increase ever so slowly (without the fertility of the species in question be likewise increased) the average number of the individuals of that species must decrease, and become finally lost. I may give a single instance of a check causing local extermination which might long have escaped discovery336; the horse, though swarming in a wild state in La Plata, and likewise under apparently the most unfavourable conditions in the scorched and alternately flooded plains of Caraccas, will not in a wild state extend beyond a certain degree of latitude into the intermediate country of Paraguay; this is owing to a certain fly depositing its eggs on the navels of the foals: as, however, man with a little care can rear horses in a tame state abundantly in Paraguay, the problem of its extinction is probably complicated by the greater exposure of the wild horse to occasional famine from the droughts, to the attacks of the jaguar and other such evils. In the Falkland Islands the check to the increase of the wild horse is said to be loss of the sucking foals337, from the stallions compelling the mares to travel across bogs and rocks in search of food: if the pasture on these islands decreased a little, the horse, perhaps, would cease to exist in a wild state, not from the absolute want of food, but from the impatience of the stallions urging the mares to travel whilst the foals were too young.

From our more intimate acquaintance with domestic animals, we cannot conceive their extinction without some glaring agency; we forget that they would undoubtedly in a state of nature (where other animals are ready to fill up their place) be acted on in some part of their lives by a destroying agency, keeping their numbers on an average constant. If the common ox was known only as a wild S. African species, we should feel no surprise at hearing that it was a very rare species; and this rarity would be a stage towards its extinction. Even in man, so infinitely better known than any other inhabitant of this world, how impossible it has been found, without statistical calculations, to judge of the proportions of births and deaths, of the duration of life, and of the increase and decrease of population; and still less of the causes of such changes: and yet, as has so often been repeated, decrease in numbers or rarity seems to be the high-road to extinction. To marvel at the extermination of a species appears to me to be the same thing as to know that illness is the road to death, – to look at illness as an ordinary event, nevertheless to conclude, when the sick man dies, that his death has been caused by some unknown and violent agency338.

In a future part of this work we shall show that, as a general rule, groups of allied species339 gradually appear and disappear, one after the other, on the face of the earth, like the individuals of the same species: and we shall then endeavour to show the probable cause of this remarkable fact.

CHAPTER VI

ON THE GEOGRAPHICAL DISTRIBUTION OF ORGANIC BEINGS IN PAST AND PRESENT TIMES

For convenience sake I shall divide this chapter into three sections340. In the first place I shall endeavour to state the laws of the distribution of existing beings, as far as our present object is concerned; in the second, that of extinct; and in the third section I shall consider how far these laws accord with the theory of allied species having a common descent.

Section First

Distribution of the inhabitants in the different continents

In the following discussion I shall chiefly refer to terrestrial mammifers, inasmuch as they are better known; their differences in different countries, strongly marked; and especially as the necessary means of their transport are more evident, and confusion, from the accidental conveyance by man of a species from one district to another district, is less likely to arise. It is known that all mammifers (as well as all other organisms) are united in one great system; but that the different species, genera, or families of the same order inhabit different quarters of the globe. If we divide the land341 into two divisions, according to the amount of difference, and disregarding the numbers of the terrestrial mammifers inhabiting them, we shall have first Australia including New Guinea; and secondly the rest of the world: if we make a three-fold division, we shall have Australia, S. America, and the rest of the world; I must observe that North America is in some respects neutral land, from possessing some S. American forms, but I believe it is more closely allied (as it certainly is in its birds, plants and shells) with Europe. If our division had been four-fold, we should have had Australia, S. America, Madagascar (though inhabited by few mammifers) and the remaining land: if five-fold, Africa, especially the southern eastern parts, would have to be separated from the remainder of the world. These differences in the mammiferous inhabitants of the several main divisions of the globe cannot, it is well known, be explained by corresponding differences in their conditions342; how similar are parts of tropical America and Africa; and accordingly we find some analogous resemblances, – thus both have monkeys, both large feline animals, both large Lepidoptera, and large dung-feeding beetles; both have palms and epiphytes; and yet the essential difference between their productions is as great as between those of the arid plains of the Cape of Good Hope and the grass-covered savannahs of La Plata343. Consider the distribution of the Marsupialia, which are eminently characteristic of Australia, and in a lesser degree of S. America; when we reflect that animals of this division, feeding both on animal and vegetable matter, frequent the dry open or wooded plains and mountains of Australia, the humid impenetrable forests of New Guinea and Brazil; the dry rocky mountains of Chile, and the grassy plains of Banda Oriental, we must look to some other cause, than the nature of the country, for their absence in Africa and other quarters of the world.

Furthermore it may be observed that all the organisms inhabiting any country are not perfectly adapted to it344; I mean by not being perfectly adapted, only that some few other organisms can generally be found better adapted to the country than some of the aborigines. We must admit this when we consider the enormous number of horses and cattle which have run wild during the three last centuries in the uninhabited parts of St Domingo, Cuba, and S. America; for these animals must have supplanted some aboriginal ones. I might also adduce the same fact in Australia, but perhaps it will be objected that 30 or 40 years has not been a sufficient period to test this power of struggling «with» and overcoming the aborigines. We know the European mouse is driving before it that of New Zealand, like the Norway rat has driven before it the old English species in England. Scarcely an island can be named, where casually introduced plants have not supplanted some of the native species: in La Plata the Cardoon covers square leagues of country on which some S. American plants must once have grown: the commonest weed over the whole of India is an introduced Mexican poppy. The geologist who knows that slow changes are in progress, replacing land and water, will easily perceive that even if all the organisms of any country had originally been the best adapted to it, this could hardly continue so during succeeding ages without either extermination, or changes, first in the relative proportional numbers of the inhabitants of the country, and finally in their constitutions and structure.