The Origin of Species by Means of Natural Selection

This greater variability in mongrels than in hybrids does not seem at all surprising. For the parents of mongrels are varieties, and mostly domestic varieties (very few experiments having been tried on natural varieties), and this implies that there has been recent variability; which would often continue and would augment that arising from the act of crossing. The slight variability of hybrids in the first generation, in contrast with that in the succeeding generations, is a curious fact and deserves attention. For it bears on the view which I have taken of one of the causes of ordinary variability; namely, that the reproductive system, from being eminently sensitive to changed conditions of life, fails under these circumstances to perform its proper function of producing offspring closely similar in all respects to the parent-form. Now, hybrids in the first generation are descended from species (excluding those long cultivated) which have not had their reproductive systems in any way affected, and they are not variable; but hybrids themselves have their reproductive systems seriously affected, and their descendants are highly variable.

But to return to our comparison of mongrels and hybrids: Gartner states that mongrels are more liable than hybrids to revert to either parent form; but this, if it be true, is certainly only a difference in degree. Moreover, Gartner expressly states that the hybrids from long cultivated plants are more subject to reversion than hybrids from species in their natural state; and this probably explains the singular difference in the results arrived at by different observers. Thus Max Wichura doubts whether hybrids ever revert to their parent forms, and he experimented on uncultivated species of willows, while Naudin, on the other hand, insists in the strongest terms on the almost universal tendency to reversion in hybrids, and he experimented chiefly on cultivated plants. Gartner further states that when any two species, although most closely allied to each other, are crossed with a third species, the hybrids are widely different from each other; whereas if two very distinct varieties of one species are crossed with another species, the hybrids do not differ much. But this conclusion, as far as I can make out, is founded on a single experiment; and seems directly opposed to the results of several experiments made by Kolreuter.

Such alone are the unimportant differences which Gartner is able to point out between hybrid and mongrel plants. On the other hand, the degrees and kinds of resemblance in mongrels and in hybrids to their respective parents, more especially in hybrids produced from nearly related species, follow, according to Gartner the same laws. When two species are crossed, one has sometimes a prepotent power of impressing its likeness on the hybrid. So I believe it to be with varieties of plants; and with animals, one variety certainly often has this prepotent power over another variety. Hybrid plants produced from a reciprocal cross generally resemble each other closely, and so it is with mongrel plants from a reciprocal cross. Both hybrids and mongrels can be reduced to either pure parent form, by repeated crosses in successive generations with either parent.

These several remarks are apparently applicable to animals; but the subject is here much complicated, partly owing to the existence of secondary sexual characters; but more especially owing to prepotency in transmitting likeness running more strongly in one sex than in the other, both when one species is crossed with another and when one variety is crossed with another variety. For instance, I think those authors are right who maintain that the ass has a prepotent power over the horse, so that both the mule and the hinny resemble more closely the ass than the horse; but that the prepotency runs more strongly in the male than in the female ass, so that the mule, which is an offspring of the male ass and mare, is more like an ass than is the hinny, which is the offspring of the female-ass and stallion.

Much stress has been laid by some authors on the supposed fact, that it is only with mongrels that the offspring are not intermediate in character, but closely resemble one of their parents; but this does sometimes occur with hybrids, yet I grant much less frequently than with mongrels. Looking to the cases which I have collected of cross-bred animals closely resembling one parent, the resemblances seem chiefly confined to characters almost monstrous in their nature, and which have suddenly appeared – such as albinism, melanism, deficiency of tail or horns, or additional fingers and toes; and do not relate to characters which have been slowly acquired through selection. A tendency to sudden reversions to the perfect character of either parent would, also, be much more likely to occur with mongrels, which are descended from varieties often suddenly produced and semi-monstrous in character, than with hybrids, which are descended from species slowly and naturally produced. On the whole, I entirely agree with Dr. Prosper Lucas, who, after arranging an enormous body of facts with respect to animals, comes to the conclusion that the laws of resemblance of the child to its parents are the same, whether the two parents differ little or much from each other, namely, in the union of individuals of the same variety, or of different varieties, or of distinct species.

Independently of the question of fertility and sterility, in all other respects there seems to be a general and close similarity in the offspring of crossed species, and of crossed varieties. If we look at species as having been specially created, and at varieties as having been produced by secondary laws, this similarity would be an astonishing fact. But it harmonises perfectly with the view that there is no essential distinction between species and varieties.

SUMMARY OF CHAPTER.

First crosses between forms, sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the most careful experimentalists have arrived at diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible to action of favourable and unfavourable conditions. The degree of sterility does not strictly follow systematic affinity, but is governed by several curious and complex laws. It is generally different, and sometimes widely different in reciprocal crosses between the same two species. It is not always equal in degree in a first cross and in the hybrids produced from this cross.

In the same manner as in grafting trees, the capacity in one species or variety to take on another, is incidental on differences, generally of an unknown nature, in their vegetative systems, so in crossing, the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. There is no more reason to think that species have been specially endowed with various degrees of sterility to prevent their crossing and blending in nature, than to think that trees have been specially endowed with various and somewhat analogous degrees of difficulty in being grafted together in order to prevent their inarching in our forests.

The sterility of first crosses and of their hybrid progeny has not been acquired through natural selection. In the case of first crosses it seems to depend on several circumstances; in some instances in chief part on the early death of the embryo. In the case of hybrids, it apparently depends on their whole organisation having been disturbed by being compounded from two distinct forms; the sterility being closely allied to that which so frequently affects pure species, when exposed to new and unnatural conditions of life. He who will explain these latter cases will be able to explain the sterility of hybrids. This view is strongly supported by a parallelism of another kind: namely, that, firstly, slight changes in the conditions of life add to the vigour and fertility of all organic beings; and secondly, that the crossing of forms, which have been exposed to slightly different conditions of life, or which have varied, favours the size, vigour and fertility of their offspring. The facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny, perhaps render it probable that some unknown bond in all cases connects the degree of fertility of first unions with that of their offspring. The consideration of these facts on dimorphism, as well as of the results of reciprocal crosses, clearly leads to the conclusion that the primary cause of the sterility of crossed species is confined to differences in their sexual elements. But why, in the case of distinct species, the sexual elements should so generally have become more or less modified, leading to their mutual infertility, we do not know; but it seems to stand in some close relation to species having been exposed for long periods of time to nearly uniform conditions of life.

It is not surprising that the difficulty in crossing any two species, and the sterility of their hybrid offspring, should in most cases correspond, even if due to distinct causes: for both depend on the amount of difference between the species which are crossed. Nor is it surprising that the facility of effecting a first cross, and the fertility of the hybrids thus produced, and the capacity of being grafted together – though this latter capacity evidently depends on widely different circumstances – should all run, to a certain extent, parallel with the systematic affinity of the forms subjected to experiment; for systematic affinity includes resemblances of all kinds.

First crosses between forms known to be varieties, or sufficiently alike to be considered as varieties, and their mongrel offspring, are very generally, but not, as is so often stated, invariably fertile. Nor is this almost universal and perfect fertility surprising, when it is remembered how liable we are to argue in a circle with respect to varieties in a state of nature; and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences, and that they have not been long exposed to uniform conditions of life. It should also be especially kept in mind, that long-continued domestication tends to eliminate sterility, and is therefore little likely to induce this same quality. Independently of the question of fertility, in all other respects there is the closest general resemblance between hybrids and mongrels, in their variability, in their power of absorbing each other by repeated crosses, and in their inheritance of characters from both parent-forms. Finally, then, although we are as ignorant of the precise cause of the sterility of first crosses and of hybrids as we are why animals and plants removed from their natural conditions become sterile, yet the facts given in this chapter do not seem to me opposed to the belief that species aboriginally existed as varieties.

CHAPTER X. ON THE IMPERFECTION OF THE GEOLOGICAL RECORD

On the absence of intermediate varieties at the present day – On the nature of extinct intermediate varieties; on their number – On the lapse of time, as inferred from the rate of denudation and of deposition number – On the lapse of time as estimated by years – On the poorness of our palaeontological collections – On the intermittence of geological formations – On the denudation of granitic areas – On the absence of intermediate varieties in any one formation – On the sudden appearance of groups of species – On their sudden appearance in the lowest known fossiliferous strata – Antiquity of the habitable earth.

In the sixth chapter I enumerated the chief objections which might be justly urged against the views maintained in this volume. Most of them have now been discussed. One, namely, the distinctness of specific forms and their not being blended together by innumerable transitional links, is a very obvious difficulty. I assigned reasons why such links do not commonly occur at the present day under the circumstances apparently most favourable for their presence, namely, on an extensive and continuous area with graduated physical conditions. I endeavoured to show, that the life of each species depends in a more important manner on the presence of other already defined organic forms, than on climate, and, therefore, that the really governing conditions of life do not graduate away quite insensibly like heat or moisture. I endeavoured, also, to show that intermediate varieties, from existing in lesser numbers than the forms which they connect, will generally be beaten out and exterminated during the course of further modification and improvement. The main cause, however, of innumerable intermediate links not now occurring everywhere throughout nature depends, on the very process of natural selection, through which new varieties continually take the places of and supplant their parent-forms. But just in proportion as this process of extermination has acted on an enormous scale, so must the number of intermediate varieties, which have formerly existed, be truly enormous. Why then is not every geological formation and every stratum full of such intermediate links? Geology assuredly does not reveal any such finely graduated organic chain; and this, perhaps, is the most obvious and serious objection which can be urged against my theory. The explanation lies, as I believe, in the extreme imperfection of the geological record.

In the first place, it should always be borne in mind what sort of intermediate forms must, on the theory, have formerly existed. I have found it difficult, when looking at any two species, to avoid picturing to myself forms DIRECTLY intermediate between them. But this is a wholly false view; we should always look for forms intermediate between each species and a common but unknown progenitor; and the progenitor will generally have differed in some respects from all its modified descendants. To give a simple illustration: the fantail and pouter pigeons are both descended from the rock-pigeon; if we possessed all the intermediate varieties which have ever existed, we should have an extremely close series between both and the rock-pigeon; but we should have no varieties directly intermediate between the fantail and pouter; none, for instance, combining a tail somewhat expanded with a crop somewhat enlarged, the characteristic features of these two breeds. These two breeds, moreover, have become so much modified, that, if we had no historical or indirect evidence regarding their origin, it would not have been possible to have determined from a mere comparison of their structure with that of the rock-pigeon, C. livia, whether they had descended from this species or from some other allied species, such as C. oenas.

So with natural species, if we look to forms very distinct, for instance to the horse and tapir, we have no reason to suppose that links directly intermediate between them ever existed, but between each and an unknown common parent. The common parent will have had in its whole organisation much general resemblance to the tapir and to the horse; but in some points of structure may have differed considerably from both, even perhaps more than they differ from each other. Hence, in all such cases, we should be unable to recognise the parent-form of any two or more species, even if we closely compared the structure of the parent with that of its modified descendants, unless at the same time we had a nearly perfect chain of the intermediate links.

It is just possible, by the theory, that one of two living forms might have descended from the other; for instance, a horse from a tapir; and in this case DIRECT intermediate links will have existed between them. But such a case would imply that one form had remained for a very long period unaltered, whilst its descendants had undergone a vast amount of change; and the principle of competition between organism and organism, between child and parent, will render this a very rare event; for in all cases the new and improved forms of life tend to supplant the old and unimproved forms.

By the theory of natural selection all living species have been connected with the parent-species of each genus, by differences not greater than we see between the natural and domestic varieties of the same species at the present day; and these parent-species, now generally extinct, have in their turn been similarly connected with more ancient forms; and so on backwards, always converging to the common ancestor of each great class. So that the number of intermediate and transitional links, between all living and extinct species, must have been inconceivably great. But assuredly, if this theory be true, such have lived upon the earth.

ON THE LAPSE OF TIME, AS INFERRED FROM THE RATE OF DEPOSITION AND EXTENT OF DENUDATION.

Independently of our not finding fossil remains of such infinitely numerous connecting links, it may be objected that time cannot have sufficed for so great an amount of organic change, all changes having been effected slowly. It is hardly possible for me to recall to the reader who is not a practical geologist, the facts leading the mind feebly to comprehend the lapse of time. He who can read Sir Charles Lyell's grand work on the Principles of Geology, which the future historian will recognise as having produced a revolution in natural science, and yet does not admit how vast have been the past periods of time, may at once close this volume. Not that it suffices to study the Principles of Geology, or to read special treatises by different observers on separate formations, and to mark how each author attempts to give an inadequate idea of the duration of each formation, or even of each stratum. We can best gain some idea of past time by knowing the agencies at work; and learning how deeply the surface of the land has been denuded, and how much sediment has been deposited. As Lyell has well remarked, the extent and thickness of our sedimentary formations are the result and the measure of the denudation which the earth's crust has elsewhere undergone. Therefore a man should examine for himself the great piles of superimposed strata, and watch the rivulets bringing down mud, and the waves wearing away the sea-cliffs, in order to comprehend something about the duration of past time, the monuments of which we see all around us.

It is good to wander along the coast, when formed of moderately hard rocks, and mark the process of degradation. The tides in most cases reach the cliffs only for a short time twice a day, and the waves eat into them only when they are charged with sand or pebbles; for there is good evidence that pure water effects nothing in wearing away rock. At last the base of the cliff is undermined, huge fragments fall down, and these remaining fixed, have to be worn away atom by atom, until after being reduced in size they can be rolled about by the waves, and then they are more quickly ground into pebbles, sand, or mud. But how often do we see along the bases of retreating cliffs rounded boulders, all thickly clothed by marine productions, showing how little they are abraded and how seldom they are rolled about! Moreover, if we follow for a few miles any line of rocky cliff, which is undergoing degradation, we find that it is only here and there, along a short length or round a promontory, that the cliffs are at the present time suffering. The appearance of the surface and the vegetation show that elsewhere years have elapsed since the waters washed their base.

We have, however, recently learned from the observations of Ramsay, in the van of many excellent observers – of Jukes, Geikie, Croll and others, that subaerial degradation is a much more important agency than coast-action, or the power of the waves. The whole surface of the land is exposed to the chemical action of the air and of the rainwater, with its dissolved carbonic acid, and in colder countries to frost; the disintegrated matter is carried down even gentle slopes during heavy rain, and to a greater extent than might be supposed, especially in arid districts, by the wind; it is then transported by the streams and rivers, which, when rapid deepen their channels, and triturate the fragments. On a rainy day, even in a gently undulating country, we see the effects of subaerial degradation in the muddy rills which flow down every slope. Messrs. Ramsay and Whitaker have shown, and the observation is a most striking one, that the great lines of escarpment in the Wealden district and those ranging across England, which formerly were looked at as ancient sea-coasts, cannot have been thus formed, for each line is composed of one and the same formation, while our sea-cliffs are everywhere formed by the intersection of various formations. This being the case, we are compelled to admit that the escarpments owe their origin in chief part to the rocks of which they are composed, having resisted subaerial denudation better than the surrounding surface; this surface consequently has been gradually lowered, with the lines of harder rock left projecting. Nothing impresses the mind with the vast duration of time, according to our ideas of time, more forcibly than the conviction thus gained that subaerial agencies, which apparently have so little power, and which seem to work so slowly, have produced great results.

When thus impressed with the slow rate at which the land is worn away through subaerial and littoral action, it is good, in order to appreciate the past duration of time, to consider, on the one hand, the masses of rock which have been removed over many extensive areas, and on the other hand the thickness of our sedimentary formations. I remember having been much struck when viewing volcanic islands, which have been worn by the waves and pared all round into perpendicular cliffs of one or two thousand feet in height; for the gentle slope of the lava-streams, due to their formerly liquid state, showed at a glance how far the hard, rocky beds had once extended into the open ocean. The same story is told still more plainly by faults – those great cracks along which the strata have been upheaved on one side, or thrown down on the other, to the height or depth of thousands of feet; for since the crust cracked, and it makes no great difference whether the upheaval was sudden, or, as most geologists now believe, was slow and effected by many starts, the surface of the land has been so completely planed down that no trace of these vast dislocations is externally visible. The Craven fault, for instance, extends for upward of thirty miles, and along this line the vertical displacement of the strata varies from 600 to 3,000 feet. Professor Ramsay has published an account of a downthrow in Anglesea of 2,300 feet; and he informs me that he fully believes that there is one in Merionethshire of 12,000 feet; yet in these cases there is nothing on the surface of the land to show such prodigious movements; the pile of rocks on either side of the crack having been smoothly swept away.

On the other hand, in all parts of the world the piles of sedimentary strata are of wonderful thickness. In the Cordillera, I estimated one mass of conglomerate at ten thousand feet; and although conglomerates have probably been accumulated at a quicker rate than finer sediments, yet from being formed of worn and rounded pebbles, each of which bears the stamp of time, they are good to show how slowly the mass must have been heaped together. Professor Ramsay has given me the maximum thickness, from actual measurement in most cases, of the successive formations in DIFFERENT parts of Great Britain; and this is the result: —

that is, very nearly thirteen and three-quarters British miles. Some of these formations, which are represented in England by thin beds, are thousands of feet in thickness on the Continent. Moreover, between each successive formation we have, in the opinion of most geologists, blank periods of enormous length. So that the lofty pile of sedimentary rocks in Britain gives but an inadequate idea of the time which has elapsed during their accumulation. The consideration of these various facts impresses the mind almost in the same manner as does the vain endeavour to grapple with the idea of eternity.