At one spot (_Rothwand_) much exposed to the sun, and difficult of access, I remarked two closely allied forms, so nearly related to _H. villosum_ that this would seem to be an intermediary form between the two. One of these (_H. villosissimum_) is distinguished by its tongue and thick p.u.b.escence, its tolerably large capitula, and by the lengthened and separated scales of the involucrum; the other, on the contrary (_H. elongatum_), is less p.u.b.escent, has smaller capitula, and more compact scales on the involucrum than _H. villosum_. Both are finally distinguishable from the type by their longer stalks, which are more decidedly aphyllous, and by their later flowering. At the spot where I found them the two forms were closely intermingled, and each was represented by a considerable number of plants. I did not find them anywhere else on the mountain, nor could I find at the spot where these were growing a single specimen of the true _H. villosum_, nor a single hybrid from these two.
I concluded that these two new forms had, by joining their forces, expelled the _H. villosum_ from its primitive abode, but had not succeeded in displacing one another. As to their origin, they had evidently developed in two different directions from a common point of departure, namely _H. villosum_. They had succeeded, not only in separating themselves from the original form, but also in preventing any intermediary form from interposing. I thought myself therefore justified in considering this as a case of varieties which have come into existence subsequently to the Glacial epoch.
The morphological characteristics of the three forms are sufficiently distinct for them to be designated as species by a good many writers. They are better defined than some of MM. Frolich and Fries" weaker species, and as well defined as some of MM. Koch and Grisebach"s (p. 222).
Now it is clear, without comment, that all this is exactly as it ought to be, if allied species have been differentiated on common areas by selective fertility. For if, as Nageli elsewhere says, "one meets forms in nature a.s.sociated with one another, and severally distinguished by every possible degree of differentiation," not only as Nageli adds, does this general fact lead to the inference that species are (usually) developed when plants grow intimately a.s.sociated together; but as certainly it leads to the further inference that such development must be due to a prior development of cross-infertility between the diverging varietal forms, cross-infertility which is therefore afterwards so characteristic of the allied species, when these are found, in their fully differentiated condition, still occupying the same area in large and intimately mingled populations.
To my mind there could not be any inference more strongly grounded than this, because, with the one exception of the physiological form, no other form of h.o.m.ogamy can be conceived which shall account for the origin and permanence of these synoical varieties, in all degrees of differentiation up to well-defined synoical species. Least of all, as we have seen, can natural selection alone have had anything to do with such a state of matters; while, as we have likewise seen, in all its details it is exactly the state of matters which the theory of physiological selection requires.
Nevertheless, although this inference is so strongly grounded, we ought to remember that it is only an inference. In order fully to verify the theory of physiological selection, we ought to prove by experiment the fact of cross-infertility between these synoical varieties, as we learn that it afterwards obtains between synoical species. It is to be regretted that the theory of physiological selection did not occur to the mind of Nageli, because he would then, no doubt, have ascertained this by actual experiment. As it is, the great value of his observations goes no further than establishing a strong presumption, that it _must_ be selective fertility which causes the progressive differentiation of synoical varieties; and also that, if so, this _must_ be the princ.i.p.al factor in the differentiation of vegetable species, seeing that some ninety-five per cent. are of synoical origin.
_Evidence from Experimental Research._
My paper on _Physiological Selection_ pointed out that the whole theory would have to stand or fall with the experimental proof of the presence or the absence of cross-infertility between varieties of the same species growing on common areas. From the facts and considerations which we have hitherto been dealing with, it did indeed appear to me that there was the strongest conceivable ground for inferring that cross-infertility between such varieties would be found by experiment to be a phenomenon of highly general occurrence--amply sufficient ground to prove that allied species on common areas for the most part owed their origin to this character of mutual sterility, and not vice versa as previously supposed. At that time I was not aware that any experiments had been made in this direction. Soon after the paper was published, however, my attention was directed to a laborious research which had been directed to this very point, and carried on for more than thirty years, by M. Jordan[25]. This had not attracted the general notice which it undoubtedly deserved; and I have since ascertained that even Darwin began to look into it only a few months before his death.
[25] _Remarques sur le fait de l"existence en societe a l"etat sauvage des especes vegetales affines et sur d"autres faits relatifs a la question de l"espece_, par Alexis Jordan; lues au congres de l"a.s.sociation Francaise pour l"Avancemeat des Sciences, 2^me session, Lyon, seance de 28 Aout, 1873.
Having devoted his life to closely observing in divers stations mult.i.tudes of different species of plants--annuals and perennials, bulbous and aquatic, trees and shrubs--M. Jordan has been able to satisfy himself, and the French school of botanists to which this line of observation has given rise, that in most cases (or "nearly everywhere"), when a Linnean species is indigenous to a country and is there of common occurrence, this species within that district is represented by more or less numerous and perfectly constant varieties.
These varieties are const.i.tuted by such minute differences of morphological character that their very existence eluded the observation of botanists, until M. Jordan began to search specially for them as the special objects of his scrutiny. Moreover, these varieties of a Linnean species occupy common areas, and there grow in intimate a.s.sociation with one another, or as M. Jordan says, "_pele-mele_." So far, be it noticed, Jordan was proceeding on exactly the same lines as Nageli; only he carried his observations over a still wider range of species on the one hand, and into a still minuter search for varieties on the other. But the all-important point for us is, that he further proceeded to test by experiment the physiological relations between these morphological varieties; and found, in many hundreds of cases, that they not only came true to seed (i. e. are hereditary and not merely climatic), but likewise cross-sterile _inter se_. For these reasons, M. Jordan, who is opposed to the theory of evolution, regards all such varieties as separately created species; and the inspiring motive of his prolonged investigations has been a desire to multiply these proofs of creative energy. But it clearly makes no difference, so far as evolutionists are concerned with them, whether all this mult.i.tude of s.e.xually isolated forms be denominated species or varieties.
The points which are of importance to evolutionists--and of the first order of importance in the present connexion--may be briefly summarized as follows:--
(1) The research embraces large numbers of species, belonging to very numerous and very varied orders of plants; (2) in the majority of cases--although not all--indigenous species which are of common occurrence present constant varieties; (3) these varieties, nevertheless, may be morphologically so slight as to be almost imperceptible; (4) they occupy common areas and grow in intimate a.s.sociation; (5) although many of them have undergone so small an amount of morphological change, they have undergone a surprising amount of physiological change; for (6) not only do very many of these varieties come true to seed; but, (7) when they do, they are always more or less cross-infertile _inter se_.
Now, it is self-evident that every one of these seven points is exactly what the theory of physiological selection requires, while there is not one of them which it does not require. For if the theory be sound, we should expect to find large numbers of species belonging to numerous and varied orders of plants presenting constant varieties on common areas; we should expect this to be a highly general, though not a universal, rule; and we should expect it to apply only to species which are indigenous. Moreover, we should expect these varieties, although but slightly differentiated morphologically, to present a great differentiation physiologically--and this in the special direction of selective fertility, combined, of course, with heredity.
On the other hand, as I have said, this catalogue of evidences leaves nothing to be supplied. It gives us all the facts--and no more than all the facts--which my paper on _Physiological Selection_ antic.i.p.ated as the eventual result of a prolonged experimental research. And if I have to regret my ignorance of these facts when that paper was published, at any rate it now furnishes the best proof that my antic.i.p.ations were not guided by the results of a verification which had already been supplied.
These antic.i.p.ations were deduced exclusively from the theory itself, as representing what _ought_ to be the case if the theory were true; and, I must confess, if I had then been told that they had already been realized--that it had actually been found to be a general rule that endemic species present constant and hereditary varieties, intimately commingled on common areas, morphologically almost indistinguishable, but physiologically isolated by selective fertility--I should have felt that the theory had been verified in advance. For there are only two alternatives: either these things are due to physiological selection, or else they are due--as M. Jordan himself believes--to special creation.
Which is equivalent to saying that, for evolutionists, the facts must be held to verify the former theory in as complete a manner as it is logically possible for the theory to be verified.
_Evidence from Prepotency._
We have now to consider the bearing of what is called "prepotency" on the theory of physiological selection.
Speaking of the vast number of species of Compositae, Darwin says:--
There can be no doubt that if the pollen of all these species could be simultaneously or successively placed on the stigma of any one species, this one would elect with unerring certainty its own pollen. This elective capacity is all the more wonderful, as it must have been acquired since the many species of this great group of plants branched off from a common progenitor.
Darwin is here speaking of elective affinity in its fully developed form, as absolute cross-sterility between fully differentiated species.
But we meet with all lower degrees of cross-infertility--sometimes between "incipient species," or permanent varieties, and at other times between closely allied species. It is then known as "prepotency" of the pollen belonging to the same variety or species over the pollen of the other variety or species, when both sets of pollen are applied to the same stigma. Although in the absence of the prepotent pollen the less potent will fertilize the seed, yet, such is the appetency for the more appropriate pollen, that even if this be applied to the stigma some considerable time after the other, it will outstrip or overcome the other in fertilizing the ovules, and therefore produce the same result on the next generation as if it had been applied to the mother plant without any admixture of the less potent pollen, although in some cases such incipient degrees of cross-infertility are further shown by the number or quality of the seeds being fewer or inferior.
Now, in different varieties and in different allied species, all degrees of such prepotency have been noticed by many observers, from the faintest perceptible amount up to complete impotency of the alien pollen--when, of course, there is absolute sterility between the two varieties or allied species. The inference is obvious. In this graduated scale of prepotency--beginning with an experimentally almost imperceptible amount of s.e.xual differentiation between two varieties, and ending in an absolute part.i.tioning of two allied species--we have the only remaining fact that is required to complete the case in favour of the present theory. We are here brought back to the very earliest stages of physiological differentiation or to the stages which lie behind Jordan"s "Physiological Species"; and therefore, when taken in conjunction with his results, the phenomena of prepotency may be said to give us the complete and final demonstration of one continuous development, which, beginning in an almost imperceptible amount of cross-infertility, ends in absolute cross-sterility. The "elective capacity" to which Darwin alludes as having been "acquired" by all the species of Compositae since they "branched off from a common progenitor," is thus seen among innumerable other species actually in process of acquisition; and so we can perfectly well understand, what is otherwise unintelligible, that closely allied species of plants occur, in ninety-five per cent. of cases, intimately a.s.sociated on common areas, while exhibiting towards one another the character of mutual sterility.
But more than this. The importance of the widespread phenomena of prepotency to the theory of physiological selection does not consist merely in thus supplying the last link in the chain of evidence touching the origin of species by selective fertility, or "elective capacity."
These phenomena are of further importance as showing how in plants, at all events, physiological selection appears to be frequently capable of differentiating specific types without the necessary a.s.sistance of any other form of h.o.m.ogamy. In my original statement of the theory, I was careful to insist upon the great value, as differentiating agents, of even small degrees of other forms of h.o.m.ogamy when co-operating with physiological selection. But I also stated my belief that in many cases selective fertility is presumably of itself capable of splitting a specific type; and the reason why I still believe this is, that I do not otherwise understand these phenomena of prepotency. I cannot believe that in all the innumerable cases where they arise, they have been super-induced by some prior morphological changes going on in some other part of the organism, or by "prolonged exposure to uniform conditions of life," on the part of two well-nigh identical forms which have arisen intimately commingled in exactly the same environment, and under the operation of a previously universal intercrossing. Even if such a thing could be imagined as happening occasionally, I feel it difficult to imagine that it can happen habitually, and yet this view must be held by those who would attribute prepotency to natural selection.
It must never be forgotten that the relatively enormous changes as to size, structure, habit, &c., which are presented by our domesticated plants as results of artificial selection, do not entail the physiological character of cross-sterility in any degree, save possibly in some small number of cases. Although in wild species any correspondingly small percentage of cases (where natural selection happens to hit upon parts of the organism modifications of which produce the physiological change by way of correlation) would doubtless be the ones to survive on common areas, still it is surely incredible that such an accidental a.s.sociation between natural selection and cross-infertility is so habitually the means of specific differentiation as the facts of prepotency (together with the observations of Jordan and Nageli) would necessarily demand.
Moreover, this view of the matter is still further corroborated by certain other facts and considerations. For example, the phenomena of prepotency (whether as between varieties or between closely allied species) are found to occur when the two forms occupy a common area, i.e. are growing intermingled with one another. Therefore, but for this physiological differentiation, there could be absolutely nothing to prevent free intercrossing. Yet the fact that hybrids are so comparatively rare in a state of nature--a fact which Sir Joseph Hooker has pointed out to me as otherwise inexplicable--proves the efficacy of even a low degree of such differentiation in preventing the physiologically-differentiated forms from intercrossing. Even in cases where there is no difficulty in producing artificial hybrids or mongrels between species or varieties growing on common areas, it is perfectly astonishing what an extremely small percentage of the hybrid or mongrel forms are found to occur in nature. And there can be no question that this is due to the very efficient manner in which prepotency does its work--efficient, I mean, from the point of view of the new theory; for upon any other theory prepotency is a meaningless phenomenon, which, notwithstanding its frequent occurrence, plays no part whatever in the process of organic evolution.
I attach considerable importance to the phenomena of prepotency in view of the contrast which is presented between plants and animals in the relation of their species to physical barriers. For animals--and especially the higher animals--appear to depend for their specific differentiations upon such barriers much more than in the case with plants. This is no more than we should expect; for, in accordance with our theory, selective fertility is not so likely to work alone in the case of the higher animals which mate together, as in plants which are fertilized through the agency of wind or insects. In the former case there is no opportunity given for the first rise of cross-infertility, in the form of prepotency; and even where selective fertility has gained a footing in other ways, the chances against the suitable mating of "physiological complements" must be much greater than it is in the latter case. Hence, among the higher animals, selective fertility ought much more frequently to be found in a.s.sociation with other forms of h.o.m.ogamy than it is among plants. And this is exactly what we find. Thus it seems to me that this contrast between the comparative absence and presence of physical barriers, where allied species of plants and of higher animals are respectively concerned, is ent.i.tled to be taken as a further corroboration of our theory. For while it displays exactly such a general correlation as this theory would expect, the correlation is one which cannot possibly be explained on any other theory. It is just where physiological selection can be seen to have the best opportunity of acting (viz. in the vegetable kingdom) that we find the most unequivocal evidence of its action; while, on the other hand, it is just where it can be seen to have the least opportunity of a.s.serting itself (viz. among the higher animals) that we find it most a.s.sociated with, and therefore a.s.sisted by, other forms of h.o.m.ogamy, i. e. not only geographical isolation, but also by s.e.xual preference in pairing, and the several other forms of h.o.m.ogamy, which Mr. Gulick has shown to arise in different places as the result of intelligence.
_Evidence from Special Cases._
Hitherto I have been considering, from the most general point of view, the most widespread facts and broadest principles which serve to substantiate the theory of physiological selection. I now pa.s.s to the consideration of one of those special cases in which the theory appears to have been successfully applied.
Professor Le Conte has adduced the fossil snails of Steinheim as serving to corroborate the theory of physiological selection[26].
[26] _Evolution and its Relations to Religious Thought_, &c. pp.
236-7.
The facts are these. The snail population of this lake remain for a long time uniform and unchanged. Then a small percentage of individuals suddenly began to vary as regards the form of their sh.e.l.ls, and this in two or three directions at the same time, each affected individual, however, only presenting one of the variations. But after all these variations had begun to affect a proportionally large number of individuals, some individuals occur in which two or more of the variations are blended together, evidently, as Weismann says, by intercrossing of the varieties so blended. Later still, both the separate varieties and their blended progeny became more and more numerous, and eventually a single blended type, comprising in itself all the initial varieties, supplanted the parent form. Then another long period of stability ensued until another eruption of new variations took place; and these variations, after having affected a greater and greater number of individuals, eventually blended together by intercrossing and supplanted their parent form. So the process went on, comparatively short periods of variation alternating with comparatively long periods of stability, the variations, moreover, always occurring suddenly in crops, then multiplying, blending together, and in their finally blended type eventually supplanting their parent form.
Now, the remarkable fact here is that whenever the variations arose, they only intercrossed between themselves, they did not intercross with their parent form; for, if they had, not only could they never have survived (having been at first so few in number and there having been no geographical barriers in the small lake), but we should have found evidence of the fact in the half-bred progeny. Moreover, natural selection can have had nothing to do with the process, because not only are the variations in the form of the sh.e.l.ls of no imaginable use in themselves; but it would be preposterous to suppose that at each of these "variation periods" several different variations should always have occurred simultaneously, all of which were of some hidden use, although no one of them ever occurred during any of the prolonged periods of stability. How, then, are we to explain the fact that the individuals composing each crop of varieties, while able to breed among themselves, never crossed with their parent form? These varieties, each time that they arose, were intimately commingled with their parent form, and would certainly have been reabsorbed into it had intercrossing in that direction been possible. With Professor Le Conte, therefore, I conclude that there is only one conceivable answer to this question.
Each crop of varieties must have been _protected from intercrossing with their parent form_.
They must have been the result of a variation, which rendered the affected individuals sterile with their parent form, whilst leaving them fertile amongst themselves. The progeny of these individuals would then have dispersed through the lake, physiologically isolated from the parent population, and especially p.r.o.ne to develop secondary variations as a direct result of the primary variation. Thus, as we might expect, two or three variations arose simultaneously, as expressions of so many different lines of family descent from the original or physiological variety; these were everywhere prevented from intercrossing with their parent form, yet capable of blending whenever they or their ever-increasing progeny happened to meet. Thus, without going into further details, we are able by the theory of physiological selection to give an explanation of all these facts, which otherwise remain inexplicable.
In view of the evidence which has now been presented, I will now ask five questions which must be suitably answered by critics of the theory of physiological selection.
1. Can you doubt that the hitherto insoluble problem of inter-specific sterility would be solved, supposing cross-infertility were proved to arise before or during the process of specific differentiation, instead of after that process had been fully completed?
2. Can you doubt, after duly considering the circ.u.mstances under which allied species of plants have been differentiated--viz. in ninety-five per cent. of cases intimately commingled on common areas, and therefore under identical environments--that cross-infertility _must_ have arisen before or during the specific differentiation?
3. Can you doubt, after duly considering the facts of prepotency on the one hand and those of Jordan"s physiological varieties on the other, that cross-infertility _does_ arise before or during the specific differentiation?
4. If you cannot express a doubt upon any of these points, can you explain why you refuse to accept the theory of the origin of species by means of physiological selection, together with the explanation which this theory affords of the continued cross-fertility of domesticated varieties?
5. Supposing this theory to be true, can you conceive of any other cla.s.ses of facts which, either quant.i.tatively or qualitatively, could more directly or more effectually prove its truth than those which have now been adduced?
On these five heads I entertain no doubt. I am convinced that the theory of physiological selection is the only one that can explain the facts of inter-specific sterility on the one hand, and, on the other hand, the contrast which these facts display to the unimpaired fertility of our domesticated varieties.
In conclusion, it seems desirable once more to insist that there is no antagonism or rivalry between the theories of natural and of physiological selection. For which purpose I will quote the final paragraph of my original paper.
So much, then, for the resemblances and the differences between the two theories. It only remains to add that the two are complementary. I have already shown some of the respects in which the newer theory comes to the a.s.sistance of the older, and this in the places where the older has stood most in need of a.s.sistance. In particular, I have shown that segregation of the fit entirely relieves survival of the fittest from the difficulty under which it has. .h.i.therto laboured of explaining why it is that sterility is so constantly found between species, while so rarely found between varieties which differ from one another even more than many species; why so many features of specific distinction are useless to the species presenting them; and why it is that incipient varieties are not obliterated by intercrossing with parent forms.
Again, we have seen that physiological selection, by preventing such intercrossing, enables natural selection to promote diversity of character, and thus to evolve species in ramifying branches instead of in linear series--a work which I cannot see how natural selection could possibly perform unless thus aided by physiological selection. Moreover, we have seen that although natural selection alone could not induce sterility between allied types, yet when this sterility is given by physiological selection, the forms which present it would be favoured in the struggle for existence; and thus again the two principles are found playing, as it were, into each other"s hands. And here, as elsewhere, I believe that the co-operation enables the two principles to effect very much more in the way of species-making than either of them could effect if working separately. On the one hand, without the a.s.sistance of physiological selection, natural selection would, I believe, be all but overcome by the adverse influences of free intercrossing--influences all the more potent under the very conditions which are required for the multiplication of species by divergence of character. On the other hand, without natural selection, physiological selection would be powerless to create any differences of specific type, other than those of mutual sterility and trivial details of structure, form, and colour--differences wholly without meaning from a utilitarian point of view. But in their combination these two principles appear to me able to accomplish what neither can accomplish alone--namely, a full and satisfactory explanation of the origin of species.
CHAPTER VI.
A BRIEF HISTORY OF OPINIONS ON ISOLATION AS A FACTOR OF ORGANIC EVOLUTION.