Chapter VI · A Brief History of Opinions on Isolation as a Factor of Organic Evolution.

This historical sketch must begin with a consideration of Darwin's opinions on the subject; but as these were considerably modified from time to time during a period of thirty years by the publications of other naturalists, it will be impossible to avoid cross-references as between his writings and theirs. It may also be observed that the Life and Letters of Charles Darwin was not published until the year 1887, so that the various opinions which I shall quote from the letters, and which show some considerable approximation in his later years to the views which have been put forward by Mr. Gulick and myself, were not before us at the time when our papers were read.

The earliest allusion that I can find to geographical isolation in the writings of Darwin occurs in a correspondence with Sir Joseph Hooker, as far back as 1844. He there says:—

I cannot give my reasons in detail; but the most general conclusion which the geographical distribution of all organic beings appears to me to indicate is, that isolation is the chief concomitant or cause of the appearance of new forms (I well know there are some staring exceptions)[27].

And again:—

With respect to original creation or production of new forms, I have said that isolation appears the chief element[28].

Next, in the earlier editions of the Origin of Species this view is abandoned, and in its stead we meet with the opinion that geographical isolation lends a certain amount of assistance to natural selection, by preventing free intercrossing. But here we must note two things. First, the distinction between monotypic and polytypic evolution is not defined. Secondly, the levelling effect of free intercrossing in nature, and hence its antagonism to divergence of character by natural selection, is not sufficiently recognized; while, on the other hand, and in consequence of this, the importance of isolation as a factor of evolution is underrated—not only in its geographical, but likewise in all its other forms.

Taking these two points separately, the only passages in Darwin's writings, so far at least as I can find, in which any distinction is drawn between evolution as monotypic and polytypic, are those in which he deals with a somewhat analogous distinction between artificial selection as intentional and unconscious. He says, for example:—

In the case of methodical selection, a breeder selects for some definite object, and if the individuals be allowed freely to intercross, his work will completely fail. But when many men, without intending to alter the breed, have a nearly common standard of perfection, and all try to procure and breed from the best animals, improvement surely but slowly follows from this unconscious process of selection, notwithstanding that there is no separation of selected individuals. Thus it will be under nature[29].

Here we have what may perhaps be regarded as a glimmering of the distinction between monotypic and polytypic evolution. But that it is only a glimmering is proved by the immediately ensuing sentences, which apply this analogy of unconscious selection not to the case of monotypic, but to that of polytypic evolution. So likewise, in the succeeding discussion on "divergence of character," the analogy is again resorted to for the purpose of showing how polytypic evolution may occur in nature.

Thus far, then, it may be said that we have scarcely so much as a glimmering of the distinction between monotypic and polytypic evolution; and as the same discussion (with but a few verbal alterations) runs through all the editions of the Origin, it may well be asked why I should have alluded to such passages in the present connexion. Well, I have done so because it is apparent that, during the last years of his life, the distinction between selection as "methodical" and "unconscious" enabled Darwin much more clearly to perceive that between evolution as monotypic and polytypic. Thus in 1868 he wrote to Moritz Wagner (who, as we shall presently see, entirely failed to distinguish between monotypic and polytypic evolution), expressing his belief—

That in many large areas all the individuals of the same species have been slowly modified, in the same manner, for instance, as the English racehorse has been improved, that is, by the continued selection of the fleetest individuals, without any separation. But I admit that by this process two or more new species could hardly be formed within the same limited area[30].

Again, in 1876 he wrote another letter to Wagner, in which the following passage occurs:—

I believe that all the individuals of a species can be slowly modified within the same district, in nearly the same manner as man effects by what I have called the process of unconscious selection. I do not believe that one species will give birth to two or more new species as long as they are mingled together within the same district[31].

Two years later he wrote to Professor Semper:—

There are two different classes of cases, it appears to me, viz. those in which species becomes slowly modified in the same country, and those cases in which a species splits into two, or three, or more new species; and, in the latter case, I should think nearly perfect separation would greatly aid in their "specification," to coin a new word[32].

Now, these passages show a very much clearer perception of the all-important distinction between monotypic and polytypic evolution than any which occur in the Origin of Species; and they likewise show that he was led to this perception through what he supposed to be a somewhat analogous distinction between "unconscious" and "methodical" selection by man. The analogy, I need hardly say, is radically unsound; and it is a curious result of its unsoundness that, whereas in the Origin of Species it is adduced to illustrate the process of polytypic evolution, as previously remarked, in the letters above quoted we find it adduced to illustrate the process of monotypic evolution. But the fact of this analogy being unsound does not affect the validity of the distinction between monotypic and polytypic evolution to which it led Darwin, in his later years, so clearly to express[33].

Turning next to the second point which we have to notice, it is easy to show that in the earlier editions of his works Darwin did not sufficiently recognize the levelling effects of free intercrossing, and consequently failed to perceive the importance of isolation (in any of its forms) as a factor of organic evolution. This may be most briefly shown by quoting his own more matured opinion upon the subject. Thus, with reference to the swamping effects of intercrossing, he wrote to Mr. Wallace in 1867 as follows:—

I must have expressed myself atrociously: I meant to say exactly the reverse of what you have understood. F. Jenkin argued in the North British Review against single variations being perpetuated, and has convinced me, though not in quite so broad a manner as here put. I always thought individual differences more important; but I was blind, and thought that single variations might be preserved much oftener than I now see is possible or probable. I mentioned this in my former note merely because I believed that you had come to a similar conclusion, and I like much to be in accord with you. I believe I was mainly deceived by single variations offering such simple illustrations, as when man selects [i.e. isolates][34].

Again, somewhere about the same time, he wrote to Moritz Wagner:—

Although I saw the effects of isolation in the case of islands and mountain-ranges, and knew of a few instances of rivers, yet the greater number of your facts were quite unknown to me. I now see that, from the want of knowledge, I did not make nearly sufficient use of the views which you advocate[35].

Now it would be easy to show the justice of these self-criticisms by quoting longer passages from earlier editions of the Origin of Species; but as this, in view of the above passages, is unnecessary, we may next pass on to another point.

The greatest oversight that Wagner made in his otherwise valuable essays on geographical isolation, was in not perceiving that geographical isolation is only one among a number of other forms of isolation: and, therefore, that although it is perfectly true, as he insisted, that polytypic evolution cannot be effected by natural selection alone, it is very far from true, as he further insisted, that geographical isolation is the only means whereby natural selection can be assisted in this matter. Hence it is that, when Darwin said he had not himself "made nearly sufficient use" of geographical isolation as a factor of specific divergence, he quite reasonably added that he could not go so far as Wagner did in regarding such isolation as a condition, sine qua non, to divergent evolution in all cases. Nevertheless, he adds the important words, "I almost wish I could believe in its importance to the same extent with you; for you well show, in a manner which never occurred to me, that it removes many difficulties and objections." These words are important, because they show that Darwin had come to feel the force of the "difficulties and objections" with regard to divergent evolution being possible by means of natural selection alone, and how readily they could be removed by assuming the assistance of isolation. Hence, it is much to be deplored that Wagner presented a single kind of isolation (geographical) as equivalent to the principle of isolation in general. For he thus failed to present the complete—and, therefore, the true—philosophy of the subject to Darwin's mind; and in this, as in certain other respects which I shall notice later on, served rather to confuse than to elucidate the matter as a whole.

To sum up. Although in his later years, as shown by his correspondence, Darwin came to recognize more fully the swamping effects of free intercrossing, and the consequent importance of "separation" for the prevention of these effects, and although in this connexion he likewise came more clearly to distinguish between the "two cases" of monotypic and polytypic evolution, it is evident that he never worked out any of these matters—"thinking it prudent," as he wrote with reference to them in 1878, "now I am growing old, to work at easier subjects[36]." Therefore he never clearly saw, on the one hand, that free intercrossing, far from constituting a "difficulty" to monotypic evolution by natural selection, is the very means whereby natural selection is in this case enabled to operate; or, on the other hand, that, in the case of polytypic evolution, the "difficulty" in question is so absolute as to render such evolution, by natural selection alone, absolutely impossible. Hence, although in one sentence of the Origin of Species he mentions three forms of isolation (besides the geographical form) as serving in some cases to assist natural selection in causing "divergence of character" (i. e. polytypic evolution[37]), on account of not perceiving how great and how sharp is the distinction between the two kinds or "cases" of evolution, he never realized that, where "two or more new species" are in course of differentiation, some form of isolation other than natural selection must necessarily be present, whether or not natural selection be likewise so. The nearest approach which he ever made to perceiving this necessity was in one of his letters to Wagner above quoted, where, after again appealing to the erroneous analogy between monotypic evolution and "unconscious selection," he says:—"But I admit that by this process (i. e. unconscious selection) two or more new species could hardly be formed within the same limited area: some degree of separation, if not indispensable, would be highly advantageous; and here your facts and views will be of great value." But even in this passage the context shows that by "separation" he is thinking exclusively of geographical separation, which he rightly enough concludes (as against Wagner) need certainly not be "indispensable." Had he gone a step further, he must have seen that separation, in some form or another, is "indispensable" to polytypic evolution. Instead of taking this further step, however, two years later he wrote to Semper as follows:—

I went as far as I could, perhaps too far, in agreement with Wagner [i. e. in the last edition of the Origin of Species]; since that time I have seen no reason to change my mind; but then I must add that my attention has been absorbed on other subjects[38].

And he seems to have ended by still failing to perceive that the explanation which he gives of "divergence of character" in the Origin of Species, can only hold on the unexpressed assumption that free intercrossing is in some way prevented at the commencement, and throughout the development, of each diverging type.

Lastly, we have to consider Darwin's opinion touching the important principle of "Independent Variability." This, it will be remembered, is the principle which ensures that when a portion (not too large) of a species is prevented from interbreeding with the rest of the species, sooner or later a divergence of type will result, owing to the fact that the average qualities of the separated portion at the time of its separation cannot have been exactly the same as the average qualities of the specific type as a whole. Thus the state of Amixia, being a state of what Mr. Gulick calls Independent Generation, will of itself—i.e. even if unassisted by natural selection—induce divergence of type, in a ratio that has been mathematically calculated by Delbœuf.

Darwin wrote thus to Professor Weismann in 1872:—

I have now read your essay with very great interest. Your view of the origin of local races through "Amixia" is altogether new to me, and seems to throw an important light on an obscure question[39].

And in the last edition of the Variation of Animals and Plants he adds the following paragraph:—

This view may throw some light on the fact that the domestic animals which formerly inhabited the several districts in Great Britain, and the half-wild cattle lately kept in several British parks, differed slightly from one another; for these animals were prevented from wandering over the whole country and intercrossing, but would have crossed freely within each district or park[40].

Now, although I allow that Darwin never attributed to this principle of Amixia, or Independent Variability, anything like the degree of importance to which, in the opinion of Delbœuf, Gulick, Giard, and myself, it is entitled, the above passage appears to show that, as soon as the "view" was clearly "suggested" to his mind, he was so far from being unfavourably disposed towards it, that he added a paragraph to the last edition of his Variation for the express purpose of countenancing it. Nevertheless, later on the matter appears to have entirely escaped his memory; for in 1878 he wrote to Semper, that he did "not see at all more clearly than I did before, from the numerous cases which he [Wagner] has brought forward, how and why it is that a long isolated form should almost always become slightly modified[41]." I think this shows entire forgetfulness of the principle in question, because, if the latter is good for explaining the initial divergence of type as between separated stocks of "domesticated animals," much more must it be competent to explain the further divergence of type which is "almost always" observable in the case of "a long isolated form" under nature. The very essence of the principle being that, when divergence of type has once begun, this divergence must ipso facto proceed at an ever-accelerating pace, it is manifestly inconsistent to entertain the principle as explaining the first commencement of divergence, and then to ignore it as explaining the further progress of divergence. Hence, I can only conclude that Darwin had forgotten this principle altogether when he wrote his letter to Semper in 1878—owing, no doubt, as he says in the sentence which immediately follows, to his having "not attended much of late years to such questions."

So much, then, for Darwin's opinions. Next in order of time we must consider Moritz Wagner's essays on what he called the "Law of Migration[42]." The merit of these essays was, first, the firm expression of opinion upon the swamping effects of free intercrossing; and, second, the production of a large body of facts showing the importance of geographical isolation in the prevention of these effects, and in the consequent differentiation of specific types. On the other hand, the defect of these essays was, first, not distinguishing between evolution as monotypic and polytypic; and, second, not perceiving that geographical isolation is only one among a number of other forms of isolation. From these two radical oversights—which, however, were shared by all other writers of the time, with the partial exception of Darwin himself, as previously shown—there arose the following and most lamentable errors.

Over and over again Moritz Wagner insists, as constituting the fundamental doctrine of his attempted reform of Darwinism, that evolution by natural selection is impossible, unless natural selection be assisted by geographical isolation, in order to prevent the swamping effects of intercrossing[43]. Now, if instead of "evolution" he had said "divergence of type," and if instead of "geographical isolation" he had said "prevention of intercrossing," he would have enunciated the general doctrine which it has been the joint endeavour of Mr. Gulick and myself to set forth. But by not perceiving that "evolution" is of two radically different kinds—polytypic and monotypic—he entirely failed to perceive that, while for one of its kinds the prevention of intercrossing is an absolute necessity, for the other of its kinds the permission of intercrossing is a necessity no less absolute. And, again, in missing the fact that geographical isolation is but one of the many ways whereby intercrossing may be prevented, he failed to perceive that, even as regards the case of polytypic evolution, he greatly erred in representing this one form of isolation as being universally a necessary condition to the process. The necessary condition to this process is, indeed, the prevention of intercrossing by some means or another; but his unfortunate insistence on geographical separation as the only possible means to this end—especially when coupled with his no less unfortunate disregard of monotypic evolution—caused him to hinder rather than to advance a generalization which he had only grasped in part. And this generalization is, as now so repeatedly stated, that while the form of isolation which we know as natural selection depends for its action upon the intercrossing of all the individuals which it isolates (i. e. selects), when acting alone it can produce only monotypic evolution; but that when it is supplemented by any of the other numerous forms of isolation, it is furnished with the necessary condition to producing polytypic evolution—and this in as many lines of divergent change as there may be cases of this efficient separation.

Nevertheless, while we must lament these shortcomings on the part of Wagner, we ought to remember that he rendered important services in the way of calling attention to the swamping effects of free intercrossing, and, still more, in that of showing the high importance of geographical isolation as a factor of organic evolution. Therefore, although in an elaborate criticism of his views Weismann was easily able to dispose of his generalizations in the imperfect form that they presented, I do not think it was just in Weismann to remark, "if Wagner had confined himself to the statement that geographical isolation materially assists the process of natural selection, and thus also promotes the origination of new species, he would have met with little or no opposition; but then, of course, in saying this much, he would not have been saying anything new." No doubt, as I have just shown, he ought thus (as well as in other and still more important respects not perceived by Prof. Weismann) to have limited his statement; but, had he done so, it does not follow that he would not have been saying anything new. For, in point of fact, in as far as he said what was true, he did say a great deal that was also new. Thus, most of what he said of the principle of separation (apogamy) was as new as it was true, although, as we have seen, he said it to very little purpose on account of his identifying this principle as a whole with that of but one of its forms. Again, notwithstanding this great error, or oversight, he certainly showed of the particular form in question—viz. geographical isolation—that it was of considerably more importance than had previously been acknowledged. And this was so far a valuable contribution to the general theory of descent.

Prof. Weismann's essay, to which allusion has just been made[44], was, however, in all respects a great advance upon those of Wagner. It was not only more comprehensive in its view of the whole subject of geographical isolation, but likewise much more adequate in its general treatment thereof. Its principal defects, in my judgement, were, first, the inordinately speculative character of some of its parts, and, second, the restriction of its analysis to but one form of isolation—a defect which it shares with the essays of Wagner, and in quite as high a degree. Furthermore, although this essay had the great merit of enunciating the principle of Amixia, it did so in a very inefficient manner. For not only was this principle adduced with exclusive reference to geographical isolation, but even in regard to this one kind of isolation it was presented in a highly inconsistent manner, as I will now endeavour to show.

Weismann was led to perceive the principle in question by the consideration that new specific characters, when they first appear, do not all appear together in the same individuals: they appear one in one individual, another in another, a third in a third, &c.; and it is only in the course of successive generations that they all become blended in the same individuals by free intercrossing. Hence, the eventually emerging constant or specific type is the resultant of all the transitory or varietal types, when these have been fused together by intercrossing. From which Weismann deduces what he considers a general law—namely, that "the constancy of a specific type does not arise suddenly, but gradually; and it is established by the promiscuous crossing of all individuals[45]." From which again it follows, that this constancy must cease so soon as the condition which maintains it ceases—i. e. so soon as free intercrossing is prevented by the geographical isolation of a portion of the species from its parent stock.

Now, to begin with, this statement of the principle in question is not a good statement of it. There was no need while stating the doctrine that separation induces differentiation, to found the doctrine on any such highly speculative basis. In point of fact, there is no real evidence that specific types do attain their constancy in the way supposed; nor, for the purposes of the doctrine in question, is it necessary that there should be. For this doctrine does not need to show how the constancy has been attained; it only has to show that the constancy is maintained by free intercrossing, with the result that when free intercrossing is by any means prevented, divergence of character ensues. In short, the correct way of stating the principle is that which has been adopted by Delbœuf and Gulick—namely, the average characters of a separated portion of a species are not likely to be the same as those of the whole species; with the result that divergence of type will be set up in the separated portion by intercrossing within that portion. Or the principle may be presented as I presented it under the designation of "Independent Variability"—namely, "a specific type may be regarded as the average mean of all individual variations, any considerable departure from this average mean being, however, checked by intercrossing," with the result that when intercrossing is prevented between a portion of a species and the rest of the species, "this population is permitted to develop an independent history of its own, shielded from intercrossing with its parent form[46]."

Not only, however, is Weismann's principle of "Amixia" thus very differently stated from that of my "Independent Variability" (apogamy), or Gulick's "Independent Generation"; but, apparently owing to this difference of statement, the principle itself is not the same. In particular, while Weismann holds with us that when new characters arise in virtue of the mere prevention of intercrossing with parent forms these new characters will be of non-utilitarian kind[47], he appears to think that divergence of character under such circumstances is not likely to go on to a specific value. Now, it is of importance to observe why he arrives at this conclusion, which is not only so different from that of Delbœuf, Gulick, and myself, but apparently so inconsistent with his own recognition of the diversifying effect of "Amixia" as regards the formation of permanent varieties. For, as we have already seen while considering Darwin's views on this same principle of "Amixia," it is highly inconsistent to recognize its diversifying effect up to the stage of constituting fixed varieties, and then not to recognize that, so much divergence of character having been already secured by the isolation alone, much more must further divergence continue, and continue at an ever accelerating pace—as Delbœuf and Gulick have so well shown. What, then, is the explanation of this apparent inconsistency on Weismann's part? The explanation evidently is that, owing to his erroneous statement of the principle, he misses the real essence of it. For, in the first place, he does not perceive that this essence consists in an initial difference of average characters on the part of the isolated colony as compared with the rest of their species. On the contrary, he loses himself in a maze of speculation about all species having had what he calls "variation-periods," or eruptions of general variability alternating with periods of repose—both being as unaccountable in respect of their causation as they are hypothetical in respect of their occurrence. From these speculations he concludes, that isolation of a portion of a species will then only lead to divergence of character when the isolation happens to coincide with a "variation-period" on the part of the species as a whole, and that the divergence will cease so soon as the "variation-period" ceases. Again, in the second place as previously remarked, equally with Wagner whom he is criticizing, he fails to perceive that geographical isolation is not the only kind of isolation, or the only possible means to the prevention of free intercrossing. And the result of this oversight is, that he thinks amixia can act but comparatively seldom upon sufficiently small populations to become a factor of much importance in the differentiation of species. Lastly, in the third place, owing to his favourite hypothesis that all species pass through a "variation-period," he eventually concludes that the total amount of divergence of type producible by isolation alone (even in a small population) can never be greater than that between the extremes of variation which occur within the whole species at the date of its partition (p. 75). In other words, the possibility of change due to amixia alone is taken to be limited by the range of deviation from the general specific average, as manifested by different individual variations, before the species was divided. Thus the doctrine of amixia fails to recognize the law of Delbœuf, or the cumulative nature of divergence of type when once such divergence begins in a separated section. Therefore, in this all-important—and, indeed, essential—respect, amixia differs entirely from the principle which has been severally stated by Delbœuf, Gulick, and myself.

Upon the whole, then, we must say that although Professor Weismann was the first to recognize the diversifying influence of merely indiscriminate isolation per se (apogamy), he did so only in part. He failed to distinguish the true essence of the principle, and by overlaying it with a mass of hypothetical speculation, concealed even more of it than he revealed.

The general theory of Isolation, as independently worked out by Mr. Gulick and myself, has already been so fully explained, that it will here be sufficient merely to enumerate its more distinguishing features. These are, first, drawing the sharpest possible line between evolution as monotypic and polytypic; second, showing that while for the former the peculiar kind of isolation which is presented by natural selection suffices of itself to transform a specific type, in order to work for the latter, or to branch a specific type, natural selection must necessarily be assisted by some other kind of isolation; third, that even in the absence of natural selection, other kinds of isolation may be sufficient to effect specific divergence through independent generation alone; fourth, that, nevertheless, natural selection, where present, will always accelerate the process of divergence; fifth, that monotypic evolution by natural selection depends upon the presence of intercrossing, quite as much as polytypic evolution (whether with or without natural selection) depends upon the absence of it; sixth, that, having regard to the process of evolution throughout all taxonomic divisions of organic nature, we must deem the physiological form of isolation as the most important, with the exception only of natural selection.

The only difference between Mr. Gulick's essays and my own is, that, on the one hand, he has analyzed much more fully than I have the various forms of isolation; while, on the other hand, I have considered much more fully than he has the particular form of physiological isolation which so frequently obtains between allied species. This particular form of physiological isolation I have called "physiological selection," and claim for it so large a share in the differentiation of specific types as to find in it a satisfactory explanation of the contrast between natural species and artificial varieties in respect of cross-infertility.

Mr. Wallace, in his Darwinism, has done good service by enabling all other naturalists clearly to perceive how natural selection alone produces monotypic evolution—namely, through the free intercrossing of all individuals which have not been eliminated by the isolating process of natural selection itself. For he very lucidly shows how the law of averages must always ensure that in respect of any given specific character, half the individuals living at the same time and place will present the character above, and half below its mean in the population as a whole. Consequently, if it should ever be of advantage to a species that this character should undergo either increase or decrease of its average size, form, colour, &c., there will always be, in each succeeding generation, a sufficient number of individuals—i. e. half of the whole—which present variations in the required direction, and which will therefore furnish natural selection with abundant material for its action, without the need of any other form of isolation. It is to be regretted, however, that while thus so clearly presenting the fact that free intercrossing is the very means whereby natural selection is enabled to effect monotypic evolution, he fails to perceive that such intercrossing must always and necessarily render it impossible for natural selection to effect polytypic evolution. A little thought might have shown him that the very proof which he gives of the necessity of intercrossing where the transmutation of species is concerned, furnishes, measure for measure, as good a proof of the necessity of its absence where the multiplication of species is concerned. In justice to him, however, it may be added, that this distinction between evolution as monotypic and polytypic (with the important consequence just mentioned) still continues to be ignored also by other well-known evolutionists of the "ultra-Darwinian" school. Professor Meldola, for example, has more recently said that in his opinion the "difficulty from intercrossing" has been in large part—if not altogether—removed by Mr. Wallace's proof that natural selection alone is capable of effecting [monotypic] evolution; while he regards the distinction between monotypic and polytypic evolution as mere "verbiage[48]."

It is in relation to my presentment of the impossibility of natural selection alone causing polytypic evolution, that Mr. Wallace has been at the pains to show how the permission of intercrossing (panmixia) is necessary for natural selection in its work of causing monotypic evolution. And not only has he thus failed to perceive that the "difficulty" which intercrossing raises against the view of natural selection being of itself capable of causing polytypic evolution in no way applies to the case of monotypic; but as regards this "difficulty," where it does apply, he says:—

Professor G. J. Romanes has adduced it as one of the difficulties which can alone be overcome by his theory of physiological selection[49].

This, however, is a misapprehension. I have by no means represented that the difficulty in question can alone be overcome by this theory. What I have represented is, that it can be overcome by any of the numerous forms of isolation which I named, and of which physiological selection is but one. And although, where common areas are concerned, I believe that the physiological form of isolation is the most important form, this is a very different thing from entertaining the supposition which Mr. Wallace here assigns to me.

I may take this opportunity of correcting a somewhat similar misunderstanding which has been more recently published by Professor W. A. Herdman, of Liverpool; and as the case which he gives is one of considerable interest in itself, I will quote his remarks in extenso. In his Opening Address to the Liverpool Biological Society, Professor Herdman said:—

Some of you will doubtless remember that in last year's address, while discussing Dr. Romanes' theory of physiological selection, I quoted Professor Flemming Jenkin's imaginary case of a white man wrecked upon an island inhabited by negroes, given as an illustration of the supposed swamping effect by free intercrossing of a marked variety with the parent species. I then went on to say in criticism of the result at which Jenkin arrived, viz. that the characteristics of the white man would be stamped out by intercrossing with the black:—

"Two influences have, I think, been ignored, viz. atavism, or reversion to ancestral characters, and the tendency of the members of a variety to breed with one another. Keeping to the case described above, I should imagine that the numbers of intelligent young mulattoes produced in the second, third, fourth, and few succeeding generations would to a large extent intermarry, the result of which would be that a more or less white aristocracy would be formed on the island, including the king and all the chief people, the most intelligent men and the bravest warriors. Then atavism might produce every now and then a much whiter individual—a reversal to the characteristics of the ancestral European—who, by being highly thought of in the whitish aristocracy, would have considerable influence on the colour and other characteristics of the next generation. Now such a white aristocracy would be in precisely the same circumstances as a favourable variety competing with its parent species," &c.

You may imagine then my pleasure when, a few months after writing the above, I accidentally found, in a letter[50] written by the celebrated African traveller Dr. David Livingstone to Lord Granville, and dated "Unyanyembe, July 1st, 1872," the following passage:—

"About five generations ago, a white man came to the highlands of Basañgo, which are in a line east of the watershed. He had six attendants, who all died, and eventually their headman, called Charura, was elected chief by the Basañgo. In the third generation he had sixty able-bodied spearmen as lineal descendants. This implies an equal number of the other sex. They are very light in colour, and easily known, as no one is allowed to wear coral beads such as Charura brought except the royal family. A book he brought was lost only lately. The interest of the case lies in its connexion with Mr. Darwin's celebrated theory on the 'origin of species,' for it shows that an improved variety, as we whites modestly call ourselves, is not so liable to be swamped by numbers as some have thought."

Here we have a perfect fulfilment of what I last year, in ignorance of this observation of Livingstone's, predicted as being likely to occur in such a case. We have the whitish aristocracy in a dominant condition, and evidently in a fair way to spread their characteristics over a larger area and give rise to a marked variety, and it had clearly struck Livingstone fourteen years before the theory of physiological selection had been heard of, just as it must strike us now, as an instance telling strongly against the "swamping" argument as used by Flemming Jenkin and Romanes.

Here we have a curious example of one writer supporting the statements of another, while appearing to be under the impression that he is controverting those statements. Both Professor Herdman's imaginary case, and its realization in Livingstone's account, go to show "the tendency of the members of a variety to breed with one another." This is what I have called "psychological selection," and, far from "ignoring" it, I have always laid stress upon it as an obviously important form of isolation or prevention of free intercrossing. But it is a form of isolation which can only occur in the higher animals, and, therefore, the whole of Professor Herdman's criticism is merely a restatement of my own views as already published in the paper which he is criticizing. For all that his argument goes to prove is, first, the necessity for some form of isolation if the overwhelming effects of intercrossing are to be obviated; and, secondly, the manifest consequence that where the psychological form is unavailable (as in many of the lower animals and in all plants), some other form must be present if divergent evolution is taking place on a common area.

Seeing that so much misunderstanding has been shown with reference to my views on "the swamping effects of intercrossing," and seeing also that this misunderstanding extends quite as much to Mr. Gulick's views as to my own, I will here supply brief extracts from both our original papers, for the double purpose of showing our complete agreement, and of leaving it to be judged whether we can fairly be held responsible for the misunderstanding in question. After having supplied these quotations, I will conclude this historical sketch by considering what Mr. Wallace has said in reply to the views therein presented. I will transcribe but a single passage from our papers, beginning with my own.

Any theory of the origin of species in the way of descent must be prepared with an answer to the question, Why have species multiplied? How is it that, in the course of evolution, species have not simply become transmuted in linear series instead of ramifying into branches? This question Mr. Darwin seeks to answer "from the simple circumstance that the more diversified the descendants from any one species becomes in structure, constitution, and habits, by so much will they be better enabled to seize on many and widely diversified places in the economy of nature, and so be enabled to increase in numbers." And he proceeds to illustrate this principle by means of a diagram, showing the hypothetical divergence of character undergone by the descendants of seven species. Thus, he attributes divergence of character exclusively to the influence of natural selection.

Now, this argument appears to me unassailable in all save one particular; but this is a most important particular: the argument wholly ignores the fact of intercrossing with parent forms. Granting to the argument that intercrossing with parent forms is prohibited, and nothing can be more satisfactory. The argument, however, sets out with showing that it is in limited areas, or in areas already overstocked with the specific form in question, that the advantages to be derived from diversification will be most pronounced. It is where they "jostle each other most closely" that natural selection will set a premium upon any members of the species which may depart from the common type. Now, inasmuch as this jostling or overcrowding of individuals is a needful condition to the agency of natural selection in the way of diversifying character, must we not feel that the general difficulty from intercrossing previously considered is here presented in a special and aggravated form? At all events, I know that, after having duly and impartially considered the matter, to me it does appear that unless the swamping effects of intercrossing with the parent form on an overcrowded area is in some way prevented to begin with, natural selection could never have any material supplied by which to go on with. Let it be observed that I regard Mr. Darwin's argument as perfectly sound where it treats of the divergence of species, and of their further divergence into genera; for in these cases the physiological barrier is known to be already present. But in applying the argument to explain the divergence of individuals into varieties, it seems to me that here, more than anywhere else, Mr. Darwin has strangely lost sight of the formidable difficulty in question; for in this particular case so formidable does the difficulty seem to me, that I cannot believe that natural selection alone could produce any divergence of specific character, so long as all the individuals on an overcrowded area occupy that area together. Yet, if any of them quit that area, and so escape from the unifying influence of free intercrossing, these individuals also escape from the conditions which Mr. Darwin names as those that are needed by natural selection in order to produce divergence. Therefore, it appears to me that, under the circumstances supposed, natural selection alone could not produce divergence; the most it could do would be to change the whole specific type in some one direction, and thus induce transmutation of species in a linear series, each succeeding member of which might supplant its parent form. But in order to secure diversity, multiplication, or ramification of species, it appears to me obvious that the primary condition required is that of preventing intercrossing with parent forms at the origin of each branch, whether the prevention be from the first absolute, or only partial.

Now for Mr. Gulick, a portion of whose more lengthy discussion of the subject, however, is all that I need quote:—

Having found that the evolution of the fitted is secured through the prevention of crossing between the better fitted and the less fitted, can we believe that the evolution of a special race, regularly transmitting a special kind of fitness, can be realized without any prevention of crossing with other races that have no power to transmit that special kind of fitness? Can we suppose that any advantage, derived from new powers that prevent severe competition with kindred, can be permanently transmitted through succeeding generations to one small section of the species while there is free crossing equally distributed between all the families of the species? Is it not apparent that the terms of this supposition are inconsistent with the fundamental laws of heredity? Does not inheritance follow the lines of consanguinity; and when consanguinity is widely diffused, can inheritance be closely limited? When there is free crossing between the families of one species, will not any peculiarity that appears in one family either be neutralized by crosses with families possessing the opposite quality, or, being preserved by natural selection, while the opposite quality is gradually excluded, will not the new quality gradually extend to all the branches of the species; so that, in this way or in that, increasing divergence of form will be prevented?

If the advantage of freedom from competition in any given variation depends on the possession, in some degree, of new adaptations to unappropriated resources, there must be some cause that favours the breeding together of those thus specially endowed, and interferes in some degree with their crossing with other variations, or, failing this, the special advantage will in succeeding generations be lost. As some degree of Independent Generation is necessary for the continuance of the advantage, it is evident that the same condition is necessary for the accumulation through Natural Selection of the powers on which the advantage depends. The advantage of divergence of character cannot be retained by those that fail to retain the divergent character; and divergent character cannot be retained by those that are constantly crossing with other kinds; and the prevention of free crossing between those that are equally successful is in no way secured by Natural Selection.

So much, then, as expressive of Mr. Gulick's opinion upon this subject. To exactly the same effect Professor Lloyd Morgan has recently published his judgement upon it thus:—

That perfectly free intercrossing, between any or all of the individuals of a given group of animals, is, so long as the characters of the parents are blended in the offspring, fatal to divergence of character, is undeniable. Through the elimination of less favourable variations, the swiftness, strength, and cunning of a race may be gradually improved. But no form of elimination can possibly differentiate the group into swift, strong, and cunning varieties, distinct from each other, so long as all three varieties freely interbreed, and the characters of the parents blend in the offspring. Elimination may and does give rise to progress in any given group, as a group; it does not and cannot give rise to differentiation and divergence, so long as interbreeding with consequent interblending of characters be freely permitted. Whence it inevitably follows, as a matter of simple logic, that where divergence has occurred, intercrossing and interbreeding must in some way have been lessened or prevented. Thus a new factor is introduced, that of isolation or segregation. And there is no questioning the fact that it is of great importance. Its importance, indeed, can only be denied by denying the swamping effects of intercrossing, and such denial implies the tacit assumption that interbreeding and interblending are held in check by some form of segregation. The isolation explicitly denied is implicitly assumed[51].

Similarly, and still more recently, Professor Le Conte writes:—

It is evident, then, as Romanes claims, that natural selection alone tends to monotypic evolution. Isolation of some sort seems necessary to polytypic evolution. The tree of evolution under the influence of natural selection alone grows palm-like from its terminal bud. Isolation was necessary to the starting of lateral buds, and thus for the profuse ramification which is its most conspicuous character[52].

In order to complete this historical review, it only remains to consider Mr. Wallace's utterances upon the subject.

It is needless to say that he stoutly resists the view of Weismann, Delbœuf, Gulick, and myself, that specific divergence can ever be due—or, as I understand him, even so much as assisted—by this principle of indiscriminate isolation (apogamy). It will be remembered, however, that Mr. Gulick has adduced certain general principles and certain special facts of geographical distribution, in order to prove that apogamy eventually leads to divergence of character, provided that the isolated section of the species does not contain any very large number of individuals. Now, Mr. Wallace, without making any reference to this argument of Mr. Gulick, simply states the reverse—namely, that, as a matter of fact, indiscriminate isolation is not found to be associated with divergence of character. For, he says, "there is an entire absence of change, where, if this were a vera causa, we should expect to find it[53]." But the only case which he gives is that of Ireland.

This, he says, furnishes "an excellent test case, for we know that it [Ireland] has been separated from Britain since the end of the glacial epoch: ... yet hardly one of its mammals, reptiles, or land molluscs has undergone the slightest change[54]." Here, however, Mr. Wallace shows that he has failed to understand "the views of those who, like Mr. Gulick, believe isolation itself to be a cause of modification of species"; for it belongs to the very essence of these views that the efficiency of indiscriminate isolation as a "vera causa" of organic evolution varies inversely with the number of individuals (i. e. the size of the species-section) exposed to its influence. Therefore, far from being "an excellent test case," the case of Ireland is unsatisfactory. If we are in search of excellent test cases, in the sense intended by Mr. Wallace, we ought not to choose a large island, which from the time of its isolation must have contained large bulks of each of the geographically separated species concerned: we ought to choose cases where as small a number as possible of the representatives of each species were in the first instance concerned. And, when we do this, the answer yielded by any really "excellent test case" is unequivocal.

No better test case of this kind has ever been furnished than that of Mr. Gulick's land-shells, which Mr. Wallace is specially considering in the part of his book where the sentence above quoted occurs. How, then, does he meet this case? He meets it by assuming that in all the numerous adjacent valleys of a small island there must be as many differences of environment, each of which is competent to induce slight varietal changes on the part of its occupants by way of natural selection, although in no one case can the utility of these slight changes be surmised. Now, against this explanation there are three overwhelming considerations. In the first place, it is purely gratuitous, or offered merely in order to save the hypothesis that there can be no other cause of even the most trivial change in species than that which is furnished by natural selection. In the second place, as Mr. Gulick writes to me in a private letter, "if the divergence of Sandwich Island land molluscs is wholly due to exposure to different environments, as Mr. Wallace argues on pages 147-150, then there must be completely occult influences in the environment that vary progressively with each successive mile. This is so violent an assumption that it throws doubt on any theory that requires such support." In the third place, the assumption that the changes in question must have been due to natural selection, is wholly incompatible with the facts of isolation elsewhere—namely, in those cases where (as in that of Ireland) a large section of species, instead of a small section, has been indiscriminately isolated. Mr. Wallace, as we have seen, inadvertently alludes to these "many other cases of isolation" as evidence against apogamy being per se a cause of specific change. But although, for the reason above stated, they are without relevancy in this respect, they appear to me fatal to the explanation which he gives of specific changes under apogamy where only small sections of species are concerned. For example, can it be rationally maintained that there are more differences of environment between every two of the many contiguous valleys of a small island, such as Mr. Gulick describes, than there are in the incomparably larger area of the whole of Ireland? But, if not, and if natural selection is able to work such "occult" wonders in each successive mile on the Sandwich Islands, why has it so entirely lost this magic power in the case of Ireland—or in the "many other cases of isolation" to which Mr. Wallace refers? On his theory there is no coherent answer to be given to this question, while on our theory the answer is given in the very terms of the theory itself. The facts are plainly just what the theory requires that they should be; and therefore, if they were not as they are, the theory would be deprived of that confirmation which it now derives from them.

Thus, in truth, though in an opposite way, the case of Ireland is, as Mr. Wallace says, "an excellent test case," when once the theory of apogamy as a "vera causa" of specific change is understood; and the effect of applying the test is fully to corroborate this theory, while at the same time it as fully negatives the other. For the consideration whereby Mr. Wallace seeks to explain the inactivity of natural selection in the case of Ireland is not "coherent." What he says is, "That changes have not occurred through natural selection, is perhaps due to the less severe struggle for existence, owing to the smaller number of competing species[55]." But even with regard to molluscs alone, there is a greatly larger number of species in Ireland than occurs in any one valley of the Sandwich Islands; while if we have regard to all the other classes of animal life, comparison entirely fails.

Much more to the point are certain cases which were adduced long ago by Weismann in his essay previously considered. Nevertheless, although this essay was published as far back as 1872, and, although it expressly deals with the question of divergence of character through the mere prevention of intercrossing (Amixia), Mr. Wallace nowhere alludes to these cases per contra, which are so much more weighty than his own "test case" of Ireland. Of such are four species of butterflies, belonging to three genera[56], which are identical in the polar regions and in the Alps, notwithstanding that the sparse Alpine populations have been presumably separated from their parent stocks since the glacial period; or of certain species of fresh water crustaceans (Apus), the representatives of which are compelled habitually to form small isolated colonies in widely separated ponds, and nevertheless exhibit no divergence of character, although apogamy has probably lasted for centuries. These cases are unquestionably of a very cogent nature, and appear of themselves to prove that apogamy alone is not invariably capable of inducing divergence—at any rate, so rapidly as we might expect. There appears, however, to be another factor, the presence or absence of which makes a great difference. This as stated in the text, is the degree in which a specific type is stable or unstable—liable or not liable to vary. Thus, for example, the Goose is what Darwin calls an "inflexible" type as compared with most other domesticated birds. Therefore, if a lot of geese were to be indiscriminately isolated from the rest of their species, the probability is that in a given time their descendants would not have diverged from the parent type to such an extent as would a similar lot of ducks under similar circumstances: the more stable specific type would require a longer time to change under the influence of apogamy alone. Now, the butterflies and crustaceans quoted by Weismann may be of a highly stable type, presenting but a small range of individual variability; and, if so, they would naturally require a long time to exhibit any change of type under the influence of apogamy alone. But, be this as it may, Weismann himself adduces these cases merely for the sake of showing that there are cases which seem to tell against the general principle of modification as due to apogamy alone—i.e. the general principle which, under the name amixia, he is engaged in defending. And the conclusion at which he himself arrives is, that while it would be wrong to affirm that apogamy must in all cases produce divergence, we are amply justified in affirming that in many cases it may have done so; while there is good evidence to prove that in not a few cases it has done so, and therefore should be accepted as one of the factors of organic evolution[57].

My view from the very first has been that variations in the way of cross-infertility are of frequent occurrence (how, indeed, can they be otherwise, looking to the complex conditions that have to be satisfied in every case of full fertility?); and, therefore, however many of such variations are destined to die out, whenever one arises, "under suitable conditions," "it must inevitably tend to be preserved as a new natural variety, or incipient species." Among the higher animals—which are "comparatively few in number"—I think it probable that some slight change of form, colour, habit, &c., must be usually needed either to "superinduce," or, which is quite a different thing, to coincide with the physiological change But in the case of plants and the lower invertebrata. I see no reason for any frequent concomitance of this kind; and therefore believe the physiological change to be, "as a general rule," the primordial change. At the same time, I have always been careful to insist that this opinion had nothing to do with "the essence of physiological selection"; seeing that "it was of no consequence" to the theory in what proportional number of cases the cross-sterility had begun per se, had been superinduced by morphological changes, or only enabled to survive by happening to coincide with any other form of homogamy. In short, "the essence of physiological selection" consists in all cases of the diversifying effect of cross-infertility, whensoever and howsoever it may happen in particular cases to have been caused.

Thus I emphatically reaffirm that "from the first I have always maintained that it makes no essential difference to the theory in what proportional number of cases they [the physiological variations] have arisen 'alone in an otherwise undifferentiated species'"; therefore, "even if I am wrong in supposing that physiological selection can ever act alone, the principle of physiological selection, as I have stated it, is not thereby affected. And this principle is, as Mr. Wallace has re-stated it, 'that some amount of infertility characterizes the distinct varieties which are in process of differentiation into species'—infertility whose absence, 'to obviate the effects of intercrossing, may be one of the usual causes of their failure to become developed into distinct species.'"

These last sentences are quoted from the correspondence in Nature[58], and to them Mr. Wallace replied by saying, "if this is not an absolute change of front, words have no meaning"; that "if this is 'the whole essence of physiological selection,' then physiological selection is but a re-statement and amplification of Darwin's views"; that such a "change of front" is incompatible, not only with my term "physiological selection," but also with my having "acknowledged that Mr. Catchpool had 'very clearly put forward the theory of physiological selection'"; and much more to the same effect.

Now, to begin with, it is due to Mr. Catchpool to state that his only publication upon this subject is much too brief to justify Mr. Wallace's, inference, that he supposes variations in the way of cross-infertility always to arise "alone in an otherwise undifferentiated species." What Mr. Catchpool's opinion on this point may be, I have no knowledge; but, whatever it is, he was unquestionably the first writer who "clearly stated the leading principles" of physiological selection, and this fact I am very glad to have "acknowledged." In my correspondence with Mr. Wallace, however, I not only named Mr. Catchpool: I also named—and much more prominently—Mr. Gulick. For even if I were to grant (which I am far indeed from doing) that there was any want of clearness in my own paper touching the point in question, I have now repeatedly shown that it is simply impossible for any reader of Mr. Gulick's papers to misunderstand his views with regard to it. Accordingly, I replied to Mr. Wallace in Nature by saying:—

Not only have I thus from the first fully recognized the sundry other causes of specific change with which the physiological variations may be associated; but Mr. Gulick has gone into this side of our common theory much more fully, and elaborately calculated out the high ratio in which the differentiating agency of any of these other causes must be increased when assisted by—i. e. associated with—even a moderate degree of the selective fertility, and vice versa. Therefore, it is simply impossible for Mr. Wallace to show that "our theory" differs from his in this respect. Yet it is the only respect in which his reply alleges any difference. (Vol. xliii. p. 127.)

I think it is to be regretted that, in his answer to this, Mr. Wallace alludes only to Mr. Catchpool, and entirely ignores Mr. Gulick—whose elaborate calculations above alluded to were communicated to the Linnaean Society by Mr. Wallace himself in 1887.

The time has now come to prove, by means of quotations, that I have from the first represented the "principle," or "essence," of physiological selection to consist in selective fertility furnishing a needful condition to specific differentiation, in at least a large proportional number of allied species which afterwards present the reciprocal character of cross-sterility; that I have never represented variations in the way of this selective fertility as necessarily constituting the initial variations, or as always arising "alone, in an otherwise undifferentiated species"; and that, although I have uniformly given it as my opinion that these variations do in some cases thus arise (especially among plants and lower invertebrata), I have as uniformly stated "that it makes no difference to the theory in what proportional number of cases they have done so"—or even if, as Mr. Wallace supposes, they have never done so in any case at all[59]. These statements (all of which are contradictory of the only points of difference alleged) have already been published in my article in the Monist of October, 1890. And although Mr. Wallace, in his reply to that article, ignores my references to the "original paper," it is scarcely necessary to quote the actual words of the paper itself, since the reader who is further interested in this controversy can readily refer to it in the Journal of the Linnaean Society (vol. xix. pp. 337-411).

Having arrived at these results with regard to the theory of Isolation in general and of Physiological Isolation in particular, I arrive also at the end of this work. And if, while dealing with the post-Darwinian period, I have imparted to any general reader the impression that there is still a great diversity of expert opinion; I must ask him to note that points with reference to which disagreement still exists are but very subordinate to those with regard to which complete agreement now prevails. The noise of wrangling disputations which has so filled the camp of evolutionists since the death of their captain, is apt to hide from the outside world the solid unanimity that prevails with regard to all the larger and more fundamental questions, which were similarly the subjects of warfare in the past generation. Indeed, if we take a fair and general view of the whole history of Darwinism, what must strike us as the really significant fact is the astonishing unanimity which has been so rapidly attained with regard to matters of such immeasurable importance. It is now but little more than thirty years since the publication of the Origin of Species; and in that period not only have all naturalists unequivocally embraced the doctrine of descent considered as a fact; but, in one degree or another, they have all as unequivocally embraced the theory of natural selection considered as a method. The only points with regard to which any difference of opinion still exist, have reference to the precise causation of that mighty stream of events which, under the name of organic evolution, we have now all learnt to accept as scientifically demonstrated. But it belongs to the very nature of scientific demonstration that, where matters of great intricacy as well as of high generality are concerned, the process of demonstration must be gradual, even if it be not always slow. It is only by the labours of many minds working in many directions that, in such cases, truth admits of being eventually displayed. Line upon line, precept upon precept, here a little and there a little—such is the course of a scientific revelation; and the larger the subject-matter, the more subtle and the more complex the causes, the greater must be the room for individual differences in our reading of the book of Nature. Now, if all this be true, must we not feel that in the matter of organic evolution the measure of agreement which has been attained is out of all proportion to the differences which still remain—differences which, although of importance in themselves, are insignificant when compared with those which once divided the opinions of not a few still living men? And if we are bound to feel this, are we not bound further to feel that the very intensity of our disputations over these residual matters of comparative detail, is really the best earnest that can be given of the determination of our quest—determination which, like that of our fathers, cannot fail to be speedily rewarded by the discovery of truth?

Nevertheless, so long as this noise of conflict is in the Senate, we cannot wonder if the people are perplexed. Therefore, in conclusion, I may ask it to be remembered exactly what are the questions—and the only questions—which still divide the parties.

Having unanimously agreed that organic evolution is a fact and that natural selection is a cause, or a factor in the process, the primary question in debate is whether natural selection is the only cause, or whether it has been assisted by the co-operation of other causes. The school of Weismann maintain that it is the only cause; and therefore deem it worse than useless to search for further causes. With this doctrine Wallace in effect agrees, excepting as regards the particular case of the human mind. The school of Darwin, on the other hand—to which I myself claim to belong—believe that natural selection has been to a considerable extent supplemented by other factors; and, therefore, although we further believe that it has been the "main" factor, we agree with Darwin himself in strongly reprobating all attempts to bar a priori the progress of scientific investigation touching what, if any, these other factors may be. Lastly, there are several more or less struggling schools, chiefly composed of individual members who agree with each other only to the extent of holding that the causal agency of natural selection is not so great as Darwin supposed. The Duke of Argyll, Mr. Mivart and Mr. Geddes may be named in this connexion; together with the self-styled neo-Lamarckians, who seek to magnify the Lamarckian principles at the expense of the distinctively Darwinian.

This primary difference of opinion leads deductively to certain secondary differences. For if a man starts with the premiss that natural selection must necessarily be the "exclusive" cause of organic evolution, he is likely to draw conclusions which another man would not draw who starts with the premiss that natural selection is but the "main" cause. Of these subordinate differences the most important are those which relate to the possible transmission of acquired characters, to the necessary (or only general) utility of specific characters, and to the problem touching the inter-sterility of allied species. But we may well hope that before another ten years shall have passed, even these still outstanding questions will have been finally settled; and thus that within the limits of an ordinary lifetime the theory of organic evolution will have been founded and completed in all its parts, to stand for ever in the world of men as at once the greatest achievement in the history of science, and the most splendid monument of the nineteenth century.

In the later chapters of the foregoing treatise I have sought to indicate certain matters of general principle, which many years of study specially devoted to this great movement of contemporary thought have led me to regard as almost certainly sound in themselves, and no less certainly requisite as complements of the Darwinian theory. I will now conclude by briefly summarizing these matters of general principle in the form of twelve sequent propositions. And, in doing so, I may ask it to be noticed that the system which these propositions serve to express may now claim, at the least, to be a strictly logical system. For the fact that, not merely in its main outlines, but likewise in its details, it has been independently constructed by Mr. Gulick, proves at any rate this much; seeing that, where matters of such intricacy are concerned, nothing but accurate reasoning from a common foundation of data could possibly have yielded so exact an agreement. The only difference between us is, that Mr. Gulick has gone into much further detail than I have ever attempted in the way of classifying the many and varied forms of isolation; while I have laid more special stress upon the physiological form, and found in it what appears to me a satisfactory solution of "the greatest of all the difficulties in the way of accepting the theory of natural selection as a complete explanation of the origin of species"—namely, "the remarkable difference between varieties and species when crossed."