It must be obvious to every one, that in a field of a certain size, beside the corn-plants which have been sown, a great number of weeds can exist, and, moreover, in places which could not have been occupied by corn-plants. The more dry and sterile places of the ground, in which no corn-plant would thrive, may still furnish sustenance to weeds of different kinds; and such species and individuals of weeds will more readily be able to exist in such conditions, in proportion as they are suited to adapt themselves to the different parts of the ground. It is the same with animals. It is evident that a much greater number of animal individuals can live together in one and the same limited district, if they are of various and different natures, than if they are all alike. There are trees (for example, the oak) on which a couple of hundred of different species of insects live together. Some feed on the fruits of the tree, others on the leaves, others again on the bark, the root, etc. It would be quite impossible for an equal number of individuals to live on this tree if all were of one species; if, for example, all fed on the bark, or only upon the leaves. Exactly the same is the case in human society. In one and the same small town, only a certain number of workmen can exist, even when they follow different occupations. The division of labour, which is of the greatest use to the whole community, as well as to the individual workman, is a direct consequence of the struggle for life, of natural selection; for this struggle can be sustained more easily the more the activities, and hence, also, the forms of the different individuals deviate from one another. The different function naturally produces its reaction in changing the form, and the physiological division of labour necessarily determines the morphological differentiation, that is, the “divergence of character.”[(37)]
Now, I beg the reader again to remember that all species of animals and plants are variable, and possess the capability of adapting themselves to different places or to local relations. The varieties or races of each species, according to the laws of adaptation, deviate all the more from the original primary species, the greater the difference of the new conditions to which they adapt themselves. If we imagine these varieties—which have proceeded from a common primary form—to be disposed in the shape of a branching, radiating bunch, then those varieties will be best able to exist side by side and propagate which are most distant from one another, which stand at the ends of the series, or at the opposite sides of the bunch. Those forms, on the other hand, occupying a middle position—presenting a state of transition—have the most difficult position in the struggle for life. The necessaries of life differ most in the two extremes, in the varieties most distant from one another, and consequently these will get into the least serious conflict with one another in the general struggle for life. But the intermediate forms, which have deviated less from the original primary form, require nearly the same necessaries of life as the original form, and therefore, in competing for them, they will have to struggle most with, and be most seriously threatened by, its members. Consequently, when numerous varieties of a species live side by side on the same spot of the earth, the extremes, or those forms deviating most from one another, can much more easily continue to exist beside one another than the intermediate forms which have to struggle with each of the different extremes. The intermediate forms will not be able to resist, for any length of time, the hostile influences which the extreme forms victoriously overcome. These alone maintain and propagate themselves, and at length cease to be any longer connected with the original primary species through intermediate forms of transition. Thus arise “good species” out of varieties. Thus, then, the struggle for life necessarily favours the general divergence of organic forms, that is, the constant tendency of organisms to form new species. This fact does not rest upon any mystic quality, or upon an unknown formative tendency, but upon the interaction of Inheritance and Adaptation in the struggle for life. As the intermediate forms, that is, the individuals in a state of transition, of the varieties of every species die out and become extinct, the process of divergence constantly goes further, and from the extremes forms develop which we distinguish as new species.
Although all naturalists have been obliged to acknowledge the variability and mutability of all species of animals and plants, yet most of them have hitherto denied that the modification or transformation of the organic form surpasses the original limit of the characters of the species. Our opponents cling to the proposition—“However far a species may exhibit deviations from its usual form in a collection of varieties, yet the varieties of it are never so distinct from one another as two really good species.” This assertion, which Darwin’s opponents usually place at the head of their arguments, is utterly untenable and unfounded. This will become quite clear as soon as we critically compare the various attempts to define the idea of species. No naturalist can answer the question as to what is in reality a “genuine or good species” (“bona species”); yet every systematic naturalist uses this expression every day, and whole libraries have been written on the question as to whether this or that observed form is a species or a variety, whether it is a really good or a bad species. The most general answer to this question used to be the following: “To one species belong all those individuals which agree in all essential characteristics. Essential characteristics of species are those which remain permanent or constant, and never become modified or vary.” But as soon as a case occurred in which the characteristic—which had hitherto been considered essential—did become modified, then it was said, “This characteristic is not essential to the species, for essential characteristics never vary.” Those who argued thus evidently moved in a circle, and the naïveté with which this circular method of defining species is laid down in thousands of books as an unassailable truth, and is still constantly repeated, is truly astonishing.
All other attempts which have been made to arrive at a definite and logical determination of the idea of organic “species” have, like the last, been utterly futile, and led to no results. Considering the nature of the case, it cannot be otherwise. The idea of species is just as truly a relative one and not absolute, as is the idea of variety, genus, family, order, class, etc. I have proved this in detail in the criticism of the idea of species in my “General Morphology” (Gen. Morph. ii. 323-364). I will waste no more time on this unsatisfactory discussion, and now only add a few words about the relation of species to hybridism. Formerly it was regarded as a dogma, that two good species could never produce hybrids which could reproduce themselves as such. Those who thus dogmatized almost always appealed to the hybrids of a horse and donkey, the mule and the hinny, which, truly enough, are seldom able to reproduce themselves. But the truth is that such unfruitful hybrids are rare examples, and in the majority of cases hybrids of two totally different species are fruitful and able to reproduce themselves. They can almost always fruitfully mix with one or other of the parent species, and sometimes also among themselves; and in this way completely new forms can originate according to the laws of “mixed transmission by inheritance.”
Thus, in fact, hybridism is a source of the origin of new species, distinct from the source we have hitherto considered—natural selection. I have already spoken occasionally of these hybrid species (species hybridæ), especially of the hare-rabbit (Lepus Darwinii), which has arisen from the crossing of a male hare and a female rabbit; the goat-sheep (Capra ovina), which has arisen from the pairing of a he-goat and ewe; also the different species of thistles (Cirsium), brambles (Rubus), etc. It is possible that many wild species have originated in this way, as even Linnæus assumed. At all events, these hybrid species, which can maintain and propagate themselves as well as pure species, prove that hybridism cannot serve in any way to give an absolute definition to the idea of species.
I have already mentioned (p. 47) that the many vain attempts to define the idea of species theoretically have nothing whatever to do with the practical distinction of species. The extensive practical application of the idea of species, as it is carried out in systematic zoology and botany, is very instructive as furnishing an example of human folly. Hitherto, by far the majority of zoologists and botanists, in distinguishing and describing the different forms of animals and plants, have endeavoured, above all things, to distinguish accurately kindred forms as so many “good species.” However, it has been found scarcely possible, in any group, to make an accurate and consistent distinction of such “genuine or good species.” There are no two zoologists, no two botanists, who agree in all cases as to which of the nearly related forms of a genus are good species, and which are not. All authors have different views about them. In the genus Hieracium, for example, one of the commonest genera of European plants, no less than 300 species have been distinguished in Germany alone. The botanist Fries, however, only admits 106, Koch only 52, as “good species,” and others accept scarcely 20. The differences in the species of brambles (Rubus) are equally great. Where one botanist makes more than a hundred species, a second admits only about one half of that number, a third only five or six, or even fewer species. The birds of Germany have long been very accurately known. Bechstein, in his careful “Natural History of German Birds,” has distinguished 367 species, L. Reichenbach 379, Meyer and Wolff 406, and Brehm, a clergyman learned in ornithology, distinguishes even more than 900 different species.
Thus we see that here, and, in fact, in every other domain of systematic zoology and botany, the most arbitrary proceedings prevail, and, from the nature of the case, must prevail. For it is quite impossible accurately to distinguish varieties and races from so-called “good species.” Varieties are commencing species. The variability or adaptability of species, under the influence of the struggle for life, necessitates the continual and progressive separation or differentiation of varieties, and the perpetual delimitation of new forms. Whenever these are maintained throughout a number of generations by inheritance, whilst the intermediate forms die out, they form independent “new species.” The origin of new species by division of labour, or separation, divergence, or differentiation of varieties, is therefore a necessary consequence of natural selection.[(37)]
The same kind of interest attaches to a second great law which we deduce from natural selection, and which is, indeed, closely connected with the law of Divergence, but in no way identical with it; namely, the law of Progress (progressus), or Perfecting (teleosis). (Gen. Morph. ii. 257.) This great and important law, like the law of differentiation, had long been empirically established by palæontological experience, before Darwin’s Theory of Selection gave us the key to the explanation of its cause. The most distinguished palæontologists have pointed out the law of progress as the most general result of their investigations of fossil organisms. This has been specially done by Bronn, whose investigations on the laws of construction[(18)] and the laws of the development[(19)] of organisms, although little heeded, are excellent, and deserve most careful consideration. The general results of the law of differentiation and the law of progress, at which Bronn arrived by a purely mechanical hypothesis, and by exceedingly accurate, laborious, and careful investigations, are brilliant confirmations of the truth of these two great laws which we deduce as necessary inferences from the theory of selection.
The law of progress or of perfecting establishes the exceedingly important fact, on the ground of palæontological experience, that in successive periods of this earth’s history, a continual increase in the perfection of organic formations has taken place. Since that inconceivably remote period in which life on our planet began with the spontaneous generation of Monera, organisms of all groups, both collectively as well as individually, have continually become more perfectly and highly developed. The steadily increasing variety of living forms has always been accompanied by progress in organization. The lower the strata of the earth in which the remains of extinct animals and plants lie buried, that is, the older the strata are, the more simple and imperfect are the forms which they contain. This applies to organisms collectively, as well as to every single large or small group of them, setting aside, of course, those exceptions which are due to the process of degeneration, which we shall discuss hereafter.
As a confirmation of this law I shall mention only the most important of all animal groups, the tribe of vertebrate animals. The oldest fossil remains of vertebrate animals known to us belong to the lowest class, that of Fishes. Upon these there followed later more perfect Amphibious animals, then Reptiles, and lastly, at a much later period, the most highly organized classes of vertebrate animals, Birds and Mammals. Of the latter only the lowest and most imperfect forms, without placenta, appeared at first, such as are the pouched animals (Marsupials), and afterwards, at a much later period, the more perfect mammals, with placenta. Of these, also, at first only the lower kinds appeared, the higher forms later; and not until the late tertiary period did man gradually develop out of these last.