Origin of Species by Charles Darwin
Chapter II. Variation Under Nature
The forms which possess in some considerable degree the character of species, but which are so closely similar to other forms, or are so closely linked to them by intermediate gradations, that naturalists do not like to rank them as distinct species, are in several respects the most important for us. We have every reason to believe that many of these doubtful and closely allied forms have permanently retained their characters for a long time; for as long, as far as we know, as have good and true species. Practically, when a naturalist can unite by means of intermediate links any two forms, he treats the one as a variety of the other, ranking the most common, but sometimes the one first described as the species, and the other as the variety. But cases of great difficulty, which I will not here enumerate, sometimes arise in deciding whether or not to rank one form as a variety of another, even when they are closely connected by intermediate links; nor will the commonly assumed hybrid nature of the intermediate forms always remove the difficulty. In very many cases, however, one form is ranked as a variety of another, not because the intermediate links have actually been found, but because analogy leads the observer to suppose either that they do now somewhere exist, or may formerly have existed; and here a wide door for the entry of doubt and conjecture is opened.
Hence, in determining whether a form should be ranked as a species or a variety, the opinion of naturalists having sound judgment and wide experience seems the only guide to follow. We must, however, in many cases, decide by a majority of naturalists, for few well-marked and well-known varieties can be named which have not been ranked as species by at least some competent judges.
That varieties of this doubtful nature are far from uncommon cannot be disputed. Compare the several floras of Great Britain, of France, or of the United States, drawn up by different botanists, and see what a surprising number of forms have been ranked by one botanist as good species, and by another as mere varieties. Mr. H.C. Watson, to whom I lie under deep obligation for assistance of all kinds, has marked for me 182 British plants, which are generally considered as varieties, but which have all been ranked by botanists as species; and in making this list he has omitted many trifling varieties, but which nevertheless have been ranked by some botanists as species, and he has entirely omitted several highly polymorphic genera. Under genera, including the most polymorphic forms, Mr. Babington gives 251 species, whereas Mr. Bentham gives only 112--a difference of 139 doubtful forms! Among animals which unite for each birth, and which are highly locomotive, doubtful forms, ranked by one zoologist as a species and by another as a variety, can rarely be found within the same country, but are common in separated areas. How many of the birds and insects in North America and Europe, which differ very slightly from each other, have been ranked by one eminent naturalist as undoubted species, and by another as varieties, or, as they are often called, geographical races! Mr. Wallace, in several valuable papers on the various animals, especially on the Lepidoptera, inhabiting the islands of the great Malayan Archipelago, shows that they may be classed under four heads, namely, as variable forms, as local forms, as geographical races or sub-species, and as true representative species. The first or variable forms vary much within the limits of the same island. The local forms are moderately constant and distinct in each separate island; but when all from the several islands are compared together, the differences are seen to be so slight and graduated that it is impossible to define or describe them, though at the same time the extreme forms are sufficiently distinct. The geographical races or sub-species are local forms completely fixed and isolated; but as they do not differ from each other by strongly marked and important characters, "There is no possible test but individual opinion to determine which of them shall be considered as species and which as varieties." Lastly, representative species fill the same place in the natural economy of each island as do the local forms and sub-species; but as they are distinguished from each other by a greater amount of difference than that between the local forms and sub-species, they are almost universally ranked by naturalists as true species. Nevertheless, no certain criterion can possibly be given by which variable forms, local forms, sub species and representative species can be recognised.
Many years ago, when comparing, and seeing others compare, the birds from the closely neighbouring islands of the Galapagos Archipelago, one with another, and with those from the American mainland, I was much struck how entirely vague and arbitrary is the distinction between species and varieties. On the islets of the little Madeira group there are many insects which are characterized as varieties in Mr. Wollaston's admirable work, but which would certainly be ranked as distinct species by many entomologists. Even Ireland has a few animals, now generally regarded as varieties, but which have been ranked as species by some zoologists. Several experienced ornithologists consider our British red grouse as only a strongly marked race of a Norwegian species, whereas the greater number rank it as an undoubted species peculiar to Great Britain. A wide distance between the homes of two doubtful forms leads many naturalists to rank them as distinct species; but what distance, it has been well asked, will suffice if that between America and Europe is ample, will that between Europe and the Azores, or Madeira, or the Canaries, or between the several islets of these small archipelagos, be sufficient?
Mr. B.D. Walsh, a distinguished entomologist of the United States, has described what he calls Phytophagic varieties and Phytophagic species. Most vegetable-feeding insects live on one kind of plant or on one group of plants; some feed indiscriminately on many kinds, but do not in consequence vary. In several cases, however, insects found living on different plants, have been observed by Mr. Walsh to present in their larval or mature state, or in both states, slight, though constant differences in colour, size, or in the nature of their secretions. In some instances the males alone, in other instances, both males and females, have been observed thus to differ in a slight degree. When the differences are rather more strongly marked, and when both sexes and all ages are affected, the forms are ranked by all entomologists as good species. But no observer can determine for another, even if he can do so for himself, which of these Phytophagic forms ought to be called species and which varieties. Mr. Walsh ranks the forms which it may be supposed would freely intercross, as varieties; and those which appear to have lost this power, as species. As the differences depend on the insects having long fed on distinct plants, it cannot be expected that intermediate links connecting the several forms should now be found. The naturalist thus loses his best guide in determining whether to rank doubtful forms as varieties or species. This likewise necessarily occurs with closely allied organisms, which inhabit distinct continents or islands. When, on the other hand, an animal or plant ranges over the same continent, or inhabits many islands in the same archipelago, and presents different forms in the different areas, there is always a good chance that intermediate forms will be discovered which will link together the extreme states; and these are then degraded to the rank of varieties.
Some few naturalists maintain that animals never present varieties; but then these same naturalists rank the slightest difference as of specific value; and when the same identical form is met with in two distant countries, or in two geological formations, they believe that two distinct species are hidden under the same dress. The term species thus comes to be a mere useless abstraction, implying and assuming a separate act of creation. It is certain that many forms, considered by highly competent judges to be varieties, resemble species so completely in character that they have been thus ranked by other highly competent judges. But to discuss whether they ought to be called species or varieties, before any definition of these terms has been generally accepted, is vainly to beat the air.
Many of the cases of strongly marked varieties or doubtful species well deserve consideration; for several interesting lines of argument, from geographical distribution, analogical variation, hybridism, etc., have been brought to bear in the attempt to determine their rank; but space does not here permit me to discuss them. Close investigation, in many cases, will no doubt bring naturalists to agree how to rank doubtful forms. Yet it must be confessed that it is in the best known countries that we find the greatest number of them. I have been struck with the fact that if any animal or plant in a state of nature be highly useful to man, or from any cause closely attracts his attention, varieties of it will almost universally be found recorded. These varieties, moreover, will often be ranked by some authors as species. Look at the common oak, how closely it has been studied; yet a German author makes more than a dozen species out of forms, which are almost universally considered by other botanists to be varieties; and in this country the highest botanical authorities and practical men can be quoted to show that the sessile and pedunculated oaks are either good and distinct species or mere varieties.
I may here allude to a remarkable memoir lately published by A. de Candolle, on the oaks of the whole world. No one ever had more ample materials for the discrimination of the species, or could have worked on them with more zeal and sagacity. He first gives in detail all the many points of structure which vary in the several species, and estimates numerically the relative frequency of the variations. He specifies above a dozen characters which may be found varying even on the same branch, sometimes according to age or development, sometimes without any assignable reason. Such characters are not of course of specific value, but they are, as Asa Gray has remarked in commenting on this memoir, such as generally enter into specific definitions. De Candolle then goes on to say that he gives the rank of species to the forms that differ by characters never varying on the same tree, and never found connected by intermediate states. After this discussion, the result of so much labour, he emphatically remarks: "They are mistaken, who repeat that the greater part of our species are clearly limited, and that the doubtful species are in a feeble minority. This seemed to be true, so long as a genus was imperfectly known, and its species were founded upon a few specimens, that is to say, were provisional. Just as we come to know them better, intermediate forms flow in, and doubts as to specific limits augment." He also adds that it is the best known species which present the greatest number of spontaneous varieties and sub-varieties. Thus Quercus robur has twenty-eight varieties, all of which, excepting six, are clustered round three sub- species, namely Q. pedunculata, sessiliflora and pubescens. The forms which connect these three sub-species are comparatively rare; and, as Asa Gray again remarks, if these connecting forms which are now rare were to become totally extinct the three sub-species would hold exactly the same relation to each other as do the four or five provisionally admitted species which closely surround the typical Quercus robur. Finally, De Candolle admits that out of the 300 species, which will be enumerated in his Prodromus as belonging to the oak family, at least two-thirds are provisional species, that is, are not known strictly to fulfil the definition above given of a true species. It should be added that De Candolle no longer believes that species are immutable creations, but concludes that the derivative theory is the most natural one, "and the most accordant with the known facts in palaeontology, geographical botany and zoology, of anatomical structure and classification."
When a young naturalist commences the study of a group of organisms quite unknown to him he is at first much perplexed in determining what differences to consider as specific and what as varietal; for he knows nothing of the amount and kind of variation to which the group is subject; and this shows, at least, how very generally there is some variation. But if he confine his attention to one class within one country he will soon make up his mind how to rank most of the doubtful forms. His general tendency will be to make many species, for he will become impressed, just like the pigeon or poultry fancier before alluded to, with the amount of difference in the forms which he is continually studying; and he has little general knowledge of analogical variation in other groups and in other countries by which to correct his first impressions. As he extends the range of his observations he will meet with more cases of difficulty; for he will encounter a greater number of closely-allied forms. But if his observations be widely extended he will in the end generally be able to make up his own mind; but he will succeed in this at the expense of admitting much variation, and the truth of this admission will often be disputed by other naturalists. When he comes to study allied forms brought from countries not now continuous, in which case he cannot hope to find intermediate links, he will be compelled to trust almost entirely to analogy, and his difficulties will rise to a climax.
Certainly no clear line of demarcation has as yet been drawn between species and sub-species--that is, the forms which in the opinion of some naturalists come very near to, but do not quite arrive at, the rank of species; or, again, between sub-species and well-marked varieties, or between lesser varieties and individual differences. These differences blend into each other by an insensible series; and a series impresses the mind with the idea of an actual passage.
Hence I look at individual differences, though of small interest to the systematist, as of the highest importance for us, as being the first step towards such slight varieties as are barely thought worth recording in works on natural history. And I look at varieties which are in any degree more distinct and permanent, as steps toward more strongly marked and permanent varieties; and at the latter, as leading to sub-species, and then to species. The passage from one stage of difference to another may, in many cases, be the simple result of the nature of the organism and of the different physical conditions to which it has long been exposed; but with respect to the more important and adaptive characters, the passage from one stage of difference to another may be safely attributed to the cumulative action of natural selection, hereafter to be explained, and to the effects of the increased use or disuse of parts. A well-marked variety may therefore be called an incipient species; but whether this belief is justifiable must be judged by the weight of the various facts and considerations to be given throughout this work.
It need not be supposed that all varieties or incipient species attain the rank of species. They may become extinct, or they may endure as varieties for very long periods, as has been shown to be the case by Mr. Wollaston with the varieties of certain fossil land-shells in Madeira, and with plants by Gaston de Saporta. If a variety were to flourish so as to exceed in numbers the parent species, it would then rank as the species, and the species as the variety; or it might come to supplant and exterminate the parent species; or both might co-exist, and both rank as independent species. But we shall hereafter return to this subject.
>From these remarks it will be seen that I look at the term species as one arbitrarily given, for the sake of convenience, to a set of individuals closely resembling each other, and that it does not essentially differ from the term variety, which is given to less distinct and more fluctuating forms. The term variety, again, in comparison with mere individual differences, is also applied arbitrarily, for convenience sake.