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.
4. Wide-Ranging, Much Diffused, And Common Species Vary Most
Guided by theoretical considerations, I thought that some interesting results might be obtained in regard to the nature and relations of the species which vary most, by tabulating all the varieties in several well-worked floras. At first this seemed a simple task; but Mr. H.C. Watson, to whom I am much indebted for valuable advice and assistance on this subject, soon convinced me that there were many difficulties, as did subsequently Dr. Hooker, even in stronger terms. I shall reserve for a future work the discussion of these difficulties, and the tables of the proportional numbers of the varying species. Dr. Hooker permits me to add that after having carefully read my manuscript, and examined the tables, he thinks that the following statements are fairly well established. The whole subject, however, treated as it necessarily here is with much brevity, is rather perplexing, and allusions cannot be avoided to the “struggle for existence,” “divergence of character,” and other questions, hereafter to be discussed.
Alphonse de Candolle and others have shown that plants which have very wide ranges generally present varieties; and this might have been expected, as they are exposed to diverse physical conditions, and as they come into competition (which, as we shall hereafter see, is a far more important circumstance) with different sets of organic beings. But my tables further show that, in any limited country, the species which are the most common, that is abound most in individuals, and the species which are most widely diffused within their own country (and this is a different consideration from wide range, and to a certain extent from commonness), oftenest give rise to varieties sufficiently well-marked to have been recorded in botanical works. Hence it is the most flourishing, or, as they may be called, the dominant species—those which range widely, are the most diffused in their own country, and are the most numerous in individuals—which oftenest produce well-marked varieties, or, as I consider them, incipient species. And this, perhaps, might have been anticipated; for, as varieties, in order to become in any degree permanent, necessarily have to struggle with the other inhabitants of the country, the species which are already dominant will be the most likely to yield offspring, which, though in some slight degree modified, still inherit those advantages that enabled their parents to become dominant over their compatriots. In these remarks on predominence, it should be understood that reference is made only to the forms which come into competition with each other, and more especially to the members of the same genus or class having nearly similar habits of life. With respect to the number of individuals or commonness of species, the comparison of course relates only to the members of the same group. One of the higher plants may be said to be dominant if it be more numerous in individuals and more widely diffused than the other plants of the same country, which live under nearly the same conditions. A plant of this kind is not the less dominant because some conferva inhabiting the water or some parasitic fungus is infinitely more numerous in individuals, and more widely diffused. But if the conferva or parasitic fungus exceeds its allies in the above respects, it will then be dominant within its
