CHAPTER II - VARIATION UNDER NATURE.

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 is a far more important circumstance) with different sets of organic beings. But my tables further show that, in any limited country, species which are most common, that is abound most in individuals, and species which are most widely diffused in their own country (this is a different consideration from wide range and to some 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 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 other inhabitants of the country, species which are already dominant will be 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 predominance, it should be understood that reference is made only to forms which come into competition with each other, and more especially to 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 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 other plants of the same country, which live under nearly the same conditions. A plant of this kind is not 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 own class.

Species of the larger genera in each country vary more frequently than species of smaller genera

If the plants inhabiting a country as described in any Flora, be divided into two equal masses, all those in the larger genera (i.e., those including many species) being placed on one side, and all those in the smaller genera on the other side, the former will be found to include a somewhat larger number of the very common and much diffused or dominant species. This might have been anticipated, for the mere fact of many species of the same genus inhabiting any country, shows that there is something in the organic or inorganic conditions of that country favorable to the genus; and, consequently, we might have expected to have found in the larger genera, or those including many species, a larger proportional number of dominant species. But so many causes tend to obscure this result, that I am surprised that my tables show even a small majority on the side of the larger genera. I will here allude to only two causes of obscurity. Fresh water and salt-loving plants generally have very wide ranges and are much diffused, but this seems to be connected with the nature of the stations inhabited by them, and has little or no relation to the size of the genera to which the species belong. Again, plants low in the scale of organization are generally much more widely diffused than plants higher in the scale; and here again there is no close relation to the size of the genera. The cause of lowly-organized plants ranging widely will be discussed in our chapter on Geographical Distribution.

From looking at species as only strongly marked and well-defined varieties, I was led to anticipate that the species of the larger genera in each country would oftener present varieties, than the species of the smaller genera; for wherever many closely related species (i.e., species of the same genus) have been formed, many varieties or incipient species ought, as a general rule, to be now forming. Where many large trees grow, we expect to find saplings. Where many species of a genus have been formed through variation, circumstances have been favorable for variation; and hence we might expect that the circumstances would generally still be favorable to variation. On the other hand, if we look at each species as a special act of creation, there is no apparent reason why more varieties should occur in a group having many species, than in one having few.

To test the truth of this anticipation I have arranged the plants of twelve countries, and coleopterous insects of two districts, into two nearly equal masses, the species of larger genera on one side, and those of smaller genera on the other side, and it has invariably proved to be the case that a larger proportion of the species on the side of the larger genera presented varieties, than on the side of the smaller genera. Moreover, the species of the large genera which present any varieties, invariably present a larger average number of varieties than do the species of the small genera. Both these results follow when another division is made, and when all the least genera, with from only one to four species, are altogether excluded from the tables. These facts are of plain signification on the view that species are only strongly marked and permanent varieties; for wherever many species of the same genus have been formed, or where, if we may use the expression, the manufactory of species has been active, we ought generally to find the manufactory still in action, more especially as we have every reason to believe the process of manufacturing new species to be a slow one. And this certainly holds true if varieties be looked at as incipient species; for my tables clearly show, as a general rule, that, wherever many species of a genus have been formed, the species of that genus present a number of varieties, that is, of incipient species, beyond the average. It is not that all large genera are now varying much, and are thus increasing in the number of their species, or that no small genera are now varying and increasing; for if this had been so, it would have been fatal to my theory; inasmuch as geology plainly tells us that small genera have in the lapse of time often increased greatly in size; and that large genera have often come to their maxima, declined, and disappeared. All that we want to show is, that where many species of a genus have been formed, on an average many are still forming; and this certainly holds good.

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