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CLASSIFICATION, groups subordinate to groups—Natural system— Rules and difficulties in classification, explained on the theory of descent with modification—Classification of varieties—Descent always used in classification—Analogical or adaptive characters-Affinities, general, complex, and radiating—Extinction separates and defines groups—MoRPHOLOGY, between members of the same class, between parts of the same individual— EMBRYoLoGY, laws of, explained by variations not supervening at an early age, and being inherited at a corresponding age—RUDIMENTARY or GANs; their origin explained—Summary . - • • • - • - * . Page 202



Recapitulation of the objections to the theory of Natural Selection -Recapitulation of the general and special circumstances in its favour—Causes of the general belief in the immutability of species—How far the theory of Natural Selection may be extended—Effects of its adoption on the study of Natural History —Concluding remarks . . . . . . . 267


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Distinction between the sterility of first crosses and of hybrids— Sterility various in degree, not universal, affected by close interbreeding, removed by domestication—Laws governing the sterility of hybrids—Sterility not a special endowment, but incidental on other differences, not accumulated by natural selection—Causes of the sterility of first crosses and of hybrids —Parallelism between the effects of changed conditions of life and of crossing—Dimorphism and trimorphism—Fertility of varieties when crossed and of their mongrel offspring not universal—Hybrids and mongrels compared independently of their fertility—Summary.

THE view commonly entertained by naturalists is that species, when intercrossed, have been specially endowed with sterility, in order to prevent their confusion. This view certainly seems at first highly probable, for species living together could hardly have been kept distinct had they been capable of freely crossing. The subject is in many ways important for us, more especially as the sterility of species when first crossed, and that of their hybrid offspring, cannot have been acquired, as I

shall show, by the preservation of successive profitable degrees of sterility. It is an incidental result of differences in the reproductive systems of the parentspecies. In treating this subject, two classes of facts, to a large extent fundamentally different, have generally been confounded; namely, the sterility of species when first crossed, and the sterility of the hybrids produced from them. Pure species have of course their organs of reproduction in a perfect condition, yet when intercrossed they produce either few or no offspring. Hybrids, on the other hand, have their reproductive organs functionally impotent, as may be clearly seen in the state of the male element in both plants and animals; though the formative organs themselves are perfect in structure, as far as the microscope reveals. In the first case the two sexual elements which go to form the embryo are perfect; in the second case they are either not at all developed, or are imperfectly developed. This distinction is important, when the cause of the sterility, which is common to the two cases, has to be considered. The distinction probably has been slurred over, owing to the sterility in both cases being looked on as a special endowment, beyond the province of our reasoning powers. The fertility of varieties, that is of the forms known or believed to be descended from common parents, when crossed, and likewise the fertility of their mongrel offspring, is, with reference to my theory, of equal importance with the sterility of species; for it seems to make a broad and clear distinction between varieties and species. Degrees of Sterility—First, for the sterility of species when crossed and of their hybrid offspring. It is impossible to study the several memoirs and works of those two conscientious and admirable observers, Kölreuter and Gärtner, who almost devoted their lives to this subject, without being deeply impressed with the high generality of some degree of sterility. Kölreuter makes the rule universal; but then he cuts the knot, for in ten cases in which he found two forms, considered by most authors as distinct species, quite fertile together, he unhesitatingly ranks them as varieties. Gärtner, also, makes the rule equally universal; and he disputes the entire fertility of Kölreuter's ten cases. But in these and in many other cases, Gärtner is obliged carefully to count the seeds, in order to show that there is any degree of sterility. He always compares the maximum number of seeds produced by two species when first crossed, and the maximum produced by their hybrid offspring, with the average number produced by both pure parent-species in a state of nature. But causes of serious error here intervene: a plant, to be hybridised, must be castrated, and, what is often more important, must be secluded in order to prevent pollen being brought to it by insects from other plants. Nearly all the plants experimented on by Gärtner were potted, and were kept in a chamber in his house. That these processes are often injurious to the fertility of a plant cannot be doubted; for Gärtner gives in his table about a score of cases of plants which he castrated, and artificially fertilised with their own pollen, and (excluding all cases such as the Leguminosae, in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. Moreover, as Gärtner repeatedly

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