Still, though there are exceptions, the general infertility of allied
species when crossed is a fact in strong contrast with the marked
fertility of varieties under domestication; concerning which, however,
it should be noted that our domesticated animals have been selected to a
very large extent on account of the freedom with which they breed in
confinement, and that domestication has probably a tendency to increase
fertility. The question, therefore, arises--Is the infertility between
species, and the general sterility of their male offspring, a secondary
effect of their segregation? or is their segregation the direct effect
of their differential fertility? The former is the general opinion; the
latter is held by Mr. Romanes. He contends that sterility is the primary
distinction of species, other specific characters being secondary, and
regards it as a pure assumption to say that the secondary differences
between species have been historically prior to the primary difference.
I do not propose to discuss this question. While it seems to me in the
highest degree improbable that differential fertility, apart from other
co-operating factors, has been or could be a practical mode of
segregation, it has probably been a not unimportant factor in
association with other modes of segregation or isolation. Suppose, for
example, two divergent local varieties were to arise in adjacent areas,
and were subsequently (by stress of competition or by geographical
changes) driven together into a single area: we are justified in
believing, from the analogy of the Falkland Island cattle, the Forest of
Dean deer, and other similar observed habits, that preferential
breeding, kind with kind, would tend to keep them apart. But, setting
this on one side, let us say they interbreed. If, then, their unions are
fertile, the isolation will be annulled by intercrossing--the two
varieties will form one mean or average variety. But if the unions be
infertile, the isolation will be preserved, and the two varieties will
continue separate. Suppose now, and the supposition is by no means an
improbable one, that this has taken place again and again in the
evolution of species: then it is clear that those varietal forms which
had continued to be fertile together would be swamped by intercrossing;
while those varietal forms which had become infertile would remain
isolated. Hence, in the long run, isolated forms occupying a common area
would be infertile. Or suppose, once more, that, instead of the unions
between the two varietal forms being infertile, they are fertile, but
give rise to sterile (mule) or degenerate offspring, as is said to be
the case in the unions of Japanese and Ainos: then it is clear that the
sterile or degenerate offspring of such unions would be eliminated, and
intercrossing, even though it occurred, would be inoperative while
breeding within the limits of the variety continued unchecked.
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