of crossed species does not strictly coincide with their systematic
affinity, that is, with the sum of their external resemblances; nor does it
coincide with their similarity in general constitution. But we are more
especially led to this same conclusion by considering reciprocal crosses,
in which the male of one species cannot be united, or can be united with
extreme difficulty, with the female of a second species, whilst the
converse cross can be effected with perfect facility; for this difference
in the facility of making reciprocal crosses, and in the fertility of their
offspring, must be attributed either to the male or female element in the
first species having been differentiated with reference to the sexual
element of the second species in a higher degree than in the converse case.
In so complex a subject as Hybridism it is of considerable importance thus
to arrive at a definitive conclusion, namely, that the sterility which
almost invariably follows the union of distinct {185} species depends
exclusively on differences in their sexual constitution.
* * * * *
On the principle which makes it necessary for man, whilst he is selecting
and improving his domestic varieties, to keep them separate, it would
clearly be advantageous to varieties in a state of nature, that is to
incipient species, if they could be kept from blending, either through
sexual aversion, or by becoming mutually sterile. Hence it at one time
appeared to me probable, as it has to others, that this sterility might
have been acquired through natural selection. On this view we must suppose
that a shade of lessened fertility first spontaneously appeared, like any
other modification, in certain individuals of a species when crossed with
other individuals of the same species; and that successive slight degrees
of infertility, from being advantageous, were slowly accumulated. This
appears all the more probable, if we admit that the structural differences
between the forms of dimorphic and trimorphic plants, as the length and
curvature of the pistil, &c., have been co-adapted through natural
selection; for if this be admitted, we can hardly avoid extending the same
conclusion to their mutual infertility. Sterility moreover has been
acquired through natural selection for other and widely different purposes,
as with neuter insects in reference to their social economy. In the case of
plants, the flowers on the circumference of the truss in the guelder-rose
(_Viburnum opulus_) and those on the summit of the spike in the
feather-hyacinth (_Muscari comosum_) have been rendered conspicuous, and
apparently in consequence sterile, in order that insects might easily
discover and visit the other flowers. But when we endeavour to apply the
principle of natural selection to the acquirement by distinct species of
mutual sterility, we meet with great difficulties. In the first place, it
may be remarked that separate regions are often inhabited by groups of
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