The female _bryoniae_ is totally unlike the others. The ground colour
is a full yellow, and each nervure is thickly irrorated with a brown
pigment often spreading so far as to hide the ground almost entirely
in the fore-wings. The males corresponding with these females are not
certainly distinguishable from those of our own _napi_. Both sexes have
the green veining of the underside of the hind wing fully developed,
rather more than is usual in the lowland races, but this is not really
diagnostic of the variety. The first serious difficulty arises in regard
to the second brood of _bryoniae_. It is stated that there is only one
brood,[16] but I feel fairly sure that a second brood is sometimes
produced, and that the females with a yellow ground and diminished
irroration of the veins, not very uncommon in the Italian Alps in July
to August, are generally representatives of it. Such insects would of
course be classed with _bryoniae_ in collections.
My experiments began with eggs of true _bryoniae_ females caught at
about 2,500 feet early in July. These emerged in August-September as
intermediates with yellow ground and about half as much black on the
upper surface as _bryoniae_. They are exactly like the intermediates
usually found in nature and in the light of later experience I regard
them as natural F_{1} forms, and I think the mothers had been fertilised
by _napi_ males, though I admit that in view of the rarity of natural
intermediates there is a difficulty in this suggestion. Three of these
females were mated with males raised from thorough _meridionalis_
females, and three families were produced. Two of them showed distinct
evidence of segregation, some being yellow and some white with various
intergrades, some being no blacker than _meridionalis_ and some ranging
up to a dark intermediate type. Part emerged in the same autumn; and
part overwintered, emerging as the spring _meridionalis_ or as the
peculiar type which I afterwards learnt to know as the spring F_{1}
form. The distinctions were fairly sharp between the several forms. But
the offspring of the third female gave a series practically continuous
from _meridionalis_ to the F_{1} type. The work of subsequent years
gave results similarly irregular which could only be described
adequately at great length. The outcome may however be summed up in
the statement that there is evidence that both the yellow ground and
the dark veining are due to factors, but that there are several of
these and that imperfect segregation is not uncommon, producing various
reduction-stages. The yellow ground may be due to one factor, and the
several shades may be the result of irregularities in dominance, but
the black markings when fully developed cannot I think be the result of
less than three factors, one for the basal darkening, one for general
irroration, and one for the margins. Probably also the enlargement of
the spots is produced by a fourth factor.
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