PDF - Wallace Online
PDF - Wallace Online PDF - Wallace Online
378 TROPICAL NATURENatural Selection as neutralising Sexual SelectionThere is also a general argument against Mr. Darwin'sviews on this question, founded on the nature and potencyof " natural " as opposed to " sexual " selection, which appearsto me to be of itself almost conclusive as to the whole matterat issue. Natural selection, or the survival of the fittest,the off-acts perpetually and on an enormous scale. Takingspring of each pair of birds as, on the average, only sixannually, one-third of these at most will be preserved, whilethe two-thirds which are least fitted will die. At intervals ofa few years, whenever unfavourable conditions occur, fivesixths,nine-tenths, or even a greater proportion of the wholeyearly production are weeded out, leaving only the mostperfect and best adapted to survive. Now unless these survivorsare, on the whole, the most ornamental, this rigidnatural selection must neutralise and destroy any influencethat may be exerted by female selection. The utmost thatcan be claimed for the latter is, that a small fraction of theleast ornamented do not obtain mates, while a few of themost ornamented may leave more than the average number ofoffspring. Unless, therefore, there is the strictest correlationbetween ornament and general perfection, the more brightlycoloured or ornamented varieties can obtain no permanentadvantage and if there is ; (as I maintain) such a correlation,then the sexual selection of colour or ornament, for whichthere is little or no evidence, becomes needless, becausenatural selection, which is an admitted vera causa, will itselfproduce all the results.In the case of butterflies the argument becomes evenstronger, because the fertility is so much greater than inbirds, and the weeding-out of the unfit takes place, to a greatextent, in the egg and larva state. Unless the eggs andlarvae which escaped to produce the next generation werethose which would produce the more highly-coloured butterflies,it is difficult to perceive how the slight preponderanceof colour sometimes selected by the females should not bewholly neutralised by the extremely rigid selection for otherqualities to which the offspring in every stage are exposed.The only way in which we can account for the observed facts
v COLOURS OF ANIMALS 879is, by the supposition that colour and ornament are strictlycorrelated with health, vigour, and general fitness to survive.We have shown that there is reason to believe that this isthe case, and if so, conscious sexual selection becomes asunnecessary as it would certainly be ineffective.Greater Brilliancy of some Female BirdsThere is one other very curious case of sexual colouringamong birds that, namely, in which the female is decidedlybrighter or more strongly marked than the male, as in thefighting quails (Turnix), painted snipe (Ehynchsea), twospecies of phalarope (Phalaropus), and the common cassowary(Casuarius galeatus). In all these cases it is known that themales take charge of and incubate the eggs, while the femalesare almost always larger and more pugnacious.In my "Theory of Birds' Nests" (see p. 132) I imputedthis difference of colour to the greater need for protection bythe male bird while incubating, to which Mr. Darwin hasobjected that the difference is not sufficient, and is not alwaysso distributed as to be most effective for this purpose and he;believes that it is due to reversed sexual selection that is, tothe female taking the usual rdle of the male, and being chosenfor her brighter tints. We have already seen reason forrejecting this latter theory in every case ;and I also admitthat Mr. Darwin's criticism is sound, and 'that my theory ofprotection is, in this case, only partially, if at all, applicable.But the theory now advanced, of intensity of colour beingdue to general vital energy, is quite applicable ;and the factthat the superiority of the female in this respect is quiteexceptional, and is therefore probably not in any case of veryancient date, will account for the difference of colour thusproduced being always very slight.Colour-development as illustrated by Humming-birdsOf the mode of action of the general principles of colourdevelopmentamong animals, we have an excellent example inthe humming-birds. Of all birds these are at once thesmallest, the most active, and the fullest of vital energy.When poised in the air their wings are invisible owing tothe rapidity of their motion, and when startled they dart
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378 TROPICAL NATURENatural Selection as neutralising Sexual SelectionThere is also a general argument against Mr. Darwin'sviews on this question, founded on the nature and potencyof " natural " as opposed to " sexual " selection, which appearsto me to be of itself almost conclusive as to the whole matterat issue. Natural selection, or the survival of the fittest,the off-acts perpetually and on an enormous scale. Takingspring of each pair of birds as, on the average, only sixannually, one-third of these at most will be preserved, whilethe two-thirds which are least fitted will die. At intervals ofa few years, whenever unfavourable conditions occur, fivesixths,nine-tenths, or even a greater proportion of the wholeyearly production are weeded out, leaving only the mostperfect and best adapted to survive. Now unless these survivorsare, on the whole, the most ornamental, this rigidnatural selection must neutralise and destroy any influencethat may be exerted by female selection. The utmost thatcan be claimed for the latter is, that a small fraction of theleast ornamented do not obtain mates, while a few of themost ornamented may leave more than the average number ofoffspring. Unless, therefore, there is the strictest correlationbetween ornament and general perfection, the more brightlycoloured or ornamented varieties can obtain no permanentadvantage and if there is ; (as I maintain) such a correlation,then the sexual selection of colour or ornament, for whichthere is little or no evidence, becomes needless, becausenatural selection, which is an admitted vera causa, will itselfproduce all the results.In the case of butterflies the argument becomes evenstronger, because the fertility is so much greater than inbirds, and the weeding-out of the unfit takes place, to a greatextent, in the egg and larva state. Unless the eggs andlarvae which escaped to produce the next generation werethose which would produce the more highly-coloured butterflies,it is difficult to perceive how the slight preponderanceof colour sometimes selected by the females should not bewholly neutralised by the extremely rigid selection for otherqualities to which the offspring in every stage are exposed.The only way in which we can account for the observed facts