The rule which Mr. Wallace believes, with very few exceptions, to hold good is, "that when both sexes are of strikingly gay and conspicuous colours, the nest is...such as to conceal the sitting bird; while, whenever there is a striking contrast of colours, the male being gay and conspicuous, the female dull and obscure, the nest is open, and the sitting bird exposed to view." At this time Mr. Wallace allowed considerably more influence to sexual selection (in combination with the need of protection) than in his later writings. The following extract from a letter from Mr. Wallace to Darwin (July 23rd, 1877) fixes the period at which the change in his views occurred: "I am almost afraid to tell you that in going over the subject of the colours of animals, etc., etc., for a small volume of essays, etc., I am preparing, I have come to conclusions directly opposed to voluntary sexual selection, and believe that I can explain (in a general way) all the phenomena of sexual ornaments and colours by laws of development aided by simple 'Natural Selection.'" He finally rejected Mr. Darwin's theory that colours "have been developed by the preference of the females, the more ornamented males becoming the parents of each successive generation." "Darwinism," 1889, page 285. See also Letters 442, 443, 449, 450, etc.)

Down, April 15th, [1868].

I have been deeply interested by your admirable article on birds' nests. I am delighted to see that we really differ very little,--not more than two men almost always will. You do not lay much or any stress on new characters spontaneously appearing in one sex (generally the male), and being transmitted exclusively, or more commonly only in excess, to that sex. I, on the other hand, formerly paid far too little attention to protection. I had only a glimpse of the truth; but even now I do not go quite as far as you. I cannot avoid thinking rather more than you do about the exceptions in nesting to the rule, especially the partial exceptions, i.e., when there is some little difference between the sexes in species which build concealed nests. I am not quite satisfied about the incubating males; there is so little difference in conspicuousness between the sexes. I wish with all my heart I could go the whole length with you. You seem to think that male birds probably select the most beautiful females; I must feel some doubt on this head, for I can find no evidence of it. Though I am writing so carping a note, I admire the article thoroughly.

And now I want to ask a question. When female butterflies are more brilliant than their males you believe that they have in most cases, or in all cases, been rendered brilliant so as to mimic some other species, and thus escape danger. But can you account for the males not having been rendered equally brilliant and equally protected? (440/2. See Wallace in the "Westminster Review," July, 1867, page 37, on the protection to the female insect afforded by its resemblance either to an inanimate object or to another insect protected by its unpalatableness. The cases are discussed in relation to the much greater importance (to the species as a whole) of the preservation of the female insect with her load of eggs than the male who may safely be sacrificed after pairing. See Letter 189, note.) Although it may be most for the welfare of the species that the female should be protected, yet it would be some advantage, certainly no disadvantage, for the unfortunate male to enjoy an equal immunity from danger. For my part, I should say that the female alone had happened to vary in the right manner, and that the beneficial variations had been transmitted to the same sex alone. Believing in this, I can see no improbability (but from analogy of domestic animals a strong probability) that variations leading to beauty must often have occurred in the males alone, and been transmitted to that sex alone. Thus I should account in many cases for the greater beauty of the male over the female, without the need of the protective principle.

Charles Darwin

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