darwin and modern science-第21章
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d brown in summer （Alpine hare； and the ptarmigan of the Alps）； and there are also green leaf…insects which remain green only while they are young and difficult to see on the leaf； but which become brown again in the last stage of larval life； when they have outgrown the leaf。 They then conceal themselves by day； sometimes only among withered leaves on the ground； sometimes in the earth itself。 It is interesting that in one genus； Chaerocampa； one species is brown in the last stage of larval life； another becomes brown earlier； and in many species the last stage is not wholly brown； a part remaining green。 Whether this is a case of a double adaptation； or whether the green is being gradually crowded out by the brown； the fact remains that the same species； even the same individual； can exhibit both variations。 The case is the same with many of the leaf…like Orthoptera； as； for instance； the praying mantis （Mantis religiosa） which we have already mentioned。
But the best proofs are furnished by those often…cited cases in which the insect bears a deceptive resemblance to another object。 We now know many such cases； such as the numerous imitations of green or withered leaves； which are brought about in the most diverse ways； sometimes by mere variations in the form of the insect and in its colour； sometimes by an elaborate marking； like that which occurs in the Indian leaf…butterflies； Kallima inachis。 In the single butterfly…genus Anaea； in the woods of South America； there are about a hundred species which are all gaily coloured on the upper surface； and on the reverse side exhibit the most delicate imitation of the colouring and pattern of a leaf； generally without any indication of the leaf…ribs； but extremely deceptive nevertheless。 Anyone who has seen only one such butterfly may doubt whether many of the insignificant details of the marking can really be of advantage to the insect。 Such details are for instance the apparent holes and splits in the apparently dry or half…rotten leaf； which are usually due to the fact that the scales are absent on a circular or oval patch so that the colourless wing…membrane lies bare； and one can look through the spot as through a window。 Whether the bird which is seeking or pursuing the butterflies takes these holes for dewdrops； or for the work of a devouring insect； does not affect the question； the mirror…like spot undoubtedly increases the general deceptiveness； for the same thing occurs in many leaf…butterflies； though not in all； and in some cases it is replaced in quite a peculiar manner。 In one species of Anaea （A。 divina）； the resting butterfly looks exactly like a leaf out of the outer edge of which a large semicircular piece has been eaten； possibly by a caterpillar； but if we look more closely it is obvious that there is no part of the wing absent； and that the semicircular piece is of a clear； pale yellow colour； while the rest of the wing is of a strongly contrasted dark brown。
But the deceptive resemblance may be caused in quite a different manner。 I have often speculated as to what advantage the brilliant white C could give to the otherwise dusky…coloured 〃Comma butterfly〃 （Grapta C。 album）。 Poulton's recent observations （〃Proc。 Ent。 Soc〃。； London； May 6； 1903。） have shown that this represents the imitation of a crack such as is often seen in dry leaves； and is very conspicuous because the light shines through it。
The utility obviously lies in presenting to the bird the very familiar picture of a broken leaf with a clear shining slit； and we may conclude； from the imitation of such small details； that the birds are very sharp observers and that the smallest deviation from the usual arrests their attention and incites them to closer investigation。 It is obvious that such detailedwe might almost say such subtledeceptive resemblances could only have come about in the course of long ages through the acquirement from time to time of something new which heightened the already existing resemblance。
In face of facts like these there can be no question of chance； and no one has succeeded so far in finding any other explanation to replace that by selection。 For the rest； the apparent leaves are by no means perfect copies of a leaf； many of them only represent the torn or broken piece； or the half or two…thirds of a leaf； but then the leaves themselves frequently do not present themselves to the eye as a whole； but partially concealed among other leaves。 Even those butterflies which； like the species of Kallima and Anaea； represent the whole of a leaf with stalk； ribs； apex； and the whole breadth； are not actual copies which would satisfy a botanist； there is often much wanting。 In Kallima the lateral ribs of the leaf are never all included in the markings； there are only two or three on the left side and at most four or five on the right； and in many individuals these are rather obscure； while in others they are comparatively distinct。 This furnishes us with fresh evidence in favour of their origin through processes of selection； for a botanically perfect picture could not arise in this way； there could only be a fixing of such details as heightened the deceptive resemblance。
Our postulate of origin through selection also enables us to understand why the leaf…imitation is on the lower surface of the wing in the diurnal Lepidoptera； and on the upper surface in the nocturnal forms； corresponding to the attitude of the wings in the resting position of the two groups。
The strongest of all proofs of the theory； however； is afforded by cases of true 〃mimicry；〃 those adaptations discovered by Bates in 1861； consisting in the imitation of one species by another； which becomes more and more like its model。 The model is always a species that enjoys some special protection from enemies； whether because it is unpleasant to taste； or because it is in some way dangerous。
It is chiefly among insects and especially among butterflies that we find the greatest number of such cases。 Several of these have been minutely studied； and every detail has been investigated； so that it is difficult to understand how there can still be disbelief in regard to them。 If the many and exact observations which have been carefully collected and critically discussed； for instance by Poulton （〃Essays on Evolution〃； 1889…1907； Oxford； 1908； passim； e。g。 page 269。） were thoroughly studied； the arguments which are still frequently urged against mimicry would be found untenable； we can hardly hope to find more convincing proof of the actuality of the processes of selection than these cases put into our hands。 The preliminary postulates of the theory of mimicry have been disputed； for instance； that diurnal butterflies are persecuted and eaten by birds； but observations specially directed towards this point in India； Africa； America and Europe have placed it beyond all doubt。 If it were necessary I could myself furnish an account of my own observations on this point。
In the same way it has been established by experiment and observation in the field that in all the great regions of distribution there are butterflies which are rejected by birds and lizards； their chief enemies； on account of their unpleasant smell or taste。 These butterflies are usually gaily and conspicuously coloured and thusas Wallace first interpreted itare furnished with an easily recognisable sign： a sign of unpalatableness or WARNING COLOURS。 If they were not thus recognisable easily and from a distance； they would frequently be pecked at by birds； and then rejected because of their unpleasant taste； but as it is； the insect…eaters recognise them at once as unpalatable booty and ignore them。 Such IMMUNE （The expression does not refer to all the enemies of this butterfly； against ichneumon…flies； for instance； their unpleasant smell usually gives no protection。） species； wherever they occur； are imitated by other palatable species； which thus acquire a certain degree of protection。
It is true that this explanation of the bright； conspicuous colours is only a hypothesis； but its foundations；unpalatableness； and the liability of other butterflies to be eaten；are certain； and its consequencesthe existence of mimetic p
