darwin and modern science-第16章
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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 palatable formsconfirm it in the most convincing manner。 Of the many cases now known I select one； which is especially remarkable； and which has been thoroughly investigated； Papilio dardanus （merope）； a large； beautiful； diurnal butterfly which ranges from Abyssinia throughout the whole of Africa to the south coast of Cape Colony。
The males of this form are everywhere ALMOST the same in colour and in form of wings； save for a few variations in the sparse black markings on the pale yellow ground。 But the females occur in several quite different forms and colourings； and one of these only； the Abyssinian form； is like the male； while the other three or four are MIMETIC； that is to say； they copy a butterfly of quite a different family the Danaids； which are among the IMMUNE forms。 In each region the females have thus copied two or three different immune species。 There is much that is interesting to be said in regard to these species； but it would be out of keeping with the general tenor of this paper to give details of this very complicated case of polymorphism in P。 dardanus。 Anyone who is interested in the matter will find a full and exact statement of the case in as far as we know it； in Poulton's 〃Essays on Evolution〃 （pages 373…375）。 （Professor Poulton has corrected some wrong descriptions which I had unfortunately overlooked in the Plates of my book 〃Vortrage uber Descendenztheorie〃； and which refer to Papilio dardanus （merope）。 These mistakes are of no importance as far as and understanding of the mimicry…theory is concerned； but I hope shortly to be able to correct them in a later edition。） I need only add that three different mimetic female forms have been reared from the eggs of a single female in South Africa。 The resemblance of these forms to their immune models goes so far that even the details of the LOCAL forms of the models are copied by the mimetic species。
It remains to be said that in Madagascar a butterfly； Papilio meriones； occurs； of which both sexes are very similar in form and markings to the non…mimetic male of P。 dardanus； so that it probably represents the ancestor of this latter species。
In face of such facts as these every attempt at another explanation must fail。 Similarly all the other details of the case fulfil the preliminary postulates of selection； and leave no room for any other interpretation。 That the males do not take on the protective colouring is easily explained； because they are in general more numerous； and the females are more important for the preservation of the species； and must also live longer in order to deposit their eggs。 We find the same state of things in many other species； and in one case （Elymnias undularis） in which the male is also mimetically coloured； it copies quite a differently coloured immune species from the model followed by the female。 This is quite intelligible when we consider that if there were TOO MANY false immune types； the birds would soon discover that there were palatable individuals among those with unpalatable warning colours。 Hence the imitation of different immune species by Papilio dardanus！
I regret that lack of space prevents my bringing forward more examples of mimicry and discussing them fully。 But from the case of Papilio dardanus alone there is much to be learnt which is of the highest importance for our understanding of transformations。 It shows us chiefly what I once called； somewhat strongly perhaps； THE OMNIPOTENCE OF NATURAL SELECTION in answer to an opponent who had spoken of its 〃inadequacy。〃 We here see that one and the same species is capable of producing four or five different patterns of colouring and marking； thus the colouring and marking are not； as has often been supposed； a necessary outcome of the specific nature of the species； but a true adaptation； which cannot arise as a direct effect of climatic conditions； but solely through what I may call the sorting out of the variations produced by the species； according to their utility。 That caterpillars may be either green or brown is already something more than could have been expected according to the old conception of species； but that one and the same butterfly should be now pale yellow； with black； now red with black and pure white； now deep black with large； pure white spots； and again black with a large ochreous…yellow spot； and many small white and yellow spots； that in one sub…species it may be tailed like the ancestral form； and in another tailless like its Danaid model；all this shows a far…reaching capacity for variation and adaptation that wide never have expected if we did not see the facts before us。 How it is possible that the primary colour…variations should thus be intensified and combined remains a puzzle even now； we are reminded of the modern three…colour printing；perhaps similar combinations of the primary colours take place in this case； in any case the direction of these primary variations is determined by the artist whom we know as natural selection； for there is no other conceivable way in which the model could affect the butterfly that is becoming more and more like it。 The same climate surrounds all four forms of female； they are subject to the same conditions of nutrition。 Moreover； Papilio dardanus is by no means the only species of butterfly which exhibits different kinds of colour…pattern on its wings。 Many species of the Asiatic genus Elymnias have on the upper surface a very good imitation of an immune Euploeine （Danainae）； often with a steel…blue ground…colour； while the under surface is well concealed when the butterfly is at rest； thus there are two kinds of protective coloration each with a different meaning！ The same thing may be observed in many non…mimetic butterflies； for instance in all our species of Vanessa； in which the under side shows a grey…brown or brownish…black protective coloration； but we do not yet know with certainty what may be the biological significance of the gaily coloured upper surface。
In general it may be said that mimetic butterflies are comparatively rare species； but there are exceptions； for instance Limenitis archippus in North America； of which the immune model （Danaida plexippus） also occurs in enormous numbers。
In another mimicry…category the imitators are often more numerous than the models； namely in the case of the imitation of DANGEROUS INSECTS by harmless species。 Bees and wasps are dreaded for their sting； and they are copied by harmless flies of the genera Eristalis and Syrphus； and these mimics often occur in swarms about flowering plants without damage to themselves or to their models； they are feared and are therefore left unmolested。
In regard also to the FAITHFULNESS OF THE COPY the facts are quite in harmony with the theory； according to which the resemblance must have arisen and increased BY DEGREES。 We can recognise this in many cases； for even now the mimetic species show very VARYING DEGREES OF RESEMBLANCE to their immune model。 If we compare； for instance； the many different imitators of Danaida chrysippus we find that； with their brownish…yellow ground…colour； and the position and size； and more or less sharp limitation of their clear marginal spots； they have reached very different degrees of nearness to their model。 Or compare the female of Elymnias undularis with its model Danaida genutia； there is a general resemblance； but the marking of the Danaida is very roughly imitated in Elymnias。
Another fact that bears out the theory of mimicry is； that even when the resemblance in colour…pattern is very great； the WING…VENATION； which is so constant； and so important in determining the systematic position of butterflies； is never affected by the variation。 The pursuers of the butterfly have no time to trouble about entomological intricacies。
I must not pass over a discovery of Poulton's which is of great theoretical importancethat mimetic butterflies may reach the same effect by very different means。 （〃Journ。 Linn。 Soc。 London （Zool。）〃； Vol。 XXVI。 1898； pages 598…602。） Thus the glass…like transparency of the wing of a certain Ithomiine （Methona） a
