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قراءة كتاب Mimicry in Butterflies
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more conspicuous the pattern the more readily would it be noticed by the enemy, and so it would be of advantage to the Ithomiine to possess as striking a pattern as possible. Those butterflies shewing a tendency to a more conspicuous pattern would be more immune to the attacks of birds and so would have a better chance of leaving progeny than those with a less conspicuous pattern. In this way variations in the direction of greater conspicuousness would be accumulated gradually by natural selection, and so would be built up in the Ithomiine the striking warning coloration by which it advertises its disagreeable properties. Such is the first step in the making of a mimicry case—the building up through natural selection of a conspicuous pattern in an unpalatable species by means of which it is enabled to advertise its disagreeable properties effectively and thereby secure immunity from the attacks of enemies which are able to appreciate the advertisement. Such patterns and colours are said to be of a "warning" nature. The existence of an unpalatable model in considerable numbers is the first step in the production of a mimetic resemblance through the agency of natural selection.
We come back now to our Pierine which must be assumed to shew the general characters and coloration of the family of whites to which they belong (cf. Pl. X, fig. 1). Theoretically they are not specially protected by nauseous properties from enemies and hence their conspicuous white coloration renders
them especially liable to attack. If, however, they could exchange their normal dress for one resembling that of the Ithomiines it is clear that they would have a chance of being mistaken for the latter and consequently of being left alone. Moreover, in certain cases these Pierines have managed to discard their normal dress and assume that of the Ithomiines. On theoretical grounds this must clearly be of advantage to them, and being so might conceivably have arisen through the operation of natural selection. This indeed is what is supposed to have taken place on the theory of mimicry. Those Pierines which exhibited a variation of colour in the direction of the Ithomiine "model" excited distrust in the minds of would-be devourers, who had learned from experience to associate that particular type of coloration with a disagreeable taste. Such Pierines would therefore have a rather better chance of surviving and of leaving offspring. Some of the offspring would exhibit the variation in a more marked degree and these again would in consequence have a yet better chance of surviving. Natural selection would encourage those varying in the direction of the Ithomiine model at the expense of the rest and by its continuous operation there would gradually be built up those beautiful cases of resemblance which have excited the admiration of naturalists.
Wallace was the next after Bates to interest himself in mimicry and, from his study of the butterflies of the Oriental region[4], shewed that in this part of
the world too there existed these remarkable resemblances between species belonging to different families. Perhaps the most important part of Wallace's contribution was the demonstration that in some species not only was it the female alone that "mimicked" but that there might be several different forms of female mimicking different models, and in some cases all unlike the male of their own species. One of the species studied by Wallace, Papilio polytes, is shewn on Plate V. We shall have occasion to refer to this case later on, and it is sufficient here to call attention to the three different forms of female, of which one is like the male while the other two resemble two other species of Papilio, P. hector and P. aristolochiae, which occur in the same localities. Instances where the female alone of some unprotected species mimics a model with obnoxious properties are common in all tropical countries. It has been suggested that this state of things has come about owing to the greater need of protection on the part of the female. Hampered by the disposal of the next generation the less protected female would be at a greater disadvantage as compared with the mimic than would the corresponding male whose obligations to posterity are more rapidly discharged. The view of course makes the assumption that the female transmits her peculiar properties to her daughters but not to her sons.
A few years later Trimen[5] did for Africa what Bates had done for America and Wallace for
Indo-Malaya. It was in this paper that he elucidated that most remarkable of all cases of mimicry—Papilio dardanus with his harem of different consorts, all tailless, all unlike himself, and often wonderfully similar to unpalatable forms found in the same localities (cf. p. 30).
We may now turn to one of the most ingenious developments of the theory of mimicry. Not long after Bates' original memoir appeared attention was directed to a group of cases which could not be explained on the simple hypothesis there put forward. Many striking cases of resemblance had been adduced in which both species obviously belonged to the presumably unpalatable groups. Instances of the sort had been recorded by Bates himself and are perhaps most plentiful in South America between species belonging respectively to the Ithomiinae and Heliconinae. On the theory of mimicry all the members of both of these groups must be regarded as specially protected owing to their conspicuous coloration and distasteful properties. What advantage then can an Ithomiine be supposed to gain by mimicking a Heliconine, or vice versâ? Why should a species exchange its own bright and conspicuous warning pattern for one which is neither brighter nor more conspicuous? To Fritz Müller, the well-known correspondent of Darwin, belongs the credit of having suggested a way out of the difficulty. Müller's explanation turns upon the education of birds. Every year there hatch into the world fresh generations of young birds, and each
generation has to learn afresh from experience what is pleasant to eat and what is not. They will try all things and hold fast to that which is good. They will learn to associate the gay colours of the Heliconine and the Ithomiine with an evil taste[6] and they will thenceforward avoid butterflies which advertise themselves by means of these particular colour combinations. But in a locality where there are many models, each with a different pattern and colour complex, each will have to be tested separately before the unpalatableness of each is realised. If for example a thousand young birds started their education on a population of butterflies in which there were five disagreeable species, each with a distinct warning pattern, it is clear that one thousand of each would devote their lives to the education