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Theory of Warning Colours. These differ greatly from the last class, inasmuch as they present us with a variety of brilliant hues, often of the greatest purity, and combined in striking contrasts and conspicuous patterns. Their use depends upon their boldness and visibility, not on the presence of any one colour ; hence we find among these groups some of the most exquisitely-coloured objects in nature. Many of the uncatable caterpillars are strikingly beautiful; while the Danaidie, II.iconidæ, and protected groups of Papilionidæ, comprise a series of butterflies of the most brilliant and contrasted colours. The bright colours of many of the sca-anemones and sea-slugs will probably be found to be in this sense protective, serving as a warning of their uneatableness. On our theory none of these colours offer any difficulty. Conspicuousness being useful, every variation tending to brighter and purer colours was selected ; the result being the beautiful variety and contrast we find.
Imitative Warning Colours : The Theory of Mimicry.-- We now come to those groups which gain protection solely by being mistaken for some of these brilliantly coloured but uneatable creatures, and here a difficulty really exists, and to many minds is so great as to be insuperable. It will be well therefore to endeavour to explain how the resemblance in question may have been brought about.
The most difficult case, and the one which may be taken as
a type of the whole class, is that of the genus Leptalis (a group of South American butterflies allied to our common white and yellow kinds), many of the larger species of which are still white or yellow, and which are all eatable by birds and other insectivorous
creatures. But there are also a number of species of Leptalis, which are brilliantly red, yellow, and black, and which, band for band and spot for spot, resemble some one of the Danaidee or Heliconide which inhabit the same district and which are nauseous and weatable. Now the usual disliculty is, that a slight approach to one of these protected butterflies would be of no use, while a greater sudden variation is not admissible on the theory of gradual change by indefinite slight variations. This objection depends almost wholly on the supposition that, when the first steps towards mimicry occurred, the South American Danaida were what they are now; while the ancestors of the Leptalides were like the ordinary white or yellow Pieridlæ to which they are allied. But the danaioid butterflies of South America are so immensely numerous and so greatly varied, not only in colour but in structure, that we may be sure they are of vast antiquity and have undergone great modification. A large number of them, however, are still of comparatively plain colours, often rendered extremely clegant by the delicate transparency of the wing membrane, but otherwise not at all conspicuous. Many have only dusky or purplish bands or spots; others have patches of reddish or yellowish brown--perhaps the commonest colour among butterflies; while a considerable number are tinged or spotted with yellow, also a very common colour, and one especially characteristic of the Pieridæ, the family to which Leptalis belongs. We may therefore reasonably suppose that in the early stages of the development of the Danaidæ, when they first began to acquire those nauseous sccretions which are now their protection, their colours were somewhat plain ; either
dusky with paler bands and spots, or yellowish with dark burlers, and sometimes with reddish bands or spots. At this time they had probably shorter wings and a more rapid flight, just like the other unprotected families of butterflies. But as soon as they became deciilelly unpalatable to any of their enemies, it would be an advantage to them to be readily distinguished from all the catalle kindls; anil as butterflies were no doubt alreadly very varied in colour, while all probably had wings adapted for rather quick or jerking flight, the best distinction might have been found in outline and habits; whence would arise the preservation of those varieties whose longer wings, boilies, and antennæ, as well as their slower flight, rendered them noticeablecharacters which now distinguish the whole group in every part of the world.
Now it would be at this stage, that some of the weaker-flying Pieridæ which happened to resemble some of the Danaidze around them in their yellow and dusky tints and in the general outline of their wings, would be sometimes mistaken for them by the common enemy, and would thus gain an advantage in the struggle for existence. Admitting this one step to be made, and all the rest must inevitably follow from simple variation and survival of the fittest. So soon as the nauscous butterfly varied in form or colour to such an extent that the corresponding catable butterfly no longer closely resembled it, the latter would be exposed to attacks, and only those variations would be preserved which kept up the resemblance. At the same time we may well suppose the enemies to become more acute and able to detect smaller differences than at first. This would
lead to the destruction of all adverse variations, and thus keep up in continually increasing complexity the outward mimicry which now so amazes us. During the long ages in which this process has been going on, and the Danaidie have been acquiring those specialities of colour which aid in their preservation, many a Leptulis may have become extinct from not varying sufficiently in the right direction and at the right time to keep up a protective resemblance to its neighbour ; and this well accorils with the comparatively small number of cases of true mimicry, as compared with the frequency of those protective resemblances to vegetable or inorganic objects whose forms are less definite and colours less changeable. About a dozen other genera of butterflies and moths mimic the Danaidie in various parts of the world, and exactly the same explanation will apply to all of them. They represent those species of each group which, at the time when the Danaidze first acquired their protective secretions, happened outwardly to resemble some of them, and which have, by concurrent variation aided by a rigiil selection, been able to keep up that resemblance to the present day."
Theory of Sexual Colour's.-In Jr. Darwin's celebrated work, The Descent of Man and Selection in Relation to Sex, he has treated of sexual colour in combination with other sexual characters, and has
· For fuller information on this suloject the reader should consult Mr. Bates's original paper,
Contributions to an Insect-fauna of the Amazon Valley,” in Transactions of the Linnean Society, vol. xxiii. p. 495 ; Mr. Trimen's paper in vol. xxvi. p. 497; the author's essay on “Mimicry,” &c., already referred to; and in the absence of collections of butterflies, the plates of Ileliconidze and Leptalidæ, in Hewitson's Erotic Butterflies, and Felder's Voyage of the “ Vorara," may be examined.
arrived at the conclusion that all or almost all the colours of the higher animals (including among these insects and all vertebrates) are due to voluntary or conscious sesual selection ; and that diversity of colour in the sexes is due, primarily, to the transmission of colour-variations either to one sex only or to both sexes; the difference depending on some unknown law, and not being due to vatural selection.
I have long held this portion of Jr. Darwin's theory to be erroneous; and have argued that the primary cause of sexual diversity of colour was the need of protection, repressing in the female those bright colours which are normally produced in both sexes by general laws; and I have attempted to explain many of the more difficult cases on this principle. (“ A Theory of Birds' Nests,” in Contributions, Sc., p. 231.) As I have since given much thought to this subject, and have arrived at some views which appear to me to be of considerable importance, it will be well to sketch briefly the theory I now holil, and afterwards show its application to some of the detailed cases adoluced in Mr. Darwin's work.
The very frequent superiority of the male bird or insect in brightness or intensity of colour, even when the general coloration is the same in both sexes, now seems to me to be, primarily, due to the greater vigour and activity and the higher vitality of the male. The colours of an animal usually fade during discase or weakness, while robust health and vigour adds to their intensity. This is a most important and suggestive fact, and one that appears to hold universally. In all quadrupeds a “dull coat” is indicative of ill-health or low condition ; while a glossy coat and sparkling eye