BIONOMICS.]
COLOURS
0 FANIMALS
individual enters a new environment in the course of its growth (such as larva, pupa, imago, &c.). In insects with more than one brood in the year, Seasonal Dimorphism is often seen, and the differences are sometimes appropriate to the altered condition of the environment as the seasons change. The causes of change in these and Arctic animals are insufficiently worked out: in both sets there are observations or experiments which indicate changes from within the organism, merely following the seasons and not caused by them, and other observations or experiments which prove that certain species are susceptible to the changing external influences. In certain species concealment is effected by the use of adventitious objects, which are employed as a covering. Examples of this AUocryptic defence are found in the tubes of the caddis worms (Phryganea), or the objects made use of by crabs of the genera Ilyas, Stenorhynchus, &c. Such animals are concealed in any environment. If sedentary, like the former example, they are covered up with local materials; if wandering, like the latter, they have the instinct to reclothe. Allocryptic methods may also be used for aggressive purposes, as the ant-lion larva, almost buried in sand, or the large frog Geratophrys,'which, covers its back with earth when waiting for its prey. Another form of Allocryptic defence is found in the use of the colour of the food in the digestive organs showing through the transparent body, and in certain cases the adventitious colour may be dissolved in the blood or secreted in superficial cells of the body: thus certain insects make use of the chlorophyll of their food (Poulton, Proc. Roy. Soc. vol. liv. p. 417). The most perfect Cryptic powers are possessed by those animals in which the individuals can change their colours into any tint which would be appropriate to a normal environment. This power is widely prevalent in Fish, and also occurs in Amphibia and Reptilia (the chameleon affording a wellknown example). Analogous powers exist in certain Crustacea and Cephalopoda. All these rapid changes of colour are due to changes in shape or position of superficial pigment cells controlled by the nervous system. That the latter is itself stimulated by light through the medium of the eye and optic nerve has been proved in many cases. Animals with a short life-history passed in a single environment, which, however, may be very different in the case of different individuals, may have a different form of Variable Cryptic Colouring, namely, the power of adapting their colour once for all (many pupae), or once or twice (many larvae). In these cases the effect appears to be produced through the nervous system, although the stimulus of light probably acts on the skin and not through the eyes. Particoloured surfaces do not produce particoloured pupae, probably because the antagonistic stimuli neutralize each other in the central nervous system which then disposes the superficial colours so that a neutral or intermediate effect is produced over the whole surface (Poulton, Trans. Ent. Soc. Lond., 1892, p. 293). Cryptic colouring may incidentally produce superficial resemblances between animals ; thus desert forms concealed in the same way may gain a likeness to each other, and in the same way Special Resemblances, e.g., to lichen, bark, grasses, pine-needles, &c., may sometimes lead to a tolerably close similarity between the animals which are thus concealed. Such likeness may be called Syncryptic or Common Protective (or Aggressive) Resemblance, and it is to be distinguished from Mimicry and Common Warning Colours, in which the likeness is not incidental but an end in itself. Syncryptic Resemblances have much in common with those incidentally caused by functional adaptation, such as the mole-like forms produced in the burrowing Insectivora, Rodentia, and Marsupialia. Such likeness may be called Syntechnic Resemblance, incidentally produced by dynamic
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similarity, just as Syncryptic Resemblance is produced by static similarity. Use of Colour for Warning and Signalling.—The use of colour for the purpose of warning is the exact opposite of the one which has been just described, its object being to render the animal conspicuous to its enemies, so that it can be easily seen, well remembered, and avoided in future. Warning colours are associated with some quality or weapon which renders the possessor unpleasant or dangerous, such as unpalatability, an evil odour, a sting, the poison-fang, &c. The object being to warn an enemy off, these colours are also called Aposematic. Recognition markings, on the other hand, are Episematic, assisting the individuals of the same species to keep together when their safety depends upon numbers, or easily to follow each other to a place of safety, the young and inexperienced benefiting by the example of the older. Episematic characters are far less common than Aposematic,and these than Cryptic; although, as regards the latter comparison, the opposite impression is generally produced from the very fact that concealment is so successfully attained. Warning or Aposematic Colours, together with the qualities they indicate, depend, as a rule, for their very existence upon the abundance of palatable food supplied by the animals with Cryptic colouring. Unpalatability, or even the possession of a sting, is not sufficient defence unless there is enough food of another kind to be obtained at the same time and place (Poulton, Proc. Zool. Soc., 1887, p. 191). Hence insects with Warning Colours are not seen in temperate countries except at the time when insect life as a whole is most abundant; and in warmer countries, with well-marked wet and dry seasons, it will probably be found that Warning Colours are proportionately less developed in the latter. In many species of African butterflies belonging to the genus Junonia (including Precis) the wet-season broods are distinguished by the more or less conspicuous under sides of the wings, those of the dry season being highly cryptic. Warning Colours are, like Cryptic, assisted by special adaptations of the body-form, and especially by movements which assist to render the colour as conspicuous as possible. On this account animals with Warning Colours generally move or fly slowly, and it is the rule in butterflies that the Warning patterns are similar on both upper and under sides of the wings. Many animals, when attacked or disturbed, “ sham death ” (as it is commonly but wrongly described), falling motionless to the ground. In the case of well-concealed animals this instinct gives them a second chance of escape in the earth or among the leaves, &c., when they have been once detected; animals with Warning Colours are, on the other hand, enabled to assume a position in which their characters are displayed to the full (Portschinski, Lepidopterorum Rossice Biologia, St. Petersburg, 1890, plate i. figs. 16, 17). In both cases a definite attitude is assumed, which is not that of death. Other warning characters exist in addition to colouring: thus sound is made use of by the disturbed Rattle-snake and the Indian Echis, &c. Large birds, when attacked, often adopt a threatening attitude, accompanied by a terrifying sound. The Cobra warns an intruder chiefly by attitude and the dilation of the flattened neck, the effect being heightened in some species by the “spectacles.” In such cases we often see the combination of Cryptic and Sematic methods, the animal being concealed until disturbed, when it instantly assumes an Aposematic attitude. The advantage to the animal itself is clear: a poisonous snake gains nothing by killing an animal it cannot eat; while the poison does not cause immediate death, and the enemy would have time to injure or destroy the snake. In the case of small unpalatable animals with Warning Colours the enemies would
