# 1911 Encyclopædia Britannica/Colours of Animals

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​COLOURS OF ANIMALS. Much interest attaches in modern biology to the questions involved in the colours of animals. The subject may best be considered in two divisions: (1) as regards the uses of colour in the struggle for existence and in sexual relationships; (2) as regards the chemical causation.
1. Bionomics
Use of Colour for Concealment.—Cryptic colouring is by far the commonest use of colour in the struggle for existence. It is employed for the purpose of attack (aggressive resemblance or anticryptic colouring) as well as of defence (protective resemblance or procryptic colouring). The fact that the same method, concealment, may be used both for attack and defence has been well explained by T. Belt (The Naturalist in Nicaragua, London, 1888), who suggests as an illustration the rapidity of movement which is also made use of by both pursuer and pursued, which is similarly raised to a maximum in both by the gradual dying out of the slowest through a series of generations. Cryptic colouring is commonly associated with other aids in the struggle for life. Thus well-concealed mammals and birds, when discovered, will generally endeavour to escape by speed, and will often attempt to defend themselves actively. On the other hand, small animals which have no means of active defence, such as large numbers of insects, frequently depend upon concealment alone. Protective resemblance is far commoner among animals than aggressive resemblance, in correspondence with the fact that predaceous forms are as a rule much larger and much less numerous than their prey. In the case of insectivorous Vertebrata and their prey such differences exist in an exaggerated form. Cryptic colouring, whether used for defence or attack, may be either general or special. In general resemblance the animal, in consequence of its colouring, produces the same effect as its environment, but the conditions do not require any special adaptation of shape and outline. General resemblance is especially common among the animals inhabiting some uniformly coloured expanse of the earth’s surface, such as an ocean or a desert. In the former, animals of all shapes are frequently protected by their transparent blue colour; on the latter, equally diverse forms are defended by their sandy appearance. The effect of a uniform appearance may be produced by a combination of tints in startling contrast. Thus the black and white stripes of the zebra blend together at a little distance, and “their proportion is such as exactly to match the pale tint which arid ground possesses when seen by moonlight” (F. Galton, South Africa, London, 1889). Special resemblance is far commoner than general, and is the form which is usually met with on the diversified surface of the earth, on the shores, and in shallow water, as well as on the floating masses of Algae on the surface of the ocean, such as the Sargasso Sea. In these environments the cryptic colouring of animals is usually aided by special modifications of shape, and by the instinct which leads them to assume particular attitudes. Complete stillness and the assumption of a certain attitude play an essential part in general resemblance on land; but in special resemblance the attitude is often highly specialized, and perhaps more important than any other element in the complex method by which concealment is effected. In special resemblance the combination of colouring, shape and attitude is such as to produce a more or less exact resemblance to some one of the objects in the environment, such as a leaf or twig, a patch of lichen, or flake of bark. In all cases the resemblance is to some object which is of no interest to the enemy or prey respectively. The animal is not hidden from view by becoming indistinguishable from its background, as in the cases of general resemblance, but it is mistaken for some well-known object.
In seeking the interpretation of these most interesting and elaborate adaptations, attempts have been made along two lines. First, it is sought to explain the effect as a result of the direct influence of the environment upon the individual (G. L. L. Buffon), or by the inherited effects of effort and the use and disuse of parts (J. B. P. Lamarck). Second, natural selection is believed to have produced the result, and afterwards maintained it by the survival of the best concealed in each generation. The former suggestions break down when the complex nature of numerous special resemblances is appreciated. Thus the arrangement of colours of many kinds into an appropriate pattern requires the co-operation of a suitable shape and the rigidly exact adoption of a certain elaborate attitude. The latter is instinctive, and thus depends on the central nervous system. The cryptic effect is due to the exact co-operation of all these factors; and in the present state of science the only possible hope of an interpretation lies in the theory of natural selection, which can accumulate any and every variation which tends towards survival. A few of the chief types of methods by which concealment is effected may be briefly described. The colours of large numbers of Vertebrate animals are darkest on the back, and become gradually lighter on the sides, passing into white on the belly. Abbott H. Thayer (The Auk, vol. xiii., 1896) has suggested that this gradation obliterates the appearance of solidity, which is due to shadow. The colour-harmony, which is also essential to concealment, is produced because the back is of the same tint as the environment (e.g. earth) bathed in the cold blue-white of the sky, while the belly, being cold blue-white bathed in shadow and yellow earth reflections, produces the same effect. Thayer has made models (in the natural history museums at London, Oxford and Cambridge) which support his interpretation in a very convincing manner. This method of neutralizing shadow for the purpose of concealment by increased lightness of tint was first suggested by E. B. Poulton in the case of a larva (Trans. Ent. Soc. Lond., 1887, p. 294) and a pupa (Trans. Ent. Soc. Lond., 1888, pp. 596, 597), but he did not appreciate the great importance of the principle. In an analogous method an animal in front of a background of dark shadow may have part of its body obliterated by the existence of a dark tint, the remainder resembling, e.g., a part of a leaf (W. Müller, Zool. Jahr. J. W. Spengel, Jena, 1886). This method of rendering invisible any part which would interfere with the resemblance is well known in mimicry. A common aid to concealment is the adoption by different individuals of two or more different appearances, each of which resembles some special object to which an enemy is indifferent. Thus the leaf-like butterflies (Kallima) present various types of colour and pattern on the under side of the wings, each of which closely resembles some well-known appearance presented by a dead leaf; and the common British yellow under-wing moth (Tryphaena pronuba) is similarly polymorphic on the upper side of its upper wings, which are exposed as it suddenly drops among dead leaves. Caterpillars and pupae are also commonly dimorphic, green and brown. Such differences as these extend the area which an enemy is compelled to search in order to make a living. In many cases the cryptic colouring changes appropriately ​during the course of an individual life, either seasonally, as in the ptarmigan or Alpine hare, or according as the 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 allocryptic defence are found in the tubes of the caddis worms (Phryganea), or the objects made use of by crabs of the genera Hyas, 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 Ceratophrys, 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. liv. 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 well-known 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 similarity, just as syncryptic resemblance is produced by static similarity.
Use of Colour for Warning and Signalling, or Sematic Coloration.—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 (J. Portschinsky, Lepidopterorum Rossiae 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 rattlesnake 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 only first become aware of the unpleasant quality by tasting and often destroying their prey; but the species would gain by the experience thus conveyed, even though the individual might suffer. An insect-eating animal does not come into the world with knowledge: it has to be educated by experience, and warning colours enable this education as to what to avoid to be gained by a small instead of a large waste of life. Furthermore, great tenacity of life is usually possessed by animals with warning colours. The tissues of aposematic insects generally possess great elasticity and power of resistance, so that large numbers of individuals can recover after very severe treatment.
The brilliant warning colours of many caterpillars attracted the attention of Darwin when he was thinking over his hypothesis of sexual selection, and he wrote to A. R. Wallace on the subject (C. Darwin, Life and Letters, London, 1887, iii. 93). Wallace, in reply, suggested their interpretation as warning ​colours, a suggestion since verified by experiment (Proc. Ent. Soc. Lond., 1867, p. lxxx; Trans. Ent. Soc. Lond., 1869, pp. 21 and 27). Although animals with warning colours are probably but little attacked by the ordinary enemies of their class, they have special enemies which keep the numbers down to the average. Thus the cuckoo appears to be an insectivorous bird which will freely devour conspicuously coloured unpalatable larvae. The effect of the warning colours of caterpillars is often intensified by gregarious habits. Another aposematic use of colours and structures is to divert attention from the vital parts, and thus give the animal attacked an extra chance of escape. The large, conspicuous, easily torn wings of butterflies and moths act in this way, as is found by the abundance of individuals which may be captured with notches bitten symmetrically out of both wings when they were in contact. The eye-spots and “tails” so common on the hinder part of the hind wing, and the conspicuous apex so frequently seen on the fore wing, probably have this meaning. Their position corresponds to the parts which are most offen found to be notched. In some cases (e.g. many Lycaenidae) the “tail” and eye-spot combine to suggest the appearance of a head with antennae at the posterior end of the butterfly, the deception being aided by movements of the hind wings. The flat-topped “tussocks” of hair on many caterpillars look like conspicuous fleshy projections of the body, and they are held prominently when the larva is attacked. If seized, the “tussock” comes out, and the enemy is greatly inconvenienced by the fine branched hairs. The tails of lizards, which easily break off, are to be similarly explained, the attention of the pursuer being probably still further diverted by the extremely active movements of the amputated member. Certain crabs similarly throw off their claws when attacked, and the claws continue to snap most actively. The tail of the dormouse, which easily comes off, and the extremely bushy tail of the squirrel, are probably of use in the same manner. Animals with warning colours often tend to resemble each other superficially. This fact was first pointed out by H. W. Bates in his paper on the theory of mimicry (Trans. Linn. Soc. vol. xxiii., 1862, p. 495). He showed that the conspicuous, presumably unpalatable, tropical American butterflies, belonging to very different groups, which are mimicked by others, also tend to resemble each other, the likeness being often remarkably exact. These resemblances were not explained by his theory of mimicry, and he could only suppose that they had been produced by the direct influence of a common environment. The problem was solved in 1879 by Fritz Müller (see Proc. Ent. Soc. Lond., 1879, p. xx.), who suggested that life is saved by this resemblance between warning colours, inasmuch as the education of young inexperienced enemies is facilitated. Each species which falls into a group with common warning (synaposematic) colours contributes to save the lives of the other members. It is sufficiently obvious that the amount of learning and remembering, and consequently of injury and loss of life involved in the process, are reduced when many species in one place possess the same aposematic colouring, instead of each exhibiting a different “danger-signal.” These resemblances are often described as “Müllerian mimicry,” as distinguished from true or “Batesian mimicry” described in the next section. Similar synaposematic resemblances between the specially protected groups of butterflies were afterwards shown to exist in tropical Asia, the East Indian Islands and Polynesia by F. Moore (Proc. Zool. Soc., 1883, p. 201), and in Africa by E. B. Poulton (Report Brit. Assoc., 1897, p. 688). R. Meldola (Ann. and Mag. Nat. Hist. x., 1882, p. 417) first pointed out and explained in the same manner the remarkable general uniformity of colour and pattern which runs through so many species of each of the distasteful groups of butterflies; while, still later, Poulton (Proc. Zool. Soc., 1887, p. 191) similarly extended the interpretation to the synaposematic resemblances between animals of all kinds in the same country. Thus, for example, longitudinal or circular bands of the same strongly contrasted colours are found in species of many groups with distant affinities.
Certain animals, especially the Crustacea, make use of the special defence and warning colours of other animals. Thus the English hermit-crab, Pagurus Bernhardus, commonly carries the sea-anemone, Sagartia parasitica, on its shell; while another English species, Pagurus Prideauxii, inhabits a shell which is invariably clothed by the flattened Adamsia palliata.
The white patch near the tail which is frequently seen in the gregarious Ungulates, and is often rendered conspicuous by adjacent black markings, probably assists the individuals in keeping together; and appearances with probably the same interpretation are found in many birds. The white upturned tail of the rabbit is probably of use in enabling the individuals to follow each other readily. The difference between a typical aposematic character appealing to enemies, and episematic intended for other individuals of the same species, is well seen when we compare such examples as (1) the huge banner-like white tail, conspicuously contrasted with the black or black and white body, by which the slow-moving skunk warns enemies of its power of emitting an intolerably offensive odour; (2) the small upturned white tail of the rabbit, only seen when it is likely to be of use and when the owner is moving, and, if pursued, very rapidly moving, towards safety.
Mimicry (see also Mimicry) or Pseudo-sematic Colours.—The fact that animals with distant affinities may more or less closely resemble each other was observed long before the existing explanation was possible. Its recognition is implied in a number of insect names with the termination -formis, usually given to species of various orders which more or less closely resemble the stinging Hymenoptera. The usefulness of the resemblance was suggested in Kirby and Spence’s Introduction to Entomology, London, 1817, ii. 223. H. W. Bates (Trans. Linn. Soc. vol. xxiii., 1862, p. 495) first proposed an explanation of mimicry based on the theory of natural selection. He supposed that every step in the formation and gradual improvement of the likeness occurred in consequence of its usefulness in the struggle for life. The subject is of additional interest, inasmuch as it was one of the first attempts to apply the theory of natural selection to a large class of phenomena up to that time well known but unexplained. Numerous examples of mimicry among tropical American butterflies were discussed by Bates in his paper; and in 1866 A. R. Wallace extended the hypothesis to the butterflies of the tropical East (Trans. Linn. Soc. vol. xxv., 1866, p. 19); Roland Trimen (Trans. Linn. Soc. vol. xxvi., 1870, p. 497) to those of Africa in 1870. The term mimicry is used in various senses. It is often extended, as indeed it was by Bates, to include all the superficial resemblances between animals and any part of their environment. Wallace, however, separated the cryptic resemblances already described, and the majority of naturalists have followed this convenient arrangement. In cryptic resemblance an animal resembles some object of no interest to its enemy (or prey), and in so doing is concealed; in mimicry an animal resembles some other animal which is specially disliked by its enemy, or some object which is specially attractive to its prey, and in so doing becomes conspicuous. Some naturalists have considered mimicry to include all superficial likenesses between animals, but such a classification would group together resemblances which have widely different uses. (1) The resemblance of a mollusc to the coral on which it lives, or an external parasite to the hair or skin of its host, would be procryptic; (2) that between moths which resemble lichen, syncryptic; (3) between distasteful insects, synaposematic; (4) between the Insectivor mole and the Rodent mole-rat, syntechnic; (5) the essential element in mimicry is that it is a false warning (pseud-aposematic) or false recognition (pseud-episematic) character. Some have considered that mimicry indicates resemblance to a moving object; but apart from the non-mimetic likenesses between animals classified above, there are ordinary cryptic resemblances to drifting leaves, swaying bits of twig, &c., while truly mimetic resemblances are often specially adapted for the attitude of rest. Many use the term mimicry to include synaposematic as well as pseudo-sematic resemblances, calling the former “Müllerian,” the latter “Batesian,” mimicry. The objection to this grouping is that it takes little account of the deceptive element which is essential in mimicry. In ​synaposematic colouring the warning is genuine, in pseud-aposematic it is a sham. The term mimicry has led to much misunderstanding from the fact that in ordinary speech it implies deliberate imitation. The production of mimicry in an individual animal has no more to do with consciousness or “taking thought” than any of the other processes of growth. Protective mimicry is here defined as an advantageous and superficial resemblance of one animal to another, which latter is specially defended so as to be disliked or feared by the majority of enemies of the groups to which both belong—a resemblance which appeals to the sense of sight, sometimes to that of hearing, and rarely to smell, but does not extend to deep-seated characters except when the superficial likeness is affected by them. Mutatis mutandis this definition will apply to aggressive (pseud-episematic) resemblance. The conditions under which mimicry occurs have been stated by Wallace:—“(1) that the imitative species occur in the same area and occupy the same station as the imitated; (2) that the imitators are always the more defenceless; (3) that the imitators are always less numerous in individuals; (4) that the imitators differ from the bulk of their allies; (5) that the imitation, however minute, is external and visible only, never extending to internal characters or to such as do not affect the external appearance.” It is obvious that conditions 2 and 3 do not hold in the case of Müllerian mimicry. Mimicry has been explained, independently of natural selection, by the supposition that it is the common expression of the direct action of common causes, such as climate, food, &c.; also by the supposition of independent lines of evolution leading to the same result without any selective action in consequence of advantage in the struggle; also by the operation of sexual selection.
It is proposed, in conclusion, to give an account of the broad aspects of mimicry, and attempt a brief discussion of the theories of origin of each class of facts (see Poulton, Linn. Soc. Journ. Zool., 1898, p. 558). It will be found that in many cases the argument here made use of applies equally to the origin of cryptic and sematic colours. The relationship between these classes has been explained: mimicry is, as Wallace has stated (Darwinism, London, 1889), merely “an exceptional form of protective resemblance. “Now, protective (cryptic) resemblance cannot be explained on any of the lines suggested above, except natural selection; even sexual selection fails, because cryptic resemblance is especially common in the immature stages of insect life. But it would be unreasonable to explain mimetic resemblance by one set of principles and cryptic by another and totally different set. Again, it may be plausible to explain the mimicry of one butterfly for another on one of the suggested lines, but the resemblance of a fly or moth to a wasp is by no means so easy, and here selection would be generally conceded; yet the appeal to antagonistic principles to explain such closely related cases would only be justified by much direct evidence. Furthermore, the mimetic resemblances between butterflies are not haphazard, but the models almost invariably belong only to certain sub-families, the Danainae and Acraeinae in all the warmer parts of the world, and, in tropical America, the Ithomiinae and Heliconinae as well. These groups have the characteristics of aposematic species, and no theory but natural selection explains their invariable occurrence as models wherever they exist. It is impossible to suggest, except by natural selection, any explanation of the fact that mimetic resemblances are confined to changes which produce or strengthen a superficial likeness. Very deep-seated changes are generally involved, inasmuch as the appropriate instincts as to attitude, &c., are as important as colour and marking. The same conclusion is reached when we analyse the nature of mimetic resemblance and realize how complex it really is, being made up of colours, both pigmentary and structural, pattern, form, attitude and movement. A plausible interpretation of colour may be wildly improbable when applied to some other element, and there is no explanation except natural selection which can explain all these elements. The appeal to the direct action of local conditions in common often breaks down upon the slightest investigation, the difference in habits between mimic and model in the same locality causing the most complete divergence in their conditions of life. Thus many insects produced from burrowing larvae mimic those whose larvae live in the open. Mimetic resemblance is far commoner in the female than in the male, a fact readily explicable by selection, as suggested by Wallace, for the female is compelled to fly more slowly and to expose itself while laying eggs, and hence a resemblance to the slow-flying freely exposed models is especially advantageous. The facts that mimetic species occur in the same locality, fly at the same time of the year as their models, and are day-flying species even though they may belong to nocturnal groups, are also more or less difficult to explain except on the theory of natural selection, and so also is the fact that mimetic resemblance is produced in the most varied manner. A spider resembles its model, an ant, by a modification of its body-form into a superficial resemblance, and by holding one pair of legs to represent antennae; certain bugs (Hemiptera) and beetles have also gained a shape unusual in their respective groups, a shape which superficially resembles an ant; a Locustid (Myrmecophana) has the shape of an ant painted, as it were, on its body, all other parts resembling the background and invisible; a Membracid (Homoptera) is entirely unlike an ant, but is concealed by an ant-like shield. When we further realize that in this and other examples of mimicry “the likeness is almost always detailed and remarkable, however it is attained, while the methods differ absolutely,” we recognize that natural selection is the only possible explanation hitherto suggested. In the cases of aggressive mimicry an animal resembles some object which is attractive to its prey. Examples are found in the flower-like species of Mantis, which attract the insects on which they feed. Such cases are generally described as possessing “alluring colours,” and are regarded as examples of aggressive (anticryptic) resemblance, but their logical position is here.
Colours displayed in Courtship, Secondary Sexual Characters, Epigamic Colours.—Darwin suggested the explanation of these appearances in his theory of sexual selection (The Descent of Man, London, 1874). The rivalry of the males for the possession of the females he believed to be decided by the preference of the latter for those individuals with especially bright colours, highly developed plumes, beautiful song, &c. Wallace does not accept the theory, but believes that natural selection, either directly or indirectly, accounts for all the facts. Probably the majority of naturalists follow Darwin in this respect. The subject is most difficult, and the interpretation of a great proportion of the examples in a high degree uncertain, so that a very brief account is here expedient. That selection of some kind has been operative is indicated by the diversity of the elements into which the effects can be analysed. The most complete set of observations on epigamic display was made by George W. and Elizabeth G. Peckham upon spiders of the family Attidae (Nat. Hist. Soc. of Wisconsin, vol. i., 1889). These observations afforded the authors “conclusive evidence that the females pay close attention to the love-dances of the males, and also that they have not only the power, but the will, to exercise a choice among the suitors for their favour.” Epigamic characters are often concealed except during courtship; they are found almost exclusively in species which are diurnal or semi-diurnal in their habits, and are excluded from those parts of the body which move too rapidly to be seen. They are very commonly directly associated with the nervous system; and in certain fish, and probably in other animals, an analogous heightening of effect accompanies nervous excitement other than sexual, such as that due to fighting or feeding. Although there is epigamic display in species with sexes alike, it is usually most marked in those with secondary sexual characters specially developed in the male. These are an exception to the rule in heredity, in that their appearance is normally restricted to a single sex, although in many of the higher animals they have been proved to be latent in the other, and may appear after the essential organs of sex have been removed or become functionless. This is also the case in the Aculeate Hymenoptera when the reproductive organs have been ​destroyed by the parasite Stylops. J. T. Cunningham has argued (Sexual Dimorphism in the Animal Kingdom, London, 1900) that secondary sexual characters have been produced by direct stimulation due to contests, &c., in the breeding period, and have gradually become hereditary, a hypothesis involving the assumption that acquired characters are transmitted. Wallace suggests that they are in part to be explained as “recognition characters,” in part as an indication of surplus vital activity in the male.
Authorities.—The following works may also be consulted:—T. Eimer, Orthogenesis der Schmetterlinge (Leipzig, 1898); E. B. Poulton, The Colours of Animals (London, 1890); F. E. Beddard, Animal Coloration (London, 1892); E. Haase, Researches on Mimicry (translation, London, 1896); A. R. Wallace, Natural Selection and Tropical Nature (London, 1895); Darwinism (London, 1897); A. H. Thayer and G. H. Thayer, Concealing-Coloration in the Animal Kingdom (New York, 1910). (E. B. P.)
2. Chemistry
The coloration of the surface of animals is caused either by pigments, or by a certain structure of the surface by means of which the light falling on it, or reflected through its superficial transparent layers, undergoes diffraction or other optical change. Or it may be the result of a combination of these two causes. It plays an important part in the relationship of the animal to its environment, in concealment, in mimicry, and so on; the presence of a pigment in the integument may also serve a more direct physiological purpose, such as a respiratory function. The coloration of birds’ feathers, of the skin of many fishes, of many insects, is partially at least due to structure and the action of the peculiar pigmented cells known as “chromatophores” (which W. Garstang defines as pigmented cells specialized for the discharge of the chromatic function), and is much better marked when these have for their background a “reflecting layer” such as is provided by guanin, a substance closely related to uric acid. Such a mechanism is seen to greatest advantage in fishes. Among these, guanin may be present in a finely granular form, causing the light falling on it to be scattered, thus producing a white effect; or it may be present in a peculiar crystalline form, the crystals being known as “iridocytes”; or in a layer of closely apposed needles forming a silvery sheet or mirror. In the iris of some fishes the golden red colour is produced by the light reflected from such a layer of guanin needles having to pass through a thin layer of a reddish pigment, known as a “lipochrome.” Again, in some lepidopterous insects a white or a yellow appearance is produced by the deposition of uric acid or a nearly allied substance on the surface of the wings. In many animals, but especially among invertebrates, colouring matters or pigments play an important rôle in surface coloration; in some cases such coloration may be of benefit to the animal, but in others the integument simply serves as an organ for the excretion of waste pigmentary substances. Pigments (1) may be of direct physiological importance; (2) they may be excretory; or (3) they may be introduced into the body of the animal with the food.
Of the many pigments which have been described up to the present time, very few have been subjected to elementary chemical analysis, owing to the great difficulties attending their isolation. An extremely small amount of pigment will give rise to a great amount of coloration, and the pigments are generally accompanied by impurities of various kinds which cling to them with great tenacity, so that when one has been thoroughly cleansed very little of it remains for ultimate analysis. Most of these substances have been detected by means of the spectroscope, their absorption bands serving for their recognition, but mere identity of spectrum does not necessarily mean chemical identity, and a few chemical tests have also to be applied before a conclusion can be drawn. The absorption bands are referred to certain definite parts of the spectrum, such as the Fraunhofer lines, or they may be given in wave-lengths. For this purpose the readings of the spectroscope are reduced to wave-lengths by means of interpolation curves; or if Zeiss’s microspectroscope be used, the position of bands in wave-lengths (denoted by the Greek letter λ) may be read directly.
Haemoglobin, the red colouring matter of vertebrate blood, C$_{758}$H$_{1203}$N$_{195}$S$_3$FeO$_{218}$, and its derivatives haematin, C$_{32}$H$_{30}$N$_4$FeO$_3$, and haematoporphyrin, C$_{16}$H$_{18}$N$_2$O$_3$, are colouring matters about which we possess definite chemical knowledge, as they have been isolated, purified and analysed. Most of the bile pigments of mammals have likewise been isolated and studied chemically, and all of these are fully described in the text-books of physiology and physiological chemistry. Haemoglobin, though physiologically of great importance in the respiratory process of vertebrate animals, is yet seldom used for surface pigmentation, except in the face of white races of man or in other parts in monkeys, &c. In some worms the transparent skin allows the haemoglobin of the blood to be seen through the integument, and in certain fishes also the haemoglobin is visible through the integument. It is a curious and noteworthy fact that in some invertebrate animals in which no haemoglobin occurs, we meet with its derivatives. Thus haematin is found in the so-called bile of slugs, snails, the limpet and the crayfish. In sea-anemones there is a pigment which yields some of the decomposition-products of haemoglobin, and associated with this is a green pigment apparently identical with biliverdin (C$_{16}$H$_{18}$N$_2$O$_4$), a green bile pigment. Again, haematoporphyrin is found in the integuments of star-fishes and slugs, and occurs in the “dorsal streak” of the earth-worm Lumbricus terrestris, and perhaps in other species. Haematoporphyrin and biliverdin also occur in the egg-shells of certain birds, but in this case they are derived from haemoglobin. Haemoglobin is said to be found as low down in the animal kingdom as the Echinoderms, e.g. in Ophiactis virens and Thyonella gemmata. It also occurs in the blood of Planorbis corneus and in the pharyngeal muscles of other mollusca.
A great number of other pigments have been described; for example, in the muscles and tissues of animals, both vertebrate and invertebrate, are the histohaematins, of which a special muscle pigment, myohaematin, is one. In vertebrates the latter is generally accompanied by haemoglobin, but in invertebrates—with the exception of the pharyngeal muscles of the mollusca—it occurs alone. Although closely related to haemoglobin or its derivative haemochromogen, the histohaematins are yet totally distinct, and they are found in animals where not a trace of haemoglobin can be detected. Another interesting pigment is turacin, which contains about 7% of nitrogen, found by Professor A. H. Church in the feathers of the Cape lory and other plantain-eaters, from which it can be extracted by water containing a trace of ammonia. It has been isolated, purified and analysed by Professor Church. From it may be obtained turacoporphyrin, which is identical with haematoporphyrin, and gives the band in the ultra-violet which J. L. Soret and subsequently A. Gamgee have found to be characteristic of haemoglobin and its compounds. Turacin itself gives a peculiar two-banded spectrum, and contains about 7% of copper in its molecule. Another copper-containing pigment is haemocyanin, which in the oxidized state gives a blue colour to the blood of various Mollusca and Arthropoda. Like haemoglobin, it acts as an oxygen-carrier in respiration, but it takes no part in surface coloration.
A class of pigments widely distributed among plants and animals are the lipochromes. As their name denotes, they are allied to fat and generally accompany it, being soluble in fat solvents. They play an important part in surface coloration, and may be greenish, yellow or red in colour. They contain no nitrogen. As an example of a lipochrome which has been isolated, crystallized and purified, we may mention carotin, which has recently been found in green leaves. Chlorophyll, which is so often associated with a lipochrome, has been found in some Infusoria, and in Hydra and Spongilla, &c. In some cases it is probably formed by the animal; in other cases it may be due to symbiotic algae, while in the gastric gland of many Mollusca, Crustacea and Echinodermata it is derived from food-chlorophyll. Here it is known as entero-chlorophyll. The black pigments which occur among both vertebrate and invertebrate animals often have only one attribute in common, viz. blackness, for among the discordant results of analysis one thing is certain, viz. that the melanins from vertebrate animals ​are not identical with those from invertebrate animals. The melanosis or blackening of insect blood, for instance, is due to the oxidation of a chromogen, the pigment produced being known as a uranidine. In some sponges a somewhat similar pigment has been noticed. Other pigments have been described, such as actiniochrome, echinochrome, pentacrinin, antedonin, polyperythrin (which appears to be a haematoporphyrin), the floridines, spongioporphyrin, &c., which need no mention here; all these pigments can only be distinguished by means of the spectroscope.
Most of the pigments are preceded by colourless substances known as “chromogens,” which by the action of the oxygen of the air and by other agencies become changed into the corresponding pigments. In some cases the pigments are built up in the tissues of an animal, in others they appear to be derived more or less directly from the food. Derivatives of chlorophyll and lipochromes especially, seem to be taken up from the intestine, probably by the agency of leucocytes, in which they may occur in combination with, or dissolved by, fatty matters and excreted by the integument. In worms especially, the skin seems to excrete many effete substances, pigments included. No direct connexion has been traced between the chlorophyll eaten with the food and the haemoglobin of blood and muscle. Attention may, however, be drawn to the work of Dr E. Schunck, who has shown that a substance closely resembling haematoporphyrin can be prepared from chlorophyll; this is known as phylloporphyrin. Not only does the visible spectrum of this substance resemble that of haematoporphyrin, but the invisible ultra-violet also, as shown by C. A. Schunck.
The reader may refer to E. A. Schäfer’s Text-Book of Physiology (1898) for A. Gamgee’s article “On Haemoglobin, and its Compounds”; to the writer’s papers in the Phil. Trans. and Proc. Roy. Soc. from 1881 onwards, and also Quart. Journ. Micros. Science and Journ. of Physiol.; to C. F. W. Krukenberg’s Vergleichende physiologische Studien from 1879 onwards, and to his Vorträge. Miss M. I. Newbigin collected in Colour in Nature (1898) most of the recent literature of this subject. Dr E. Schunck’s papers will be found under the heading “Contribution to the Chemistry of Chlorophyll” in Proc. Roy. Soc. from 1885 onwards; and Mr C. A. Schunck’s paper in Proc. Roy. Soc. vol. lxiii. (C. A. MacM.)