Contributions to the Theory of Natural Selection. Alfred Russel Wallace

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down upon the desert, how, it may be asked, do the elegant markings of the tiger, the jaguar, and the other large cats agree with this theory? We reply that these are generally cases of more or less special adaptation. The tiger is a jungle animal, and hides himself among tufts of grass or of bamboos, and in these positions the vertical stripes with which his body is adorned must so assimilate with the vertical stems of the bamboo, as to assist greatly in concealing him from his approaching prey. How remarkable it is that besides the lion and tiger, almost all the other large cats are arboreal in their habits, and almost all have ocellated or spotted skins, which must certainly tend to blend them with the background of foliage; while the one exception, the puma, has an ashy brown uniform fur, and has the habit of clinging so closely to a limb of a tree while waiting for his prey to pass beneath as to be hardly distinguishable from the bark.

      Among birds, the ptarmigan, already mentioned, must be considered a remarkable case of special adaptation. Another is a South-American goatsucker (Caprimulgus rupestris) which rests in the bright sunshine on little bare rocky islets in the Upper Rio Negro, where its unusually light colours so closely resemble those of the rock and sand, that it can scarcely be detected till trodden upon.

      The Duke of Argyll, in his “Reign of Law,” has pointed out the admirable adaptation of the colours of the woodcock to its protection. The various browns and yellows and pale ash-colour that occur in fallen leaves are all reproduced in its plumage, so that when according to its habit it rests upon the ground under trees, it is almost impossible to detect it. In snipes the colours are modified so as to be equally in harmony with the prevalent forms and colours of marshy vegetation. Mr. J. M. Lester, in a paper read before the Rugby School Natural History Society, observes:—“The wood-dove, when perched amongst the branches of its favourite fir, is scarcely discernible; whereas, were it among some lighter foliage, the blue and purple tints in its plumage would far sooner betray it. The robin redbreast too, although it might be thought that the red on its breast made it much easier to be seen, is in reality not at all endangered by it, since it generally contrives to get among some russet or yellow fading leaves, where the red matches very well with the autumn tints, and the brown of the rest of the body with the bare branches.”

      Reptiles offer us many similar examples. The most arboreal lizards, the iguanas, are as green as the leaves they feed upon, and the slender whip-snakes are rendered almost invisible as they glide among the foliage by a similar colouration. How difficult it is sometimes to catch sight of the little green tree-frogs sitting on the leaves of a small plant enclosed in a glass case in the Zoological Gardens; yet how much better concealed must they be among the fresh green damp foliage of a marshy forest. There is a North-American frog found on lichen-covered rocks and walls, which is so coloured as exactly to resemble them, and as long as it remains quiet would certainly escape detection. Some of the geckos which cling motionless on the trunks of trees in the tropics, are of such curiously marbled colours as to match exactly with the bark they rest upon.

      In every part of the tropics there are tree-snakes that twist among boughs and shrubs, or lie coiled up on the dense masses of foliage. These are of many distinct groups, and comprise both venomous and harmless genera; but almost all of them are of a beautiful green colour, sometimes more or less adorned with white or dusky bands and spots. There can be little doubt that this colour is doubly useful to them, since it will tend to conceal them from their enemies, and will lead their prey to approach them unconscious of danger. Dr. Gunther informs me that there is only one genus of true arboreal snakes (Dipsas) whose colours are rarely green, but are of various shades of black, brown, and olive, and these are all nocturnal reptiles, and there can be little doubt conceal themselves during the day in holes, so that the green protective tint would be useless to them, and they accordingly retain the more usual reptilian hues.

      Fishes present similar instances. Many flat fish, as for example the flounder and the skate, are exactly the colour of the gravel or sand on which they habitually rest. Among the marine flower gardens of an Eastern coral reef the fishes present every variety of gorgeous colour, while the river fish even of the tropics rarely if ever have gay or conspicuous markings. A very curious case of this kind of adaptation occurs in the sea-horses (Hippocampus) of Australia, some of which bear long foliaceous appendages resembling seaweed, and are of a brilliant red colour; and they are known to live among seaweed of the same hue, so that when at rest they must be quite invisible. There are now in the aquarium of the Zoological Society some slender green pipe-fish which fasten themselves to any object at the bottom by their prehensile tails, and float about with the current, looking exactly like some simple cylindrical algæ.

      It is, however, in the insect world that this principle of the adaptation of animals to their environment is most fully and strikingly developed. In order to understand how general this is, it is necessary to enter somewhat into details, as we shall thereby be better able to appreciate the significance of the still more remarkable phenomena we shall presently have to discuss. It seems to be in proportion to their sluggish motions or the absence of other means of defence, that insects possess the protective colouring. In the tropics there are thousands of species of insects which rest during the day clinging to the bark of dead or fallen trees; and the greater portion of these are delicately mottled with gray and brown tints, which though symmetrically disposed and infinitely varied, yet blend so completely with the usual colours of the bark, that at two or three feet distance they are quite undistinguishable. In some cases a species is known to frequent only one species of tree. This is the case with the common South American long-horned beetle (Onychocerus scorpio) which, Mr. Bates informed me, is found only on a rough-barked tree, called Tapiribá, on the Amazon. It is very abundant, but so exactly does it resemble the bark in colour and rugosity, and so closely does it cling to the branches, that until it moves it is absolutely invisible! An allied species (O. concentricus) is found only at Pará, on a distinct species of tree, the bark of which it resembles with equal accuracy. Both these insects are abundant, and we may fairly conclude that the protection they derive from this strange concealment is at least one of the causes that enable the race to flourish.

      Many of the species of Cicindela, or tiger beetle, will illustrate this mode of protection. Our common Cicindela campestris frequents grassy banks, and is of a beautiful green colour, while C. maritima, which is found only on sandy sea-shores, is of a pale bronzy yellow, so as to be almost invisible. A great number of the species found by myself in the Malay islands are similarly protected. The beautiful Cicindela gloriosa, of a very deep velvety green colour, was only taken upon wet mossy stones in the bed of a mountain stream, where it was with the greatest difficulty detected. A large brown species (C. heros) was found chiefly on dead leaves in forest paths; and one which was never seen except on the wet mud of salt marshes was of a glossy olive so exactly the colour of the mud as only to be distinguished when the sun shone, by its shadow! Where the sandy beach was coralline and nearly white, I found a very pale Cicindela; wherever it was volcanic and black, a dark species of the same genus was sure to be met with.

      There are in the East small beetles of the family Buprestidæ which generally rest on the midrib of a leaf, and the naturalist often hesitates before picking them off, so closely do they resemble pieces of bird’s dung. Kirby and Spence mention the small beetle Onthophilus sulcatus as being like the seed of an umbelliferous plant; and another small weevil, which is much persecuted by predatory beetles of the genus Harpalus, is of the exact colour of loamy soil, and was found to be particularly abundant in loam pits. Mr. Bates mentions a small beetle (Chlamys pilula) which was undistinguishable by the eye from the dung of caterpillars, while some of the Cassidæ, from their hemispherical forms and pearly gold colour, resemble glittering dew-drops upon the leaves.

      A number of our small brown and speckled weevils at the approach of any object roll off the leaf they are sitting on, at the same time drawing in their legs and antennæ, which fit so perfectly into cavities for their reception that the insect becomes a mere oval brownish lump, which it is hopeless to look for among the similarly coloured little stones and earth pellets among which it lies motionless.

      The distribution of colour in butterflies and moths respectively is very instructive from this point of view. The former have all their brilliant colouring on the upper surface

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