see that the whole problem is so intricate that it is safer to leave its solution for the future.

CHAPTER IX.

SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL KINGDOM.

These characters absent in the lowest classes-Brilliant colours-Mollusca -Annelids-Crustacea, secondary sexual characters strongly developed ; dimorphism; colour; characters not acquired before maturity-Spiders, sexual colours of; stridulation by the males-Myriapoda.

WITH animals belonging to the lower classes, the two sexes are not rarely united in the same individual, and therefore secondary sexual characters cannot be developed. In many cases where the sexes are separate, both are permanently attached to some support, and the one cannot search or struggle for the other. Moreover it is almost certain that these animals have too imperfect senses and much too low mental powers, to appreciate each other's beauty or other attractions, or to feel rivalry.

Hence in these classes or sub-kingdoms, such as the Protozoa, Cœlenterata, Echinodermata, Scolecida, secondary sexual characters, of the kind which we have to consider, do not occur; and this fact agrees with the belief that such characters in the higher classes have been acquired through sexual selection, which depends on the will, desire, and choice of either sex. Nevertheless some few apparent exceptions occur; thus, as I hear from Dr. Baird, the males of certain Entozoa, or internal parasitic worms, differ slightly in colour from the females; but we have no reason to suppose that such differences have been augmented through sexual selection. Contrivances by which the male holds the female, and which are indispensable for the propagation of the species, are independent of sexual selection, and have been acquired through ordinary selection.

Many of the lower animals, whether hermaphrodites or with separate sexes, are ornamented with the most brilliant tints, or are shaded and striped in an elegant manner; for instance, many corals and sea-anemones (Actinia), some jelly-fish (Medusa, Porpita, &c.), some Planariæ, many star-fishes, Echini, Ascidians, &c.; but we may conclude from the reasons already indicated, namely the union of the two sexes in some of these animals, the permanently affixed condition of others, and the low mental powers of all, that such colours do not serve as a sexual attraction, and have not been acquired through sexual selection.

It should be borne in mind that in no case have we sufficient evidence that colours have been thus acquired, except where one sex is much more brilliantly or conspicuously coloured than the other, and where there is no difference in habits between the sexes sufficient to account for their different colours. But the evidence is rendered as complete as it can ever be, only when the more ornamented individuals, almost always the males, voluntarily display their attractions before the other sex; for we cannot believe that such display is useless, and if it be advantageous, sexual selection will almost inevitably follow. We may, however, extend this conclusion to both sexes, when coloured alike, if their colours are plainly analogous to those of one sex alone in certain other species of the same group.

How, then, are we to account for the beautiful or even gorgeous colours of many animals in the lowest classes? It appears doubtful whether such colours often serve as a protection; but that we may easily err on this head, will be admitted by every one who reads Mr. Wallace's excellent essay on this subject. It would not, for instance, at first occur to any one that the transparency of the Medusa, or jelly-fishes, is of the highest service to them as a protection; but when we are reminded by Häckel that not only the medusa, but many floating mollusca, crustaceans, and even small oceanic fishes partake of this same glass-like appearance, often accompanied by prismatic colours, we can hardly doubt that they thus escape the notice of pelagic birds and other enemies. M. Girard is also convinced that the bright tints of certain sponges and ascidians serve as a protection. Conspicuous colours are likewise beneficial to many animals as a warning to their would-be devourers that they are distasteful, or that they possess some special means of defence; but this subject will be discussed more conveniently hereafter.

We can, in our ignorance of most of the lowest animals, only say that their bright tints result either from the chemical nature or the minute structure of their tissues, independently of any benefit thus derived. Hardly any colour is finer than that of arterial blood; but there is no reason to suppose that the colour of the blood is in itself any advantage; and though it adds to the beauty of the maiden's cheek, no one will pretend that it has been acquired for this purpose. So again with many animals, especially the lower ones, the bile is richly coloured; thus, as I am informed by Mr. Hancock, the extreme beauty of the Eolidæ (naked sea-slugs) is chiefly due to the biliary glands 1 Archives de Zoolog. Expér.,' Oct. 1872, p. 563.

being seen through the translucent integuments-this beauty being probably of no service to these animals. The tints of the decaying leaves in an American forest are described by every one as gorgeous; yet no one supposes that these tints are of the least advantage to the trees. Bearing in mind how many substances closely analogous to natural organic compounds have been recently formed by chemists, and which exhibit the most splendid colours, it would have been a strange fact if substances similarly coloured had not often originated, independently of any useful end thus gained, in the complex laboratory of living organisms.

never occur.

The sub-kingdom of the Mollusca.-Throughout this great division of the animal kingdom, as far as I can discover, secondary sexual characters, such as we are here considering, Nor could they be expected in the three lowest classes, namely in the Ascidians, Polyzoa, and Brachiopods (constituting the Molluscoida of some authors), for most of these animals are permanently affixed to a support or have their sexes united in the same individual. In the Lamellibranchiata, or bivalve shells, hermaphroditism is not rare. In the next higher class of the Gasteropoda, or univalve shells, the sexes are either united or separate. But in the latter case the males never possess special organs for finding, securing, or charming the females, or for fighting with other males. As I am informed by Mr. Gwyn Jeffreys, the sole external difference between the sexes consists in the shell sometimes differing a little in form; for instance, the shell of the male periwinkle (Littorina littorea) is narrower and has a more elongated spire than that of the female. But differences of this nature, it may be presumed, are directly connected with the act of reproduction, or with the development of the ova.

The Gasteropoda, though capable of locomotion and furnished with imperfect eyes, do not appear to be endowed with sufficient mental powers for the members of the same sex to struggle together in rivalry, and thus to acquire secondary sexual characters. Nevertheless with the pulmoniferous gasteropods, or land-snails, the pairing is preceded by courtship; for these animals, though hermaphrodites, are compelled by their structure to pair together. Agassiz remarks," "Quiconque a eu l'occasion "d'observer les amours des limaçons, ne saurait mettre en doute "la séduction déployée dans les mouvements et les allures qui "préparent et accomplissent le double embrassement de ces hermaphrodites." These animals appear also susceptible of some degree of permanent attachment: an accurate observer, 2De l'Espèce et de la Class,' &c., 1869, p. 106.

Mr. Lonsdale, informs me that he placed a pair of land-snails, (Helix pomatia), one of which was weakly, into a small and illprovided garden. After a short time the strong and healthy individual disappeared, and was traced by its track of slime over a wall into an adjoining well-stocked garden. Mr. Lonsdale concluded that it had deserted its sickly mate; but after an absence of twenty-four hours it returned, and apparently communicated the result of its successful exploration, for both then started along the same track and disappeared over the wall.

Even in the highest class of the Mollusca, the Cephalopoda or cuttlefishes, in which the sexes are separate, secondary sexual characters of the present kind do not, as far as I can discover, occur. This is a surprising circumstance, as these animals possess highly-developed sense-organs and have considerable mental powers, as will be admitted by every one who has watched their artful endeavours to escape from an enemy.3 Certain Cephalopoda, however, are characterised by one extraordinary sexual character, namely, that the male element collects within one of the arms or tentacles, which is then cast off, and clinging by its sucking-discs to the female, lives for a time an independent life. So completely does the cast-off arm resemble a separate animal, that it was described by Cuvier as a parasitic worm under the name of Hectocotyle. But this marvellous structure may be classed as a primary rather than as a secondary sexual character.

Although with the Mollusca sexual selection does not seem to have come into play; yet many univalve and bivalve shells, such as volutes, cones, scallops, &c., are beautifully coloured and shaped. The colours do not appear in most cases to be of any use as a protection; they are probably the direct result, as in the lowest classes, of the nature of the tissues; the patterns and the sculpture of the shell depending on its manner of growth. The amount of light seems to be influential to a certain extent; for although, as repeatedly stated by Mr. Gwyn Jeffreys, the shells of some species living at a profound depth are brightly coloured, yet we generally see the lower surfaces, as well as the parts covered by the mantle, less highly-coloured than the upper and exposed surfaces.

See, for instance, the account which I have given in my 'Journal of Researches,' 1845, p. 7.

I have given (Geolog. Observations on Volcanic Islands,' 1844, p. 53) a curious instance of the

In some cases, as with shells

influence of light on the colours of a frondescent incrustation, deposited by the surf on the coastrocks of Ascension, and formed by the solution of triturated sea-shells.

living amongst corals or brightly-tinted sea-weeds, the bright colours may serve as a protection. But that many of the nudibranch mollusca, or sea-slugs, are as beautifully coloured as any shells, may be seen in Messrs. Alder and Hancock's magnificent work; and from information kindly given me by Mr. Hancock, it seems extremely doubtful whether these colours usually serve as a protection. With some species this may be the case, as with one kind which lives on the green leaves of algæ, and is itself bright-green. But many brightly-coloured, white or otherwise conspicuous species, do not seek concealment; whilst again some equally conspicuous species, as well as other dull-coloured kinds, live under stones and in dark recesses. So that with these nudibranch molluscs, colour apparently does not stand in any close relation to the nature of the places which they inhabit.

These naked sea-slugs are hermaphrodites, yet they pair together, as do land-snails, many of which have extremely pretty shells. It is conceivable that two hermaphrodites, attracted by each other's greater beauty, might unite and leave offspring which would inherit their parents' greater beauty. But with such lowly-organised creatures this is extremely improbable. Nor is it at all obvious how the offspring from the more beautiful pairs of hermaphrodites would have any advantage over the offspring of the less beautiful, so as to increase in number, unless indeed vigour and beauty generally coincided. We have not here the case of a number of males becoming mature before the females, with the more beautiful males selected by the more vigorous females. If, indeed, brilliant colours were beneficial to a hermaphrodite animal in relation to its general habits of life, the more brightly-tinted individuals would succeed best and would increase in number; but this would be a case of natural and not of sexual selection.

Sub-kingdom of the Vermes: Class, Annelida (or Sea-worms).— In this class, although the sexes, when separate, sometimes differ from each other in characters of such importance that they have been placed under distinct genera or even families, yet the differences do not seem of the kind which can be safely attributed to sexual selection. These animals are often beautifully coloured, but as the sexes do not differ in this respect, we are but little concerned with them. Even the Nemertians, though so lowly organised, "vie in beauty and variety of colouring with any other group in the invertebrate series;" yet

5 Dr. Morse has lately discussed this subject in his paper on the Adaptive Coloration of Mollusca,

'Proc. Boston Soc. of Nat. Hist.' vol. xiv., April, 1871.