The Descent of Man, and Selection in Relation to Sex

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 Dr. McIntosh (6. See his beautiful monograph on 'British Annelids,' part i. 1873, p. 3.) cannot discover that these colours are of any service. The sedentary annelids become duller-coloured, according to M. Quatrefages (7. See M. Perrier: 'L'Origine de l'Homme d'après Darwin,' 'Revue Scientifique', Feb. 1873, p. 866.), after the period of reproduction; and this I presume may be attributed to their less vigorous condition at that time. All these worm- like animals apparently stand too low in the scale for the individuals of either sex to exert any choice in selecting a partner, or for the individuals of the same sex to struggle together in rivalry.

SUB-KINGDOM OF THE ARTHROPODA: CLASS, CRUSTACEA

In this great class we first meet with undoubted secondary sexual characters, often developed in a remarkable manner. Unfortunately the habits of crustaceans are very imperfectly known, and we cannot explain the uses of many structures peculiar to one sex. With the lower parasitic species the males are of small size, and they alone are furnished with perfect swimming-legs, antennae and sense-organs; the females being destitute of these organs, with their bodies often consisting of a mere distorted mass. But these extraordinary differences between the two sexes are no doubt related to their widely different habits of life, and consequently do not concern us. In various crustaceans, belonging to distinct families, the anterior antennae are furnished with peculiar thread-like bodies, which are believed to act as smelling-organs, and these are much more numerous in the males than in the females. As the males, without any unusual development of their olfactory organs, would almost certainly be able sooner or later to find the females, the increased number of the smelling-threads has probably been acquired through sexual selection, by the better provided males having been the more successful in finding partners and in producing offspring. Fritz Müller has described a remarkable dimorphic species of Tanais, in which the male is represented by two distinct forms, which never graduate into each other. In the one form the male is furnished with more numerous smelling-threads, and in the other form with more powerful and more elongated chelae or pincers, which serve to hold the female. Fritz Müller suggests that these differences between the two male forms of the same species may have originated in certain individuals having varied in the number of the smelling-threads, whilst other individuals varied in the shape and size of their chelae; so that of the former, those which were best able to find the female, and of the latter, those which were best able to hold her, have left the greatest number of progeny to inherit their respective advantages. (8. 'Facts and Arguments for Darwin,' English translat., 1869, p. 20. See the previous discussion on the olfactory threads. Sars has described a somewhat analogous case (as quoted in 'Nature,' 1870, p. 455) in a Norwegian crustacean, the Pontoporeia affinis.)

[Fig.4. Labidocera Darwinii (from Lubbock). Labelled are: a. Part of right anterior antenna of male, forming a prehensile organ. b. Posterior pair of thoracic legs of male. c. Ditto of female.]

In some of the lower crustaceans, the right anterior antenna of the male differs greatly in structure from the left, the latter resembling in its simple tapering joints the antennae of the female. In the male the modified antenna is either swollen in the middle or angularly bent, or converted (Fig. 4) into an elegant, and sometimes wonderfully complex, prehensile organ. (9. See Sir J. Lubbock in 'Annals and Mag. of Nat. Hist.' vol. xi. 1853, pl. i. and x.; and vol. xii. (1853), pl. vii. See also Lubbock in 'Transactions, Entomological Society,' vol. iv. new series, 1856-1858, p. 8. With respect to the zigzagged antennae mentioned below, see Fritz Müller, 'Facts and Arguments for Darwin,' 1869, p. 40, foot- note.) It serves, as I hear from Sir J. Lubbock, to hold the female, and for this same purpose one of the two posterior legs (b) on the same side of the body is converted into a forceps. In another family the inferior or posterior antennae are "curiously zigzagged" in the males alone.

[Fig. 5. Anterior part of body of Callianassa (from Milne-Edwards), showing the unequal and differently-constructed right and left-hand chelae of the male. N.B. – The artist by mistake has reversed the drawing, and made the left-hand chela the largest.

Fig. 6. Second leg of male Orchestia Tucuratinga (from Fritz Müller).

Fig. 7. Ditto of female.]

In the higher crustaceans the anterior legs are developed into chelae or pincers; and these are generally larger in the male than in the female, – so much so that the market value of the male edible crab (Cancer pagurus), according to Mr. C. Spence Bate, is five times as great as that of the female. In many species the chelae are of unequal size on the opposite side of the body, the right-hand one being, as I am informed by Mr. Bate, generally, though not invariably, the largest. This inequality is also often much greater in the male than in the female. The two chelae of the male often differ in structure (Figs. 5, 6, and 7), the smaller one resembling that of the female. What advantage is gained by their inequality in size on the opposite sides of the body, and by the inequality being much greater in the male than in the female; and why, when they are of equal size, both are often much larger in the male than in the female, is not known. As I hear from Mr. Bate, the chelae are sometimes of such length and size that they cannot possibly be used for carrying food to the mouth. In the males of certain fresh-water prawns (Palaemon) the right leg is actually longer than the whole body. (10. See a paper by Mr. C. Spence Bate, with figures, in 'Proceedings, Zoological Society,' 1868, p. 363; and on the nomenclature of the genus, ibid. p. 585. I am greatly indebted to Mr. Spence Bate for nearly all the above statements with respect to the chelae of the higher crustaceans.) The great size of the one leg with its chelae may aid the male in fighting with his rivals; but this will not account for their inequality in the female on the opposite sides of the body. In Gelasimus, according to a statement quoted by Milne Edwards (11. 'Hist. Nat. des Crust.' tom. ii. 1837, p. 50.), the male and the female live in the same burrow, and this shews that they pair; the male closes the mouth of the burrow with one of its chelae, which is enormously developed; so that here it indirectly serves as a means of defence. Their main use, however, is probably to seize and to secure the female, and this in some instances, as with Gammarus, is known to be the case. The male of the hermit or soldier crab (Pagurus) for weeks together, carries about the shell inhabited by the female. (12. Mr. C. Spence Bate, 'British Association, Fourth Report on the Fauna of S. Devon.') The sexes, however, of the common shore-crab (Carcinus maenas), as Mr. Bate informs me, unite directly after the female has moulted her hard shell, when she is so soft that she would be injured if seized by the strong pincers of the male; but as she is caught and carried about by the male before moulting, she could then be seized with impunity.

[Fig.8. Orchestia Darwinii (from Fritz Müller), showing the differently- constructed chelae of the two male forms.]

Fritz Müller states that certain species of Melita are distinguished from all other amphipods by the females having "the coxal lamellae of the penultimate pair of feet produced into hook-like processes, of which the males lay hold with the hands of the first pair." The development of these hook-like processes has probably followed from those females which were the most securely held during the act of reproduction, having left the largest number of offspring. Another Brazilian amphipod (see Orchestia darwinii, Fig. 8) presents a case of dimorphism, like that of Tanais; for there are two male forms, which differ in the structure of their chelae. (13. Fritz Müller, 'Facts and Arguments for Darwin,' 1869, pp. 25-28.) As either chela would certainly suffice to hold the female, – for both are now used for this purpose, – the two male forms probably originated by some having varied in one manner and some in another; both forms having derived certain special, but nearly equal advantages, from their differently shaped organs.

It is not known that male crustaceans fight together for the possession of the females, but it is probably the case; for with most animals when the male is larger than the female, he seems to owe his greater size to his ancestors having fought with other males during many generations. In most of the orders, especially in the highest or the Brachyura, the male is larger than the female; the parasitic genera, however, in which the sexes follow different habits of life, and most of the Entomostraca must be excepted. The chelae of many crustaceans are weapons well adapted for fighting. Thus when a Devil-crab (Portunus puber) was seen by a son of Mr. Bate fighting with a Carcinus maenas, the latter was soon thrown on its back, and had every limb torn from its body. When several males of a Brazilian Gelasimus, a species furnished with immense pincers, were placed together in a glass vessel by Fritz Müller, they mutilated and killed one another. Mr. Bate put a large male Carcinus maenas into a pan of water, inhabited by a female which was paired with a smaller male; but the latter was soon dispossessed. Mr. Bate adds, "if they fought, the victory was a bloodless one, for I saw no wounds." This same naturalist separated a male sand-skipper (so common on our sea-shores), Gammarus marinus, from its female, both of whom were imprisoned in the same vessel with many individuals of the same species. The female, when thus divorced, soon joined the others. After a time the male was put again into the same vessel; and he then, after swimming about for a time, dashed into the crowd, and without any fighting at once took away his wife. This fact shews that in the Amphipoda, an order low in the scale, the males and females recognise each other, and are mutually attached.

The mental powers of the Crustacea are probably higher than at first sight appears probable. Any one who tries to catch one of the shore-crabs, so common on tropical coasts, will perceive how wary and alert they are. There is a large crab (Birgus latro), found on coral islands, which makes a thick bed of the picked fibres of the cocoa-nut, at the bottom of a deep burrow. It feeds on the fallen fruit of this tree by tearing off the husk, fibre by fibre; and it always begins at that end where the three eye-like depressions are situated. It then breaks through one of these eyes by hammering with its heavy front pincers, and turning round, extracts the albuminous core with its narrow posterior pincers. But these actions are probably instinctive, so that they would be performed as well by a young animal as by an old one. The following case, however, can hardly be so considered: a trustworthy naturalist, Mr. Gardner (14. 'Travels in the Interior of Brazil,' 1846, p. 111. I have given, in my 'Journal of Researches,' p. 463, an account of the habits of the Birgus.), whilst watching a shore-crab (Gelasimus) making its burrow, threw some shells towards the hole. One rolled in, and three other shells remained within a few inches of the mouth. In about five minutes the crab brought out the shell which had fallen in, and carried it away to a distance of a foot; it then saw the three other shells lying near, and evidently thinking that they might likewise roll in, carried them to the spot where it had laid the first. It would, I think, be difficult to distinguish this act from one performed by man by the aid of reason.

Mr. Bate does not know of any well-marked case of difference of colour in the two sexes of our British crustaceans, in which respect the sexes of the higher animals so often differ. In some cases, however, the males and females differ slightly in tint, but Mr. Bate thinks not more than may be accounted for by their different habits of life, such as by the male wandering more about, and being thus more exposed to the light. Dr. Power tried to distinguish by colour the sexes of the several species which inhabit the Mauritius, but failed, except with one species of Squilla, probably S. stylifera, the male of which is described as being "of a beautiful bluish-green," with some of the appendages cherry-red, whilst the female is clouded with brown and grey, "with the red about her much less vivid than in the male." (15. Mr. Ch. Fraser, in 'Proc. Zoolog. Soc.' 1869, p. 3. I am indebted to Mr. Bate for Dr. Power's statement.) In this case, we may suspect the agency of sexual selection. From M. Bert's observations on Daphnia, when placed in a vessel illuminated by a prism, we have reason to believe that even the lowest crustaceans can distinguish colours. With Saphirina (an oceanic genus of Entomostraca), the males are furnished with minute shields or cell-like bodies, which exhibit beautiful changing colours; these are absent in the females, and in both sexes of one species. (16. Claus, 'Die freilebenden Copepoden,' 1863, s. 35.) It would, however, be extremely rash to conclude that these curious organs serve to attract the females. I am informed by Fritz Müller, that in the female of a Brazilian species of Gelasimus, the whole body is of a nearly uniform greyish-brown. In the male the posterior part of the cephalo- thorax is pure white, with the anterior part of a rich green, shading into dark brown; and it is remarkable that these colours are liable to change in the course of a few minutes – the white becoming dirty grey or even black, the green "losing much of its brilliancy." It deserves especial notice that the males do not acquire their bright colours until they become mature. They appear to be much more numerous than the females; they differ also in the larger size of their chelae. In some species of the genus, probably in all, the sexes pair and inhabit the same burrow. They are also, as we have seen, highly intelligent animals. From these various considerations it seems probable that the male in this species has become gaily ornamented in order to attract or excite the female.

It has just been stated that the male Gelasimus does not acquire his conspicuous colours until mature and nearly ready to breed. This seems a general rule in the whole class in respect to the many remarkable structural differences between the sexes. We shall hereafter find the same law prevailing throughout the great sub-kingdom of the Vertebrata; and in all cases it is eminently distinctive of characters which have been acquired through sexual selection. Fritz Müller (17. 'Facts and Arguments,' etc., p. 79.) gives some striking instances of this law; thus the male sand-hopper (Orchestia) does not, until nearly full grown, acquire his large claspers, which are very differently constructed from those of the female; whilst young, his claspers resemble those of the female.

CLASS, ARACHNIDA (SPIDERS)

The sexes do not generally differ much in colour, but the males are often darker than the females, as may be seen in Mr. Blackwall's magnificent work. (18. 'A History of the Spiders of Great Britain,' 1861-64. For the following facts, see pp. 77, 88, 102.) In some species, however, the difference is conspicuous: thus the female of Sparassus smaragdulus is dullish green, whilst the adult male has the abdomen of a fine yellow, with three longitudinal stripes of rich red. In certain species of Thomisus the sexes closely resemble each other, in others they differ much; and analogous cases occur in many other genera. It is often difficult to say which of the two sexes departs most from the ordinary coloration of the genus to which the species belong; but Mr. Blackwall thinks that, as a general rule, it is the male; and Canestrini (19. This author has recently published a valuable essay on the 'Caratteri sessuali secondarii degli Arachnidi,' in the 'Atti della Soc. Veneto-Trentina di Sc. Nat. Padova,' vol. i. Fasc. 3, 1873.) remarks that in certain genera the males can be specifically distinguished with ease, but the females with great difficulty. I am informed by Mr. Blackwall that the sexes whilst young usually resemble each other; and both often undergo great changes in colour during their successive moults, before arriving at maturity. In other cases the male alone appears to change colour. Thus the male of the above bright-coloured Sparassus at first resembles the female, and acquires his peculiar tints only when nearly adult. Spiders are possessed of acute senses, and exhibit much intelligence; as is well known, the females often shew the strongest affection for their eggs, which they carry about enveloped in a silken web. The males search eagerly for the females, and have been seen by Canestrini and others to fight for possession of them. This same author says that the union of the two sexes has been observed in about twenty species; and he asserts positively that the female rejects some of the males who court her, threatens them with open mandibles, and at last after long hesitation accepts the chosen one. From these several considerations, we may admit with some confidence that the well-marked differences in colour between the sexes of certain species are the results of sexual selection; though we have not here the best kind of evidence, – the display by the male of his ornaments. From the extreme variability of colour in the male of some species, for instance of Theridion lineatum, it would appear that these sexual characters of the males have not as yet become well fixed. Canestrini draws the same conclusion from the fact that the males of certain species present two forms, differing from each other in the size and length of their jaws; and this reminds us of the above cases of dimorphic crustaceans.

The male is generally much smaller than the female, sometimes to an extraordinary degree (20. Aug. Vinson ('Araneides des Iles de la Reunion,' pl. vi. figs. 1 and 2) gives a good instance of the small size of the male, in Epeira nigra. In this species, as I may add, the male is testaceous and the female black with legs banded with red. Other even more striking cases of inequality in size between the sexes have been recorded ('Quarterly Journal of Science,' July 1868, p. 429); but I have not seen the original accounts.), and he is forced to be extremely cautious in making his advances, as the female often carries her coyness to a dangerous pitch. De Geer saw a male that "in the midst of his preparatory caresses was seized by the object of his attentions, enveloped by her in a web and then devoured, a sight which, as he adds, filled him with horror and indignation." (21. Kirby and Spence, 'Introduction to Entomology,' vol. i. 1818, p. 280.) The Rev. O.P. Cambridge (22. 'Proceedings, Zoological Society,' 1871, p. 621.) accounts in the following manner for the extreme smallness of the male in the genus Nephila. "M. Vinson gives a graphic account of the agile way in which the diminutive male escapes from the ferocity of the female, by gliding about and playing hide and seek over her body and along her gigantic limbs: in such a pursuit it is evident that the chances of escape would be in favour of the smallest males, while the larger ones would fall early victims; thus gradually a diminutive race of males would be selected, until at last they would dwindle to the smallest possible size compatible with the exercise of their generative functions, – in fact, probably to the size we now see them, i.e., so small as to be a sort of parasite upon the female, and either beneath her notice, or too agile and too small for her to catch without great difficulty."

Westring has made the interesting discovery that the males of several species of Theridion (23. Theridion (Asagena, Sund.) serratipes, 4- punctatum et guttatum; see Westring, in Kroyer, 'Naturhist. Tidskrift,' vol. iv. 1842-1843, p. 349; and vol. ii. 1846-1849, p. 342. See, also, for other species, 'Araneae Suecicae,' p. 184.) have the power of making a stridulating sound, whilst the females are mute. The apparatus consists of a serrated ridge at the base of the abdomen, against which the hard hinder part of the thorax is rubbed; and of this structure not a trace can be detected in the females. It deserves notice that several writers, including the well-known arachnologist Walckenaer, have declared that spiders are attracted by music. (24. Dr. H.H. van Zouteveen, in his Dutch translation of this work (vol. i. p. 444), has collected several cases.) From the analogy of the Orthoptera and Homoptera, to be described in the next chapter, we may feel almost sure that the stridulation serves, as Westring also believes, to call or to excite the female; and this is the first case known to me in the ascending scale of the animal kingdom of sounds emitted for this purpose. (25. Hilgendorf, however, has lately called attention to an analogous structure in some of the higher crustaceans, which seems adapted to produce sound; see 'Zoological Record,' 1869, p. 603.)

CLASS, MYRIAPODA

In neither of the two orders in this class, the millipedes and centipedes, can I find any well-marked instances of such sexual differences as more particularly concern us. In Glomeris limbata, however, and perhaps in some few other species, the males differ slightly in colour from the females; but this Glomeris is a highly variable species. In the males of the Diplopoda, the legs belonging either to one of the anterior or of the posterior segments of the body are modified into prehensile hooks which serve to secure the female. In some species of Iulus the tarsi of the male are furnished with membranous suckers for the same purpose. As we shall see when we treat of Insects, it is a much more unusual circumstance, that it is the female in Lithobius, which is furnished with prehensile appendages at the extremity of her body for holding the male. (26. Walckenaer et P. Gervais, 'Hist. Nat. des Insectes: Apteres,' tom. iv. 1847, pp. 17, 19, 68.)

CHAPTER X

SECONDARY SEXUAL CHARACTERS OF INSECTS

Diversified structures possessed by the males for seizing the females – Differences between the sexes, of which the meaning is not understood – Difference in size between the sexes – Thysanura – Diptera – Hemiptera – Homoptera, musical powers possessed by the males alone – Orthoptera, musical instruments of the males, much diversified in structure; pugnacity; colours – Neuroptera, sexual differences in colour – Hymenoptera, pugnacity and odours – Coleoptera, colours; furnished with great horns, apparently as an ornament; battles, stridulating organs generally common to both sexes.

In the immense class of insects the sexes sometimes differ in their locomotive-organs, and often in their sense-organs, as in the pectinated and beautifully plumose antennae of the males of many species. In Chloeon, one of the Ephemerae, the male has great pillared eyes, of which the female is entirely destitute. (1. Sir J. Lubbock, 'Transact. Linnean Soc.' vol. xxv, 1866, p. 484. With respect to the Mutillidae see Westwood, 'Modern Class. of Insects,' vol. ii. p. 213.) The ocelli are absent in the females of certain insects, as in the Mutillidae; and here the females are likewise wingless. But we are chiefly concerned with structures by which one male is enabled to conquer another, either in battle or courtship, through his strength, pugnacity, ornaments, or music. The innumerable contrivances, therefore, by which the male is able to seize the female, may be briefly passed over. Besides the complex structures at the apex of the abdomen, which ought perhaps to be ranked as primary organs (2. These organs in the male often differ in closely-allied species, and afford excellent specific characters. But their importance, from a functional point of view, as Mr. R. MacLachlan has remarked to me, has probably been overrated. It has been suggested, that slight differences in these organs would suffice to prevent the intercrossing of well-marked varieties or incipient species, and would thus aid in their development. That this can hardly be the case, we may infer from the many recorded cases (see, for instance, Bronn, 'Geschichte der Natur,' B. ii. 1843, s. 164; and Westwood, 'Transact. Ent. Soc.' vol. iii. 1842, p. 195) of distinct species having been observed in union. Mr. MacLachlan informs me (vide 'Stett. Ent. Zeitung,' 1867, s. 155) that when several species of Phryganidae, which present strongly-pronounced differences of this kind, were confined together by Dr. Aug. Meyer, THEY COUPLED, and one pair produced fertile ova.), "it is astonishing," as Mr. B.D. Walsh (3. 'The Practical Entomologist,' Philadelphia, vol. ii. May 1867, p. 88.) has remarked, "how many different organs are worked in by nature for the seemingly insignificant object of enabling the male to grasp the female firmly." The mandibles or jaws are sometimes used for this purpose; thus the male Corydalis cornutus (a neuropterous insect in some degree allied to the Dragon flies, etc.) has immense curved jaws, many times longer than those of the female; and they are smooth instead of being toothed, so that he is thus enabled to seize her without injury. (4. Mr. Walsh, ibid. p. 107.) One of the stag-beetles of North America (Lucanus elaphus) uses his jaws, which are much larger than those of the female, for the same purpose, but probably likewise for fighting. In one of the sand- wasps (Ammophila) the jaws in the two sexes are closely alike, but are used for widely different purposes: the males, as Professor Westwood observes, "are exceedingly ardent, seizing their partners round the neck with their sickle-shaped jaws" (5. 'Modern Classification of Insects,' vol. ii. 1840, pp. 205, 206. Mr. Walsh, who called my attention to the double use of the jaws, says that he has repeatedly observed this fact.); whilst the females use these organs for burrowing in sand-banks and making their nests.