[Fig. 30. Xiphophorus Hellerii. Upper figure, male; lower figure, female.]
In the very distinct family of the Cyprinodontidae—inhabitants of the fresh waters of foreign lands—the sexes sometimes differ much in various characters. In the male of the Mollienesia petenensis (16. With respect to this and the following species I am indebted to Dr. Gunther for information: see also his paper on the ‘Fishes of Central America,’ in ‘Transact. Zoological Soc.’ vol. vi. 1868, p. 485.), the dorsal fin is greatly developed and is marked with a row of large, round, ocellated, bright-coloured spots; whilst the same fin in the female is smaller, of a different shape, and marked only with irregularly curved brown spots. In the male the basal margin of the anal fin is also a little produced and dark coloured. In the male of an allied form, the Xiphophorus Hellerii (Fig. 30), the inferior margin of the caudal fin is developed into a long filament, which, as I hear from Dr. Gunther, is striped with bright colours. This filament does not contain any muscles, and apparently cannot be of any direct use to the fish. As in the case of the Callionymus, the males whilst young resemble the adult females in colour and structure. Sexual differences such as these may be strictly compared with those which are so frequent with gallinaceous birds. (17. Dr. Gunther makes this remark; ‘Catalogue of Fishes in the British Museum,’ vol. iii. 1861, p. 141.)
[Fig.31. Plecostomus barbatus. Upper figure, head of male; lower figure, female.]
In a siluroid fish, inhabiting the fresh waters of South America, the Plecostomus barbatus (18. See Dr. Gunther on this genus, in ‘Proceedings of the Zoological Society,’ 1868, p. 232.) (Fig. 31), the male has its mouth and inter-operculum fringed with a beard of stiff hairs, of which the female shows hardly a trace. These hairs are of the nature of scales. In another species of the same genus, soft flexible tentacles project from the front part of the head of the male, which are absent in the female. These tentacles are prolongations of the true skin, and therefore are not homologous with the stiff hairs of the former species; but it can hardly be doubted that both serve the same purpose. What this purpose may be, it is difficult to conjecture; ornament does not here seem probable, but we can hardly suppose that stiff hairs and flexible filaments can be useful in any ordinary way to the males alone. In that strange monster, the Chimaera monstrosa, the male has a hook-shaped bone on the top of the head, directed forwards, with its end rounded and covered with sharp spines; in the female “this crown is altogether absent,” but what its use may be to the male is utterly unknown. (19. F. Buckland, in ‘Land and Water,’ July 1868, p. 377, with a figure. Many other cases could be added of structures peculiar to the male, of which the uses are not known.)
The structures as yet referred to are permanent in the male after he has arrived at maturity; but with some Blennies, and in another allied genus (20. Dr. Gunther, ‘Catalogue of Fishes,’ vol. iii. pp. 221 and 240.), a crest is developed on the head of the male only during the breeding-season, and the body at the same time becomes more brightly-coloured. There can be little doubt that this crest serves as a temporary sexual ornament, for the female does not exhibit a trace of it. In other species of the same genus both sexes possess a crest, and in at least one species neither sex is thus provided. In many of the Chromidae, for instance in Geophagus and especially in Cichla, the males, as I hear from Professor Agassiz (21. See also ‘A Journey in Brazil,’ by Prof. and Mrs. Agassiz, 1868, p. 220.), have a conspicuous protuberance on the forehead, which is wholly wanting in the females and in the young males. Professor Agassiz adds, “I have often observed these fishes at the time of spawning when the protuberance is largest, and at other seasons when it is totally wanting, and the two sexes shew no difference whatever in the outline of the profile of the head. I never could ascertain that it subserves any special function, and the Indians on the Amazon know nothing about its use.” These protuberances resemble, in their periodical appearance, the fleshy carbuncles on the heads of certain birds; but whether they serve as ornaments must remain at present doubtful.
I hear from Professor Agassiz and Dr. Gunther, that the males of those fishes, which differ permanently in colour from the females, often become more brilliant during the breeding-season. This is likewise the case with a multitude of fishes, the sexes of which are identical in colour at all other seasons of the year. The tench, roach, and perch may be given as instances. The male salmon at this season is “marked on the cheeks with orange-coloured stripes, which give it the appearance of a Labrus, and the body partakes of a golden orange tinge. The females are dark in colour, and are commonly called black-fish.” (22. Yarrell, ‘History of British Fishes,’ vol. ii. 1836, pp. 10, 12, 35.) An analogous and even greater change takes place with the Salmo eriox or bull trout; the males of the char (S. umbla) are likewise at this season rather brighter in colour than the females. (23. W. Thompson, in ‘Annals and Magazine of Natural History,’ vol. vi. 1841, p. 440.) The colours of the pike (Esox reticulatus) of the United States, especially of the male, become, during the breeding-season, exceedingly intense, brilliant, and iridescent. (24. ‘The American Agriculturalist,’ 1868, p. 100.) Another striking instance out of many is afforded by the male stickleback (Gasterosteus leiurus), which is described by Mr. Warington (25. ‘Annals and Mag. of Nat. Hist.’ Oct. 1852.), as being then “beautiful beyond description.” The back and eyes of the female are simply brown, and the belly white. The eyes of the male, on the other hand, are “of the most splendid green, having a metallic lustre like the green feathers of some humming-birds. The throat and belly are of a bright crimson, the back of an ashy-green, and the whole fish appears as though it were somewhat translucent and glowed with an internal incandescence.” After the breeding season these colours all change, the throat and belly become of a paler red, the back more green, and the glowing tints subside.
With respect to the courtship of fishes, other cases have been observed since the first edition of this book appeared, besides that already given of the stickleback. Mr. W.S. Kent says that the male of the Labrus mixtus, which, as we have seen, differs in colour from the female, makes “a deep hollow in the sand of the tank, and then endeavours in the most persuasive manner to induce a female of the same species to share it with him, swimming backwards and forwards between her and the completed nest, and plainly exhibiting the greatest anxiety for her to follow.” The males of Cantharus lineatus become, during the breeding-season, of deep leaden-black; they then retire from the shoal, and excavate a hollow as a nest. “Each male now mounts vigilant guard over his respective hollow, and vigorously attacks and drives away any other fish of the same sex. Towards his companions of the opposite sex his conduct is far different; many of the latter are now distended with spawn, and these he endeavours by all the means in his power to lure singly to his prepared hollow, and there to deposit the myriad ova with which they are laden, which he then protects and guards with the greatest care.” (26. ‘Nature,’ May 1873, p. 25.)
A more striking case of courtship, as well as of display, by the males of a Chinese Macropus has been given by M. Carbonnier, who carefully observed these fishes under confinement. (27. ‘Bulletin de la Societé d’Acclimat.’ Paris, July 1869, and Jan. 1870.) The males are most beautifully coloured, more so than the females. During the breeding-season they contend for the possession of the females; and, in the act of courtship, expand their fins, which are spotted and ornamented with brightly coloured rays, in the same manner, according to M. Carbonnier, as the peacock. They then also bound about the females with much vivacity, and appear by “l’étalage de leurs vives couleurs chercher a attirer l’attention des femelles, lesquelles ne paraissaient indifférentes a ce manège, elles nageaient avec une molle lenteur vers les males et semblaient se complaire dans leur voisinage.” After the male has won his bride, he makes a little disc of froth by blowing air and mucus out of his mouth. He then collects the fertilised ova, dropped by the female, in his mouth; and this caused M. Carbonnier much alarm, as he thought that they were going to be devoured. But the male soon deposits them in the disc of froth, afterwards guarding them, repairing the froth, and taking care of the young when hatched. I mention these particulars because, as we shall presently see, there are fishes, the males of which hatch their eggs in their mouths; and those who do not believe in the principle of gradual evolution might ask how could such a habit have originated; but the difficulty is much diminished when we know that there are fishes which thus collect and carry the eggs; for if delayed by any cause in depositing them, the habit of hatching them in their mouths might have been acquired.
To return to our more immediate subject. The case stands thus: female fishes, as far as I can learn, never willingly spawn except in the presence of the males; and the males never fertilise the ova except in the presence of the females. The males fight for the possession of the females. In many species, the males whilst young resemble the females in colour; but when adult become much more brilliant, and retain their colours throughout life. In other species the males become brighter than the females and otherwise more highly ornamented, only during the season of love. The males sedulously court the females, and in one case, as we have seen, take pains in displaying their beauty before them. Can it be believed that they would thus act to no purpose during their courtship? And this would be the case, unless the females exert some choice and select those males which please or excite them most. If the female exerts such choice, all the above facts on the ornamentation of the males become at once intelligible by the aid of sexual selection.
We have next to inquire whether this view of the bright colours of certain male fishes having been acquired through sexual selection can, through the law of the equal transmission of characters to both sexes, be extended to those groups in which the males and females are brilliant in the same, or nearly the same degree and manner. In such a genus as Labrus, which includes some of the most splendid fishes in the world—for instance, the Peacock Labrus (L. pavo), described (28. Bory Saint Vincent, in ‘Dict. Class. d’Hist. Nat.’ tom. ix. 1826, p. 151.), with pardonable exaggeration, as formed of polished scales of gold, encrusting lapis-lazuli, rubies, sapphires, emeralds, and amethysts—we may, with much probability, accept this belief; for we have seen that the sexes in at least one species of the genus differ greatly in colour. With some fishes, as with many of the lowest animals, splendid colours may be the direct result of the nature of their tissues and of the surrounding conditions, without the aid of selection of any kind. The gold-fish (Cyprinus auratus), judging from the analogy of the golden variety of the common carp, is perhaps a case in point, as it may owe its splendid colours to a single abrupt variation, due to the conditions to which this fish has been subjected under confinement. It is, however, more probable that these colours have been intensified through artificial selection, as this species has been carefully bred in China from a remote period. (29. Owing to some remarks on this subject, made in my work ‘On the Variation of Animals under Domestication,’ Mr. W.F. Mayers (‘Chinese Notes and Queries,’ Aug. 1868, p. 123) has searched the ancient Chinese encyclopedias. He finds that gold-fish were first reared in confinement during the Sung Dynasty, which commenced A.D. 960. In the year 1129 these fishes abounded. In another place it is said that since the year 1548 there has been “produced at Hangchow a variety called the fire-fish, from its intensely red colour. It is universally admired, and there is not a household where it is not cultivated, IN RIVALRY AS TO ITS COLOUR, and as a source of profit.”) Under natural conditions it does not seem probable that beings so highly organised as fishes, and which live under such complex relations, should become brilliantly coloured without suffering some evil or receiving some benefit from so great a change, and consequently without the intervention of natural selection.