1 1 the DESCENT OF MAN SELECTION IN RELATION TO SEX. BY CHARLES DARWIN, M.A., F.R.S., Etc. WITH ILLUSTRATIONS. IN TWO VOLUMES. — Vol. II. NEW YOE K : D. APPLET ON ANT) COMPANY, 549 & 551 B ROADWAY. 1871. library of Jhe ^Wh Collene of Hawaii H J* 'gr/ m CONTENTS. PART n. SEXUAL SELECTION— continued. CHAPTER XII. SECONDARY SEXUAL CHARACTERS OP FISHES, AMPHIBIANS, AND REPTILES. Fishes : Courtship and Battles of the Males. — Larger Size of. the Females. — Males, Bright Colors and Ornamental Appendages; other Strange Characters. — Colors and Appendages acquired by the Males during the Breeding-season alone. — Fishes with both Sexes brilliantly colored. — Protective Colors. — The less Conspicuous Colors of the Female cannot bo accounted for on the Principle of Protection. — Male Fishes building Nests, and taking Charge of the Ova and Young. Amphibians : Differ- ences in Structure and Color between the Sexes. — Vocal Organs. Rep- tiles : Chelonians. — Crocodiles. — Snakes, Colors in some Cases pro- tective.— Lizards, Battles of. — Ornamental Appendages. — Strange Dif- ferences in Structure between the Sexes. — Colors. — Sexual Differences almost as great ^as with Birds . . . . . p. 1 CIIAPTEK XIII. SECONDARY SEXUAL CHARACTERS OP BIRDS. Sexual Differences. — Law of Battle— Special "Weapons. — Vocal Organs.— Instrumental Music. — Love-Antics and Dances.— Decorations, lYrma- nont and Seasonal. — Double and Single Annual Moults. — Display of Ornaments by the Males ...... 30 CH APTEE XIV. niuns — continued. Choice exerted by the Female. — Length of Courtship. — Unpaired Birds. — Mental Qualities and Taste for the Beautiful.- Preference or Antip- athy shown by the Female foi Partloulai Males.- - Yai nihility of 20111 iv CONTEXTS. Birds.— Variations sometimes abrupt.— Laws of Variation.— Formation of Ocelli. — Gradations of Character. — Case of Peacock, Argus Pheas- ant, and Urosticte . . . * • • . p. 95 CHAPTER XV. bi uns — continued. Discussion why the Males alone of some Species, and both Seles of other Species, are brightly colored. — On Sexually-limited Inheritance, as ap- plied to Various Struetures and to Bri^htl v-celon .l Plumairo. — Nidifi- cation in Relation to Color. — Lobs of Nuptial Plumage during tho Winter 147 CHAPTER XVI. birds — concluded. . The Immature Plumage in Relation to the Character of tho Plumage in both Sexes when Adult. — Six Classes of Cases. — Sexual Differences between the Males of Closely-allied or Representative Species. — The Female assuming the Characters of tho Male. — Plumage of tho Young in Relation to the Summer and "Winter Plumage of tho Adults. — Ou the Increase of Beauty in tho Birds of tho World. — Protective Color- ing.— Conspicuously-colored Birds. — Novelty appreeiated. — Summary of the Four Cliapters on Birds ..... 175 CHAPTER XVU, SECONDARY SEXUAL CHARACTERS OF MAMMALS. The Law of Battle— Special Weapons, confined to tho Males. — Cause of Absence of Weapons in the Female. — Weapons common to both Sexes, yet primarily acquired by the Male. — Other Uses of such Weap- ons.— Their High Importance. — Greater Sixe of the Male. — Means of Defence.— On tho Preference shown by either Sex in the Pairing of Quadrupeds ....... 228 CHAPTER XVin. SECOND ART 8EXCAL CHARACTERS Of MAMMALS — COtUin Ufd. Voice. — Remarkable Sexual Peculiarities in Seals. — Odor. — Development of 'the Hair.— Color of the Hair and Skin.— Anomalous Case of the Female being more ornamented than the Male. — Color and Ornaments due to Sexual Selection. — Color acquired for tho Sake of Protection. — Color, though common to both Sexes, often due to Sexual Selection. — CONTENTS. v On the Disappearance of Spots and Stripes in Adult Quadrupeds. — On the Colors and Ornaments of the Quadrumana. — Summary p. 261 CHAPTEK XIX. SECONDARY SEXUAL CHARACTERS OP MAN. Differences "between Man and "Woman. — Causes of such Differences and of Certain Characters common to both Sexes. — Law of Battle. — Differ- ences in Mental Powers — and Voice. — On the Influence of Beauty in determining the Marriages of Mankind. — Attention paid by Savages to Ornaments. — Their Ideas of Beauty in Woman. — The Tendency to exaggerate each Natural Peculiarity . . . . 301 CHAPTER XX. SECONDARY SEXUAL CHARACTERS OF MAN — Continued- On the Effects of the Continued Selection of "Women according to a Different Standard of Beauty in each Race. — On the Causes which interfere with Sexual Selection in Civilized and Sayags Nations. — Conditions favorable to Sexual Selection during Primeval Times. — On the Manner of Action of Sexual Selection with Mankind.— On tho "Women in Savage Tribes having some Power to chooso their Hus- bands.— Absence of Hair on the Body, and Development of the Beard. — Color of the Skin.— Summary ..... 339 CHAPTER XXL GKNKHAL SUMMARY AND CONCLUSION. Main Conclusion that Mania descended from BOme Lower Form. — Man- ner of Development. — Genealogy of Man.— Intellectual and Moral Faculties. — Scxuul Selection. — Concluding Remarks . . 808 POSTSCBIP T. Vol. I., pp. 287-289. — I have fallen into a serious and unfortunate error, in relation to the sexual differences of animals, in attempting to explain what seemed to me a singular coincidence in the late period of life at which the necessary variations have arisen in many cases, and the late period at which sexual selection acts. The ex- planation given is wholly erroneous, as I have discovered by working out an illustration in figures. Moreover, the supposed coincidence of period is far from general, and is not remarkable ; for, as I have elsewhere attempted to show, variations arising early in life have often been accumulated' through sexual selection, being then com- monly transmitted to both sexes. On the other hand, variations arising late in life cannot fail to coincide ap- proximately in period with that of the process of sexual selection. SEXUAL SELECTION". CHAPTER XII. Secondary Sextjal Characters of Fishes, Amphibi- ans, and Reptiles. FisSes : Courtship and Battles of the Males. — Larger Size of the Females. — Males, Bright Colors and Ornamental Appendages ; other Strange Characters. — Colors and Appendages acquired by the Males during the Breeding-season alone. — Fishes with both Sexes brilliantly colored. — Protective Colors. — The less Conspicuous Cojors of the Female cannot be accounted for on the Principle of Protection. — Male Fishes building Nests, and taking Charge of the Ova and Young. Amphibians : Differ- ences in Structure and Color between the Sexes. — Vocal Organs. Rep- tiles : Chelonians. — Crocodiles. — Snakes, Colors in some Cases pro- tective.— Lizards, Battles of. — Ornamental Appendages. — Strange Dif- ferences in Structure between the Sexes. — Colors. — Sexual Ditlerenees almost as great as with Birds. We have now arrived at the great sub-kingdom bf the Vertebrata, and will commence with the lowest class, namely, Fishes. The males of Plagiostomous fishes (sharks, rays) and of Chinueroid lishes are provided with claspers which serve to retain the female, like the various Structures possessed by so many of the lower animals. Besides the claspers, the males of many rays have clus- ters of strong sharp spines on their heads, and several rows along "the upper outer surface of their pectoral fins." These are present in the males of somo species, which have the other parts of their bodies smooth. They 2 SEXUAL SELECTION. [Part II. are only temporarily developed during the breeding-sea- son ; and Dr. Gtlnther suspects that they are brought into action as prehensile organs by the doubling inward and downward of the two sides of the body. It is a remark- able fact that the females and not the males of some spe- cies, as of Raid clavata, have their backs studded with large hook-formed spines.1 Owing to the element which fishes inhabit, little is known about their courtship, and not much about their battles. The male stickleback ( Gasterosteus leiurus) has been described as " mad with delight " when the female comes out of her hiding-place and surveys the nest which he has made for her. " He darts round her in every direction, then to his accumulated materials for the nest, then back again in an instant ; and as she does not ad- vance he endeavors to push her with his snout, and then tries to pull her by the tail and side-spine to the nest." a The males are said to be polygamists ; 8 they are extraor- dinarily bold and pugnacious, while "the females are quite pacific." Their battles are at times desperate ; " for these puny combatants fasten tight on each other for sev- eral seconds, tumbling over and over again, until their strength appears completely exhausted." With the rough-tailed stickleback (G. trachurus) the males while fighting swim round and round each other, biting and en- deavoring to pierce each other with their raised lateral spines. The same writer adds : * " The bite of these little furies is very severe. They also use their lateral spines 1 Yarrell, 'Hist, of British Fishes,' vol. ii. 1836, pp. 417, 425, 436. Dr. Giinther informs me that the spines in R. clavata arc peculiar to the female. * See Mr. R. Warington's interesting articles in 1 Annals and Mag. of Nat. Hist.' Oct. 1852 and Nov. 1855. • Noel Humphreys, 'River Gardens,' 1857. 4 Loudon's * Mag. of Natural History,' vol. iii. 1830, p. 831. Chap. XII.J FISHES. 3 with such fatal effect, that I have seen one during a bat- tle absolutely rip his opponent quite open, so that he sank to the bottom and died." When a fish is conquered, "his gallant bearing forsakes him ; his gay colors fade away ; and he hides his disgrace among his peaceable compan- ions, but is for some time the constant object of his con- queror's persecution." The male salmon^s as pugnacious as the little stickle- back ; and so is the male trout, as I hear from Dr. Gtin- ther. Mr. Shaw saw a violent contest between two male salmons which lasted the whole day ; and Mr. R. Buist, Superintendent of Fisheries, informs me that he has often watched from the bridge at Perth the males driving away their rivals while the females were spawning. The males " are constantly fighting and tearing each other on the spawning-beds, and many so injure each other as to cause the death of numbers, many being seen swimming near the banks of the river in a state of exhaustion, and apparently in a dying state." 5 The keeper of the Stor- montfield breeding-ponds visited, as Mr. Buist informs me, in June, 1868, the northern Tyne, and found about 300 dead salmon, all of which with one exception were males ; and he was convinced that they had lost their lives by fighting. The most curious point about the male salmon is that during the breeding-season, besides a slight change in color, " the lower jaw elongates, and a cartilaginous pro- jection turns upward from the point, which, when the jaws arc closed, occupies a deep cavity between " the in- termaxillary bones of the upper jaw." 0 (Figs. 20 and 8 'The Field,' Juno 29, 1807. For Mr. Shaw's statement, Bee 1 l-Min- burgh Review,' l k i :j. Another experienced observer (Sorope'a 1 Days <>t' Salmon Fishing,' p. CO) remarks that the male worn1.!, ,1 lie <>"M, keep, like the Btag, all other males away, • Yarrell, « History oflJritisli Fishes,' vol. ii. ls:;r,, p. 10, 4 SEXUAL SELECTION. [Pakt II. 27.) In our salmon this change of structure lasts only during the breeding-season ; but in the Salmo lycaodon of Northwestern America the change, as Mr J. K. Lord 7 Fig. 26.— Head of male of common salmon (Salmo salar) during the breeding- season. [This drawing, as well aa all the others in the present chapter, have been ex- ecuted by the well-known artist, Mr. Q. Ford, under the kind superintendence of Dr. Gunther, from specimens in the British Museum]. 7 'The Naturalist in Vancouver's Island,' vol. i. 1866, p. 54. Chap. XII.] FISHES. 5 believes, is permanent and best marked in the older males which have previously ascended the rivers. In these old males the jaws become developed into immense hook-like Fig. 27.— Head of female salmon. projections, and the teeth grow into regular fangs, often more than half an inch in length. With the European salmon, according to Mr. Lloyd/ the temporary hook-like Structure serves to Strengthen and protect the jaw s, when 8 4 Scandinavian Adventures,' vol. i. 185-1, |>p. lol, 101. 6 SEXUAL SELECTION. [Part II. one inali' diaries another with wonderful violence ; but the greatly developed teeth of the male American salmon may be compared with the tusks of many male mf*"TO«rtp^ ami they indicate an offensive rather than a protective purpose. The salmon is not the only fish in which the teeth differ in the two sexes. This is the case witli many rajs. In the thornback (Raia clavata) the adult male has sharp, pointed teeth, directed backward, while those of the fe- male are broad aiyl flat, forming a pavement; so that these teeth differ in the two sexes of the same species more than is usual in distinct genera of the same family. The teeth of the male become sharp only when he is adult : while young they are broad and flat like those of the female. As so frequently occurs with secondary sex- ual characters, both sexes of some species of rays, for in- stance II* batis, possess, when adult, sharp, pointed teeth ; and here a character, proper to and primarily gained by the male, appears to have been transmitted to the off- spring of both sexes. The teeth are likewise pointed in both sexes of It. macnkUa, but only when completely adult ; the males acquiring them at an earlier age than the females. We shall hereafter meet with analogous cases with certain birds, in which the male acquires the plu- mage common to both adult sexes, at a somewhat earlier age than the female. With other species of rays the males even when old never possess sharp teeth, and con- sequently both sexes when adult are provided with broad, flat teeth, like those of the young, and of the mature fe- males of the above-mentioned species.* As the rays are bold, strong, and voracious fishes, we may suspect that the males require their sharp teeth for fighting with their rivals; but as they possess many parts modified and • See Yarrcll's account of the Rays in his • Hist of British Fishes,' ▼oL U. 1836, p. 416, with an excellent figure, and pp. 422, 432. Chap. XII.] FISHES. 7 adapted for the prehension of the female, it is possible that their teeth may be used for this purpose. In regard to size, M. Carbonnier 10 maintains that with almost all fishes the female is larger than the male ; and Dr. Gunther does not know of a single instance in which the male is actually larger than the female. With some Cyprinodonts the male is not even half as large as the female. As with many kinds of fishes, the males habitu- ally fight together ; it is surprising that th^y have not generally become through the effects of sexual selection larger and stronger than the females. The males suffer from their small size, for according to M. Carbonnier they are liable to be devoured by the females of their own spe- cies when carnivorous, and no doubt by other species. Increased size must be in some manner of more impor- tance to the females, than strength and size are to the males for fighting with other males ; and this perhaps is to allow of the production of a vast number of ova. In many species the male alone is ornamented with bright colors; or these arc much brighter in the male than in the female. The male, also, is sometimes provided with appendages which appear to be of no more use to him for the ordinary purposes of life than are the tail-feat hers to the peacock. I am indebted for most of the following facts to the great kindness of Dr. (iunther. There is rea- son to suspect that many tropical lishes differ sexually in color and structure; and there are some striking oases with our British fishes. The male Callionymus Jyra has been palled "the gemmeous dragoiiet " from its brilliant, gem-like colors." Whenfreshly taken from tin' sea the body is yellow of various shades, striped and Bpotted with vivid blue on the head; the dorsal lins are pale brown with dark longitudinal hands; the ventral, caudal, and anal lins being bluish-black. The female, or sordid drag* • 10 As quoted in 'The luinurr,' |Nt',M, \> ".i'.'.i. 8 SEXUAL SELECTION. [Part II. onet, was considered by Linnaeus and by many subse- quent naturalists as a distinct species; it is of a dingy Fig. 28.— Callionymns lyra. Upper figure, male ; lower figure, female. reddish-brown, with the dorsal fin brown and the other fins white. The* sexes differ also in the proportional size of the head and mouth, and in the position of the eyes ; 11 but the most striking difference is the extraordinary elon- gation in the male (Fig. 28) of the dorsal fin. The young males resemble, in structure and color, the adult females. Throughout the genus Callionymus,13 the male is gener- 11 I have drawn up this description from YarrelTs 'British Fishes,' vol. i. 1836, pp. 261, 266. 18 1 Catalogue of Acanth. Fishes in the British Museum,' by Dr. Giinther, 1861, pp. 138-151. Chap. XII.] FISHES. 9 ally much more brightly spotted than the female, and in several species, not only the dorsal, but the anal fin of the male, is much elongated. The male of the Cottus scorpius, or sea-scorpion, is more slender antl smaller than the female. There is also a great difference in color between them. It is difficult, as Mr. Lloyd 13 remarks, " for any one, who has not seen this fish during the spawning-season, when its hues are brightest, to conceive the admixture of brilliant colors with which it, in other respects so ill-favored, is at that time adorned." Both sexes of the Ldbrus rnixtus, al- though very different in color, are beautiful ; the male being orange with bright-blue stripes, and the female bright-red with some black spots on the back. In the very distinct family of the Cyprinodontidge — inhabitants of the fresh waters of foreign lands — the sexes sometimes differ much in various characters. In the male v of the Mollienesia petenensis,1* the dorsal fin is greatly developed, and is marked with a row of large, round, ocel- lated, bright-colored spots ; while the same fin in the fe- male 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- colored. In the male of an allied form, the Xiphophorus Hellerii (Fig. 29), the inferior margin of the anal fin is developed into a long filament, which is striped, as I hear from Dr. Gllnther,with bright colors. This filament does not contain any muscles, and apparently cannot be of any direct use to the fish. As in the case of the CMKonymus, the males while young resemble in color and structure (he adult females. Sexual differences such as these may be 18 1 Game IVirds of Sweden,' etc., 1807, p. 400. 14 With respect to this and (lie following species I am indebted to Dr. Giinthcr for information; sec also his paper on the Fishes of Central America, in 'Transact. Zoolog. Soc.' vol. vi. lNt;s( p. I S5. 10 SEXUAL SELECTION [Part EC strictly compared with those which are so frequent with gallinaceous birds.16 Pio. 29.— Xiptaophorus Hellerii. Upper figure, male; lower figure, female. In a siluroid fish, inhabiting the fresh waters of South America, namely the Pltcostomus barbatus " (Fig. 30), the male has its mouth and interoperculum 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 15 Dr. Gunther makes this remark : ' Catalogue of Fishes in the Brit- ish Museum,' vol. iii. 1861, p. 141. M See Dr. Gunther on this genus, in 'Proc. Zoolog. Soc.' 1868, p. 232. Chap. XII.] FISHES. . H 12 SEXUAL SELECTION. [Part II. that stiff hairs and flexible filaments can be useful in any- ordinary way to the males alone. The Monacanthus scopas, which was shown to me in the British Museum by Dr. Giinther, presents a nearly analogous case. The male has a cluster of stiff, straight spines, like those of a comb, on the sides of the tail ; and these in a specimen six inches long were nearly an inch and a half in length ; the female has on the same place a cluster of bristles, which may be compared with those of a tooth-brush. In another species, the M. peronii, the male has a brush like that possessed by the female of the last species, while the sides of the tail in the female are smooth. In some other species the same part of the tail can be perceived to be a little rough- ened in the male and perfectly smooth in the female ; and lastly, in others, both sexes have smooth sides. In that strange monster, the Chimcera monstrosa, the male has a hook-shaped bone on the top of the head, directed for- ward, with its rounded end covered with sharp spines ; in the female " this crown is altogether absent," but what its use may be is utterly unknown.17 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 18 a crest is developed on the head of the male only during the breeding-season, and their bodies at the same time become more brightly- colored. 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 this case and in that of the Monacanthus, we have good instances to how great an extent the sexual characters of closely-allied forms may 17 F. Buckland, in 'Land and Water/ July, 1868, p. 377, with a figure. 18 Dr. Giinther, 'Catalogue of Fishes,' vol. hi. pp. 221, 240. Chap. XII.J FISHES. 13 differ. In many of the Chromidae, for instance, in Geo- phagus and especially in Cichla, the males, as I hear from Prof. Agassiz,19 have a conspicuous protuberance on the forehead, which is wholly wanting in the females and in the young males. Prof. 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 show 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 Amazons know nothing about its use." These protuberances in their periodical appearance resem- ble the fleshy caruncles on the heads of certain birds ; but whether they serve as ornaments must remain at present doubtful. The males of those fishes, which differ permanently in color from the females, often become more brilliant, as I hear from Prof. Agassiz and Dr. Gunther, during the breeding-season. This is likewise the case with a multi- tude of fishes, the sexes of which at all other seasons of the year are identical in color. The tench, roach, and perch, may be given as instances. The male salmon at this season is " marked on the cheeks witli orange-colored stripes, which gave it the appearance of a Labrus, and the body partakes of a golden-orange tinge. The females are dark in color, and are commonly called blacktish." 30 An analogous and even greater change takes place with the tfalmo eriox, or bull-trout ; the males of the char (S. umbla) arc likewise at this season rather brighter in color than the females.31 The colors of the pike (Esox re- 19 Sec also 'A Journey in Brazil,' by Prof, and Mrs. Agassi/, IST.S, p. 220. 90 Yarrcll, 'British Fishes,' vol. ii. is:!)',, pp. 10, 12, :tr.. " W. Thompson, in ' Annals and Ma^. of Nat. History,' vol. \ i. ISM, p. 440. 14 SEXUAL SELECTION. [Part II. tieulatus), of the United States, especially of the male, become, during the breeding-season, exceedingly intense, brilliant, and iridescent." Another striking instance out of many is afforded by the male stickleback ( Gasterosteus leiurus), which is described by Mr. Warington," 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 trans- lucent, and glowed with an internal incandescence." After the breeding-season these colors all change, the throat and belly become of a paler red, the back more green, and the glowing tints subside. That with fishes there exists some close relation be- tween their colors and their sexual functions we can clearly see — firstly, from the adult males of certain species being differently colored from the females, and often much more brilliantly; secondly, from the same males, while im- mature, resembling the mature females ; and, lastly, from the males, even of those species which at all other times of the year are identical in color with the females, often acquiring brilliant tints during the spawning-season. We know that the males are ardent in their courtship, and sometimes fight desperately together. If we may assume that the females have the power of exerting a choice and of selecting the more highly-ornamented males, all the above facts become intelligible through the principle of sexual selection. On the other hand, if the females ha- bitually deposited and left their ova to be fertilized by the first male whicli chanced to approach, this fact would be w 'The American Agriculturist,' 1868, p. 100. n 1 Annals and Mag.' of Nat. Hist.' Oct. 1852. Chap XII.] FISHES. 15 • fatal to the efficiency of sexual selection ; for there could be no choice of a partner. But, as far as is known, the female never willingly spawns except in the close presence of a male, and the male never fertilizes the ova except in the close presence of a female. It is obviously difficult to obtain direct evidence with respect to female fishes select- ing their partners. An excellent observe^24 who carefully watched the spawning of minnows (Cyprinus phoxinus)9 remarks that owing to the males, which were ten times as numerous as the females, crowding closely round them, he could " speak only doubtfully on their operations. When a female came among a number of males they immediately pursued her ; if she was not ready for shedding her spawn, she made a precipitate retreat ; but if she was ready, she came boldly in among them, and was immediately pressed closely by a male on each side ; and when they had been in that situation a short time, were superseded by other two, who wedged themselves in between them and the female, who appeared to treat all her lovers with the same kindness." Notwithstanding this last statement, I can- not, from the several previous considerations, give up the belief that the males which are the most attractive to the females, from their brighter colors or other ornaments, are commonly preferred by them ; and that the males have thus been rendered more beautiful in the course of ages. We have next to inquire whether tliis view can be ex- tended, through the law of the equal transmission of char- acters to both sexes, 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 (£. pavo), described," with M Loudon's 'Mag. of Nat. Hist.' vol. v. 1832, p. 081. ••Bory . 84 Prof. Wyman, in ' Proo. Boston Sue of Nat. Hist.' Sept. L5, is;>7. Also, W. Turner, in 'Journal of Anatomy and Phys.' Nov. l, L866, p. 78. Dr. Gttnther has likewise described other eases. 20 SEXUAL SELECTION. [Part IL Prof. Agassiz, " not only are generally brighter than the females, but the difterence is greater at the spawning-sea- son than at any other time." The species of Geophagus act in the same manner ; and in this genus, a conspicuous protuberance becomes developed on the forehead of the males during the breeding-season. With the various spe- cies of Chromids, as Prof. Agassiz likewise informs me, sexual differences in color may be observed, " whether they lay their eggs in the water among aquatic plants, or deposit them in holes, leaving them to come out without further care, or build shallow nests in the river-mud, over which they sit, as our Promotis does. It ought also to be observed that these sitters are among the brightest spe- cies in their respective families ; for instance, Hygrogonus is bright green, with large black ocelli, encircled with the most brilliant red." Whether with all the species of Chromids it is the male alone which sits on the eggs is not known. It is, however, manifest that the fact of the eggs being protected or unprotected, has had little or no influence on the differences in color between the sexes. It is further manifestpn all the cases in which the males take exclusive charge of the nests and young, that the destruction of the brighter-colored males would be far more influential on the character of the race, than the de- struction of the brighter-colored females ; for the death of the male during the period of incubation or nursing would entail the death of the young, so that these could not inherit his peculiarities ; yet, in many of these very cases tlfe males are more conspicuously colored than the females. In most of the Lophobranchii (Pipe-fish, Hippocampi, etc.) the males have cither marsupial sacks or hemispheri- cal depressions on the abdomen, in which the ova laid by the female are hatched. The males also show great .at- tachment to their young." The sexes do not commonly 85 Yarrell, 4 Ilist. of British Fishes,' voL ii. 1836, pp. 829, 888. Chap. XII.] FISHES. 21 differ much in color ; but Dr. Gunther believes that the male Hippocampi are rather brighter than the females. The genus Solenostoma, however, offers a very curious ex- ceptional case,36 for the female is much more vividly col- ored and spotted than the male, and she alone has a mar- supial sack and hatches the eggs ; so that the female of Solenostoma differs from all the other Lophobranchii in this latter respect, and from almost all other fishes, in being more brightly colored than the male. It is improb- able that this remarkable double inversion of character in the female should be an accidental coincidence. As the males of several fishes which take exclusive charge of the eggs and young are more brightly colored than the fe- males, and as here the female Solenostoma takes the same charge and is brighter than the male, it might be argued that the conspicuous colors of the sex which is the most important of the two for the welfare of the offspring must serve, in some mauner, as a protection. But from the multitude of fishes, the males of which are either perma- nently or periodically brighter than the* females, but whose life is not at all more important than that of the female for the welfare of the species, this view can hardly be maintained. When we treat of birds we shall meet with analogous cases in which there has been a Complete inversion of the usual attributes of the two sexes, and we shall then give what appears to be the probable explana- tion, namely, that the males have selected the more at- tractive females, instead of the latter having selected, in accordance with the usual rule throughout the animal kingdom, the more attractive males. On the whole, we may conclude that, with most fishes, in which the sexes differ in color or in other ornamental M Dr. Giinthcr, sineo publishing un account of this species in ' The Fishes of Zanzibar,' by Colonel Play fair, lKCC, p. l.'tT, has rccxamincil the specimens, and bus given me the above information. 22 SEXUAL SELECTION. [Part II. characters, the males originally varied, with their varia- tions transmitted to the same Sex, and accumulated through sexual selection hy attracting or exciting the fe- males. In many cases, however, such characters have been transferred, either partially or completely, to the females. In other cases, again, both sexes have been col- ored alike for the sake of protection ; but in no instance does it appear that the female alone has had her colors or other characters specially modified for this purpose. The last point which need be noticed is that in many parts of the world fishes are known to make peculiar noises, which are described in some cases as being musi- cal. Very little has been ascertained with respect to the means by which such sounds are produced, and even less about their purpose. The drumming of the Umbrinas in the European seas is said to be audible from a depth of twenty fathoms. The fishermen of Rochelle assert " that the males alone make the noise during the spawning-time ; and that it is possible, by imitating it, to take them with- out bait." '7 'If this statement is trustworthy, we have BO instance in this, the lowest class of the Vertebrata, of what we shall find prevailing throughout the other verte- brate classes, and which prevails, as we have already seen, with insects and spiders ; namely, that vocal and instru- mental sounds so commonly serve as a love-call or as a love-charm, that the power of producing them was proba- bly first developed in connection with the propagation of the species. Amphibians. Urodela. — First for the tailed amphibians. The sexes of salamanders or newts often differ much both in color and structure. In some species prehensile claws are de- veloped on the fore-legs of the males during the breeding- * The Rev. C. Kingsley, in 4 Nature,' May, 1870, p. 40. Chap. XII.] AMPHIBIANS. 23 season ; and at this season in the male Triton palmipes the hind-feet are provided with a swimming web, which is almost completely absorbed during the winter; so that their feet then resemble those of the female.38 This struct- Fig. 31.— Triton cristatns (half natural size, from Bell's 'British Reptiles'). Upper figure, male during the hreeding-season ; lower figure, female. ure no doubt aids the male in his eager search and pursuit of the female. With our common newts (Triton puno tatus and cristatus) a deep, much indented crest is devel- oped along the back and tail of the male during the breed- ing-season, being absorbed during the winter. It is not furnished, as Mr. St. George Mivart informs me, with muscles, and therefore cannot be used for locomotion. As during the season of courtship it becomes edged with bright colors, it serves, there can hardly be a doubt, as :i masculine ornament. In many species the body promts strongly-contrasted though lurid tints; and these become more vivid during the breeding-season. The male, lor instance, of our common little newt (Triton />inift. For the Indian (alotes, see ' Reptiles of Uritish India,' h\ Dr. tliinther, p. 14:;. 86 SEXUAL SELECTION. [Pakt II. CHAPTER XIII. Secondary Sexual Characters of Birds. Sexual Differences. — Law of Battle. — Special Weapons. — Vocal Organs.— Instrumental Music. — Love-Antics and Dances.— Decorations, Perma-* nent and Seasonal. — Double and Single Annual Moults. — Display of Ornaments by the Males. Secondary sexual characters are more diversified and conspicuous in birds, though not perhaps entailing more important changes of structure, than in any other class of animals. I shall, therefore, treat the subject at consider- able length. Male birds sometimes, though rarely, pos- sess special weapons for fighting with each other. They charm the females by vocal or instrumental music of the most varied kinds. They are ornamented by all sorts of combs, wattles, protuberances, horns, air-distended sacs, topknots, naked shafts, plumes and lengthened feathers gracefully springing from all parts of the body. The beak and naked skin about the head and the feathers are often gorgeously colored. The males sometimes pay their court by dancing, or by fantastic antics performed either on the ground or in the air. In one instance, at least, the male emits a musky odor which we may suppose serves to charm or excite the female ; for that excellent observ- er, Mr. Ramsay,1 says of the Australian musk-duck (Bizv- ura lobata) that " the smell which the male emits during the summer months is confined to that sex, and in some individuals is retained throughout the year ; I have never, even in the breeding-season, shot a female which had any 1 'Ibis,' vol. iii. (new aeries) 1867, p. 414. Chap. XIII.] BIRDS. 37 smell of musk." So powerful is this odor during the pair- ing-season, that it can be detected long before the bird can be seen.2 On the whole, birds appear to be the most aesthetic of all animals, excepting of course man, and they have nearly the same taste for the beautiful as we have. This is shown by our enjoyment of the singing of birds, and by our women, both civilized and savage, decking their heads with borrowed plumes, and using gems which are hardly more brilliantly colored than the naked skin and wattles of certain birds. Before treating of the characters with which we are here more particularly concerned, I may just allude to certain differences between the sexes which apparently depend on differences in their habits of life ; for such cases, though common in the lower, are rare in the higher classes. Two humming-birds belonging to the genus Eustephanus, which inhabit the island of Juan Fernandez, were long thought to be specifically distinct, but are now known, as Mr. Gould informs me, to be the sexes of the same species, and they differ slightly in the form of the beak. In another genus of humming-birds (Grypus), the beak of the male is serrated along the margin and hooked at the extremity, thus differing much from that of the female. In the curious Neomorpha of New Zealand, there is a still wider difference in the form of the beak ; and Mr. Gould has been informed that the male with his "straight and stout beak" tears off the bark of trees, in order that the female may feed on the uncovered larva- with her weaker and more curved beak. Something of the same kind may he observed with our goldfinch (Car- duclix elegans), for I am assured by Mr. .1. .Tenner Weir that the bird-catohers can distinguish the males bythefar slightly longer beaks. The Hocks of males, :is an old and trustworthy bird-catcher asserted, are commonly found * (JouM, ' llaml-l>ool< to the llii.lt ill' Australia,' lst.:., vol. li. |». .is:; 38 SEXUAL SELECTION: BIRDS. [Part II. feeding on the seeds of the teazle (Dipsacus) which they can reach with their elongated beaks, while the females more commonly feed on the seeds of the betony or Scrophularia. With a slight difference of this nature as a foundation, we can see how the beaks of the two sexes might be made to differ greatly through natural selection. In all these cases, however, especially in that of the quar- relsome humming-birds, it is possible that the differences in the beaks may have been first acquired by the males in relation to their battles, and afterward led to slightly changed habits of life. Law of Battle. — Almost all male birds are extremely pugnacious, using their beaks, wings, and legs, for fighting together. We see this every spring with our robins and sparrows. The smallest of all birds, namely, the hum- ming-bird, is one of the most quarrelsome. Mr. Gosse * describes a battle, in which a pair of humming-birds seized hold of each other's beaks, and whirled round and round, till they almost fell to the ground ; and M. Montes de Oca, in speaking of another genus, says that two males rarely meet without a fierce aerial encounter : when kept in cages "their fighting has mostly ended in the splitting of the tongue of one of the two, which then surely dies from being unable to feed." * With Waders, the males of the common water-hen (Gallinula chloropus) "when pairing, fight violently for the females : they stand nearly upright in the water and strike with their feet." Two were seen to be thus engaged- for halt* an hour, until one got hold of the head of the other, which would have been killed had not the observer interfered ; the female all the time looking on as a quiet spectator.* The males of an allied * Quoted by Mr. Gould, 4 Introduction to the Trochilidae,' 1861, p. 29. * Gould, ibid. p. 62. » W. Thompson, 4 Nat Hist, of Ireland : Birds,' vol. ii. 1850, p. 327. Chap. XII1.J LAW OF BATTLE. 39 bird {Gallicrex cristatus), as Mr. Blyth informs me, are one-third larger than the females, and are so pugnacious during the breeding-season, that they are kept by the natives of Eastern Bengal for the sake of fighting. Various other birds are kept in India for the same pur- pose, for instance, the Bulbuls (Pycnonotus hcemorrhous) which " fight with great spirit." 6 The polygamous Ruff {Machetes pugnax, Fig. 37) is notorious for his extreme pugnacity ; and in the spring, the males, which are considerably larger than the females, congregate day after day at a particular spot, where the females propose to lay their eggs. The fowlers discover these spots by the turf being trampled somewhat bare. Here they fight very much like game-cocks, seizing each other with their beaks and striking with their wings. The great ruff of feathers round the neck is then erected, and according to Colonel Montagu " sweeps the ground as a shield to defend the more tender parts ; " and this is the only instance known to me in the case of birds, of any structure serving as a shield. The ruff of feathers, how- ever, from its varied and rich colors probably serves in chief part as an ornament. Like most pugnacious birds, they seem always ready to fight, and when closely confined often kill each other; but Montagu observed that their pugnacity becomes greater during the spring, when the long feathers on their necks are fully developed ; and at this period the least movement by any one bird provokes a general battle.7 Of the pugnacity of web-footed birds, two instances will suffice : in Guiana " bloody fights occur ; vol. li. p S88. 11 Mr. Hewitt in tin; ' Poultry Hook by Top'tiiu-irr,' I *•'•<'., p. 137. AS SEXUAL SELECTION: BIRDS. [Part II. garue-hen accompanied by her chickens, when the cock rushed to the rescue and drove his spur right through the eye and skull of the aggressor. The spur was with diffi- culty drawn from the skull, and as the kite though dead retained his grasp, the two birds were firmly locked to- gether; but the cock when disentangled was very little injured. The invincible courage of the game-cock is noto- rious: a gentleman who long ago witnessed the following brutal scene, told me that a bird had both its legs broke n by some accident in the cockpit, and the owner laid a wager that if the legs could be spliced so that the bird could stand upright, he would continue fighting. This was effected on the spot, and the bird fought with un- daunted courage until he received his death-stroke. In Ceylon a closely-allied and wild species, the GaUus Stan- leyi, is known to fight desperately "in defence of his seraglio," so that one of the combatants is frequently found dead.13 An Indian partridge ( Ortygornis gularis), the male of which is furnished with strong and sharp spurs, is so quarrelsome, " that the scars of former fights disfigure the breast of almost every bird you kill." 11 The males of almost all gallinaceous birds, even those which are not furnished with spurs, engage during the breeding-season in fierce conflicts. The Capercailzie and Blackcock (Tetrao urog alius and T. tetrix), which are both polygamists, have regular appointed places, where during many weeks they congregate in numbers to fight together and to display their charms before the females. M. W. Kowalevsky informs me that in Russia lie has seen the snow all bloody on the arenas where the Capercailzie have fought ; and the Blackcocks " make the feathers fly in every direction," when several "engage in a battle royal." The elder Brehm gives a curious account of the " Layard, 'Annals and Mag. of Nat. Hist.' vol. xiv. 1854, p. 68. u Jerdon, ' Birds of India,' vol. Hi. p. 674. Chap. XIII.] LAW OF BATTLE. 43 Balz, as the love-dance and love-song of the Blackcock is called in Germany. The bird utters almost continuously the most strange noises : " He holds his tail up and spreads it out like a fan, he lifts up his head and neck with all the feathers erect, and stretches his wings from the body. Then he takes a few jumps in different directions, some- times in a circle, and presses the under part of his beak so hard against the ground that the chin-feathers are rubbed off. During these movements he beats his wings and turns round and round. The more ardent he grows the more lively he becomes, until at last the bird appears like a frantic creature." At such times the blackcocks are so absorbed that they become almost blind and deaf, but less so than the capercailzie : hence bird after bird may be shot on the same spot, or even caught by the hand. After performing these antics the males begin to fight : and the same blackcock, in order to prove his strength over several antagonists, will visit in the course of one morning several Balz-places, which remain the same dur- ing successive years.14 The peacock with his long train appears more like a dandy than a warrior, but he sometimes engages in fierce contests : the Rev. W. Darwin Fox informs me that two peacocks became so excited while fighting at some little distance from Chester, that they flew over the whole city, still fighting, until they alighted on the top of St. John's tower. The spur, in those gallinaceous birds which are thus provided, is generally single ; but Polyplectron (see Pig. 51, p. 90) has two or more on each leg; and oik* of the l>loo