males suffer from their small size, for according to M. Carbonnier, they are liable to be devoured by the females of their own species when carnivorous, and no doubt by other species. Increased size must be in some manner of more importance 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.
[Fig. 29. Callionymus lyra.
Upper figure, male;
lower figure, female.
N.B. The lower figure is more reduced than the upper.]
In many species the male alone is ornamented with bright colours; or these are much brighter in the male than 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 feathers to the peacock. I am indebted for most of the following facts to the kindness of Dr. Gunther. There is reason to suspect that many tropical fishes differ sexually in colour and structure; and there are some striking cases with our British fishes. The male Callionymus lyra has been called the gemmeous dragonet “from its brilliant gem-like colours.” When fresh caught from the sea the body is yellow of various shades, striped and spotted with vivid blue on the head; the dorsal fins are pale brown with dark longitudinal bands; the ventral, caudal, and anal fins being bluish-black. The female, or sordid dragonet, was considered by Linnaeus, and by many subsequent naturalists, as a distinct species; it is of a dingy 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 (12. I have drawn up this description from Yarrell’s ‘British Fishes,’ vol. i. 1836, pp. 261 and 266.); but the most striking difference is the extraordinary elongation in the male (Fig. 29) of the dorsal fin. Mr. W. Saville Kent remarks that this “singular appendage appears from my observations of the species in confinement, to be subservient to the same end as the wattles, crests, and other abnormal adjuncts of the male in gallinaceous birds, for the purpose of fascinating their mates.” (13. ‘Nature,’ July 1873, p. 264.) The young males resemble the adult females in structure and colour. Throughout the genus Callionymus (14. ‘Catalogue of Acanth. Fishes in the British Museum,’ by Dr. Gunther, 1861, pp. 138- 151.), the male is generally much more brightly spotted than the female, and in several species, not only the dorsal, but the anal fin is much elongated in the males.
The male of the Cottus scorpius, or sea-scorpion, is slenderer and smaller than the female. There is also a great difference in colour between them. It is difficult, as Mr. Lloyd (15. ‘Game Birds of Sweden,’ etc., 1867, p. 466.) 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 colours with which it, in other respects so ill-favoured, is at that time adorned.” Both sexes of the Labrus mixtus, although very different in colour, are beautiful; the male being orange with bright blue stripes, and the female bright red with some black spots on the back.
[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 Societe 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’etalage de leurs vives couleurs chercher a attirer l’attention des femelles, lesquelles ne paraissaient indifferentes a ce manege, 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.
What, then, are we to conclude in regard to the many fishes, both sexes of which are splendidly coloured? Mr. Wallace (30. ‘Westminster Review,’ July 1867, p. 7.) believes that the species which frequent reefs, where corals and other brightly-coloured organisms abound, are brightly coloured in order to escape detection by their enemies; but according to my recollection they were thus rendered highly conspicuous. In the fresh- waters of the tropics there are no brilliantly-coloured corals or other organisms for the fishes to resemble; yet many species in the Amazons are beautifully coloured, and many of the carnivorous Cyprinidae in India are ornamented with “bright longitudinal lines of various tints.” (31. ‘Indian Cyprinidae,’ by Mr. M’Clelland, ‘Asiatic Researches,’ vol. xix. part ii. 1839, p. 230.) Mr. M’Clelland, in describing these fishes, goes so far as to suppose that “the peculiar brilliancy of their colours” serves as “a better mark for king-fishers, terns, and other birds which are destined to keep the number of these fishes in check”; but at the present day few naturalists will admit that any animal has been made conspicuous as an aid to its own destruction. It is possible that certain fishes may have been rendered conspicuous in order to warn birds and beasts of prey that they were unpalatable, as explained when treating of caterpillars; but it is not, I believe, known that any fish, at least any fresh-water fish, is rejected from being distasteful to fish-devouring animals. On the whole, the most probable view in regard to the fishes, of which both sexes are brilliantly coloured, is that their colours were acquired by the males as a sexual ornament, and were transferred equally, or nearly so, to the other sex.
We have now to consider whether, when the male differs in a marked manner from the female in colour or in other ornaments, he alone has been modified, the variations being inherited by his male offspring alone; or whether the female has been specially modified and rendered inconspicuous for the sake of protection, such modifications being inherited only by the females. It is impossible to doubt that colour has been gained by many fishes as a protection: no one can examine the speckled upper surface of a flounder, and overlook its resemblance to the sandy bed of the sea on which it lives. Certain fishes, moreover, can through the action of the nervous system change their colours in adaptation to surrounding objects, and that within a short time. (32. G. Pouchet, ‘L’Institut.’ Nov. 1, 1871, p. 134.) One of the most striking instances ever recorded of an animal being protected by its colour (as far as it can be judged of in preserved specimens), as well as by its form, is that given by Dr. Gunther (33. ‘Proc. Zoolog. Soc.’ 1865, p. 327, pl. xiv. and xv.) of a pipe-fish, which, with its reddish streaming filaments, is hardly distinguishable from the sea-weed to which it clings with its prehensile tail. But the question now under consideration is whether the females alone have been modified for this object. We can see that one sex will not be modified through natural selection for the sake of protection more than the other, supposing both to vary, unless one sex is exposed for a longer period to danger, or has less power of escaping from such danger than the other; and it does not appear that with fishes the sexes differ in these respects. As far as there is any difference, the males, from being generally smaller and from wandering more about, are exposed to greater danger than the females; and yet, when the sexes differ, the males are almost always the more conspicuously coloured. The ova are fertilised immediately after being deposited; and when this process lasts for several days, as in the case of the salmon (34. Yarrell, ‘British Fishes,’ vol. ii. p. 11.), the female, during the whole time, is attended by the male. After the ova are fertilised they are, in most cases, left unprotected by both parents, so that the males and females, as far as oviposition is concerned, are equally exposed to danger, and both are equally important for the production of fertile ova; consequently the more or less brightly-coloured individuals of either sex would be equally liable to be destroyed or preserved, and both would have an equal influence on the colours of their offspring.
Certain fishes, belonging to several families, make nests, and some of them take care of their young when hatched. Both sexes of the bright coloured Crenilabrus massa and melops work together in building their nests with sea-weed, shells, etc. (35. According to the observations of M. Gerbe; see Gunther’s ‘Record of Zoolog. Literature,’ 1865, p. 194.) But the males of certain fishes do all the work, and afterwards take exclusive charge of the young. This is the case with the dull-coloured gobies (36. Cuvier, ‘Regne Animal,’ vol. ii. 1829, p. 242.), in which the sexes are not known to differ in colour, and likewise with the sticklebacks (Gasterosteus), in which the males become brilliantly coloured during the spawning season. The male of the smooth-tailed stickleback (G. leiurus) performs the duties of a nurse with exemplary care and vigilance during a long time, and is continually employed in gently leading back the young to the nest, when they stray too far. He courageously drives away all enemies including the females of his own species. It would indeed be no small relief to the male, if the female, after depositing her eggs, were immediately devoured by some enemy, for he is forced incessantly to drive her from the nest. (37. See Mr. Warington’s most interesting description of the habits of the Gasterosteus leiurus in ‘Annals and Magazine of Nat. History,’ November 1855.)
The males of certain other fishes inhabiting South America and Ceylon, belonging to two distinct Orders, have the extraordinary habit of hatching within their mouths, or branchial cavities, the eggs laid by the females. (38. Prof. Wyman, in ‘Proc. Boston Soc. of Nat. Hist.’ Sept. 15, 1857. Also Prof. Turner, in ‘Journal of Anatomy and Physiology,’ Nov. 1, 1866, p. 78. Dr. Gunther has likewise described other cases.) I am informed by Professor Agassiz that the males of the Amazonian species which follow this habit, “not only are generally brighter than the females, but the difference is greater at the spawning-season 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 species of Chromids, as Professor Agassiz likewise informs me, sexual differences in colour 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 Pomotis does. It ought also to be observed that these sitters are among the brightest species 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 by the parents, has had little or no influence on the differences in colour between the sexes. It is further manifest, in all the cases in which the males take exclusive charge of the nests and young, that the destruction of the brighter-coloured males would be far more influential on the character of the race, than the destruction of the brighter-coloured females; for the death of the male during the period of incubation or nursing would entail the death of the young, so that they could not inherit his peculiarities; yet, in many of these very cases the males are more conspicuously coloured than the females.
In most of the Lophobranchii (Pipe-fish, Hippocampi, etc.) the males have either marsupial sacks or hemispherical depressions on the abdomen, in which the ova laid by the female are hatched. The males also shew great attachment to their young. (39. Yarrell, ‘History of British Fishes,’ vol. ii. 1836, pp. 329, 338.) The sexes do not commonly differ much in colour; but Dr. Gunther believes that the male Hippocampi are rather brighter than the females. The genus Solenostoma, however, offers a curious exceptional case (40. Dr. Gunther, since publishing an account of this species in ‘The Fishes of Zanzibar,’ by Col. Playfair, 1866, p. 137, has re-examined the specimens, and has given me the above information.), for the female is much more vividly-coloured and spotted than the male, and she alone has a marsupial 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-coloured than the male. It is improbable 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 coloured than the females, and as here the female Solenostoma takes the same charge and is brighter than the male, it might be argued that the conspicuous colours of that sex which is the more important of the two for the welfare of the offspring, must be in some manner protective. But from the large number of fishes, of which the males are either permanently or periodically brighter than the females, but whose life is not at all more important for the welfare of the species than that of the female, this view can hardly be maintained. When we treat of birds we shall meet with analogous cases, where 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 explanation, namely, that the males have selected the more attractive 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 colour or in other ornamental characters, the males originally varied, with their variations transmitted to the same sex, and accumulated through sexual selection by attracting or exciting the females. In many cases, however, such characters have been transferred, either partially or completely, to the females. In other cases, again, both sexes have been coloured alike for the sake of protection; but in no instance does it appear that the female alone has had her colours or other characters specially modified for this latter purpose.
The last point which need be noticed is that fishes are known to make various noises, some of which are described as being musical. Dr. Dufosse, who has especially attended to this subject, says that the sounds are voluntarily produced in several ways by different fishes: by the friction of the pharyngeal bones–by the vibration of certain muscles attached to the swim bladder, which serves as a resounding board–and by the vibration of the intrinsic muscles of the swim bladder. By this latter means the Trigla produces pure and long-drawn sounds which range over nearly an octave. But the most interesting case for us is that of two species of Ophidium, in which the males alone are provided with a sound-producing apparatus, consisting of small movable bones, with proper muscles, in connection with the swim bladder. (41. ‘Comptes-Rendus,’ tom. xlvi. 1858, p. 353; tom. xlvii. 1858, p. 916; tom. liv. 1862, p. 393. The noise made by the Umbrinas (Sciaena aquila), is said by some authors to be more like that of a flute or organ, than drumming: Dr. Zouteveen, in the Dutch translation of this work (vol. ii. p. 36), gives some further particulars on the sounds made by fishes.) The drumming of the Umbrinas in the European seas is said to be audible from a depth of twenty fathoms; and 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 without bait.” (42. The Rev. C. Kingsley, in ‘Nature,’ May 1870, p. 40.) From this statement, and more especially from the case of Ophidium, it is almost certain that in this, the lowest class of the Vertebrata, as with so many insects and spiders, sound-producing instruments have, at least in some cases, been developed through sexual selection, as a means for bringing the sexes together.
AMPHIBIANS.
URODELA.
[Fig. 32. Triton cristatus (half natural size, from Bell’s ‘British Reptiles’).
Upper figure, male during the breeding season; lower figure, female.]
I will begin with the tailed amphibians. The sexes of salamanders or newts often differ much both in colour and structure. In some species prehensile claws are developed on the fore-legs of the males during the breeding- 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. (43. Bell, ‘History of British Reptiles,’ 2nd ed., 1849, pp. 156-159.) This structure no doubt aids the male in his eager search and pursuit of the female. Whilst courting her he rapidly vibrates the end of his tail. With our common newts (Triton punctatus and cristatus) a deep, much indented crest is developed along the back and tail of the male during the breeding- season, which disappears during the winter. Mr. St. George Mivart informs me that it is not furnished with muscles, and therefore cannot be used for locomotion. As during the season of courtship it becomes edged with bright colours, there can hardly be a doubt that it is a masculine ornament. In many species the body presents strongly contrasted, though lurid tints, and these become more vivid during the breeding-season. The male, for instance, of our common little newt (Triton punctatus) is “brownish-grey above, passing into yellow beneath, which in the spring becomes a rich bright orange, marked everywhere with round dark spots.” The edge of the crest also is then tipped with bright red or violet. The female is usually of a yellowish-brown colour with scattered brown dots, and the lower surface is often quite plain. (44. Bell, ‘History of British Reptiles,’ 2nd ed., 1849, pp. 146, 151.) The young are obscurely tinted. The ova are fertilised during the act of deposition, and are not subsequently tended by either parent. We may therefore conclude that the males have acquired their strongly-marked colours and ornamental appendages through sexual selection; these being transmitted either to the male offspring alone, or to both sexes.
ANURA OR BATRACHIA.
With many frogs and toads the colours evidently serve as a protection, such as the bright green tints of tree frogs and the obscure mottled shades of many terrestrial species. The most conspicuously-coloured toad which I ever saw, the Phryniscus nigricans (45. ‘Zoology of the Voyage of the “Beagle,”‘ 1843. Bell, ibid. p. 49.), had the whole upper surface of the body as black as ink, with the soles of the feet and parts of the abdomen spotted with the brightest vermilion. It crawled about the bare sandy or open grassy plains of La Plata under a scorching sun, and could not fail to catch the eye of every passing creature. These colours are probably beneficial by making this animal known to all birds of prey as a nauseous mouthful.
In Nicaragua there is a little frog “dressed in a bright livery of red and blue” which does not conceal itself like most other species, but hops about during the daytime, and Mr. Belt says (46. ‘The Naturalist in Nicaragua,’ 1874, p. 321.) that as soon as he saw its happy sense of security, he felt sure that it was uneatable. After several trials he succeeded in tempting a young duck to snatch up a young one, but it was instantly rejected; and the duck “went about jerking its head, as if trying to throw off some unpleasant taste.”
With respect to sexual differences of colour, Dr. Gunther does not know of any striking instance either with frogs or toads; yet he can often distinguish the male from the female by the tints of the former being a little more intense. Nor does he know of any striking difference in external structure between the sexes, excepting the prominences which become developed during the breeding-season on the front legs of the male, by which he is enabled to hold the female. (47. The male alone of the Bufo sikimmensis (Dr. Anderson, ‘Proc. Zoolog. Soc.’ 1871, p. 204) has two plate-like callosities on the thorax and certain rugosities on the fingers, which perhaps subserve the same end as the above-mentioned prominences.) It is surprising that these animals have not acquired more strongly-marked sexual characters; for though cold-blooded their passions are strong. Dr. Gunther informs me that he has several times found an unfortunate female toad dead and smothered from having been so closely embraced by three or four males. Frogs have been observed by Professor Hoffman in Giessen fighting all day long during the breeding-season, and with so much violence that one had its body ripped open.
Frogs and toads offer one interesting sexual difference, namely, in the musical powers possessed by the males; but to speak of music, when applied to the discordant and overwhelming sounds emitted by male bull-frogs and some other species, seems, according to our taste, a singularly inappropriate expression. Nevertheless, certain frogs sing in a decidedly pleasing manner. Near Rio Janeiro I used often to sit in the evening to listen to a number of little Hylae, perched on blades of grass close to the water, which sent forth sweet chirping notes in harmony. The various sounds are emitted chiefly by the males during the breeding-season, as in the case of the croaking of our common frog. (48. Bell, ‘History British Reptiles,’ 1849, p. 93.) In accordance with this fact the vocal organs of the males are more highly-developed than those of the females. In some genera the males alone are provided with sacs which open into the larynx. (49. J. Bishop, in ‘Todd’s Cyclopaedia of Anatomy and Physiology,’ vol. iv. p. 1503.) For instance, in the edible frog (Rana esculenta) “the sacs are peculiar to the males, and become, when filled with air in the act of croaking, large globular bladders, standing out one on each side of the head, near the corners of the mouth.” The croak of the male is thus rendered exceedingly powerful; whilst that of the female is only a slight groaning noise. (50. Bell, ibid. pp. 112-114.) In the several genera of the family the vocal organs differ considerably in structure, and their development in all cases may be attributed to sexual selection.
REPTILES.
CHELONIA.
Tortoises and turtles do not offer well-marked sexual differences. In some species, the tail of the male is longer than that of the female. In some, the plastron or lower surface of the shell of the male is slightly concave in relation to the back of the female. The male of the mud-turtle of the United States (Chrysemys picta) has claws on its front feet twice as long as those of the female; and these are used when the sexes unite. (51. Mr. C.J. Maynard, ‘The American Naturalist,’ Dec. 1869, p. 555.) With the huge tortoise of the Galapagos Islands (Testudo nigra) the males are said to grow to a larger size than the females: during the pairing-season, and at no other time, the male utters a hoarse bellowing noise, which can be heard at the distance of more than a hundred yards; the female, on the other hand, never uses her voice. (52. See my ‘Journal of Researches during the Voyage of the “Beagle,”‘ 1845, p. 384.)
With the Testudo elegans of India, it is said “that the combats of the males may be heard at some distance, from the noise they produce in butting against each other.” (53. Dr. Gunther, ‘Reptiles of British India,’ 1864, p. 7.)
CROCODILIA.
The sexes apparently do not differ in colour; nor do I know that the males fight together, though this is probable, for some kinds make a prodigious display before the females. Bartram (54. ‘Travels through Carolina,’ etc., 1791, p. 128.) describes the male alligator as striving to win the female by splashing and roaring in the midst of a lagoon, “swollen to an extent ready to burst, with its head and tail lifted up, he springs or twirls round on the surface of the water, like an Indian chief rehearsing his feats of war.” During the season of love, a musky odour is emitted by the submaxiliary glands of the crocodile, and pervades their haunts. (55. Owen, ‘Anatomy of Vertebrates,’ vol. i. 1866, p. 615.)
OPHIDIA.
Dr. Gunther informs me that the males are always smaller than the females, and generally have longer and slenderer tails; but he knows of no other difference in external structure. In regard to colour, be can almost always distinguish the male from the female, by his more strongly- pronounced tints; thus the black zigzag band on the back of the male English viper is more distinctly defined than in the female. The difference is much plainer in the rattle-snakes of N. America, the male of which, as the keeper in the Zoological Gardens shewed me, can at once be distinguished from the female by having more lurid yellow about its whole body. In S. Africa the Bucephalus capensis presents an analogous difference, for the female “is never so fully variegated with yellow on the sides as the male.” (56. Sir Andrew Smith, ‘Zoology of S. Africa: Reptilia,’ 1849, pl. x.) The male of the Indian Dipsas cynodon, on the other hand, is blackish-brown, with the belly partly black, whilst the female is reddish or yellowish-olive, with the belly either uniform yellowish or marbled with black. In the Tragops dispar of the same country the male is bright green, and the female bronze-coloured. (57. Dr. A. Gunther, ‘Reptiles of British India,’ Ray Soc., 1864, pp. 304, 308.) No doubt the colours of some snakes are protective, as shewn by the green tints of tree-snakes, and the various mottled shades of the species which live in sandy places; but it is doubtful whether the colours of many kinds, for instance of the common English snake and viper, serve to conceal them; and this is still more doubtful with the many foreign species which are coloured with extreme elegance. The colours of certain species are very different in the adult and young states. (58. Dr. Stoliczka, ‘Journal of Asiatic Society of Bengal,’ vol. xxxix, 1870, pp. 205, 211.)
During the breeding-season the anal scent-glands of snakes are in active function (59. Owen, ‘Anatomy of Vertebrates,’ vol. i. 1866, p. 615.); and so it is with the same glands in lizards, and as we have seen with the submaxiliary glands of crocodiles. As the males of most animals search for the females, these odoriferous glands probably serve to excite or charm the female, rather than to guide her to the spot where the male may be found. Male snakes, though appearing so sluggish, are amorous; for many have been observed crowding round the same female, and even round her dead body. They are not known to fight together from rivalry. Their intellectual powers are higher than might have been anticipated. In the Zoological Gardens they soon learn not to strike at the iron bar with which their cages are cleaned; and Dr. Keen of Philadelphia informs me that some snakes which he kept learned after four or five times to avoid a noose, with which they were at first easily caught. An excellent observer in Ceylon, Mr. E. Layard, saw (60. ‘Rambles in Ceylon,’ in ‘Annals and Magazine of Natural History,’ 2nd series, vol. ix. 1852, p. 333.) a cobra thrust its head through a narrow hole and swallow a toad. “With this encumbrance he could not withdraw himself; finding this, he reluctantly disgorged the precious morsel, which began to move off; this was too much for snake philosophy to bear, and the toad was again seized, and again was the snake, after violent efforts to escape, compelled to part with its prey. This time, however, a lesson had been learnt, and the toad was seized by one leg, withdrawn, and then swallowed in triumph.”
The keeper in the Zoological Gardens is positive that certain snakes, for instance Crotalus and Python, distinguish him from all other persons. Cobras kept together in the same cage apparently feel some attachment towards each other. (61. Dr. Gunther, ‘Reptiles of British India,’ 1864, p. 340.)
It does not, however, follow because snakes have some reasoning power, strong passions and mutual affection, that they should likewise be endowed with sufficient taste to admire brilliant colours in their partners, so as to lead to the adornment of the species through sexual selection. Nevertheless, it is difficult to account in any other manner for the extreme beauty of certain species; for instance, of the coral-snakes of S. America, which are of a rich red with black and yellow transverse bands. I well remember how much surprise I felt at the beauty of the first coral- snake which I saw gliding across a path in Brazil. Snakes coloured in this peculiar manner, as Mr. Wallace states on the authority of Dr. Gunther (62. ‘Westminster Review,’ July 1st, 1867, p. 32.), are found nowhere else in the world except in S. America, and here no less than four genera occur. One of these, Elaps, is venomous; a second and widely-distinct genus is doubtfully venomous, and the two others are quite harmless. The species belonging to these distinct genera inhabit the same districts, and are so like each other that no one “but a naturalist would distinguish the harmless from the poisonous kinds.” Hence, as Mr. Wallace believes, the innocuous kinds have probably acquired their colours as a protection, on the principle of imitation; for they would naturally be thought dangerous by their enemies. The cause, however, of the bright colours of the venomous Elaps remains to be explained, and this may perhaps be sexual selection.
Snakes produce other sounds besides hissing. The deadly Echis carinata has on its sides some oblique rows of scales of a peculiar structure with serrated edges; and when this snake is excited these scales are rubbed against each other, which produces “a curious prolonged, almost hissing sound.” (63. Dr. Anderson, ‘Proc. Zoolog. Soc.’ 1871, p. 196.) With respect to the rattling of the rattle-snake, we have at last some definite information: for Professor Aughey states (64. The ‘American Naturalist,’ 1873, p. 85.), that on two occasions, being himself unseen, he watched from a little distance a rattle-snake coiled up with head erect, which continued to rattle at short intervals for half an hour: and at last he saw another snake approach, and when they met they paired. Hence he is satisfied that one of the uses of the rattle is to bring the sexes together. Unfortunately he did not ascertain whether it was the male or the female which remained stationary and called for the other. But it by no means follows from the above fact that the rattle may not be of use to these snakes in other ways, as a warning to animals which would otherwise attack them. Nor can I quite disbelieve the several accounts which have appeared of their thus paralysing their prey with fear. Some other snakes also make a distinct noise by rapidly vibrating their tails against the surrounding stalks of plants; and I have myself heard this in the case of a Trigonocephalus in S. America.
LACERTILIA.
The males of some, probably of many kinds of lizards, fight together from rivalry. Thus the arboreal Anolis cristatellus of S. America is extremely pugnacious: “During the spring and early part of the summer, two adult males rarely meet without a contest. On first seeing one another, they nod their heads up and down three or four times, and at the same time expanding the frill or pouch beneath the throat; their eyes glisten with rage, and after waving their tails from side to side for a few seconds, as if to gather energy, they dart at each other furiously, rolling over and over, and holding firmly with their teeth. The conflict generally ends in one of the combatants losing his tail, which is often devoured by the victor.” The male of this species is considerably larger than the female (65. Mr. N.L. Austen kept these animals alive for a considerable time; see ‘Land and Water,’ July 1867, p. 9.); and this, as far as Dr. Gunther has been able to ascertain, is the general rule with lizards of all kinds. The male alone of the Cyrtodactylus rubidus of the Andaman Islands possesses pre-anal pores; and these pores, judging from analogy, probably serve to emit an odour. (66. Stoliczka, ‘Journal of the Asiatic Society of Bengal,’ vol. xxxiv. 1870, p. 166.)
[Fig.33. Sitana minor.
Male with the gular pouch expanded (from Gunther’s ‘Reptiles of India’)’]
The sexes often differ greatly in various external characters. The male of the above-mentioned Anolis is furnished with a crest which runs along the back and tail, and can be erected at pleasure; but of this crest the female does not exhibit a trace. In the Indian Cophotis ceylanica, the female has a dorsal crest, though much less developed than in the male; and so it is, as Dr. Gunther informs me, with the females of many Iguanas, Chameleons, and other lizards. In some species, however, the crest is equally developed in both sexes, as in the Iguana tuberculata. In the genus Sitana, the males alone are furnished with a large throat pouch (Fig. 33), which can be folded up like a fan, and is coloured blue, black, and red; but these splendid colours are exhibited only during the pairing-season. The female does not possess even a rudiment of this appendage. In the Anolis cristatellus, according to Mr. Austen, the throat pouch, which is bright red marbled with yellow, is present in the female, though in a rudimental condition. Again, in certain other lizards, both sexes are equally well provided with throat pouches. Here we see with species belonging to the same group, as in so many previous cases, the same character either confined to the males, or more largely developed in them than in the females, or again equally developed in both sexes. The little lizards of the genus Draco, which glide through the air on their rib- supported parachutes, and which in the beauty of their colours baffle description, are furnished with skinny appendages to the throat “like the wattles of gallinaceous birds.” These become erected when the animal is excited. They occur in both sexes, but are best developed when the male arrives at maturity, at which age the middle appendage is sometimes twice as long as the head. Most of the species likewise have a low crest running along the neck; and this is much more developed in the full-grown males than in the females or young males. (67. All the foregoing statements and quotations, in regard to Cophotis, Sitana and Draco, as well as the following facts in regard to Ceratophora and Chamaeleon, are from Dr. Gunther himself, or from his magnificent work on the ‘Reptiles of British India,’ Ray Soc., 1864, pp. 122, 130, 135.)
A Chinese species is said to live in pairs during the spring; “and if one is caught, the other falls from the tree to the ground, and allows itself to be captured with impunity”–I presume from despair. (68. Mr. Swinhoe, ‘Proc. Zoolog. Soc.’ 1870, p. 240.)
[Fig. 34. Ceratophora Stoddartii.
Upper figure;
lower figure, female.]
There are other and much more remarkable differences between the sexes of certain lizards. The male of Ceratophora aspera bears on the extremity of his snout an appendage half as long as the head. It is cylindrical, covered with scales, flexible, and apparently capable of erection: in the female it is quite rudimental. In a second species of the same genus a terminal scale forms a minute horn on the summit of the flexible appendage; and in a third species (C. Stoddartii, fig. 34) the whole appendage is converted into a horn, which is usually of a white colour, but assumes a purplish tint when the animal is excited. In the adult male of this latter species the horn is half an inch in length, but it is of quite minute size in the female and in the young. These appendages, as Dr. Gunther has remarked to me, may be compared with the combs of gallinaceous birds, and apparently serve as ornaments.
[Fig. 35. Chamaeleo bifurcus.
Upper figure, male;
lower figure, female.
Fig. 36. Chamaeleo Owenii.
Upper figure, male;
lower figure, female.]
In the genus Chamaeleon we come to the acme of difference between the sexes. The upper part of the skull of the male C. bifurcus (Fig. 35), an inhabitant of Madagascar, is produced into two great, solid, bony projections, covered with scales like the rest of the head; and of this wonderful modification of structure the female exhibits only a rudiment. Again, in Chamaeleo Owenii (Fig. 36), from the West Coast of Africa, the male bears on his snout and forehead three curious horns, of which the female has not a trace. These horns consist of an excrescence of bone covered with a smooth sheath, forming part of the general integuments of the body, so that they are identical in structure with those of a bull, goat, or other sheath-horned ruminant. Although the three horns differ so much in appearance from the two great prolongations of the skull in C. bifurcus, we can hardly doubt that they serve the same general purpose in the economy of these two animals. The first conjecture, which will occur to every one, is that they are used by the males for fighting together; and as these animals are very quarrelsome (69. Dr. Buchholz, ‘Monatsbericht K. Preuss. Akad.’ Jan. 1874, p. 78.), this is probably a correct view. Mr. T.W. Wood also informs me that he once watched two individuals of C. pumilus fighting violently on the branch of a tree; they flung their heads about and tried to bite each other; they then rested for a time and afterwards continued their battle.
With many lizards the sexes differ slightly in colour, the tints and stripes of the males being brighter and more distinctly defined than in the females. This, for instance, is the case with the above Cophotis and with the Acanthodactylus capensis of S. Africa. In a Cordylus of the latter country, the male is either much redder or greener than the female. In the Indian Calotes nigrilabris there is a still greater difference; the lips also of the male are black, whilst those of the female are green. In our common little viviparous lizard (Zootoca vivipara) “the under side of the body and base of the tail in the male are bright orange, spotted with black; in the female these parts are pale-greyish-green without spots.” (70. Bell, ‘History of British Reptiles,’ 2nd ed., 1849, p. 40.) We have seen that the males alone of Sitana possess a throat-pouch; and this is splendidly tinted with blue, black, and red. In the Proctotretus tenuis of Chile the male alone is marked with spots of blue, green, and coppery-red. (71. For Proctotretus, see ‘Zoology of the Voyage of the “Beagle”; Reptiles,’ by Mr. Bell, p. 8. For the Lizards of S. Africa, see ‘Zoology of S. Africa: Reptiles,’ by Sir Andrew Smith, pl. 25 and 39. For the Indian Calotes, see ‘Reptiles of British India,’ by Dr. Gunther, p. 143.) In many cases the males retain the same colours throughout the year, but in others they become much brighter during the breeding-season; I may give as an additional instance the Calotes maria, which at this season has a bright red head, the rest of the body being green. (72. Gunther in ‘Proceedings, Zoological Society,’ 1870, p. 778, with a coloured figure.)
Both sexes of many species are beautifully coloured exactly alike; and there is no reason to suppose that such colours are protective. No doubt with the bright green kinds which live in the midst of vegetation, this colour serves to conceal them; and in N. Patagonia I saw a lizard (Proctotretus multimaculatus) which, when frightened, flattened its body, closed its eyes, and then from its mottled tints was hardly distinguishable from the surrounding sand. But the bright colours with which so many lizards are ornamented, as well as their various curious appendages, were probably acquired by the males as an attraction, and then transmitted either to their male offspring alone, or to both sexes. Sexual selection, indeed, seems to have played almost as important a part with reptiles as with birds; and the less conspicuous colours of the females in comparison with the males cannot be accounted for, as Mr. Wallace believes to be the case with birds, by the greater exposure of the females to danger during incubation.
CHAPTER XIII.
SECONDARY SEXUAL CHARACTERS OF BIRDS.
Sexual differences–Law of battle–Special weapons–Vocal organs– Instrumental music–Love-antics and dances–Decorations, permanent 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 considerable length. Male birds sometimes, though rarely, possess special weapons for fighting with each other. They charm the female by vocal or instrumental music of the most varied kinds. They are ornamented by all sorts of combs, wattles, protuberances, horns, air-distended sacks, top- knots, 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 coloured. 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 odour, which we may suppose serves to charm or excite the female; for that excellent observer, Mr. Ramsay (1. ‘Ibis,’ vol. iii. (new series), 1867, p. 414.), says of the Australian musk-duck (Biziura 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 smell of musk.” So powerful is this odour during the pairing-season, that it can be detected long before the bird can be seen. (2. Gould, ‘Handbook of the Birds of Australia,’ 1865, vol. ii. p. 383.) 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 shewn by our enjoyment of the singing of birds, and by our women, both civilised and savage, decking their heads with borrowed plumes, and using gems which are hardly more brilliantly coloured than the naked skin and wattles of certain birds. In man, however, when cultivated, the sense of beauty is manifestly a far more complex feeling, and is associated with various intellectual ideas.
Before treating of the sexual 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 male and female 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 Neomorpha of New Zealand, there is, as we have seen, a still wider difference in the form of the beak in relation to the manner of feeding of the two sexes. Something of the same kind has been observed with the goldfinch (Carduelis elegans), for I am assured by Mr. J. Jenner Weir that the bird-catchers can distinguish the males by their slightly longer beaks. The flocks of males are often found feeding on the seeds of the teazle (Dipsacus), which they can reach with their elongated beaks, whilst the females more commonly feed on the seeds of the betony or Scrophularia. With a slight difference of this kind as a foundation, we can see how the beaks of the two sexes might be made to differ greatly through natural selection. In some of the above cases, however, it is possible that the beaks of the males may have been first modified in relation to their contests with other males; and that this afterwards 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 humming-bird, is one of the most quarrelsome. Mr. Gosse (3. Quoted by Mr. Gould, ‘Introduction to the Trochilidae,’ 1861, page 29.) describes a battle in which a pair 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 or another genus of humming-bird, 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.” (4. Gould, ibid. p. 52.) 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 half 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. (5. W. Thompson, ‘Natural History of Ireland: Birds,’ vol. ii. 1850, p. 327.) Mr. Blyth informs me that the males of an allied bird (Gallicrex cristatus) are a 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 purpose, for instance, the bulbuls (Pycnonotus hoemorrhous) which “fight with great spirit.” (6. Jerdon, ‘Birds of India,’ 1863, vol. ii. p. 96.)
[Fig. 37. The Ruff or Machetes pugnax (from Brehm’s ‘Thierleben’).]
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 Col. 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, however, from its varied and rich colours 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. Macgillivray, ‘History of British Birds,’ vol. iv. 1852, pp. 177-181.) Of the pugnacity of web-footed birds, two instances will suffice: in Guiana “bloody fights occur during the breeding-season between the males of the wild musk-duck (Cairina moschata); and where these fights have occurred the river is covered for some distance with feathers.” (8. Sir R. Schomburgk, in ‘Journal of Royal Geographic Society,’ vol. xiii. 1843, p. 31.) Birds which seem ill-adapted for fighting engage in fierce conflicts; thus the stronger males of the pelican drive away the weaker ones, snapping with their huge beaks and giving heavy blows with their wings. Male snipe fight together, “tugging and pushing each other with their bills in the most curious manner imaginable.” Some few birds are believed never to fight; this is the case, according to Audubon, with one of the woodpeckers of the United States (Picu sauratus), although “the hens are followed by even half a dozen of their gay suitors.” (9. ‘Ornithological Biography,’ vol. i. p. 191. For pelicans and snipes, see vol. iii. pp. 138, 477.)
The males of many birds are larger than the females, and this no doubt is the result of the advantage gained by the larger and stronger males over their rivals during many generations. The difference in size between the two sexes is carried to an extreme point in several Australian species; thus the male musk-duck (Biziura), and the male Cincloramphus cruralis (allied to our pipits) are by measurement actually twice as large as their respective females. (10. Gould, ‘Handbook of Birds of Australia,’ vol. i. p. 395; vol. ii. p. 383.) With many other birds the females are larger than the males; and, as formerly remarked, the explanation often given, namely, that the females have most of the work in feeding their young, will not suffice. In some few cases, as we shall hereafter see, the females apparently have acquired their greater size and strength for the sake of conquering other females and obtaining possession of the males.
The males of many gallinaceous birds, especially of the polygamous kinds, are furnished with special weapons for fighting with their rivals, namely spurs, which can be used with fearful effect. It has been recorded by a trustworthy writer (11. Mr. Hewitt, in the ‘Poultry Book’ by Tegetmeier, 1866, p. 137.) that in Derbyshire a kite struck at a game-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 difficulty drawn from the skull, and as the kite, though dead, retained his grasp, the two birds were firmly locked together; but the cock when disentangled was very little injured. The invincible courage of the game- cock is notorious: a gentleman who long ago witnessed the brutal scene, told me that a bird had both its legs broken 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 undaunted courage until he received his death-stroke. In Ceylon a closely allied, wild species, the Gallus Stanleyi, is known to fight desperately “in defence of his seraglio,” so that one of the combatants is frequently found dead. (12. Layard, ‘Annals and Magazine of Natural History,’ vol. xiv. 1854, p. 63.) 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.” (13. Jerdon, ‘Birds of India,’ vol. iii. p. 574.)
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 Black-cock (Tetrao urogallus 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. Dr. W. Kovalevsky informs me that in Russia he has seen the snow all bloody on the arenas where the capercailzie have fought; and the black-cocks “make the feathers fly in every direction,” when several “engage in a battle royal.” The elder Brehm gives a curious account of the Balz, as the love-dances and love-songs of the Black-cock are called in Germany. The bird utters almost continuously the strangest 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, sometimes 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 black-cocks 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 black-cock, in order to prove his strength over several antagonists, will visit in the course of one morning several Balz-places, which remain the same during successive years. (14. Brehm, ‘Thierleben,’ 1867, B. iv. s. 351. Some of the foregoing statements are taken from L. Lloyd, ‘The Game Birds of Sweden,’ etc., 1867, p. 79.)
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 at some little distance from Chester two peacocks became so excited whilst fighting, that they flew over the whole city, still engaged, 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 (Fig. 51) has two or more on each leg; and one of the Blood-pheasants (Ithaginis cruentus) has been seen with five spurs. The spurs are generally confined to the male, being represented by mere knobs or rudiments in the female; but the females of the Java peacock (Pavo muticus) and, as I am informed by Mr. Blyth, of the small fire-backed pheasant (Euplocamus erythrophthalmus) possess spurs. In Galloperdix it is usual for the males to have two spurs, and for the females to have only one on each leg. (15. Jerdon, ‘Birds of India’: on Ithaginis, vol. iii. p. 523; on Galloperdix, p. 541.) Hence spurs may be considered as a masculine structure, which has been occasionally more or less transferred to the females. Like most other secondary sexual characters, the spurs are highly variable, both in number and development, in the same species.
[Fig.38. Palamedea cornuta (from Brehm), shewing the double wing-spurs, and the filament on the head.]
Various birds have spurs on their wings. But the Egyptian goose (Chenalopex aegyptiacus) has only “bare obtuse knobs,” and these probably shew us the first steps by which true spurs have been developed in other species. In the spur-winged goose, Plectropterus gambensis, the males have much larger spurs than the females; and they use them, as I am informed by Mr. Bartlett, in fighting together, so that, in this case, the wing-spurs serve as sexual weapons; but according to Livingstone, they are chiefly used in the defence of the young. The Palamedea (Fig. 38) is armed with a pair of spurs on each wing; and these are such formidable weapons that a single blow has been known to drive a dog howling away. But it does not appear that the spurs in this case, or in that of some of the spur-winged rails, are larger in the male than in the female. (16. For the Egyptian goose, see Macgillivray, ‘British Birds,’ vol. iv. p. 639. For Plectropterus, Livingstone’s ‘Travels,’ p. 254. For Palamedea, Brehm’s ‘Thierleben,’ B. iv. s. 740. See also on this bird Azara, ‘Voyages dans l’Amerique merid.’ tom. iv. 1809, pp. 179, 253.) In certain plovers, however, the wing-spurs must be considered as a sexual character. Thus in the male of our common peewit (Vanellus cristatus) the tubercle on the shoulder of the wing becomes more prominent during the breeding-season, and the males fight together. In some species of Lobivanellus a similar tubercle becomes developed during the breeding-season “into a short horny spur.” In the Australian L. lobatus both sexes have spurs, but these are much larger in the males than in the females. In an allied bird, the Hoplopterus armatus, the spurs do not increase in size during the breeding- season; but these birds have been seen in Egypt to fight together, in the same manner as our peewits, by turning suddenly in the air and striking sideways at each other, sometimes with fatal results. Thus also they drive away other enemies. (17. See, on our peewit, Mr. R. Carr in ‘Land and Water,’ Aug. 8th, 1868, p. 46. In regard to Lobivanellus, see Jerdon’s ‘Birds of India,’ vol. iii. p. 647, and Gould’s ‘Handbook of Birds of Australia,’ vol. ii. p. 220. For the Hoplopterus, see Mr. Allen in the ‘Ibis,’ vol. v. 1863, p. 156.)
The season of love is that of battle; but the males of some birds, as of the game-fowl and ruff, and even the young males of the wild turkey and grouse (18. Audubon, ‘Ornithological Biography,’ vol. ii. p. 492; vol. i. pp. 4-13.), are ready to fight whenever they meet. The presence of the female is the teterrima belli causa. The Bengali baboos make the pretty little males of the amadavat (Estrelda amandava) fight together by placing three small cages in a row, with a female in the middle; after a little time the two males are turned loose, and immediately a desperate battle ensues. (19. Mr. Blyth, ‘Land and Water,’ 1867, p. 212.) When many males congregate at the same appointed spot and fight together, as in the case of grouse and various other birds, they are generally attended by the females (20. Richardson on Tetrao umbellus, ‘Fauna Bor. Amer.: Birds,’ 1831, p. 343. L. Lloyd, ‘Game Birds of Sweden,’ 1867, pp. 22, 79, on the capercailzie and black-cock. Brehm, however, asserts (‘Thierleben,’ B. iv. s. 352) that in Germany the grey-hens do not generally attend the Balzen of the black-cocks, but this is an exception to the common rule; possibly the hens may lie hidden in the surrounding bushes, as is known to be the case with the gray-hens in Scandinavia, and with other species in N. America.), which afterwards pair with the victorious combatants. But in some cases the pairing precedes instead of succeeding the combat: thus according to Audubon (21. ‘Ornithological Biography,’ vol. ii. p. 275.), several males of the Virginian goat-sucker (Caprimulgus virgianus) “court, in a highly entertaining manner the female, and no sooner has she made her choice, than her approved gives chase to all intruders, and drives them beyond his dominions.” Generally the males try to drive away or kill their rivals before they pair. It does not, however, appear that the females invariably prefer the victorious males. I have indeed been assured by Dr. W. Kovalevsky that the female capercailzie sometimes steals away with a young male who has not dared to enter the arena with the older cocks, in the same manner as occasionally happens with the does of the red-deer in Scotland. When two males contend in presence of a single female, the victor, no doubt, commonly gains his desire; but some of these battles are caused by wandering males trying to distract the peace of an already mated pair. (22. Brehm, ‘Thierleben,’ etc., B. iv. 1867, p. 990. Audubon, ‘Ornithological Biography,’ vol. ii. p. 492.)
Even with the most pugnacious species it is probable that the pairing does not depend exclusively on the mere strength and courage of the male; for such males are generally decorated with various ornaments, which often become more brilliant during the breeding-season, and which are sedulously displayed before the females. The males also endeavour to charm or excite their mates by love-notes, songs, and antics; and the courtship is, in many instances, a prolonged affair. Hence it is not probable that the females are indifferent to the charms of the opposite sex, or that they are invariably compelled to yield to the victorious males. It is more probable that the females are excited, either before or after the conflict, by certain males, and thus unconsciously prefer them. In the case of Tetrao umbellus, a good observer (23. ‘Land and Water,’ July 25, 1868, p. 14.) goes so far as to believe that the battles of the male “are all a sham, performed to show themselves to the greatest advantage before the admiring females who assemble around; for I have never been able to find a maimed hero, and seldom more than a broken feather.” I shall have to recur to this subject, but I may here add that with the Tetrao cupido of the United States, about a score of males assemble at a particular spot, and, strutting about, make the whole air resound with their extraordinary noises. At the first answer from a female the males begin to fight furiously, and the weaker give way; but then, according to Audubon, both the victors and vanquished search for the female, so that the females must either then exert a choice, or the battle must be renewed. So, again, with one of the field-starlings of the United States (Sturnella ludoviciana) the males engage in fierce conflicts, “but at the sight of a female they all fly after her as if mad.” (24. Audubon’s ‘Ornithological Biography;’ on Tetrao cupido, vol. ii. p. 492; on the Sturnus, vol. ii. p. 219.)
VOCAL AND INSTRUMENTAL MUSIC.
With birds the voice serves to express various emotions, such as distress, fear, anger, triumph, or mere happiness. It is apparently sometimes used to excite terror, as in the case of the hissing noise made by some nestling-birds. Audubon (25. ‘Ornithological Biography,’ vol. v. p. 601.), relates that a night-heron (Ardea nycticorax, Linn.), which he kept tame, used to hide itself when a cat approached, and then “suddenly start up uttering one of the most frightful cries, apparently enjoying the cat’s alarm and flight.” The common domestic cock clucks to the hen, and the hen to her chickens, when a dainty morsel is found. The hen, when she has laid an egg, “repeats the same note very often, and concludes with the sixth above, which she holds for a longer time” (26. The Hon. Daines Barrington, ‘Philosophical Transactions,’ 1773, p. 252.); and thus she expresses her joy. Some social birds apparently call to each other for aid; and as they flit from tree to tree, the flock is kept together by chirp answering chirp. During the nocturnal migrations of geese and other water-fowl, sonorous clangs from the van may be heard in the darkness overhead, answered by clangs in the rear. Certain cries serve as danger signals, which, as the sportsman knows to his cost, are understood by the same species and by others. The domestic cock crows, and the humming-bird chirps, in triumph over a defeated rival. The true song, however, of most birds and various strange cries are chiefly uttered during the breeding- season, and serve as a charm, or merely as a call-note, to the other sex.
Naturalists are much divided with respect to the object of the singing of birds. Few more careful observers ever lived than Montagu, and he maintained that the “males of song-birds and of many others do not in general search for the female, but, on the contrary, their business in the spring is to perch on some conspicuous spot, breathing out their full and armorous notes, which, by instinct, the female knows, and repairs to the spot to choose her mate.” (27. ‘Ornithological Dictionary,’ 1833, p. 475.) Mr. Jenner Weir informs me that this is certainly the case with the nightingale. Bechstein, who kept birds during his whole life, asserts, “that the female canary always chooses the best singer, and that in a state of nature the female finch selects that male out of a hundred whose notes please her most.” (28. ‘Naturgeschichte der Stubenvogel,’ 1840, s. 4. Mr. Harrison Weir likewise writes to me:–“I am informed that the best singing males generally get a mate first, when they are bred in the same room.”) There can be no doubt that birds closely attend to each other’s song. Mr. Weir has told me of the case of a bullfinch which had been taught to pipe a German waltz, and who was so good a performer that he cost ten guineas; when this bird was first introduced into a room where other birds were kept and he began to sing, all the others, consisting of about twenty linnets and canaries, ranged themselves on the nearest side of their cages, and listened with the greatest interest to the new performer. Many naturalists believe that the singing of birds is almost exclusively “the effect of rivalry and emulation,” and not for the sake of charming their mates. This was the opinion of Daines Barrington and White of Selborne, who both especially attended to this subject. (29. ‘Philosophical Transactions,’ 1773, p. 263. White’s ‘Natural History of Selborne,’ 1825, vol. i. p. 246.) Barrington, however, admits that “superiority in song gives to birds an amazing ascendancy over others, as is well known to bird- catchers.”
It is certain that there is an intense degree of rivalry between the males in their singing. Bird-fanciers match their birds to see which will sing longest; and I was told by Mr. Yarrell that a first-rate bird will sometimes sing till he drops down almost dead, or according to Bechstein (30. ‘Naturgesch. der Stubenvogel,’ 1840, s. 252.), quite dead from rupturing a vessel in the lungs. Whatever the cause may be, male birds, as I hear from Mr. Weir, often die suddenly during the season of song. That the habit of singing is sometimes quite independent of love is clear, for a sterile, hybrid canary-bird has been described (31. Mr. Bold, ‘Zoologist,’ 1843-44, p. 659.) as singing whilst viewing itself in a mirror, and then dashing at its own image; it likewise attacked with fury a female canary, when put into the same cage. The jealousy excited by the act of singing is constantly taken advantage of by bird-catchers; a male, in good song, is hidden and protected, whilst a stuffed bird, surrounded by limed twigs, is exposed to view. In this manner, as Mr. Weir informs me, a man has in the course of a single day caught fifty, and in one instance, seventy, male chaffinches. The power and inclination to sing differ so greatly with birds that although the price of an ordinary male chaffinch is only sixpence, Mr. Weir saw one bird for which the bird-catcher asked three pounds; the test of a really good singer being that it will continue to sing whilst the cage is swung round the owner’s head.
That male birds should sing from emulation as well as for charming the female, is not at all incompatible; and it might have been expected that these two habits would have concurred, like those of display and pugnacity. Some authors, however, argue that the song of the male cannot serve to charm the female, because the females of some few species, such as of the canary, robin, lark, and bullfinch, especially when in a state of widowhood, as Bechstein remarks, pour forth fairly melodious strains. In some of these cases the habit of singing may be in part attributed to the females having been highly fed and confined (32. D. Barrington, ‘Philosophical Transactions,’ 1773, p. 262. Bechstein, ‘Stubenvogel,’ 1840, s. 4.), for this disturbs all the functions connected with the reproduction of the species. Many instances have already been given of the partial transference of secondary masculine characters to the female, so that it is not at all surprising that the females of some species should possess the power of song. It has also been argued, that the song of the male cannot serve as a charm, because the males of certain species, for instance of the robin, sing during the autumn. (33. This is likewise the case with the water-ouzel; see Mr. Hepburn in the ‘Zoologist,’ 1845-46, p. 1068.) But nothing is more common than for animals to take pleasure in practising whatever instinct they follow at other times for some real good. How often do we see birds which fly easily, gliding and sailing through the air obviously for pleasure? The cat plays with the captured mouse, and the cormorant with the captured fish. The weaver-bird (Ploceus), when confined in a cage, amuses itself by neatly weaving blades of grass between the wires of its cage. Birds which habitually fight during the breeding-season are generally ready to fight at all times; and the males of the capercailzie sometimes hold their Balzen or leks at the usual place of assemblage during the autumn. (34. L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 25.) Hence it is not at all surprising that male birds should continue singing for their own amusement after the season for courtship is over.
As shewn in a previous chapter, singing is to a certain extent an art, and is much improved by practice. Birds can be taught various tunes, and even the unmelodious sparrow has learnt to sing like a linnet. They acquire the song of their foster parents (35. Barrington, ibid. p. 264, Bechstein, ibid. s. 5.), and sometimes that of their neighbours. (36. Dureau de la Malle gives a curious instance (‘Annales des Sc. Nat.’ 3rd series, Zoolog., tom. x. p. 118) of some wild blackbirds in his garden in Paris, which naturally learnt a republican air from a caged bird.) All the common songsters belong to the Order of Insessores, and their vocal organs are much more complex than those of most other birds; yet it is a singular fact that some of the Insessores, such as ravens, crows, and magpies, possess the proper apparatus (37. Bishop, in ‘Todd’s Cyclopaedia of Anatomy and Physiology,’ vol. iv. p. 1496.), though they never sing, and do not naturally modulate their voices to any great extent. Hunter asserts (38. As stated by Barrington in ‘Philosophical Transactions,’ 1773, p. 262.) that with the true songsters the muscles of the larynx are stronger in the males than in the females; but with this slight exception there is no difference in the vocal organs of the two sexes, although the males of most species sing so much better and more continuously than the females.
It is remarkable that only small birds properly sing. The Australian genus Menura, however, must be excepted; for the Menura Alberti, which is about the size of a half-grown turkey, not only mocks other birds, but “its own whistle is exceedingly beautiful and varied.” The males congregate and form “corroborying places,” where they sing, raising and spreading their tails like peacocks, and drooping their wings. (39. Gould, ‘Handbook to the Birds of Australia,’ vol. i. 1865, pp. 308-310. See also Mr. T.W. Wood in the ‘Student,’ April 1870, p. 125.) It is also remarkable that birds which sing well are rarely decorated with brilliant colours or other ornaments. Of our British birds, excepting the bullfinch and goldfinch, the best songsters are plain-coloured. The kingfisher, bee-eater, roller, hoopoe, woodpeckers, etc., utter harsh cries; and the brilliant birds of the tropics are hardly ever songsters. (40. See remarks to this effect in Gould’s ‘Introduction to the Trochilidae,’ 1861, p. 22.) Hence bright colours and the power of song seem to replace each other. We can perceive that if the plumage did not vary in brightness, or if bright colours were dangerous to the species, other means would be employed to charm the females; and melody of voice offers one such means.
[Fig. 39. Tetrao cupido: male. (T.W. Wood.)]
In some birds the vocal organs differ greatly in the two sexes. In the Tetrao cupido (Fig. 39) the male has two bare, orange-coloured sacks, one on each side of the neck; and these are largely inflated when the male, during the breeding-season, makes his curious hollow sound, audible at a great distance. Audubon proved that the sound was intimately connected with this apparatus (which reminds us of the air-sacks on each side of the mouth of certain male frogs), for he found that the sound was much diminished when one of the sacks of a tame bird was pricked, and when both were pricked it was altogether stopped. The female has “a somewhat similar, though smaller naked space of skin on the neck; but this is not capable of inflation.” (41. ‘The Sportsman and Naturalist in Canada,’ by Major W. Ross King, 1866, pp. 144-146. Mr. T.W. Wood gives in the ‘Student’ (April 1870, p. 116) an excellent account of the attitude and habits of this bird during its courtship. He states that the ear-tufts or neck-plumes are erected, so that they meet over the crown of the head. See his drawing, Fig. 39.) The male of another kind of grouse (Tetrao urophasianus), whilst courting the female, has his “bare yellow oesophagus inflated to a prodigious size, fully half as large as the body”; and he then utters various grating, deep, hollow tones. With his neck-feathers erect, his wings lowered, and buzzing on the ground, and his long pointed tail spread out like a fan, he displays a variety of grotesque attitudes. The oesophagus of the female is not in any way remarkable. (42. Richardson, ‘Fauna Bor. American: Birds,’ 1831, p. 359. Audubon, ibid. vol. iv. p. 507.)
[Fig. 40. The Umbrella-bird or Cephalopterus ornatus, male (from Brehm).]
It seems now well made out that the great throat pouch of the European male bustard (Otis tarda), and of at least four other species, does not, as was formerly supposed, serve to hold water, but is connected with the utterance during the breeding-season of a peculiar sound resembling “oak.” (43. The following papers have been lately written on this subject: Prof. A. Newton, in the ‘Ibis,’ 1862, p. 107; Dr. Cullen, ibid. 1865, p. 145; Mr. Flower, in ‘Proc. Zool. Soc.’ 1865, p. 747; and Dr. Murie, in ‘Proc. Zool. Soc.’ 1868, p. 471. In this latter paper an excellent figure is given of the male Australian Bustard in full display with the sack distended. It is a singular fact that the sack is not developed in all the males of the same species.) A crow-like bird inhabiting South America (see Cephalopterus ornatus, Fig. 40) is called the umbrella-bird, from its immense top knot, formed of bare white quills surmounted by dark-blue plumes, which it can elevate into a great dome no less than five inches in diameter, covering the whole head. This bird has on its neck a long, thin, cylindrical fleshy appendage, which is thickly clothed with scale-like blue feathers. It probably serves in part as an ornament, but likewise as a resounding apparatus; for Mr. Bates found that it is connected “with an unusual development of the trachea and vocal organs.” It is dilated when the bird utters its singularly deep, loud and long sustained fluty note. The head- crest and neck-appendage are rudimentary in the female. (44. Bates, ‘The Naturalist on the Amazons,’ 1863, vol. ii. p. 284; Wallace, in ‘Proceedings, Zoological Society,’ 1850, p. 206. A new species, with a still larger neck-appendage (C. penduliger), has lately been discovered, see ‘Ibis,’ vol. i. p. 457.)
The vocal organs of various web-footed and wading birds are extraordinarily complex, and differ to a certain extent in the two sexes. In some cases the trachea is convoluted, like a French horn, and is deeply embedded in the sternum. In the wild swan (Cygnus ferus) it is more deeply embedded in the adult male than in the adult female or young male. In the male Merganser the enlarged portion of the trachea is furnished with an additional pair of muscles. (45. Bishop, in Todd’s ‘Cyclopaedia of Anatomy and Physiology,’ vol. iv. p. 1499.) In one of the ducks, however, namely Anas punctata, the bony enlargement is only a little more developed in the male than in the female. (46. Prof. Newton, ‘Proc. Zoolog. Soc.’ 1871, p. 651.) But the meaning of these differences in the trachea of the two sexes of the Anatidae is not understood; for the male is not always the more vociferous; thus with the common duck, the male hisses, whilst the female utters a loud quack. (47. The spoonbill (Platalea) has its trachea convoluted into a figure of eight, and yet this bird (Jerdon, ‘Birds of India,’ vol. iii. p. 763) is mute; but Mr. Blyth informs me that the convolutions are not constantly present, so that perhaps they are now tending towards abortion.) In both sexes of one of the cranes (Grus virgo) the trachea penetrates the sternum, but presents “certain sexual modifications.” In the male of the black stork there is also a well-marked sexual difference in the length and curvature of the bronchi. (48. ‘Elements of Comparative Anatomy,’ by R. Wagner, Eng. translat. 1845, p. 111. With respect to the swan, as given above, Yarrell’s ‘History of British Birds,’ 2nd edition, 1845, vol. iii. p. 193.) Highly important structures have, therefore, in these cases been modified according to sex.
It is often difficult to conjecture whether the many strange cries and notes uttered by male birds during the breeding-season serve as a charm or merely as a call to the female. The soft cooing of the turtle-dove and of many pigeons, it may be presumed, pleases the female. When the female of the wild turkey utters her call in the morning, the male answers by a note which differs from the gobbling noise made, when with erected feathers, rustling wings and distended wattles, he puffs and struts before her. (49. C.L. Bonaparte, quoted in the ‘Naturalist Library: Birds,’ vol. xiv. p. 126.) The spel of the black-cock certainly serves as a call to the female, for it has been known to bring four or five females from a distance to a male under confinement; but as the black-cock continues his spel for hours during successive days, and in the case of the capercailzie “with an agony of passion,” we are led to suppose that the females which are present are thus charmed. (50. L. Lloyd, ‘The Game Birds of Sweden,’ etc., 1867, pp. 22, 81.) The voice of the common rook is known to alter during the breeding-season, and is therefore in some way sexual. (51. Jenner, ‘Philosophical Transactions,’ 1824, p. 20.) But what shall we say about the harsh screams of, for instance, some kinds of macaws; have these birds as bad taste for musical sounds as they apparently have for colour, judging by the inharmonious contrast of their bright yellow and blue plumage? It is indeed possible that without any advantage being thus gained, the loud voices of many male birds may be the result of the inherited effects of the continued use of their vocal organs when excited by the strong passions of love, jealousy and rage; but to this point we shall recur when we treat of quadrupeds.
We have as yet spoken only of the voice, but the males of various birds practise, during their courtship, what may be called instrumental music. Peacocks and Birds of Paradise rattle their quills together. Turkey-cocks scrape their wings against the ground, and some kinds of grouse thus produce a buzzing sound. Another North American grouse, the Tetrao umbellus, when with his tail erect, his ruffs displayed, “he shows off his finery to the females, who lie hid in the neighbourhood,” drums by rapidly striking his wings together above his back, according to Mr. R. Haymond, and not, as Audubon thought, by striking them against his sides. The sound thus produced is compared by some to distant thunder, and by others to the quick roll of a drum. The female never drums, “but flies directly to the place where the male is thus engaged.” The male of the Kalij-pheasant, in the Himalayas, often makes a singular drumming noise with his wings, not unlike the sound produced by shaking a stiff piece of cloth.” On the west coast of Africa the little black-weavers (Ploceus?) congregate in a small party on the bushes round a small open space, and sing and glide through the air with quivering wings, “which make a rapid whirring sound like a child’s rattle.” One bird after another thus performs for hours together, but only during the courting-season. At this season, and at no other time, the males of certain night-jars (Caprimulgus) make a strange booming noise with their wings. The various species of woodpeckers strike a sonorous branch with their beaks, with so rapid a vibratory movement that “the head appears to be in two places at once.” The sound thus produced is audible at a considerable distance but cannot be described; and I feel sure that its source would never be conjectured by any one hearing it for the first time. As this jarring sound is made chiefly during the breeding-season, it has been considered as a love-song; but it is perhaps more strictly a love- call. The female, when driven from her nest, has been observed thus to call her mate, who answered in the same manner and soon appeared. Lastly, the male hoopoe (Upupa epops) combines vocal and instrumental music; for during the breeding-season this bird, as Mr. Swinhoe observed, first draws in air, and then taps the end of its beak perpendicularly down against a stone or the trunk of a tree, “when the breath being forced down the tubular bill produces the correct sound.” If the beak is not thus struck against some object, the sound is quite different. Air is at the same time swallowed, and the oesophagus thus becomes much swollen; and this probably acts as a resonator, not only with the hoopoe, but with pigeons and other birds. (52. For the foregoing facts see, on Birds of Paradise, Brehm, ‘Thierleben,’ Band iii. s. 325. On Grouse, Richardson, ‘Fauna Bor. Americ.: Birds,’ pp. 343 and 359; Major W. Ross King, ‘The Sportsman in Canada,’ 1866, p. 156; Mr. Haymond, in Prof. Cox’s ‘Geol. Survey of Indiana,’ p. 227; Audubon, ‘American Ornitholog. Biograph.’ vol. i. p. 216. On the Kalij-pheasant, Jerdon, ‘Birds of India,’ vol. iii. p. 533. On the Weavers, Livingstone’s ‘Expedition to the Zambesi,’ 1865, p. 425. On Woodpeckers, Macgillivray, ‘Hist. of British Birds,’ vol. iii. 1840, pp. 84, 88, 89, and 95. On the Hoopoe, Mr. Swinhoe, in ‘Proc. Zoolog. Soc.’ June 23, 1863 and 1871, p. 348. On the Night-jar, Audubon, ibid. vol. ii. p. 255, and ‘American Naturalist,’ 1873, p. 672. The English Night-jar likewise makes in the spring a curious noise during its rapid flight.)
[Fig. 41. Outer tail-feather of Scolopax gallinago (from ‘Proc. Zool. Soc.’ 1858).
Fig. 42. Outer tail-feather of Scolopax frenata.
Fig. 43. Outer tail-feather of Scolopax javensis.]
In the foregoing cases sounds are made by the aid of structures already present and otherwise necessary; but in the following cases certain feathers have been specially modified for the express purpose of producing sounds. The drumming, bleating, neighing, or thundering noise (as expressed by different observers) made by the common snipe (Scolopax gallinago) must have surprised every one who has ever heard it. This bird, during the pairing-season, flies to “perhaps a thousand feet in height,” and after zig-zagging about for a time descends to the earth in a curved line, with outspread tail and quivering pinions, and surprising velocity. The sound is emitted only during this rapid descent. No one was able to explain the cause until M. Meves observed that on each side of the tail the outer feathers are peculiarly formed (Fig. 41), having a stiff sabre-shaped shaft with the oblique barbs of unusual length, the outer webs being strongly bound together. He found that by blowing on these feathers, or by fastening them to a long thin stick and waving them rapidly through the air, he could reproduce the drumming noise made by the living bird. Both sexes are furnished with these feathers, but they are generally larger in the male than in the female, and emit a deeper note. In some species, as in S. frenata (Fig. 42), four feathers, and in S. javensis (Fig. 43), no less than eight on each side of the tail are greatly modified. Different tones are emitted by the feathers of the different species when waved through the air; and the Scolopax Wilsonii of the United States makes a switching noise whilst descending rapidly to the earth. (53. See M. Meves’ interesting paper in ‘Proc. Zool. Soc.’ 1858, p. 199. For the habits of the snipe, Macgillivray, ‘History of British Birds,’ vol. iv. p. 371. For the American snipe, Capt. Blakiston, ‘Ibis,’ vol. v. 1863, p. 131.)
[Fig. 44. Primary wing-feather of a Humming-bird, the Selasphorus platycercus (from a sketch by Mr. Salvin). Upper figure, that of male;
lower figure, corresponding feather of female.]
In the male of the Chamaepetes unicolor (a large gallinaceous bird of America), the first primary wing-feather is arched towards the tip and is much more attenuated than in the female. In an allied bird, the Penelope nigra, Mr. Salvin observed a male, which, whilst it flew downwards “with outstretched wings, gave forth a kind of crashing rushing noise,” like the falling of a tree. (54. Mr. Salvin, in ‘Proceedings, Zoological Society,’ 1867, p. 160. I am much indebted to this distinguished ornithologist for sketches of the feathers of the Chamaepetes, and for other information.) The male alone of one of the Indian bustards (Sypheotides auritus) has its primary wing-feathers greatly acuminated; and the male of an allied species is known to make a humming noise whilst courting the female. (55. Jerdon, ‘Birds of India,’ vol. iii. pp. 618, 621.) In a widely different group of birds, namely Humming-birds, the males alone of certain kinds have either the shafts of their primary wing-feathers broadly dilated, or the webs abruptly excised towards the extremity. The male, for instance, of Selasphorus platycercus, when adult, has the first primary wing-feather (Fig. 44), thus excised. Whilst flying from flower to flower he makes “a shrill, almost whistling noise” (56. Gould, ‘Introduction to the Trochilidae,’ 1861, p. 49. Salvin, ‘Proceedings, Zoological Society,’ 1867, p. 160.); but it did not appear to Mr. Salvin that the noise was intentionally made.
[Fig. 45. Secondary wing-feathers of Pipra deliciosa (from Mr. Sclater, in ‘Proc. Zool. Soc.’ 1860).
The three upper feathers, a, b, c, from the male; the three lower corresponding feathers, d, e, f, from the female. a and d, fifth secondary wing-feather of male and female, upper surface. b and e, sixth secondary, upper surface. c and f, seventh secondary, lower surface.]
Lastly, in several species of a sub-genus of Pipra or Manakin, the males, as described by Mr. Sclater, have their SECONDARY wing-feathers modified in a still more remarkable manner. In the brilliantly-coloured P. deliciosa the first three secondaries are thick-stemmed and curved towards the body; in the fourth and fifth (Fig. 45, a) the change is greater; and in the sixth and seventh (b, c) the shaft “is thickened to an extraordinary degree, forming a solid horny lump.” The barbs also are greatly changed in shape, in comparison with the corresponding feathers (d, e, f) in the female. Even the bones of the wing, which support these singular feathers in the male, are said by Mr. Fraser to be much thickened. These little birds make an extraordinary noise, the first “sharp note being not unlike the crack of a whip.” (57. Sclater, in ‘Proceedings, Zoological Society,’ 1860, p. 90, and in ‘Ibis,’ vol. iv. 1862, p. 175. Also Salvin, in ‘Ibis,’ 1860, p. 37.)
The diversity of the sounds, both vocal and instrumental, made by the males of many birds during the breeding-season, and the diversity of the means for producing such sounds, are highly remarkable. We thus gain a high idea of their importance for sexual purposes, and are reminded of the conclusion arrived at as to insects. It is not difficult to imagine the steps by which the notes of a bird, primarily used as a mere call or for some other purpose, might have been improved into a melodious love song. In the case of the modified feathers, by which the drumming, whistling, or roaring noises are produced, we know that some birds during their courtship flutter, shake, or rattle their unmodified feathers together; and if the females were led to select the best performers, the males which possessed the strongest or thickest, or most attenuated feathers, situated on any part of the body, would be the most successful; and thus by slow degrees the feathers might be modified to almost any extent. The females, of course, would not notice each slight successive alteration in shape, but only the sounds thus produced. It is a curious fact that in the same class of animals, sounds so different as the drumming of the snipe’s tail, the tapping of the woodpecker’s beak, the harsh trumpet-like cry of certain water-fowl, the cooing of the turtle-dove, and the song of the nightingale, should all be pleasing to the females of the several species. But we must not judge of the tastes of distinct species by a uniform standard; nor must we judge by the standard of man’s taste. Even with man, we should remember what discordant noises, the beating of tom-toms and the shrill notes of reeds, please the ears of savages. Sir S. Baker remarks (58. ‘The Nile Tributaries of Abyssinia,’ 1867, p. 203.), that “as the stomach of the Arab prefers the raw meat and reeking liver taken hot from the animal, so does his ear prefer his equally coarse and discordant music to all other.”
LOVE ANTICS AND DANCES.
The curious love gestures of some birds have already been incidentally noticed; so that little need here be added. In Northern America large numbers of a grouse, the Tetrao phasianellus, meet every morning during the breeding-season on a selected level spot, and here they run round and round in a circle of about fifteen or twenty feet in diameter, so that the ground is worn quite bare, like a fairy-ring. In these Partridge-dances, as they are called by the hunters, the birds assume the strangest attitudes, and run round, some to the left and some to the right. Audubon describes the males of a heron (Ardea herodias) as walking about on their long legs with great dignity before the females, bidding defiance to their rivals. With one of the disgusting carrion-vultures (Cathartes jota) the same naturalist states that “the gesticulations and parade of the males at the beginning of the love-season are extremely ludicrous.” Certain birds perform their love-antics on the wing, as we have seen with the black African weaver, instead of on the ground. During the spring our little white-throat (Sylvia cinerea) often rises a few feet or yards in the air above some bush, and “flutters with a fitful and fantastic motion, singing all the while, and then drops to its perch.” The great English bustard throws himself into indescribably odd attitudes whilst courting the female, as has been figured by Wolf. An allied Indian bustard (Otis bengalensis) at such times “rises perpendicularly into the air with a hurried flapping of his wings, raising his crest and puffing out the feathers of his neck and breast, and then drops to the ground;” he repeats this manoeuvre several times, at the same time humming in a peculiar tone. Such females as happen to be near “obey this saltatory summons,” and when they approach he trails his wings and spreads his tail like a turkey-cock. (59. For Tetrao phasianellus, see Richardson, ‘Fauna, Bor. America,’ p. 361, and for further particulars Capt. Blakiston, ‘Ibis,’ 1863, p. 125. For the Cathartes and Ardea, Audubon, ‘Ornithological Biography,’ vol. ii. p. 51, and vol. iii. p. 89. On the White-throat, Macgillivray, ‘History of British Birds,’ vol. ii. p. 354. On the Indian Bustard, Jerdon, ‘Birds of India,’ vol. iii. p. 618.)
[Fig. 46. Bower-bird, Chlamydera maculata, with bower (from Brehm).]
But the most curious case is afforded by three allied genera of Australian birds, the famous Bower-birds,–no doubt the co-descendants of some ancient species which first acquired the strange instinct of constructing bowers for performing their love-antics. The bowers (Fig. 46), which, as we shall hereafter see, are decorated with feathers, shells, bones, and leaves, are built on the ground for the sole purpose of courtship, for their nests are formed in trees. Both sexes assist in the erection of the bowers, but the male is the principal workman. So strong is this instinct that it is practised under confinement, and Mr. Strange has described (60. Gould, ‘Handbook to the Birds of Australia,’ vol. i. pp. 444, 449, 455. The bower of the Satin Bower-bird may be seen in the Zoological Society’s Gardens, Regent’s Park.) the habits of some Satin Bower-birds which he kept in an aviary in New South Wales. “At times the male will chase the female all over the aviary, then go to the bower, pick up a gay feather or a large leaf, utter a curious kind of note, set all his feathers erect, run round the bower and become so excited that his eyes appear ready to start from his bead; he continues opening first one wing then the other, uttering a low, whistling note, and, like the domestic cock, seems to be picking up something from the ground, until at last the female goes gently towards him.” Captain Stokes has described the habits and “play-houses” of another species, the Great Bower-bird, which was seen “amusing itself by flying backwards and forwards, taking a shell alternately from each side, and carrying it through the archway in its mouth.” These curious structures, formed solely as halls of assemblage, where both sexes amuse themselves and pay their court, must cost the birds much labour. The bower, for instance, of the Fawn-breasted species, is nearly four feet in length, eighteen inches in height, and is raised on a thick platform of sticks.
DECORATION.
I will first discuss the cases in which the males are ornamented either exclusively or in a much higher degree than the females, and in a succeeding chapter those in which both sexes are equally ornamented, and finally the rare cases in which the female is somewhat more brightly- coloured than the male. As with the artificial ornaments used by savage and civilised men, so with the natural ornaments of birds, the head is the chief seat of decoration. (61. See remarks to this effect, on the ‘Feeling of Beauty among Animals,’ by Mr. J. Shaw, in the ‘Athenaeum,’ Nov. 24th, 1866, p. 681.) The ornaments, as mentioned at the commencement of this chapter, are wonderfully diversified. The plumes on the front or back of the head consist of variously-shaped feathers, sometimes capable of erection or expansion, by which their beautiful colours are fully displayed. Elegant ear-tufts (Fig. 39) are occasionally present. The head is sometimes covered with velvety down, as with the pheasant; or is naked and vividly coloured. The throat, also, is sometimes ornamented with a beard, wattles, or caruncles. Such appendages are generally brightly- coloured, and no doubt serve as ornaments, though not always ornamental in our eyes; for whilst the male is in the act of courting the female, they often swell and assume vivid tints, as in the male turkey. At such times the fleshy appendages about the head of the male Tragopan pheasant (Ceriornis Temminckii) swell into a large lappet on the throat and into two horns, one on each side of the splendid top-knot; and these are then coloured of the most intense blue which I have ever beheld. (62. See Dr. Murie’s account with coloured figures in ‘Proceedings, Zoological Society,’ 1872, p. 730.) The African hornbill (Bucorax abyssinicus) inflates the scarlet bladder-like wattle on its neck, and with its wings drooping and tail expanded “makes quite a grand appearance.” (63. Mr. Monteiro, ‘Ibis,’ vol. iv. 1862, p. 339.) Even the iris of the eye is sometimes more brightly-coloured in the male than in the female; and this is frequently the case with the beak, for instance, in our common blackbird. In Buceros corrugatus, the whole beak and immense casque are coloured more conspicuously in the male than in the female; and “the oblique grooves upon the sides of the lower mandible are peculiar to the male sex.” (64. ‘Land and Water,’ 1868, p. 217.)
The head, again, often supports fleshy appendages, filaments, and solid protuberances. These, if not common to both sexes, are always confined to the males. The solid protuberances have been described in detail by Dr. W. Marshall (65. ‘Ueber die Schadelhocker,’ etc., ‘Niederland. Archiv. fur Zoologie,’ B. I. Heft 2, 1872.), who shews that they are formed either of cancellated bone coated with skin, or of dermal and other tissues. With mammals true horns are always supported on the frontal bones, but with birds various bones have been modified for this purpose; and in species of the same group the protuberances may have cores of bone, or be quite destitute of them, with intermediate gradations connecting these two extremes. Hence, as Dr. Marshall justly remarks, variations of the most different kinds have served for the development through sexual selection of these ornamental appendages. Elongated feathers or plumes spring from almost every part of the body. The feathers on the throat and breast are sometimes developed into beautiful ruffs and collars. The tail-feathers are frequently increased in length; as we see in the tail-coverts of the peacock, and in the tail itself of the Argus pheasant. With the peacock even the bones of the tail have been modified to support the heavy tail- coverts. (66. Dr. W. Marshall, ‘Uber den Vogelschwanz,’ ibid. B. I. Heft 2, 1872.) The body of the Argus is not larger than that of a fowl; yet the length from the end of the beak to the extremity of the tail is no less than five feet three inches (67. Jardine’s ‘Naturalist Library: Birds,’ vol. xiv. p. 166.), and that of the beautifully ocellated secondary wing- feathers nearly three feet. In a small African night-jar (Cosmetornis vexillarius) one of the primary wing-feathers, during the breeding-season, attains a length of twenty-six inches, whilst the bird itself is only ten inches in length. In another closely-allied genus of night-jars, the shafts of the elongated wing-feathers are naked, except at the extremity, where there is a disc. (68. Sclater, in the ‘Ibis,’ vol. vi. 1864, p. 114; Livingstone, ‘Expedition to the Zambesi,’ 1865, p. 66.) Again, in another genus of night-jars, the tail-feathers are even still more prodigiously developed. In general the feathers of the tail are more often elongated than those of the wings, as any great elongation of the latter impedes flight. We thus see that in closely-allied birds ornaments of the same kind have been gained by the males through the development of widely different feathers.
It is a curious fact that the feathers of species belonging to very distinct groups have been modified in almost exactly the same peculiar manner. Thus the wing-feathers in one of the above-mentioned night-jars are bare along the shaft, and terminate in a disc; or are, as they are sometimes called, spoon or racket-shaped. Feathers of this kind occur in the tail of a motmot (Eumomota superciliaris), of a king-fisher, finch, humming-bird, parrot, several Indian drongos (Dicrurus and Edolius, in one of which the disc stands vertically), and in the tail of certain birds of paradise. In these latter birds, similar feathers, beautifully ocellated, ornament the head, as is likewise the case with some gallinaceous birds. In an Indian bustard (Sypheotides auritus) the feathers forming the ear- tufts, which are about four inches in length, also terminate in discs. (69. Jerdon, ‘Birds of India,’ vol. iii. p. 620.) It is a most singular fact that the motmots, as Mr. Salvin has clearly shewn (70. ‘Proceedings, Zoological Society,’ 1873, p. 429.), give to their tail feathers the racket-shape by biting off the barbs, and, further, that this continued mutilation has produced a certain amount of inherited effect.
[Fig. 47. Paradisea Papuana (T.W. Wood).]
Again, the barbs of the feathers in various widely-distinct birds are filamentous or plumose, as with some herons, ibises, birds of paradise, and Gallinaceae. In other cases the barbs disappear, leaving the shafts bare from end to end; and these in the tail of the Paradisea apoda attain a length of thirty-four inches (71. Wallace, in ‘Annals and Magazine of Natural History,’ vol. xx. 1857, p. 416, and in his ‘Malay Archipelago,’ vol. ii. 1869, p. 390.): in P. Papuana (Fig. 47) they are much shorter and thin. Smaller feathers when thus denuded appear like bristles, as on the breast of the turkey-cock. As any fleeting fashion in dress comes to be admired by man, so with birds a change of almost any kind in the structure or colouring of the feathers in the male appears to have been admired by the female. The fact of the feathers in widely distinct groups having been modified in an analogous manner no doubt depends primarily on all the feathers having nearly the same structure and manner of development, and consequently tending to vary in the same manner. We often see a tendency to analogous variability in the plumage of our domestic breeds belonging to distinct species. Thus top-knots have appeared in several species. In an extinct variety of the turkey, the top-knot consisted of bare quills surmounted with plumes of down, so that they somewhat resembled the racket- shaped feathers above described. In certain breeds of the pigeon and fowl the feathers are plumose, with some tendency in the shafts to be naked. In the Sebastopol goose the scapular feathers are greatly elongated, curled, or even spirally twisted, with the margins plumose. (72. See my work on ‘The Variation of Animals and Plants under Domestication,’ vol. i. pp. 289, 293.)
In regard to colour, hardly anything need here be said, for every one knows how splendid are the tints of many birds, and how harmoniously they are combined. The colours are often metallic and iridescent. Circular spots are sometimes surrounded by one or more differently shaded zones, and are thus converted into ocelli. Nor need much be said on the wonderful difference between the sexes of many birds. The common peacock offers a striking instance. Female birds of paradise are obscurely coloured and destitute of all ornaments, whilst the males are probably the most highly decorated of all birds, and in so many different ways that they must be seen to be appreciated. The elongated and golden-orange plumes which spring from beneath the wings of the Paradisea apoda, when vertically erected and made to vibrate, are described as forming a sort of halo, in the centre of which the head “looks like a little emerald sun with its rays formed by the two plumes.” (73. Quoted from M. de Lafresnaye in ‘Annals and Mag. of Natural History,’ vol. xiii. 1854, p. 157: see also Mr. Wallace’s much fuller account in vol. xx. 1857, p. 412, and in his ‘Malay Archipelago.’S) In another most beautiful species the head is bald, “and of a rich cobalt blue, crossed by several lines of black velvety feathers.” (74. Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 405.)
[Fig. 48. Lophornis ornatus, male and female (from Brehm).
Fig. 49. Spathura underwoodi, male and female (from Brehm).]
Male humming-birds (Figs. 48 and 49) almost vie with birds of paradise in their beauty, as every one will admit who has seen Mr. Gould’s splendid volumes, or his rich collection. It is very remarkable in how many different ways these birds are ornamented. Almost every part of their plumage has been taken advantage of, and modified; and the modifications have been carried, as Mr. Gould shewed me, to a wonderful extreme in some species belonging to nearly every sub-group. Such cases are curiously like those which we see in our fancy breeds, reared by man for the sake of ornament; certain individuals originally varied in one character, and other individuals of the same species in other characters; and these have been seized on by man and much augmented–as shewn by the tail of the fantail- pigeon, the hood of the jacobin, the beak and wattle of the carrier, and so forth. The sole difference between these cases is that in the one, the result is due to man’s selection, whilst in the other, as with humming- birds, birds of paradise, etc., it is due to the selection by the females of the more beautiful males.
I will mention only one other bird, remarkable from the extreme contrast in colour between the sexes, namely the famous bell-bird (Chasmorhynchus niveus) of S. America, the note of which can be distinguished at the distance of nearly three miles, and astonishes every one when first hearing it. The male is pure white, whilst the female is dusky-green; and white is a very rare colour in terrestrial species of moderate size and inoffensive habits. The male, also, as described by Waterton, has a spiral tube, nearly three inches in length, which rises from the base of the beak. It is jet-black, dotted over with minute downy feathers. This tube can be inflated with air, through a communication with the palate; and when not inflated hangs down on one side. The genus consists of four species, the males of which are very distinct, whilst the females, as described by Mr. Sclater in a very interesting paper, closely resemble each other, thus offering an excellent instance of the common rule that within the same group the males differ much more from each other than do the females. In a second species (C. nudicollis) the male is likewise snow-white, with the exception of a large space of naked skin on the throat and round the eyes, which during the breeding-season is of a fine green colour. In a third species (C. tricarunculatus) the head and neck alone of the male are white, the rest of the body being chestnut-brown, and the male of this species is provided with three filamentous projections half as long as the body–one rising from the base of the beak, and the two others from the corners of the mouth. (75. Mr. Sclater, ‘Intellectual Observer,’ Jan. 1867. Waterton’s ‘Wanderings,’ p. 118. See also Mr. Salvin’s interesting paper, with a plate, in the ‘Ibis,’ 1865, p. 90.)
The coloured plumage and certain other ornaments of the adult males are either retained for life, or are periodically renewed during the summer and breeding-season. At this same season the beak and naked skin about the head frequently change colour, as with some herons, ibises, gulls, one of the bell-birds just noticed, etc. In the white ibis, the cheeks, the inflatable skin of the throat, and the basal portion of the beak then become crimson. (76. ‘Land and Water,’ 1867, p. 394.) In one of the rails, Gallicrex cristatus, a large red caruncle is developed during this period on the head of the male. So it is with a thin horny crest on the beak of one of the pelicans, P. erythrorhynchus; for, after the breeding- season, these horny crests are shed, like horns from the heads of stags, and the shore of an island in a lake in Nevada was found covered with these curious exuviae. (77. Mr. D.G. Elliot, in ‘Proc. Zool. Soc.’ 1869, p. 589.)
Changes of colour in the plumage according to the season depend, firstly on a double annual moult, secondly on an actual change of colour in the feathers themselves, and thirdly on their dull-coloured margins being periodically shed, or on these three processes more or less combined. The shedding of the deciduary margins may be compared with the shedding of their down by very young birds; for the down in most cases arises from the summits of the first true feathers. (78. Nitzsch’s ‘Pterylography,’ edited by P.L. Sclater, Ray Society, 1867, p. 14.)
With respect to the birds which annually undergo a double moult, there are, firstly, some kinds, for instance snipes, swallow-plovers (Glareolae), and curlews, in which the two sexes resemble each other, and do not change colour at any season. I do not know whether the winter plumage is thicker and warmer than the summer plumage, but warmth seems the most probable end attained of a double moult, where there is no change of colour. Secondly, there are birds, for instance, certain species of Totanus and other Grallatores, the sexes of which resemble each other, but in which the summer and winter plumage differ slightly in colour. The difference, however, in these cases is so small that it can hardly be an advantage to them; and it may, perhaps, be attributed to the direct action of the different conditions to which the birds are exposed during the two seasons. Thirdly, there are many other birds the sexes of which are alike, but which are widely different in their summer and winter plumage. Fourthly, there are birds the sexes of which differ from each other in colour; but the females, though moulting twice, retain the same colours throughout the year, whilst the males undergo a change of colour, sometimes a great one, as with certain bustards. Fifthly and lastly, there are birds the sexes of which differ from each other in both their summer and winter plumage; but the male undergoes a greater amount of change at each recurrent season than the female–of which the ruff (Machetes pugnax) offers a good instance.
With respect to the cause or purpose of the differences in colour between the summer and winter plumage, this may in some instances, as with the ptarmigan (79. The brown mottled summer plumage of the ptarmigan is of as much importance to it, as a protection, as the white winter plumage; for in Scandinavia during the spring, when the snow has disappeared, this bird is known to suffer greatly from birds of prey, before it has acquired its summer dress: see Wilhelm von Wright, in Lloyd, ‘Game Birds of Sweden,’ 1867, p. 125.), serve during both seasons as a protection. When the difference between the two plumages is slight it may perhaps be attributed, as already remarked, to the direct action of the conditions of life. But with many birds there can hardly be a doubt that the summer plumage is ornamental, even when both sexes are alike. We may conclude that this is the case with many herons, egrets, etc., for they acquire their beautiful plumes only during the breeding-season. Moreover, such plumes, top-knots, etc., though possessed by both sexes, are occasionally a little more developed in the male than in the female; and they resemble the plumes and ornaments possessed by the males alone of other birds. It is also known that confinement, by affecting the reproductive system of male birds, frequently checks the development of their secondary sexual characters, but has no immediate influence on any other characters; and I am informed by Mr. Bartlett that eight or nine specimens of the Knot (Tringa canutus) retained their unadorned winter plumage in the Zoological Gardens throughout the year, from which fact we may infer that the summer plumage, though common to both sexes, partakes of the nature of the exclusively masculine plumage of many other birds. (80. In regard to the previous statements on moulting, see, on snipes, etc., Macgillivray, ‘Hist. Brit. Birds,’ vol. iv. p. 371; on Glareolae, curlews, and bustards, Jerdon, ‘Birds of India,’ vol. iii. pp. 615, 630, 683; on Totanus, ibid. p. 700; on the plumes of herons, ibid. p. 738, and Macgillivray, vol. iv. pp. 435 and 444, and Mr. Stafford Allen, in the ‘Ibis,’ vol. v. 1863, p. 33.)
From the foregoing facts, more especially from neither sex of certain birds changing colour during either annual moult, or changing so slightly that the change can hardly be of any service to them, and from the females of other species moulting twice yet retaining the same colours throughout the year, we may conclude that the habit of annually moulting twice has not been acquired in order that the male should assume an ornamental character during the breeding-season; but that the double moult, having been originally acquired for some distinct purpose, has subsequently been taken advantage of in certain cases for gaining a nuptial plumage.
It appears at first sight a surprising circumstance that some closely- allied species should regularly undergo a double annual moult, and others only a single one. The ptarmigan, for instance, moults twice or even thrice in the year, and the blackcock only once: some of the splendidly coloured honey-suckers (Nectariniae) of India and some sub-genera of obscurely coloured pipits (Anthus) have a double, whilst others have only a single annual moult. (81. On the moulting of the ptarmigan, see Gould’s ‘Birds of Great Britain.’ On the honey-suckers, Jerdon, ‘Birds of India,’ vol. i. pp. 359, 365, 369. On the moulting of Anthus, see Blyth, in ‘Ibis,’ 1867, p. 32.) But the gradations in the manner of moulting, which are known to occur with various birds, shew us how species, or whole groups, might have originally acquired their double annual moult, or having once gained the habit, have again lost it. With certain bustards and plovers the vernal moult is far from complete, some feathers being renewed, and some changed in colour. There is also reason to believe that with certain bustards and rail-like birds, which properly undergo a double moult, some of the older males retain their nuptial plumage throughout the year. A few highly modified feathers may merely be added during the spring to the plumage, as occurs with the disc-formed tail-feathers of certain drongos (Bhringa) in India, and with the elongated feathers on the back, neck, and crest of certain herons. By such steps as these, the vernal moult might be rendered more and more complete, until a perfect double moult was acquired. Some of the birds of paradise retain their nuptial feathers throughout the year, and thus have only a single moult; others cast them directly after the breeding-season, and thus have a double moult; and others again cast them at this season during the first year, but not afterwards; so that these latter species are intermediate in their manner of moulting. There is also a great difference with many birds in the length of time during which the two annual plumages are retained; so that the one might come to be retained for the whole year, and the other completely lost. Thus in the spring Machetes pugnax retains his ruff for barely two months. In Natal the male widow-bird (Chera progne) acquires his fine plumage and long tail-feathers in December or January, and loses them in March; so that they are retained only for about three months. Most species, which undergo a double moult, keep their ornamental feathers for about six months. The male, however, of the wild Gallus bankiva retains his neck-hackles for nine or ten months; and when these are cast off, the underlying black feathers on the neck are fully exposed to view. But with the domesticated descendant of this species, the neck-hackles of the male are immediately replaced by new ones; so that we here see, as to part of the plumage, a double moult changed under domestication into a single moult. (82. For the foregoing statements in regard to partial moults, and on old males retaining their nuptial plumage, see Jerdon, on bustards and plovers, in ‘Birds of India,’ vol. iii. pp. 617, 637, 709, 711. Also Blyth in ‘Land and Water,’ 1867, p. 84. On the moulting of Paradisea, see an interesting article by Dr. W. Marshall, ‘Archives Neerlandaises,’ tom. vi. 1871. On the Vidua, ‘Ibis,’ vol. iii. 1861, p. 133. On the Drongo- shrikes, Jerdon, ibid. vol. i. p. 435. On the vernal moult of the Herodias bubulcus, Mr. S.S. Allen, in ‘Ibis,’ 1863, p. 33. On Gallus bankiva, Blyth, in ‘Annals and Mag. of Natural History,’ vol. i. 1848, p. 455; see, also, on this subject, my ‘Variation of Animals under Domestication,’ vol. i. p. 236.)
The common drake (Anas boschas), after the breeding-season, is well known to lose his male plumage for a period of three months, during which time he assumes that of the female. The male pin-tail duck (Anas acuta) loses his plumage for the shorter period of six weeks or two months; and Montagu remarks that “this double moult within so short a time is a most extraordinary circumstance, that seems to bid defiance to all human reasoning.” But the believer in the gradual modification of species will be far from feeling surprise at finding gradations of all kinds. If the male pin-tail were to acquire his new plumage within a still shorter period, the new male feathers would almost necessarily be mingled with the old, and both with some proper to the female; and this apparently is the case with the male of a not distantly-allied bird, namely the Merganser serrator, for the males are said to “undergo a change of plumage, which assimilates them in some measure to the female.” By a little further acceleration in the process, the double moult would be completely lost. (83. See Macgillivray, ‘Hist. British Birds’ (vol. v. pp. 34, 70, and 223), on the moulting of the Anatidae, with quotations from Waterton and Montagu. Also Yarrell, ‘History of British Birds,’ vol. iii. p. 243.)
Some male birds, as before stated, become more brightly coloured in the spring, not by a vernal moult, but either by an actual change of colour in the feathers, or by their obscurely-coloured deciduary margins being shed. Changes of colour thus caused may last for a longer or shorter time. In the Pelecanus onocrotalus a beautiful rosy tint, with lemon-coloured marks on the breast, overspreads the whole plumage in the spring; but these tints, as Mr. Sclater states, “do not last long, disappearing generally in about six weeks or two months after they have been attained.” Certain finches shed the margins of their feathers in the spring, and then become brighter coloured, while other finches undergo no such change. Thus the Fringilla tristis of the United States (as well as many other American species) exhibits its bright colours only when the winter is past, whilst our goldfinch, which exactly represents this bird in habits, and our siskin, which represents it still more closely in structure, undergo no such annual change. But a difference of this kind in the plumage of allied species is not surprising, for with the common linnet, which belongs to the same family, the crimson forehead and breast are displayed only during the summer in England, whilst in Madeira these colours are retained throughout the year. (84. On the pelican, see Sclater, in ‘Proc. Zool. Soc.’ 1868, p. 265. On the American finches, see Audubon, ‘Ornithological Biography,’ vol. i. pp. 174, 221, and Jerdon, ‘Birds of India,’ vol. ii. p. 383. On the Fringilla cannabina of Madeira, Mr. E. Vernon Harcourt, ‘Ibis,’ vol. v. 1863, p. 230.)
DISPLAY BY MALE BIRDS OF THEIR PLUMAGE.
Ornaments of all kinds, whether permanently or temporarily gained, are sedulously displayed by the males, and apparently serve to excite, attract, or fascinate the females. But the males will sometimes display their ornaments, when not in the presence of the females, as occasionally occurs with grouse at their balz-places, and as may be noticed with the peacock; this latter bird, however, evidently wishes for a spectator of some kind, and, as I have often seen, will shew off his finery before poultry, or even pigs. (85. See also ‘Ornamental Poultry,’ by Rev. E.S. Dixon, 1848, p. 8.) All naturalists who have closely attended to the habits of birds,