The Descent of Man, and Selection in Relation to Sex

Of CATTLE I have received returns from nine gentlemen of 982 births, too few to be trusted; these consisted of 477 bull-calves and 505 cow-calves; i.e., in the proportion of 94.4 males to 100 females. The Rev. W.D. Fox informs me that in 1867 out of 34 calves born on a farm in Derbyshire only one was a bull. Mr. Harrison Weir has enquired from several breeders of PIGS, and most of them estimate the male to the female births as about 7 to 6. This same gentleman has bred RABBITS for many years, and has noticed that a far greater number of bucks are produced than does. But estimations are of little value.

Of mammalia in a state of nature I have been able to learn very little. In regard to the common rat, I have received conflicting statements. Mr. R. Elliot, of Laighwood, informs me that a rat-catcher assured him that he had always found the males in great excess, even with the young in the nest. In consequence of this, Mr. Elliot himself subsequently examined some hundred old ones, and found the statement true. Mr. F. Buckland has bred a large number of white rats, and he also believes that the males greatly exceed the females. In regard to Moles, it is said that "the males are much more numerous than the females" (60. Bell, 'History of British Quadrupeds,' p. 100.): and as the catching of these animals is a special occupation, the statement may perhaps be trusted. Sir A. Smith, in describing an antelope of S. Africa (61. 'Illustrations of the Zoology of S. Africa,' 1849, pl. 29.) (Kobus ellipsiprymnus), remarks, that in the herds of this and other species, the males are few in number compared with the females: the natives believe that they are born in this proportion; others believe that the younger males are expelled from the herds, and Sir A. Smith says, that though he has himself never seen herds consisting of young males alone, others affirm that this does occur. It appears probable that the young when expelled from the herd, would often fall a prey to the many beasts of prey of the country.

BIRDS

With respect to the FOWL, I have received only one account, namely, that out of 1001 chickens of a highly-bred stock of Cochins, reared during eight years by Mr. Stretch, 487 proved males and 514 females; i.e., as 94.7 to 100. In regard to domestic pigeons there is good evidence either that the males are produced in excess, or that they live longer; for these birds invariably pair, and single males, as Mr. Tegetmeier informs me, can always be purchased cheaper than females. Usually the two birds reared from the two eggs laid in the same nest are a male and a female; but Mr. Harrison Weir, who has been so large a breeder, says that he has often bred two cocks from the same nest, and seldom two hens; moreover, the hen is generally the weaker of the two, and more liable to perish.

With respect to birds in a state of nature, Mr. Gould and others (62. Brehm ('Thierleben,' B. iv. s. 990) comes to the same conclusion.) are convinced that the males are generally the more numerous; and as the young males of many species resemble the females, the latter would naturally appear to be the more numerous. Large numbers of pheasants are reared by Mr. Baker of Leadenhall from eggs laid by wild birds, and he informs Mr. Jenner Weir that four or five males to one female are generally produced. An experienced observer remarks (63. On the authority of L. Lloyd, 'Game Birds of Sweden,' 1867, pp. 12, 132.), that in Scandinavia the broods of the capercailzie and black-cock contain more males than females; and that with the Dal-ripa (a kind of ptarmigan) more males than females attend the leks or places of courtship; but this latter circumstance is accounted for by some observers by a greater number of hen birds being killed by vermin. From various facts given by White of Selborne (64. 'Nat. Hist. of Selborne,' letter xxix. edit. of 1825, vol. i. p. 139.), it seems clear that the males of the partridge must be in considerable excess in the south of England; and I have been assured that this is the case in Scotland. Mr. Weir on enquiring from the dealers, who receive at certain seasons large numbers of ruffs (Machetes pugnax), was told that the males are much the more numerous. This same naturalist has also enquired for me from the birdcatchers, who annually catch an astonishing number of various small species alive for the London market, and he was unhesitatingly answered by an old and trustworthy man, that with the chaffinch the males are in large excess: he thought as high as 2 males to 1 female, or at least as high as 5 to 3. (65. Mr. Jenner Weir received similar information, on making enquiries during the following year. To shew the number of living chaffinches caught, I may mention that in 1869 there was a match between two experts, and one man caught in a day 62, and another 40, male chaffinches. The greatest number ever caught by one man in a single day was 70.) The males of the blackbird, he likewise maintained, were by far the more numerous, whether caught by traps or by netting at night. These statements may apparently be trusted, because this same man said that the sexes are about equal with the lark, the twite (Linaria montana), and goldfinch. On the other hand, he is certain that with the common linnet, the females preponderate greatly, but unequally during different years; during some years he has found the females to the males as four to one. It should, however, be borne in mind, that the chief season for catching birds does not begin till September, so that with some species partial migrations may have begun, and the flocks at this period often consist of hens alone. Mr. Salvin paid particular attention to the sexes of the humming-birds in Central America, and is convinced that with most of the species the males are in excess; thus one year he procured 204 specimens belonging to ten species, and these consisted of 166 males and of only 38 females. With two other species the females were in excess: but the proportions apparently vary either during different seasons or in different localities; for on one occasion the males of Campylopterus hemileucurus were to the females as 5 to 2, and on another occasion (66. 'Ibis,' vol. ii. p. 260, as quoted in Gould's 'Trochilidae,' 1861, p. 52. For the foregoing proportions, I am indebted to Mr. Salvin for a table of his results.) in exactly the reversed ratio. As bearing on this latter point, I may add, that Mr. Powys found in Corfu and Epirus the sexes of the chaffinch keeping apart, and "the females by far the most numerous"; whilst in Palestine Mr. Tristram found "the male flocks appearing greatly to exceed the female in number." (67. 'Ibis,' 1860, p. 137; and 1867, p. 369.) So again with the Quiscalus major, Mr. G. Taylor says, that in Florida there were "very few females in proportion to the males," (68. 'Ibis,' 1862, p. 187.) whilst in Honduras the proportion was the other way, the species there having the character of a polygamist.

FISH

With fish the proportional numbers of the sexes can be ascertained only by catching them in the adult or nearly adult state; and there are many difficulties in arriving at any just conclusion. (69. Leuckart quotes Bloch (Wagner, 'Handwörterbuch der Phys.' B. iv. 1853, s. 775), that with fish there are twice as many males as females.) Infertile females might readily be mistaken for males, as Dr. Gunther has remarked to me in regard to trout. With some species the males are believed to die soon after fertilising the ova. With many species the males are of much smaller size than the females, so that a large number of males would escape from the same net by which the females were caught. M. Carbonnier (70. Quoted in the 'Farmer,' March 18, 1869, p. 369.), who has especially attended to the natural history of the pike (Esox lucius), states that many males, owing to their small size, are devoured by the larger females; and he believes that the males of almost all fish are exposed from this same cause to greater danger than the females. Nevertheless, in the few cases in which the proportional numbers have been actually observed, the males appear to be largely in excess. Thus Mr. R. Buist, the superintendent of the Stormontfield experiments, says that in 1865, out of 70 salmon first landed for the purpose of obtaining the ova, upwards of 60 were males. In 1867 he again "calls attention to the vast disproportion of the males to the females. We had at the outset at least ten males to one female." Afterwards females sufficient for obtaining ova were procured. He adds, "from the great proportion of the males, they are constantly fighting and tearing each other on the spawning-beds." (71. 'The Stormontfield Piscicultural Experiments,' 1866, p. 23. The 'Field' newspaper, June 29, 1867.) This disproportion, no doubt, can be accounted for in part, but whether wholly is doubtful, by the males ascending the rivers before the females. Mr. F. Buckland remarks in regard to trout, that "it is a curious fact that the males preponderate very largely in number over the females. It INVARIABLY happens that when the first rush of fish is made to the net, there will be at least seven or eight males to one female found captive. I cannot quite account for this; either the males are more numerous than the females, or the latter seek safety by concealment rather than flight." He then adds, that by carefully searching the banks sufficient females for obtaining ova can be found. (72. 'Land and Water,' 1868, p. 41.) Mr. H. Lee informs me that out of 212 trout, taken for this purpose in Lord Portsmouth's park, 150 were males and 62 females.

The males of the Cyprinidae likewise seem to be in excess; but several members of this Family, viz., the carp, tench, bream and minnow, appear regularly to follow the practice, rare in the animal kingdom, of polyandry; for the female whilst spawning is always attended by two males, one on each side, and in the case of the bream by three or four males. This fact is so well known, that it is always recommended to stock a pond with two male tenches to one female, or at least with three males to two females. With the minnow, an excellent observer states, that on the spawning-beds the males are ten times as numerous as the females; when a female comes amongst the males, "she is immediately pressed closely by a male on each side; and when they have been in that situation for a time, are superseded by other two males." (73. Yarrell, 'Hist. British Fishes,' vol. i. 1826, p. 307; on the Cyprinus carpio, p. 331; on the Tinca vulgaris, p. 331; on the Abramis brama, p. 336. See, for the minnow (Leuciscus phoxinus), 'Loudon's Magazine of Natural History,' vol. v. 1832, p. 682.)

INSECTS

In this great Class, the Lepidoptera almost alone afford means for judging of the proportional numbers of the sexes; for they have been collected with special care by many good observers, and have been largely bred from the egg or caterpillar state. I had hoped that some breeders of silk-moths might have kept an exact record, but after writing to France and Italy, and consulting various treatises, I cannot find that this has ever been done. The general opinion appears to be that the sexes are nearly equal, but in Italy, as I hear from Professor Canestrini, many breeders are convinced that the females are produced in excess. This same naturalist, however, informs me, that in the two yearly broods of the Ailanthus silk-moth (Bombyx cynthia), the males greatly preponderate in the first, whilst in the second the two sexes are nearly equal, or the females rather in excess.

In regard to Butterflies in a state of nature, several observers have been much struck by the apparently enormous preponderance of the males. (74. Leuckart quotes Meinecke (Wagner, 'Handwörterbuch der Phys.' B. iv. 1853, s. 775) that the males of Butterflies are three or four times as numerous as the females.) Thus Mr. Bates (75. 'The Naturalist on the Amazons,' vol. ii. 1863, pp. 228, 347.), in speaking of several species, about a hundred in number, which inhabit the upper Amazons, says that the males are much more numerous than the females, even in the proportion of a hundred to one. In North America, Edwards, who had great experience, estimates in the genus Papilio the males to the females as four to one; and Mr. Walsh, who informed me of this statement, says that with P. turnus this is certainly the case. In South Africa, Mr. R. Trimen found the males in excess in 19 species (76. Four of these cases are given by Mr. Trimen in his 'Rhopalocera Africae Australis.'); and in one of these, which swarms in open places, he estimated the number of males as fifty to one female. With another species, in which the males are numerous in certain localities, he collected only five females during seven years. In the island of Bourbon, M. Maillard states that the males of one species of Papilio are twenty times as numerous as the females. (77. Quoted by Trimen, 'Transactions of the Ent. Society,' vol. v. part iv. 1866, p. 330.) Mr. Trimen informs me that as far as he has himself seen, or heard from others, it is rare for the females of any butterfly to exceed the males in number; but three South African species perhaps offer an exception. Mr. Wallace (78. 'Transactions, Linnean Society,' vol. xxv. p. 37.) states that the females of Ornithoptera croesus, in the Malay archipelago, are more common and more easily caught than the males; but this is a rare butterfly. I may here add, that in Hyperythra, a genus of moths, Guenee says, that from four to five females are sent in collections from India for one male.

When this subject of the proportional numbers of the sexes of insects was brought before the Entomological Society (79. 'Proceedings, Entomological Society,' Feb. 17, 1868.), it was generally admitted that the males of most Lepidoptera, in the adult or imago state, are caught in greater numbers than the females: but this fact was attributed by various observers to the more retiring habits of the females, and to the males emerging earlier from the cocoon. This latter circumstance is well known to occur with most Lepidoptera, as well as with other insects. So that, as M. Personnat remarks, the males of the domesticated Bombyx Yamamai, are useless at the beginning of the season, and the females at the end, from the want of mates. (80. Quoted by Dr. Wallace in 'Proceedings, Entomological Society,' 3rd series, vol. v. 1867, p. 487.) I cannot, however, persuade myself that these causes suffice to explain the great excess of males, in the above cases of certain butterflies which are extremely common in their native countries. Mr. Stainton, who has paid very close attention during many years to the smaller moths, informs me that when he collected them in the imago state, he thought that the males were ten times as numerous as the females, but that since he has reared them on a large scale from the caterpillar state, he is convinced that the females are the more numerous. Several entomologists concur in this view. Mr. Doubleday, however, and some others, take an opposite view, and are convinced that they have reared from the eggs and caterpillars a larger proportion of males than of females.

Besides the more active habits of the males, their earlier emergence from the cocoon, and in some cases their frequenting more open stations, other causes may be assigned for an apparent or real difference in the proportional numbers of the sexes of Lepidoptera, when captured in the imago state, and when reared from the egg or caterpillar state. I hear from Professor Canestrini, that it is believed by many breeders in Italy, that the female caterpillar of the silk-moth suffers more from the recent disease than the male; and Dr. Staudinger informs me that in rearing Lepidoptera more females die in the cocoon than males. With many species the female caterpillar is larger than the male, and a collector would naturally choose the finest specimens, and thus unintentionally collect a larger number of females. Three collectors have told me that this was their practice; but Dr. Wallace is sure that most collectors take all the specimens which they can find of the rarer kinds, which alone are worth the trouble of rearing. Birds when surrounded by caterpillars would probably devour the largest; and Professor Canestrini informs me that in Italy some breeders believe, though on insufficient evidence, that in the first broods of the Ailanthus silk-moth, the wasps destroy a larger number of the female than of the male caterpillars. Dr. Wallace further remarks that female caterpillars, from being larger than the males, require more time for their development, and consume more food and moisture: and thus they would be exposed during a longer time to danger from ichneumons, birds, etc., and in times of scarcity would perish in greater numbers. Hence it appears quite possible that in a state of nature, fewer female Lepidoptera may reach maturity than males; and for our special object we are concerned with their relative numbers at maturity, when the sexes are ready to propagate their kind.

The manner in which the males of certain moths congregate in extraordinary numbers round a single female, apparently indicates a great excess of males, though this fact may perhaps be accounted for by the earlier emergence of the males from their cocoons. Mr. Stainton informs me that from twelve to twenty males, may often be seen congregated round a female Elachista rufocinerea. It is well known that if a virgin Lasiocampa quercus or Saturnia carpini be exposed in a cage, vast numbers of males collect round her, and if confined in a room will even come down the chimney to her. Mr. Doubleday believes that he has seen from fifty to a hundred males of both these species attracted in the course of a single day by a female in confinement. In the Isle of Wight Mr. Trimen exposed a box in which a female of the Lasiocampa had been confined on the previous day, and five males soon endeavoured to gain admittance. In Australia, Mr. Verreaux, having placed the female of a small Bombyx in a box in his pocket, was followed by a crowd of males, so that about 200 entered the house with him. (81. Blanchard, 'Metamorphoses, Moeurs des Insectes,' 1868, pp. 225-226.)

Mr. Doubleday has called my attention to M. Staudinger's (82. 'Lepidopteren-Doubletten Liste,' Berlin, No. x. 1866.) list of Lepidoptera, which gives the prices of the males and females of 300 species or well- marked varieties of butterflies (Rhopalocera). The prices for both sexes of the very common species are of course the same; but in 114 of the rarer species they differ; the males being in all cases, excepting one, the cheaper. On an average of the prices of the 113 species, the price of the male to that of the female is as 100 to 149; and this apparently indicates that inversely the males exceed the females in the same proportion. About 2000 species or varieties of moths (Heterocera) are catalogued, those with wingless females being here excluded on account of the difference in habits between the two sexes: of these 2000 species, 141 differ in price according to sex, the males of 130 being cheaper, and those of only 11 being dearer than the females. The average price of the males of the 130 species, to that of the females, is as 100 to 143. With respect to the butterflies in this priced list, Mr. Doubleday thinks (and no man in England has had more experience), that there is nothing in the habits of the species which can account for the difference in the prices of the two sexes, and that it can be accounted for only by an excess in the number of the males. But I am bound to add that Dr. Staudinger informs me, that he is himself of a different opinion. He thinks that the less active habits of the females and the earlier emergence of the males will account for his collectors securing a larger number of males than of females, and consequently for the lower prices of the former. With respect to specimens reared from the caterpillar-state, Dr. Staudinger believes, as previously stated, that a greater number of females than of males die whilst confined to the cocoons. He adds that with certain species one sex seems to preponderate over the other during certain years.

Of direct observations on the sexes of Lepidoptera, reared either from eggs or caterpillars, I have received only the few following cases: (See following table.)

So that in these eight lots of cocoons and eggs, males were produced in excess. Taken together the proportion of males is as 122.7 to 100 females. But the numbers are hardly large enough to be trustworthy.

On the whole, from these various sources of evidence, all pointing in the same direction, I infer that with most species of Lepidoptera, the mature males generally exceed the females in number, whatever the proportions may be at their first emergence from the egg.

Males Females

The Rev. J. Hellins* of Exeter reared, during

1868, imagos of 73 species, which

consisted of 153 137

Mr. Albert Jones of Eltham reared, during 1868, imagos of 9 species, which consisted of 159 126

During 1869 he reared imagos from 4 species consisting of 114 112

Mr. Buckler of Emsworth, Hants, during 1869, reared imagos from 74 species, consisting of 180 169

Dr. Wallace of Colchester reared from one

brood of Bombyx cynthia 52 48

Dr. Wallace raised, from cocoons of Bombyx

Pernyi sent from China, during 1869 224 123

Dr. Wallace raised, during 1868 and 1869, from

two lots of cocoons of Bombyx yamamai 52 46

Total 934 761

(*83. This naturalist has been so kind as to send me some results from former years, in which the females seemed to preponderate; but so many of the figures were estimates, that I found it impossible to tabulate them.)

With reference to the other Orders of insects, I have been able to collect very little reliable information. With the stag-beetle (Lucanus cervus) "the males appear to be much more numerous than the females"; but when, as Cornelius remarked during 1867, an unusual number of these beetles appeared in one part of Germany, the females appeared to exceed the males as six to one. With one of the Elateridae, the males are said to be much more numerous than the females, and "two or three are often found united with one female (84. Gunther's 'Record of Zoological Literature,' 1867, p. 260. On the excess of female Lucanus, ibid, p. 250. On the males of Lucanus in England, Westwood,' 'Modern Classification of Insects,' vol. i. p. 187. On the Siagonium, ibid. p. 172.); so that here polyandry seems to prevail." With Siagonium (Staphylinidae), in which the males are furnished with horns, "the females are far more numerous than the opposite sex." Mr. Janson stated at the Entomological Society that the females of the bark feeding Tomicus villosus are so common as to be a plague, whilst the males are so rare as to be hardly known.

It is hardly worth while saying anything about the proportion of the sexes in certain species and even groups of insects, for the males are unknown or very rare, and the females are parthenogenetic, that is, fertile without sexual union; examples of this are afforded by several of the Cynipidae. (85. Walsh in 'The American Entomologist,' vol. i. 1869, p. 103. F. Smith, 'Record of Zoological Lit.' 1867, p. 328.) In all the gall-making Cynipidae known to Mr. Walsh, the females are four or five times as numerous as the males; and so it is, as he informs me, with the gall-making Cecidomyiidae (Diptera). With some common species of Saw-flies (Tenthredinae) Mr. F. Smith has reared hundreds of specimens from larvae of all sizes, but has never reared a single male; on the other hand, Curtis says (86. 'Farm Insects,' pp. 45-46.), that with certain species (Athalia), bred by him, the males were to the females as six to one; whilst exactly the reverse occurred with the mature insects of the same species caught in the fields. In the family of bees, Hermann Müller (87. 'Anwendung der Darwin'schen Lehre,' Verh. d. n. Jahrg., xxiv.), collected a large number of specimens of many species, and reared others from the cocoons, and counted the sexes. He found that the males of some species greatly exceeded the females in number; in others the reverse occurred; and in others the two sexes were nearly equal. But as in most cases the males emerge from the cocoons before the females, they are at the commencement of the breeding-season practically in excess. Müller also observed that the relative number of the two sexes in some species differed much in different localities. But as H. Müller has himself remarked to me, these remarks must be received with some caution, as one sex might more easily escape observation than the other. Thus his brother Fritz Müller has noticed in Brazil that the two sexes of the same species of bee sometimes frequent different kinds of flowers. With respect to the Orthoptera, I know hardly anything about the relative number of the sexes: Korte (88. 'Die Strich, Zug oder Wanderheuschrecke,' 1828, p. 20.), however, says that out of 500 locusts which he examined, the males were to the females as five to six. With the Neuroptera, Mr. Walsh states that in many, but by no means in all the species of the Odonatous group, there is a great overplus of males: in the genus Hetaerina, also, the males are generally at least four times as numerous as the females. In certain species in the genus Gomphus the males are equally in excess, whilst in two other species, the females are twice or thrice as numerous as the males. In some European species of Psocus thousands of females may be collected without a single male, whilst with other species of the same genus both sexes are common. (89. 'Observations on N. American Neuroptera,' by H. Hagen and B.D. Walsh, 'Proceedings, Ent. Soc. Philadelphia,' Oct. 1863, pp. 168, 223, 239.) In England, Mr. MacLachlan has captured hundreds of the female Apatania muliebris, but has never seen the male; and of Boreus hyemalis only four or five males have been seen here. (90. 'Proceedings, Ent. Soc. London,' Feb. 17, 1868.) With most of these species (excepting the Tenthredinae) there is at present no evidence that the females are subject to parthenogenesis; and thus we see how ignorant we are of the causes of the apparent discrepancy in the proportion of the two sexes.