Part 2 — CHAPTER VIII. (continued 2)

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.386 Infertile females might readily be mistaken for males, as Dr. Günther 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,387 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 the 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 sufficient females 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.”388 This disproportion, no doubt, can be accounted for in part, but whether wholly is very 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.389 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.

With the Cyprinidæ the males 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.”390

INSECTS.

In this class, the Lepidoptera alone afford the means of 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. The 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.391 Thus Mr. Bates,392 in speaking of the species, no less than 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;393 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 during seven years only five females. In the island of Bourbon, M. Maillard states that the males of one species of Papilio are twenty times as numerous as the females.394 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 in number the males; but this is perhaps the case with three South African species. Mr. Wallace395 states that the females of Ornithoptera crœsus, 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, Guenée 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,396 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 lost at the beginning of the season, and the females at the end, from the want of mates.397 I cannot however persuade myself that these causes suffice to explain the great excess of males in the cases, above given, of butterflies which are extremely common in their native countries. Mr. Stainton, who has paid such 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 most 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 egg and caterpillar states a larger proportion of males than of females.

Besides the more active habits of the males, their earlier emergence from the cocoon, and their frequenting in some cases 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. It is believed by many breeders in Italy, as I hear from Professor Canestrini, 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 brood 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, &c., 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 the 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 under confinement. Mr. Trimen exposed in the Isle of Wight a box in which a female of the Lasiocampa had been confined on the previous day, and five males soon endeavoured to gain admittance. M. Verreaux, in Australia, 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.398

Mr. Doubleday has called my attention to Dr. Staudinger’s399 list of Lepidoptera, which gives the prices of the males and females of 300 species or well-marked varieties of (Rhopalocera) butterflies. The prices for both sexes of the very common species are of course the same; but with 114 of the rarer species they differ; the males being in all cases, excepting one, the cheapest. 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 number 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 of the two sexes: of these 2000 species, 141 differ in price according to sex, the males of 130 being cheaper, and the males 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 numbers of the males. But I am bound to add that Dr. Staudinger himself, as he informs me, is 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 under confinement in 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:—

Males. Females.

The Rev. J. Hellins400 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

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

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 so one. With one of the Elateridæ, the males are said to be much more numerous than the females, and “two or three are often found united with one female;”401 so that here polyandry seems to prevail. With Siagonium (Staphylinidæ), 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. In other Orders, from unknown causes, but apparently in some instances owing to parthenogenesis, the males of certain species have never been discovered or are excessively rare, as with several of the Cynipidæ.402 In all the gall-making Cynipidæ 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 Cecidomyiiæ (Diptera). With some common species of Saw-flies (Tenthredinæ) Mr. F. Smith has reared hundreds of specimens from larvæ of all sizes, but has never reared a single male: on the other hand Curtis says,403 that with certain species (Athalia), bred by him, the males to the females were as six to one; whilst exactly the reverse occurred with the mature insects of the same species caught in the fields. With the Neuroptera, Mr. Walsh states that in many, but by no means in all, the species of the Odonatous groups (Ephemerina), there is a great overplus of males: in the genus Hetærina, 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 numerous, 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.404 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 here seen.405 With most of these species (excepting, as I have heard, with the Tenthredinæ) there is no reason to suppose that the females are subject to parthenogenesis; and thus we see how ignorant we are on the causes of the apparent discrepancy in the proportional numbers of the two sexes.

In the other Classes of the Articulata I have been able to collect still less information. With Spiders, Mr. Blackwall, who has carefully attended to this class during many years, writes to me that the males from their more erratic habits are more commonly seen, and therefore appear to be the more numerous. This is actually the case with a few species; but he mentions several species in six genera, in which the females appear to be much more numerous than the males.406 The small size of the males in comparison with the females, which is sometimes carried to an extreme degree, and their widely different appearance, may account in some instances for their rarity in collections.407

Some of the lower Crustaceans are able to propagate their kind asexually, and this will account for the extreme rarity of the males. With some other forms (as with Tanais and Cypris) there is reason to believe, as Fritz Müller informs me, that the male is much shorter-lived than the female, which, supposing the two sexes to be at first equal in number, would explain the scarcity of the males. On the other hand this same naturalist has invariably taken, on the shores of Brazil, far more males than females of the Diastylidæ and of Cypridina; thus with a species in the latter genus, 63 specimens caught the same day, included 57 males; but he suggests that this preponderance may be due to some unknown difference in the habits of the two sexes. With one of the higher Brazilian crabs, namely a Gelasimus, Fritz Müller found the males to be more numerous than the females. The reverse seems to be the case, according to the large experience of Mr. C. Spence Bate, with six common British crabs, the names of which he has given me.

On the Power of Natural Selection to regulate the proportional Numbers of the Sexes, and General Fertility.—In some peculiar cases, an excess in the number of one sex over the other might be a great advantage to a species, as with the sterile females of social insects, or with those animals in which more than one male is requisite to fertilise the female, as with certain cirripedes and perhaps certain fishes. An inequality between the sexes in these cases might have been acquired through natural selection, but from their rarity they need not here be further considered. In all ordinary cases an inequality would be no advantage or disadvantage to certain individuals more than to others; and therefore it could hardly have resulted from natural selection. We must attribute the inequality to the direct action of those unknown conditions, which with mankind lead to the males being born in a somewhat larger excess in certain countries than in others, or which cause the proportion between the sexes to differ slightly in legitimate and illegitimate births.

Let us now take the case of a species producing from the unknown causes just alluded to, an excess of one sex—we will say of males—these being superfluous and useless, or nearly useless. Could the sexes be equalised through natural selection? We may feel sure, from all characters being variable, that certain pairs would produce a somewhat less excess of males over females than other pairs. The former, supposing the actual number of the offspring to remain constant, would necessarily produce more females, and would therefore be more productive. On the doctrine of chances a greater number of the offspring of the more productive pairs would survive; and these would inherit a tendency to procreate fewer males and more females. Thus a tendency towards the equalisation of the sexes would be brought about. But our supposed species would by this process be rendered, as just remarked, more productive; and this would in many cases be far from an advantage; for whenever the limit to the numbers which exist, depends, not on destruction by enemies, but on the amount of food, increased fertility will lead to severer competition and to most of the survivors being badly fed. In this case, if the sexes were equalised by an increase in the number of the females, a simultaneous decrease in the total number of the offspring would be beneficial, or even necessary, for the existence of the species; and this, I believe, could be effected through natural selection in the manner hereafter to be described. The same train of reasoning is applicable in the above, as well as in the following case, if we assume that females instead of males are produced in excess, for such females from not uniting with males would be superfluous and useless. So it would be with polygamous species, if we assume the excess of females to be inordinately great.

An excess of either sex, we will again say of the males, could, however, apparently be eliminated through natural selection in another and indirect manner, namely by an actual diminution of the males, without any increase of the females, and consequently without any increase in the productiveness of the species. From the variability of all characters, we may feel assured that some pairs, inhabiting any locality, would produce a rather smaller excess of superfluous males, but an equal number of productive females. When the offspring from the more and the less male-productive parents were all mingled together, none would have any direct advantage over the others; but those that produced few superfluous males would have one great indirect advantage, namely that their ova or embryos would probably be larger and finer, or their young better nurtured in the womb and afterwards. We see this principle illustrated with plants; as those which bear a vast number of seed produce small ones; whilst those which bear comparatively few seeds, often produce large ones well-stocked with nutriment for the use of the seedlings.408 Hence the offspring of the parents which had wasted least force in producing superfluous males would be the most likely to survive, and would inherit the same tendency not to produce superfluous males, whilst retaining their full fertility in the production of females. So it would be with the converse case of the female sex. Any slight excess, however, of either sex could hardly be checked in so indirect a manner. Nor indeed has a considerable inequality between the sexes been always prevented, as we have seen in some of the cases given in the previous discussion. In these cases the unknown causes which determine the sex of the embryo, and which under certain conditions lead to the production of one sex in excess over the other, have not been mastered by the survival of those varieties which were subjected to the least waste of organised matter and force by the production of superfluous individuals of either sex. Nevertheless we may conclude that natural selection will always tend, though sometimes inefficiently, to equalise the relative numbers of the two sexes.

Having said this much on the equalisation of the sexes, it may be well to add a few remarks on the regulation through natural selection of the ordinary fertility of species. Mr. Herbert Spencer has shewn in an able discussion409 that with all organisms a ratio exists between what he calls individuation and genesis; whence it follows that beings which consume much matter or force in their growth, complicated structure or activity, or which produce ova and embryos of large size, or which expend much energy in nurturing their young, cannot be so productive as beings of an opposite nature. Mr. Spencer further shews that minor differences in fertility will be regulated through natural selection. Thus the fertility of each species will tend to increase, from the more fertile pairs producing a larger number of offspring, and these from their mere number will have the best chance of surviving, and will transmit their tendency to greater fertility. The only check to a continued augmentation of fertility in each organism seems to be either the expenditure of more power and the greater risks run by the parents that produce a more numerous progeny, or the contingency of very numerous eggs and young being produced of smaller size, or less vigorous, or subsequently not so well nurtured. To strike a balance in any case between the disadvantages which follow from the production of a numerous progeny, and the advantages (such as the escape of at least some individuals from various dangers) is quite beyond our power of judgment.

When an organism has once been rendered extremely fertile, how its fertility can be reduced through natural selection is not so clear as how this capacity was first acquired. Yet it is obvious that if individuals of a species, from a decrease of their natural enemies, were habitually reared in larger numbers than could be supported, all the members would suffer. Nevertheless the offspring from the less fertile parents would have no direct advantage over the offspring from the more fertile parents, when all were mingled together in the same district. All the individuals would mutually tend to starve each other. The offspring indeed of the less fertile parents would lie under one great disadvantage, for from the simple fact of being produced in smaller numbers, they would be the most liable to extermination. Indirectly, however, they would partake of one great advantage; for under the supposed condition of severe competition, when all were pressed for food, it is extremely probable that those individuals which from some variation in their constitution produced fewer eggs or young, would produce them of greater size or vigour; and the adults reared from such eggs or young would manifestly have the best chance of surviving, and would inherit a tendency towards lessened fertility. The parents, moreover, which had to nourish or provide for fewer offspring would themselves be exposed to a less severe strain in the struggle for existence, and would have a better chance of surviving. By these steps, and by no others as far as I can see, natural selection under the above conditions of severe competition for food, would lead to the formation of a new race less fertile, but better adapted for survival, than the parent-race.