THE PROPORTION OF THE SEXES IN RELATION TO NATURAL SELECTION.

There is reason to suspect that in some cases man has by selection indirectly influenced his own sex-producing powers. Certain women tend to produce during their whole lives more children of one sex than of the other: and the same holds good of many animals, for instance, cows and horses; thus Mr. Wright of Yeldersley House informs me that one of his Arab mares, though put seven times to different horses, produced seven fillies. Though I have very little evidence on this head, analogy would lead to the belief, that the tendency to produce either sex would be inherited like almost every other peculiarity, for instance, that of producing twins; and concerning the above tendency a good authority, Mr. J. Downing, has communicated to me facts which seem to prove that this does occur in certain families of short-horn cattle. Col. Marshall (94. 'The Todas,' 1873, pp. 100, 111, 194, 196.) has recently found on careful examination that the Todas, a hill-tribe of India, consist of 112 males and 84 females of all ages—that is in a ratio of 133.3 males to 100 females. The Todas, who are polyandrous in their marriages, during former times invariably practised female infanticide; but this practice has now been discontinued for a considerable period. Of the children born within late years, the males are more numerous than the females, in the proportion of 124 to 100. Colonel Marshall accounts for this fact in the following ingenious manner. "Let us for the purpose of illustration take three families as representing an average of the entire tribe; say that one mother gives birth to six daughters and no sons; a second mother has six sons only, whilst the third mother has three sons and three daughters. The first mother, following the tribal custom, destroys four daughters and preserves two. The second retains her six sons. The third kills two daughters and keeps one, as also her three sons. We have then from the three families, nine sons and three daughters, with which to continue the breed. But whilst the males belong to families in which the tendency to produce sons is great, the females are of those of a converse inclination. Thus the bias strengthens with each generation, until, as we find, families grow to have habitually more sons than daughters."

That this result would follow from the above form of infanticide seems almost certain; that is if we assume that a sex-producing tendency is inherited. But as the above numbers are so extremely scanty, I have searched for additional evidence, but cannot decide whether what I have found is trustworthy; nevertheless the facts are, perhaps, worth giving. The Maories of New Zealand have long practised infanticide; and Mr. Fenton (95. 'Aboriginal Inhabitants of New Zealand: Government Report,' 1859, p. 36.) states that he "has met with instances of women who have destroyed four, six, and even seven children, mostly females. However, the universal testimony of those best qualified to judge, is conclusive that this custom has for many years been almost extinct. Probably the year 1835 may be named as the period of its ceasing to exist." Now amongst the New Zealanders, as with the Todas, male births are considerably in excess. Mr. Fenton remarks (p. 30), "One fact is certain, although the exact period of the commencement of this singular condition of the disproportion of the sexes cannot be demonstratively fixed, it is quite clear that this course of decrease was in full operation during the years 1830 to 1844, when the non-adult population of 1844 was being produced, and has continued with great energy up to the present time." The following statements are taken from Mr. Fenton (p. 26), but as the numbers are not large, and as the census was not accurate, uniform results cannot be expected. It should be borne in mind in this and the following cases, that the normal state of every population is an excess of women, at least in all civilised countries, chiefly owing to the greater mortality of the male sex during youth, and partly to accidents of all kinds later in life. In 1858, the native population of New Zealand was estimated as consisting of 31,667 males and 24,303 females of all ages, that is in the ratio of 130.3 males to 100 females. But during this same year, and in certain limited districts, the numbers were ascertained with much care, and the males of all ages were here 753 and the females 616; that is in the ratio of 122.2 males to 100 females. It is more important for us that during this same year of 1858, the NON-ADULT males within the same district were found to be 178, and the NON-ADULT females 142, that is in the ratio of 125.3 to 100. It may be added that in 1844, at which period female infanticide had only lately ceased, the NON-ADULT males in one district were 281, and the NON- ADULT females only 194, that is in the ratio of 144.8 males to 100 females.

In the Sandwich Islands, the males exceed the females in number. Infanticide was formerly practised there to a frightful extent, but was by no means confined to female infants, as is shewn by Mr. Ellis (96. 'Narrative of a Tour through Hawaii,' 1826, p. 298.), and as I have been informed by Bishop Staley and the Rev. Mr. Coan. Nevertheless, another apparently trustworthy writer, Mr. Jarves (97. 'History of the Sandwich Islands,' 1843, p. 93.), whose observations apply to the whole archipelago, remarks:—"Numbers of women are to be found, who confess to the murder of from three to six or eight children," and he adds, "females from being considered less useful than males were more often destroyed." From what is known to occur in other parts of the world, this statement is probable; but must be received with much caution. The practice of infanticide ceased about the year 1819, when idolatry was abolished and missionaries settled in the Islands. A careful census in 1839 of the adult and taxable men and women in the island of Kauai and in one district of Oahu (Jarves, p. 404), gives 4723 males and 3776 females; that is in the ratio of 125.08 to 100. At the same time the number of males under fourteen years in Kauai and under eighteen in Oahu was 1797, and of females of the same ages 1429; and here we have the ratio of 125.75 males to 100 females.

In a census of all the islands in 1850 (98. This is given in the Rev. H.T. Cheever's 'Life in the Sandwich Islands,' 1851, p. 277.), the males of all ages amount to 36,272, and the females to 33,128, or as 109.49 to 100. The males under seventeen years amounted to 10,773, and the females under the same age to 9593, or as 112.3 to 100. From the census of 1872, the proportion of males of all ages (including half-castes) to females, is as 125.36 to 100. It must be borne in mind that all these returns for the Sandwich Islands give the proportion of living males to living females, and not of the births; and judging from all civilised countries the proportion of males would have been considerably higher if the numbers had referred to births. (99. Dr. Coulter, in describing ('Journal R. Geograph. Soc.' vol. v. 1835, p. 67) the state of California about the year 1830, says that the natives, reclaimed by the Spanish missionaries, have nearly all perished, or are perishing, although well treated, not driven from their native land, and kept from the use of spirits. He attributes this, in great part, to the undoubted fact that the men greatly exceed the women in number; but he does not know whether this is due to a failure of female offspring, or to more females dying during early youth. The latter alternative, according to all analogy, is very improbable. He adds that "infanticide, properly so called, is not common, though very frequent recourse is had to abortion." If Dr. Coulter is correct about infanticide, this case cannot be advanced in support of Colonel Marshall's view. From the rapid decrease of the reclaimed natives, we may suspect that, as in the cases lately given, their fertility has been diminished from changed habits of life.

I had hoped to gain some light on this subject from the breeding of dogs; inasmuch as in most breeds, with the exception, perhaps, of greyhounds, many more female puppies are destroyed than males, just as with the Toda infants. Mr. Cupples assures me that this is usual with Scotch deer- hounds. Unfortunately, I know nothing of the proportion of the sexes in any breed, excepting greyhounds, and there the male births are to the females as 110.1 to 100. Now from enquiries made from many breeders, it seems that the females are in some respects more esteemed, though otherwise troublesome; and it does not appear that the female puppies of the best- bred dogs are systematically destroyed more than the males, though this does sometimes take place to a limited extent. Therefore I am unable to decide whether we can, on the above principles, account for the preponderance of male births in greyhounds. On the other hand, we have seen that with horses, cattle, and sheep, which are too valuable for the young of either sex to be destroyed, if there is any difference, the females are slightly in excess.)

From the several foregoing cases we have some reason to believe that infanticide practised in the manner above explained, tends to make a male- producing race; but I am far from supposing that this practice in the case of man, or some analogous process with other species, has been the sole determining cause of an excess of males. There may be some unknown law leading to this result in decreasing races, which have already become somewhat infertile. Besides the several causes previously alluded to, the greater facility of parturition amongst savages, and the less consequent injury to their male infants, would tend to increase the proportion of live-born males to females. There does not, however, seem to be any necessary connection between savage life and a marked excess of males; that is if we may judge by the character of the scanty offspring of the lately existing Tasmanians and of the crossed offspring of the Tahitians now inhabiting Norfolk Island.

As the males and females of many animals differ somewhat in habits and are exposed in different degrees to danger, it is probable that in many cases, more of one sex than of the other are habitually destroyed. But as far as I can trace out the complication of causes, an indiscriminate though large destruction of either sex would not tend to modify the sex-producing power of the species. With strictly social animals, such as bees or ants, which produce a vast number of sterile and fertile females in comparison with the males, and to whom this preponderance is of paramount importance, we can see that those communities would flourish best which contained females having a strong inherited tendency to produce more and more females; and in such cases an unequal sex-producing tendency would be ultimately gained through natural selection. With animals living in herds or troops, in which the males come to the front and defend the herd, as with the bisons of North America and certain baboons, it is conceivable that a male- producing tendency might be gained by natural selection; for the individuals of the better defended herds would leave more numerous descendants. In the case of mankind the advantage arising from having a preponderance of men in the tribe is supposed to be one chief cause of the practice of female infanticide.

In no case, as far as we can see, would an inherited tendency to produce both sexes in equal numbers or to produce one sex in excess, be a direct advantage or disadvantage to certain individuals more than to others; for instance, an individual with a tendency to produce more males than females would not succeed better in the battle for life than an individual with an opposite tendency; and therefore a tendency of this kind could not be gained through natural selection. Nevertheless, there are certain animals (for instance, fishes and cirripedes) in which two or more males appear to be necessary for the fertilisation of the female; and the males accordingly largely preponderate, but it is by no means obvious how this male-producing tendency could have been acquired. I formerly thought that when a tendency to produce the two sexes in equal numbers was advantageous to the species, it would follow from natural selection, but I now see that the whole problem is so intricate that it is safer to leave its solution for the future.

CHAPTER IX.

SECONDARY SEXUAL CHARACTERS IN THE LOWER CLASSES OF THE ANIMAL KINGDOM.

These characters absent in the lowest classes—Brilliant colours—Mollusca —Annelids—Crustacea, secondary sexual characters strongly developed; dimorphism; colour; characters not acquired before maturity—Spiders, sexual colours of; stridulation by the males—Myriapoda.

With animals belonging to the lower classes, the two sexes are not rarely united in the same individual, and therefore secondary sexual characters cannot be developed. In many cases where the sexes are separate, both are permanently attached to some support, and the one cannot search or struggle for the other. Moreover it is almost certain that these animals have too imperfect senses and much too low mental powers to appreciate each other's beauty or other attractions, or to feel rivalry.

Hence in these classes or sub-kingdoms, such as the Protozoa, Coelenterata, Echinodermata, Scolecida, secondary sexual characters, of the kind which we have to consider, do not occur: and this fact agrees with the belief that such characters in the higher classes have been acquired through sexual selection, which depends on the will, desire, and choice of either sex. Nevertheless some few apparent exceptions occur; thus, as I hear from Dr. Baird, the males of certain Entozoa, or internal parasitic worms, differ slightly in colour from the females; but we have no reason to suppose that such differences have been augmented through sexual selection. Contrivances by which the male holds the female, and which are indispensable for the propagation of the species, are independent of sexual selection, and have been acquired through ordinary selection.

Many of the lower animals, whether hermaphrodites or with separate sexes, are ornamented with the most brilliant tints, or are shaded and striped in an elegant manner; for instance, many corals and sea-anemones (Actiniae), some jelly-fish (Medusae, Porpita, etc.), some Planariae, many star-fishes, Echini, Ascidians, etc.; but we may conclude from the reasons already indicated, namely, the union of the two sexes in some of these animals, the permanently affixed condition of others, and the low mental powers of all, that such colours do not serve as a sexual attraction, and have not been acquired through sexual selection. It should be borne in mind that in no case have we sufficient evidence that colours have been thus acquired, except where one sex is much more brilliantly or conspicuously coloured than the other, and where there is no difference in habits between the sexes sufficient to account for their different colours. But the evidence is rendered as complete as it can ever be, only when the more ornamented individuals, almost always the males, voluntarily display their attractions before the other sex; for we cannot believe that such display is useless, and if it be advantageous, sexual selection will almost inevitably follow. We may, however, extend this conclusion to both sexes, when coloured alike, if their colours are plainly analogous to those of one sex alone in certain other species of the same group.

How, then, are we to account for the beautiful or even gorgeous colours of many animals in the lowest classes? It appears doubtful whether such colours often serve as a protection; but that we may easily err on this head, will be admitted by every one who reads Mr. Wallace's excellent essay on this subject. It would not, for instance, at first occur to any one that the transparency of the Medusae, or jelly-fish, is of the highest service to them as a protection; but when we are reminded by Haeckel that not only the Medusae, but many floating Mollusca, crustaceans, and even small oceanic fishes partake of this same glass-like appearance, often accompanied by prismatic colours, we can hardly doubt that they thus escape the notice of pelagic birds and other enemies. M. Giard is also convinced (1. 'Archives de Zoolog. Exper.' Oct. 1872, p. 563.) that the bright tints of certain sponges and ascidians serve as a protection. Conspicuous colours are likewise beneficial to many animals as a warning to their would-be devourers that they are distasteful, or that they possess some special means of defence; but this subject will be discussed more conveniently hereafter.

We can, in our ignorance of most of the lowest animals, only say that their bright tints result either from the chemical nature or the minute structure of their tissues, independently of any benefit thus derived. Hardly any colour is finer than that of arterial blood; but there is no reason to suppose that the colour of the blood is in itself any advantage; and though it adds to the beauty of the maiden's cheek, no one will pretend that it has been acquired for this purpose. So again with many animals, especially the lower ones, the bile is richly coloured; thus, as I am informed by Mr. Hancock, the extreme beauty of the Eolidae (naked sea-slugs) is chiefly due to the biliary glands being seen through the translucent integuments—this beauty being probably of no service to these animals. The tints of the decaying leaves in an American forest are described by every one as gorgeous; yet no one supposes that these tints are of the least advantage to the trees. Bearing in mind how many substances closely analogous to natural organic compounds have been recently formed by chemists, and which exhibit the most splendid colours, it would have been a strange fact if substances similarly coloured had not often originated, independently of any useful end thus gained, in the complex laboratory of living organisms.

THE SUB-KINGDOM OF THE MOLLUSCA.

Throughout this great division of the animal kingdom, as far as I can discover, secondary sexual characters, such as we are here considering, never occur. Nor could they be expected in the three lowest classes, namely, in the Ascidians, Polyzoa, and Brachiopods (constituting the Molluscoida of some authors), for most of these animals are permanently affixed to a support or have their sexes united in the same individual. In the Lamellibranchiata, or bivalve shells, hermaphroditism is not rare. In the next higher class of the Gasteropoda, or univalve shells, the sexes are either united or separate. But in the latter case the males never possess special organs for finding, securing, or charming the females, or for fighting with other males. As I am informed by Mr. Gwyn Jeffreys, the sole external difference between the sexes consists in the shell sometimes differing a little in form; for instance, the shell of the male periwinkle (Littorina littorea) is narrower and has a more elongated spire than that of the female. But differences of this nature, it may be presumed, are directly connected with the act of reproduction, or with the development of the ova.

The Gasteropoda, though capable of locomotion and furnished with imperfect eyes, do not appear to be endowed with sufficient mental powers for the members of the same sex to struggle together in rivalry, and thus to acquire secondary sexual characters. Nevertheless with the pulmoniferous gasteropods, or land-snails, the pairing is preceded by courtship; for these animals, though hermaphrodites, are compelled by their structure to pair together. Agassiz remarks, "Quiconque a eu l'occasion d'observer les amours des limacons, ne saurait mettre en doute la seduction deployee dans les mouvements et les allures qui preparent et accomplissent le double embrassement de ces hermaphrodites." (2. 'De l'Espece et de la Class.' etc., 1869, p. 106.) These animals appear also susceptible of some degree of permanent attachment: an accurate observer, Mr. Lonsdale, informs me that he placed a pair of land-snails, (Helix pomatia), one of which was weakly, into a small and ill-provided garden. After a short time the strong and healthy individual disappeared, and was traced by its track of slime over a wall into an adjoining well-stocked garden. Mr. Lonsdale concluded that it had deserted its sickly mate; but after an absence of twenty-four hours it returned, and apparently communicated the result of its successful exploration, for both then started along the same track and disappeared over the wall.

Even in the highest class of the Mollusca, the Cephalopoda or cuttle- fishes, in which the sexes are separate, secondary sexual characters of the present kind do not, as far as I can discover, occur. This is a surprising circumstance, as these animals possess highly-developed sense-organs and have considerable mental powers, as will be admitted by every one who has watched their artful endeavours to escape from an enemy. (3. See, for instance, the account which I have given in my 'Journal of Researches,' 1845, p. 7.) Certain Cephalopoda, however, are characterised by one extraordinary sexual character, namely that the male element collects within one of the arms or tentacles, which is then cast off, and clinging by its sucking-discs to the female, lives for a time an independent life. So completely does the cast-off arm resemble a separate animal, that it was described by Cuvier as a parasitic worm under the name of Hectocotyle. But this marvellous structure may be classed as a primary rather than as a secondary sexual character.

Although with the Mollusca sexual selection does not seem to have come into play; yet many univalve and bivalve shells, such as volutes, cones, scallops, etc., are beautifully coloured and shaped. The colours do not appear in most cases to be of any use as a protection; they are probably the direct result, as in the lowest classes, of the nature of the tissues; the patterns and the sculpture of the shell depending on its manner of growth. The amount of light seems to be influential to a certain extent; for although, as repeatedly stated by Mr. Gwyn Jeffreys, the shells of some species living at a profound depth are brightly coloured, yet we generally see the lower surfaces, as well as the parts covered by the mantle, less highly-coloured than the upper and exposed surfaces. (4. I have given ('Geological Observations on Volcanic Islands,' 1844, p. 53) a curious instance of the influence of light on the colours of a frondescent incrustation, deposited by the surf on the coast-rocks of Ascension and formed by the solution of triturated sea-shells.) In some cases, as with shells living amongst corals or brightly-tinted seaweeds, the bright colours may serve as a protection. (5. Dr. Morse has lately discussed this subject in his paper on the 'Adaptive Coloration of Mollusca,' 'Proc. Boston Soc. of Nat. Hist.' vol. xiv. April 1871.) But that many of the nudibranch Mollusca, or sea-slugs, are as beautifully coloured as any shells, may be seen in Messrs. Alder and Hancock's magnificent work; and from information kindly given me by Mr. Hancock, it seems extremely doubtful whether these colours usually serve as a protection. With some species this may be the case, as with one kind which lives on the green leaves of algae, and is itself bright-green. But many brightly-coloured, white, or otherwise conspicuous species, do not seek concealment; whilst again some equally conspicuous species, as well as other dull-coloured kinds live under stones and in dark recesses. So that with these nudibranch molluscs, colour apparently does not stand in any close relation to the nature of the places which they inhabit.

These naked sea-slugs are hermaphrodites, yet they pair together, as do land-snails, many of which have extremely pretty shells. It is conceivable that two hermaphrodites, attracted by each other's greater beauty, might unite and leave offspring which would inherit their parents' greater beauty. But with such lowly-organised creatures this is extremely improbable. Nor is it at all obvious how the offspring from the more beautiful pairs of hermaphrodites would have any advantage over the offspring of the less beautiful, so as to increase in number, unless indeed vigour and beauty generally coincided. We have not here the case of a number of males becoming mature before the females, with the more beautiful males selected by the more vigorous females. If, indeed, brilliant colours were beneficial to a hermaphrodite animal in relation to its general habits of life, the more brightly-tinted individuals would succeed best and would increase in number; but this would be a case of natural and not of sexual selection.

SUB-KINGDOM OF THE VERMES: CLASS, ANNELIDA (OR SEA-WORMS).

In this class, although the sexes, when separate, sometimes differ from each other in characters of such importance that they have been placed under distinct genera or even families, yet the differences do not seem of the kind which can be safely attributed to sexual selection. These animals are often beautifully coloured, but as the sexes do not differ in this respect, we are but little concerned with them. Even the Nemertians, though so lowly organised, "vie in beauty and variety of colouring with any other group in the invertebrate series"; yet Dr. McIntosh (6. See his beautiful monograph on 'British Annelids,' part i. 1873, p. 3.) cannot discover that these colours are of any service. The sedentary annelids become duller-coloured, according to M. Quatrefages (7. See M. Perrier: 'L'Origine de l'Homme d'apres Darwin,' 'Revue Scientifique', Feb. 1873, p. 866.), after the period of reproduction; and this I presume may be attributed to their less vigorous condition at that time. All these worm- like animals apparently stand too low in the scale for the individuals of either sex to exert any choice in selecting a partner, or for the individuals of the same sex to struggle together in rivalry.

SUB-KINGDOM OF THE ARTHROPODA: CLASS, CRUSTACEA.

In this great class we first meet with undoubted secondary sexual characters, often developed in a remarkable manner. Unfortunately the habits of crustaceans are very imperfectly known, and we cannot explain the uses of many structures peculiar to one sex. With the lower parasitic species the males are of small size, and they alone are furnished with perfect swimming-legs, antennae and sense-organs; the females being destitute of these organs, with their bodies often consisting of a mere distorted mass. But these extraordinary differences between the two sexes are no doubt related to their widely different habits of life, and consequently do not concern us. In various crustaceans, belonging to distinct families, the anterior antennae are furnished with peculiar thread-like bodies, which are believed to act as smelling-organs, and these are much more numerous in the males than in the females. As the males, without any unusual development of their olfactory organs, would almost certainly be able sooner or later to find the females, the increased number of the smelling-threads has probably been acquired through sexual selection, by the better provided males having been the more successful in finding partners and in producing offspring. Fritz Muller has described a remarkable dimorphic species of Tanais, in which the male is represented by two distinct forms, which never graduate into each other. In the one form the male is furnished with more numerous smelling-threads, and in the other form with more powerful and more elongated chelae or pincers, which serve to hold the female. Fritz Muller suggests that these differences between the two male forms of the same species may have originated in certain individuals having varied in the number of the smelling-threads, whilst other individuals varied in the shape and size of their chelae; so that of the former, those which were best able to find the female, and of the latter, those which were best able to hold her, have left the greatest number of progeny to inherit their respective advantages. (8. 'Facts and Arguments for Darwin,' English translat., 1869, p. 20. See the previous discussion on the olfactory threads. Sars has described a somewhat analogous case (as quoted in 'Nature,' 1870, p. 455) in a Norwegian crustacean, the Pontoporeia affinis.)

[Fig.4. Labidocera Darwinii (from Lubbock). Labelled are: a. Part of right anterior antenna of male, forming a prehensile organ. b. Posterior pair of thoracic legs of male. c. Ditto of female.]

In some of the lower crustaceans, the right anterior antenna of the male differs greatly in structure from the left, the latter resembling in its simple tapering joints the antennae of the female. In the male the modified antenna is either swollen in the middle or angularly bent, or converted (Fig. 4) into an elegant, and sometimes wonderfully complex, prehensile organ. (9. See Sir J. Lubbock in 'Annals and Mag. of Nat. Hist.' vol. xi. 1853, pl. i. and x.; and vol. xii. (1853), pl. vii. See also Lubbock in 'Transactions, Entomological Society,' vol. iv. new series, 1856-1858, p. 8. With respect to the zigzagged antennae mentioned below, see Fritz Muller, 'Facts and Arguments for Darwin,' 1869, p. 40, foot- note.) It serves, as I hear from Sir J. Lubbock, to hold the female, and for this same purpose one of the two posterior legs (b) on the same side of the body is converted into a forceps. In another family the inferior or posterior antennae are "curiously zigzagged" in the males alone.

[Fig. 5. Anterior part of body of Callianassa (from Milne-Edwards), showing the unequal and differently-constructed right and left-hand chelae of the male. N.B.—The artist by mistake has reversed the drawing, and made the left-hand chela the largest.

Fig. 6. Second leg of male Orchestia Tucuratinga (from Fritz Muller).

Fig. 7. Ditto of female.]

In the higher crustaceans the anterior legs are developed into chelae or pincers; and these are generally larger in the male than in the female,—so much so that the market value of the male edible crab (Cancer pagurus), according to Mr. C. Spence Bate, is five times as great as that of the female. In many species the chelae are of unequal size on the opposite side of the body, the right-hand one being, as I am informed by Mr. Bate, generally, though not invariably, the largest. This inequality is also often much greater in the male than in the female. The two chelae of the male often differ in structure (Figs. 5, 6, and 7), the smaller one resembling that of the female. What advantage is gained by their inequality in size on the opposite sides of the body, and by the inequality being much greater in the male than in the female; and why, when they are of equal size, both are often much larger in the male than in the female, is not known. As I hear from Mr. Bate, the chelae are sometimes of such length and size that they cannot possibly be used for carrying food to the mouth. In the males of certain fresh-water prawns (Palaemon) the right leg is actually longer than the whole body. (10. See a paper by Mr. C. Spence Bate, with figures, in 'Proceedings, Zoological Society,' 1868, p. 363; and on the nomenclature of the genus, ibid. p. 585. I am greatly indebted to Mr. Spence Bate for nearly all the above statements with respect to the chelae of the higher crustaceans.) The great size of the one leg with its chelae may aid the male in fighting with his rivals; but this will not account for their inequality in the female on the opposite sides of the body. In Gelasimus, according to a statement quoted by Milne Edwards (11. 'Hist. Nat. des Crust.' tom. ii. 1837, p. 50.), the male and the female live in the same burrow, and this shews that they pair; the male closes the mouth of the burrow with one of its chelae, which is enormously developed; so that here it indirectly serves as a means of defence. Their main use, however, is probably to seize and to secure the female, and this in some instances, as with Gammarus, is known to be the case. The male of the hermit or soldier crab (Pagurus) for weeks together, carries about the shell inhabited by the female. (12. Mr. C. Spence Bate, 'British Association, Fourth Report on the Fauna of S. Devon.') The sexes, however, of the common shore-crab (Carcinus maenas), as Mr. Bate informs me, unite directly after the female has moulted her hard shell, when she is so soft that she would be injured if seized by the strong pincers of the male; but as she is caught and carried about by the male before moulting, she could then be seized with impunity.

[Fig.8. Orchestia Darwinii (from Fritz Muller), showing the differently- constructed chelae of the two male forms.]

Fritz Muller states that certain species of Melita are distinguished from all other amphipods by the females having "the coxal lamellae of the penultimate pair of feet produced into hook-like processes, of which the males lay hold with the hands of the first pair." The development of these hook-like processes has probably followed from those females which were the most securely held during the act of reproduction, having left the largest number of offspring. Another Brazilian amphipod (see Orchestia darwinii, Fig. 8) presents a case of dimorphism, like that of Tanais; for there are two male forms, which differ in the structure of their chelae. (13. Fritz Muller, 'Facts and Arguments for Darwin,' 1869, pp. 25-28.) As either chela would certainly suffice to hold the female,—for both are now used for this purpose,—the two male forms probably originated by some having varied in one manner and some in another; both forms having derived certain special, but nearly equal advantages, from their differently shaped organs.

It is not known that male crustaceans fight together for the possession of the females, but it is probably the case; for with most animals when the male is larger than the female, he seems to owe his greater size to his ancestors having fought with other males during many generations. In most of the orders, especially in the highest or the Brachyura, the male is larger than the female; the parasitic genera, however, in which the sexes follow different habits of life, and most of the Entomostraca must be excepted. The chelae of many crustaceans are weapons well adapted for fighting. Thus when a Devil-crab (Portunus puber) was seen by a son of Mr. Bate fighting with a Carcinus maenas, the latter was soon thrown on its back, and had every limb torn from its body. When several males of a Brazilian Gelasimus, a species furnished with immense pincers, were placed together in a glass vessel by Fritz Muller, they mutilated and killed one another. Mr. Bate put a large male Carcinus maenas into a pan of water, inhabited by a female which was paired with a smaller male; but the latter was soon dispossessed. Mr. Bate adds, "if they fought, the victory was a bloodless one, for I saw no wounds." This same naturalist separated a male sand-skipper (so common on our sea-shores), Gammarus marinus, from its female, both of whom were imprisoned in the same vessel with many individuals of the same species. The female, when thus divorced, soon joined the others. After a time the male was put again into the same vessel; and he then, after swimming about for a time, dashed into the crowd, and without any fighting at once took away his wife. This fact shews that in the Amphipoda, an order low in the scale, the males and females recognise each other, and are mutually attached.

The mental powers of the Crustacea are probably higher than at first sight appears probable. Any one who tries to catch one of the shore-crabs, so common on tropical coasts, will perceive how wary and alert they are. There is a large crab (Birgus latro), found on coral islands, which makes a thick bed of the picked fibres of the cocoa-nut, at the bottom of a deep burrow. It feeds on the fallen fruit of this tree by tearing off the husk, fibre by fibre; and it always begins at that end where the three eye-like depressions are situated. It then breaks through one of these eyes by hammering with its heavy front pincers, and turning round, extracts the albuminous core with its narrow posterior pincers. But these actions are probably instinctive, so that they would be performed as well by a young animal as by an old one. The following case, however, can hardly be so considered: a trustworthy naturalist, Mr. Gardner (14. 'Travels in the Interior of Brazil,' 1846, p. 111. I have given, in my 'Journal of Researches,' p. 463, an account of the habits of the Birgus.), whilst watching a shore-crab (Gelasimus) making its burrow, threw some shells towards the hole. One rolled in, and three other shells remained within a few inches of the mouth. In about five minutes the crab brought out the shell which had fallen in, and carried it away to a distance of a foot; it then saw the three other shells lying near, and evidently thinking that they might likewise roll in, carried them to the spot where it had laid the first. It would, I think, be difficult to distinguish this act from one performed by man by the aid of reason.

Mr. Bate does not know of any well-marked case of difference of colour in the two sexes of our British crustaceans, in which respect the sexes of the higher animals so often differ. In some cases, however, the males and females differ slightly in tint, but Mr. Bate thinks not more than may be accounted for by their different habits of life, such as by the male wandering more about, and being thus more exposed to the light. Dr. Power tried to distinguish by colour the sexes of the several species which inhabit the Mauritius, but failed, except with one species of Squilla, probably S. stylifera, the male of which is described as being "of a beautiful bluish-green," with some of the appendages cherry-red, whilst the female is clouded with brown and grey, "with the red about her much less vivid than in the male." (15. Mr. Ch. Fraser, in 'Proc. Zoolog. Soc.' 1869, p. 3. I am indebted to Mr. Bate for Dr. Power's statement.) In this case, we may suspect the agency of sexual selection. From M. Bert's observations on Daphnia, when placed in a vessel illuminated by a prism, we have reason to believe that even the lowest crustaceans can distinguish colours. With Saphirina (an oceanic genus of Entomostraca), the males are furnished with minute shields or cell-like bodies, which exhibit beautiful changing colours; these are absent in the females, and in both sexes of one species. (16. Claus, 'Die freilebenden Copepoden,' 1863, s. 35.) It would, however, be extremely rash to conclude that these curious organs serve to attract the females. I am informed by Fritz Muller, that in the female of a Brazilian species of Gelasimus, the whole body is of a nearly uniform greyish-brown. In the male the posterior part of the cephalo- thorax is pure white, with the anterior part of a rich green, shading into dark brown; and it is remarkable that these colours are liable to change in the course of a few minutes—the white becoming dirty grey or even black, the green "losing much of its brilliancy." It deserves especial notice that the males do not acquire their bright colours until they become mature. They appear to be much more numerous than the females; they differ also in the larger size of their chelae. In some species of the genus, probably in all, the sexes pair and inhabit the same burrow. They are also, as we have seen, highly intelligent animals. From these various considerations it seems probable that the male in this species has become gaily ornamented in order to attract or excite the female.

It has just been stated that the male Gelasimus does not acquire his conspicuous colours until mature and nearly ready to breed. This seems a general rule in the whole class in respect to the many remarkable structural differences between the sexes. We shall hereafter find the same law prevailing throughout the great sub-kingdom of the Vertebrata; and in all cases it is eminently distinctive of characters which have been acquired through sexual selection. Fritz Muller (17. 'Facts and Arguments,' etc., p. 79.) gives some striking instances of this law; thus the male sand-hopper (Orchestia) does not, until nearly full grown, acquire his large claspers, which are very differently constructed from those of the female; whilst young, his claspers resemble those of the female.

CLASS, ARACHNIDA (SPIDERS).

The sexes do not generally differ much in colour, but the males are often darker than the females, as may be seen in Mr. Blackwall's magnificent work. (18. 'A History of the Spiders of Great Britain,' 1861-64. For the following facts, see pp. 77, 88, 102.) In some species, however, the difference is conspicuous: thus the female of Sparassus smaragdulus is dullish green, whilst the adult male has the abdomen of a fine yellow, with three longitudinal stripes of rich red. In certain species of Thomisus the sexes closely resemble each other, in others they differ much; and analogous cases occur in many other genera. It is often difficult to say which of the two sexes departs most from the ordinary coloration of the genus to which the species belong; but Mr. Blackwall thinks that, as a general rule, it is the male; and Canestrini (19. This author has recently published a valuable essay on the 'Caratteri sessuali secondarii degli Arachnidi,' in the 'Atti della Soc. Veneto-Trentina di Sc. Nat. Padova,' vol. i. Fasc. 3, 1873.) remarks that in certain genera the males can be specifically distinguished with ease, but the females with great difficulty. I am informed by Mr. Blackwall that the sexes whilst young usually resemble each other; and both often undergo great changes in colour during their successive moults, before arriving at maturity. In other cases the male alone appears to change colour. Thus the male of the above bright-coloured Sparassus at first resembles the female, and acquires his peculiar tints only when nearly adult. Spiders are possessed of acute senses, and exhibit much intelligence; as is well known, the females often shew the strongest affection for their eggs, which they carry about enveloped in a silken web. The males search eagerly for the females, and have been seen by Canestrini and others to fight for possession of them. This same author says that the union of the two sexes has been observed in about twenty species; and he asserts positively that the female rejects some of the males who court her, threatens them with open mandibles, and at last after long hesitation accepts the chosen one. From these several considerations, we may admit with some confidence that the well-marked differences in colour between the sexes of certain species are the results of sexual selection; though we have not here the best kind of evidence,— the display by the male of his ornaments. From the extreme variability of colour in the male of some species, for instance of Theridion lineatum, it would appear that these sexual characters of the males have not as yet become well fixed. Canestrini draws the same conclusion from the fact that the males of certain species present two forms, differing from each other in the size and length of their jaws; and this reminds us of the above cases of dimorphic crustaceans.

The male is generally much smaller than the female, sometimes to an extraordinary degree (20. Aug. Vinson ('Araneides des Iles de la Reunion,' pl. vi. figs. 1 and 2) gives a good instance of the small size of the male, in Epeira nigra. In this species, as I may add, the male is testaceous and the female black with legs banded with red. Other even more striking cases of inequality in size between the sexes have been recorded ('Quarterly Journal of Science,' July 1868, p. 429); but I have not seen the original accounts.), and he is forced to be extremely cautious in making his advances, as the female often carries her coyness to a dangerous pitch. De Geer saw a male that "in the midst of his preparatory caresses was seized by the object of his attentions, enveloped by her in a web and then devoured, a sight which, as he adds, filled him with horror and indignation." (21. Kirby and Spence, 'Introduction to Entomology,' vol. i. 1818, p. 280.) The Rev. O.P. Cambridge (22. 'Proceedings, Zoological Society,' 1871, p. 621.) accounts in the following manner for the extreme smallness of the male in the genus Nephila. "M. Vinson gives a graphic account of the agile way in which the diminutive male escapes from the ferocity of the female, by gliding about and playing hide and seek over her body and along her gigantic limbs: in such a pursuit it is evident that the chances of escape would be in favour of the smallest males, while the larger ones would fall early victims; thus gradually a diminutive race of males would be selected, until at last they would dwindle to the smallest possible size compatible with the exercise of their generative functions,— in fact, probably to the size we now see them, i.e., so small as to be a sort of parasite upon the female, and either beneath her notice, or too agile and too small for her to catch without great difficulty."

Westring has made the interesting discovery that the males of several species of Theridion (23. Theridion (Asagena, Sund.) serratipes, 4- punctatum et guttatum; see Westring, in Kroyer, 'Naturhist. Tidskrift,' vol. iv. 1842-1843, p. 349; and vol. ii. 1846-1849, p. 342. See, also, for other species, 'Araneae Suecicae,' p. 184.) have the power of making a stridulating sound, whilst the females are mute. The apparatus consists of a serrated ridge at the base of the abdomen, against which the hard hinder part of the thorax is rubbed; and of this structure not a trace can be detected in the females. It deserves notice that several writers, including the well-known arachnologist Walckenaer, have declared that spiders are attracted by music. (24. Dr. H.H. van Zouteveen, in his Dutch translation of this work (vol. i. p. 444), has collected several cases.) From the analogy of the Orthoptera and Homoptera, to be described in the next chapter, we may feel almost sure that the stridulation serves, as Westring also believes, to call or to excite the female; and this is the first case known to me in the ascending scale of the animal kingdom of sounds emitted for this purpose. (25. Hilgendorf, however, has lately called attention to an analogous structure in some of the higher crustaceans, which seems adapted to produce sound; see 'Zoological Record,' 1869, p. 603.)

CLASS, MYRIAPODA.

In neither of the two orders in this class, the millipedes and centipedes, can I find any well-marked instances of such sexual differences as more particularly concern us. In Glomeris limbata, however, and perhaps in some few other species, the males differ slightly in colour from the females; but this Glomeris is a highly variable species. In the males of the Diplopoda, the legs belonging either to one of the anterior or of the posterior segments of the body are modified into prehensile hooks which serve to secure the female. In some species of Iulus the tarsi of the male are furnished with membranous suckers for the same purpose. As we shall see when we treat of Insects, it is a much more unusual circumstance, that it is the female in Lithobius, which is furnished with prehensile appendages at the extremity of her body for holding the male. (26. Walckenaer et P. Gervais, 'Hist. Nat. des Insectes: Apteres,' tom. iv. 1847, pp. 17, 19, 68.)

CHAPTER X.

SECONDARY SEXUAL CHARACTERS OF INSECTS.

Diversified structures possessed by the males for seizing the females— Differences between the sexes, of which the meaning is not understood— Difference in size between the sexes—Thysanura—Diptera—Hemiptera— Homoptera, musical powers possessed by the males alone—Orthoptera, musical instruments of the males, much diversified in structure; pugnacity; colours—Neuroptera, sexual differences in colour—Hymenoptera, pugnacity and odours—Coleoptera, colours; furnished with great horns, apparently as an ornament; battles, stridulating organs generally common to both sexes.

In the immense class of insects the sexes sometimes differ in their locomotive-organs, and often in their sense-organs, as in the pectinated and beautifully plumose antennae of the males of many species. In Chloeon, one of the Ephemerae, the male has great pillared eyes, of which the female is entirely destitute. (1. Sir J. Lubbock, 'Transact. Linnean Soc.' vol. xxv, 1866, p. 484. With respect to the Mutillidae see Westwood, 'Modern Class. of Insects,' vol. ii. p. 213.) The ocelli are absent in the females of certain insects, as in the Mutillidae; and here the females are likewise wingless. But we are chiefly concerned with structures by which one male is enabled to conquer another, either in battle or courtship, through his strength, pugnacity, ornaments, or music. The innumerable contrivances, therefore, by which the male is able to seize the female, may be briefly passed over. Besides the complex structures at the apex of the abdomen, which ought perhaps to be ranked as primary organs (2. These organs in the male often differ in closely-allied species, and afford excellent specific characters. But their importance, from a functional point of view, as Mr. R. MacLachlan has remarked to me, has probably been overrated. It has been suggested, that slight differences in these organs would suffice to prevent the intercrossing of well-marked varieties or incipient species, and would thus aid in their development. That this can hardly be the case, we may infer from the many recorded cases (see, for instance, Bronn, 'Geschichte der Natur,' B. ii. 1843, s. 164; and Westwood, 'Transact. Ent. Soc.' vol. iii. 1842, p. 195) of distinct species having been observed in union. Mr. MacLachlan informs me (vide 'Stett. Ent. Zeitung,' 1867, s. 155) that when several species of Phryganidae, which present strongly-pronounced differences of this kind, were confined together by Dr. Aug. Meyer, THEY COUPLED, and one pair produced fertile ova.), "it is astonishing," as Mr. B.D. Walsh (3. 'The Practical Entomologist,' Philadelphia, vol. ii. May 1867, p 88.) has remarked, "how many different organs are worked in by nature for the seemingly insignificant object of enabling the male to grasp the female firmly." The mandibles or jaws are sometimes used for this purpose; thus the male Corydalis cornutus (a neuropterous insect in some degree allied to the Dragon flies, etc.) has immense curved jaws, many times longer than those of the female; and they are smooth instead of being toothed, so that he is thus enabled to seize her without injury. (4. Mr. Walsh, ibid. p. 107.) One of the stag-beetles of North America (Lucanus elaphus) uses his jaws, which are much larger than those of the female, for the same purpose, but probably likewise for fighting. In one of the sand- wasps (Ammophila) the jaws in the two sexes are closely alike, but are used for widely different purposes: the males, as Professor Westwood observes, "are exceedingly ardent, seizing their partners round the neck with their sickle-shaped jaws" (5. 'Modern Classification of Insects,' vol. ii. 1840, pp. 205, 206. Mr. Walsh, who called my attention to the double use of the jaws, says that he has repeatedly observed this fact.); whilst the females use these organs for burrowing in sand-banks and making their nests.

[Fig. 9. Crabro cribrarius. Upper figure, male; lower figure, female.]

The tarsi of the front-legs are dilated in many male beetles, or are furnished with broad cushions of hairs; and in many genera of water-beetles they are armed with a round flat sucker, so that the male may adhere to the slippery body of the female. It is a much more unusual circumstance that the females of some water-beetles (Dytiscus) have their elytra deeply grooved, and in Acilius sulcatus thickly set with hairs, as an aid to the male. The females of some other water-beetles (Hydroporus) have their elytra punctured for the same purpose. (6. We have here a curious and inexplicable case of dimorphism, for some of the females of four European species of Dytiscus, and of certain species of Hydroporus, have their elytra smooth; and no intermediate gradations between the sulcated or punctured, and the quite smooth elytra have been observed. See Dr. H. Schaum, as quoted in the 'Zoologist,' vols. v.-vi. 1847-48, p. 1896. Also Kirby and Spence, 'Introduction to Entomology,' vol. iii. 1826, p. 305.) In the male of Crabro cribrarius (Fig. 9), it is the tibia which is dilated into a broad horny plate, with minute membraneous dots, giving to it a singular appearance like that of a riddle. (7. Westwood, 'Modern Class.' vol. ii. p. 193. The following statement about Penthe, and others in inverted commas, are taken from Mr. Walsh, 'Practical Entomologist,' Philadelphia, vol. iii. p. 88.) In the male of Penthe (a genus of beetles) a few of the middle joints of the antennae are dilated and furnished on the inferior surface with cushions of hair, exactly like those on the tarsi of the Carabidae, "and obviously for the same end." In male dragon-flies, "the appendages at the tip of the tail are modified in an almost infinite variety of curious patterns to enable them to embrace the neck of the female." Lastly, in the males of many insects, the legs are furnished with peculiar spines, knobs or spurs; or the whole leg is bowed or thickened, but this is by no means invariably a sexual character; or one pair, or all three pairs are elongated, sometimes to an extravagant length. (8. Kirby and Spence, 'Introduct.' etc., vol. iii. pp. 332-336.)

[Fig. 10. Taphroderes distortus (much enlarged). Upper figure, male; lower figure, female.]

The sexes of many species in all the orders present differences, of which the meaning is not understood. One curious case is that of a beetle (Fig. 10), the male of which has left mandible much enlarged; so that the mouth is greatly distorted. In another Carabidous beetle, Eurygnathus (9. 'Insecta Maderensia,' 1854, page 20.), we have the case, unique as far as known to Mr. Wollaston, of the head of the female being much broader and larger, though in a variable degree, than that of the male. Any number of such cases could be given. They abound in the Lepidoptera: one of the most extraordinary is that certain male butterflies have their fore-legs more or less atrophied, with the tibiae and tarsi reduced to mere rudimentary knobs. The wings, also, in the two sexes often differ in neuration (10. E. Doubleday, 'Annals and Mag. of Nat. Hist.' vol. i. 1848, p. 379. I may add that the wings in certain Hymenoptera (see Shuckard, 'Fossorial Hymenoptera,' 1837, pp. 39-43) differ in neuration according to sex.), and sometimes considerably in outline, as in the Aricoris epitus, which was shewn to me in the British Museum by Mr. A. Butler. The males of certain South American butterflies have tufts of hair on the margins of the wings, and horny excrescences on the discs of the posterior pair. (11. H.W. Bates, in 'Journal of Proc. Linn. Soc.' vol. vi. 1862, p. 74. Mr. Wonfor's observations are quoted in 'Popular Science Review,' 1868, p. 343.) In several British butterflies, as shewn by Mr. Wonfor, the males alone are in parts clothed with peculiar scales.

The use of the bright light of the female glow-worm has been subject to much discussion. The male is feebly luminous, as are the larvae and even the eggs. It has been supposed by some authors that the light serves to frighten away enemies, and by others to guide the male to the female. At last, Mr. Belt (12. 'The Naturalist in Nicaragua,' 1874, pp. 316-320. On the phosphorescence of the eggs, see 'Annals and Magazine of Natural History,' Nov. 1871, p. 372.) appears to have solved the difficulty: he finds that all the Lampyridae which he has tried are highly distasteful to insectivorous mammals and birds. Hence it is in accordance with Mr. Bates' view, hereafter to be explained, that many insects mimic the Lampyridae closely, in order to be mistaken for them, and thus to escape destruction. He further believes that the luminous species profit by being at once recognised as unpalatable. It is probable that the same explanation may be extended to the Elaters, both sexes of which are highly luminous. It is not known why the wings of the female glow-worm have not been developed; but in her present state she closely resembles a larva, and as larvae are so largely preyed on by many animals, we can understand why she has been rendered so much more luminous and conspicuous than the male; and why the larvae themselves are likewise luminous.

DIFFERENCE IN SIZE BETWEEN THE SEXES.

With insects of all kinds the males are commonly smaller than the females; and this difference can often be detected even in the larval state. So considerable is the difference between the male and female cocoons of the silk-moth (Bombyx mori), that in France they are separated by a particular mode of weighing. (13. Robinet, 'Vers a Soie,' 1848, p. 207.) In the lower classes of the animal kingdom, the greater size of the females seems generally to depend on their developing an enormous number of ova; and this may to a certain extent hold good with insects. But Dr. Wallace has suggested a much more probable explanation. He finds, after carefully attending to the development of the caterpillars of Bombyx cynthia and yamamai, and especially to that of some dwarfed caterpillars reared from a second brood on unnatural food, "that in proportion as the individual moth is finer, so is the time required for its metamorphosis longer; and for this reason the female, which is the larger and heavier insect, from having to carry her numerous eggs, will be preceded by the male, which is smaller and has less to mature." (14. 'Transact. Ent. Soc.' 3rd series, vol. v. p. 486.) Now as most insects are short-lived, and as they are exposed to many dangers, it would manifestly be advantageous to the female to be impregnated as soon as possible. This end would be gained by the males being first matured in large numbers ready for the advent of the females; and this again would naturally follow, as Mr. A.R. Wallace has remarked (15. 'Journal of Proc. Ent. Soc.' Feb. 4, 1867, p. lxxi.), through natural selection; for the smaller males would be first matured, and thus would procreate a large number of offspring which would inherit the reduced size of their male parents, whilst the larger males from being matured later would leave fewer offspring.

There are, however, exceptions to the rule of male insects being smaller than the females: and some of these exceptions are intelligible. Size and strength would be an advantage to the males, which fight for the possession of the females; and in these cases, as with the stag-beetle (Lucanus), the males are larger than the females. There are, however, other beetles which are not known to fight together, of which the males exceed the females in size; and the meaning of this fact is not known; but in some of these cases, as with the huge Dynastes and Megasoma, we can at least see that there would be no necessity for the males to be smaller than the females, in order to be matured before them, for these beetles are not short-lived, and there would be ample time for the pairing of the sexes. So again, male dragon-flies (Libellulidae) are sometimes sensibly larger, and never smaller, than the females (16. For this and other statements on the size of the sexes, see Kirby and Spence, ibid. vol. iii. p. 300; on the duration of life in insects, see p. 344.); and as Mr. MacLachlan believes, they do not generally pair with the females until a week or fortnight has elapsed, and until they have assumed their proper masculine colours. But the most curious case, shewing on what complex and easily-overlooked relations, so trifling a character as difference in size between the sexes may depend, is that of the aculeate Hymenoptera; for Mr. F. Smith informs me that throughout nearly the whole of this large group, the males, in accordance with the general rule, are smaller than the females, and emerge about a week before them; but amongst the Bees, the males of Apis mellifica, Anthidium manicatum, and Anthophora acervorum, and amongst the Fossores, the males of the Methoca ichneumonides, are larger than the females. The explanation of this anomaly is that a marriage flight is absolutely necessary with these species, and the male requires great strength and size in order to carry the female through the air. Increased size has here been acquired in opposition to the usual relation between size and the period of development, for the males, though larger, emerge before the smaller females.

We will now review the several Orders, selecting such facts as more particularly concern us. The Lepidoptera (Butterflies and Moths) will be retained for a separate chapter.

ORDER, THYSANURA.

The members of this lowly organised order are wingless, dull-coloured, minute insects, with ugly, almost misshapen heads and bodies. Their sexes do not differ, but they are interesting as shewing us that the males pay sedulous court to the females even low down in the animal scale. Sir J. Lubbock (17. 'Transact. Linnean Soc.' vol. xxvi. 1868, p. 296.) says: "it is very amusing to see these little creatures (Smynthurus luteus) coquetting together. The male, which is much smaller than the female, runs round her, and they butt one another, standing face to face and moving backward and forward like two playful lambs. Then the female pretends to run away and the male runs after her with a queer appearance of anger, gets in front and stands facing her again; then she turns coyly round, but he, quicker and more active, scuttles round too, and seems to whip her with his antennae; then for a bit they stand face to face, play with their antennae, and seem to be all in all to one another."

ORDER, DIPTERA (FLIES).

The sexes differ little in colour. The greatest difference, known to Mr. F. Walker, is in the genus Bibio, in which the males are blackish or quite black, and the females obscure brownish-orange. The genus Elaphomyia, discovered by Mr. Wallace (18. 'The Malay Archipelago,' vol. ii. 1869, p. 313.) in New Guinea, is highly remarkable, as the males are furnished with horns, of which the females are quite destitute. The horns spring from beneath the eyes, and curiously resemble those of a stag, being either branched or palmated. In one of the species, they equal the whole body in length. They might be thought to be adapted for fighting, but as in one species they are of a beautiful pink colour, edged with black, with a pale central stripe, and as these insects have altogether a very elegant appearance, it is perhaps more probable that they serve as ornaments. That the males of some Diptera fight together is certain; Prof. Westwood (19. 'Modern Classification of Insects,' vol. ii. 1840, p. 526.) has several times seen this with the Tipulae. The males of other Diptera apparently try to win the females by their music: H. Muller (20. 'Anwendung,' etc., 'Verh. d. n. V. Jahrg.' xxix. p. 80. Mayer, in 'American Naturalist,' 1874, p. 236.) watched for some time two males of an Eristalis courting a female; they hovered above her, and flew from side to side, making a high humming noise at the same time. Gnats and mosquitoes (Culicidae) also seem to attract each other by humming; and Prof. Mayer has recently ascertained that the hairs on the antennae of the male vibrate in unison with the notes of a tuning-fork, within the range of the sounds emitted by the female. The longer hairs vibrate sympathetically with the graver notes, and the shorter hairs with the higher ones. Landois also asserts that he has repeatedly drawn down a whole swarm of gnats by uttering a particular note. It may be added that the mental faculties of the Diptera are probably higher than in most other insects, in accordance with their highly- developed nervous system. (21. See Mr. B.T. Lowne's interesting work, 'On the Anatomy of the Blow-fly, Musca vomitoria,' 1870, p. 14. He remarks (p. 33) that, "the captured flies utter a peculiar plaintive note, and that this sound causes other flies to disappear.")

ORDER, HEMIPTERA (FIELD-BUGS).

Mr. J.W. Douglas, who has particularly attended to the British species, has kindly given me an account of their sexual differences. The males of some species are furnished with wings, whilst the females are wingless; the sexes differ in the form of their bodies, elytra, antennae and tarsi; but as the signification of these differences are unknown, they may be here passed over. The females are generally larger and more robust than the males. With British, and, as far as Mr. Douglas knows, with exotic species, the sexes do not commonly differ much in colour; but in about six British species the male is considerably darker than the female, and in about four other species the female is darker than the male. Both sexes of some species are beautifully coloured; and as these insects emit an extremely nauseous odour, their conspicuous colours may serve as a signal that they are unpalatable to insectivorous animals. In some few cases their colours appear to be directly protective: thus Prof. Hoffmann informs me that he could hardly distinguish a small pink and green species from the buds on the trunks of lime-trees, which this insect frequents.

Some species of Reduvidae make a stridulating noise; and, in the case of Pirates stridulus, this is said (22. Westwood, 'Modern Classification of Insects,' vol. ii. p. 473.) to be effected by the movement of the neck within the pro-thoracic cavity. According to Westring, Reduvius personatus also stridulates. But I have no reason to suppose that this is a sexual character, excepting that with non-social insects there seems to be no use for sound-producing organs, unless it be as a sexual call.

ORDER: HOMOPTERA.

Every one who has wandered in a tropical forest must have been astonished at the din made by the male Cicadae. The females are mute; as the Grecian poet Xenarchus says, "Happy the Cicadas live, since they all have voiceless wives." The noise thus made could be plainly heard on board the "Beagle," when anchored at a quarter of a mile from the shore of Brazil; and Captain Hancock says it can be heard at the distance of a mile. The Greeks formerly kept, and the Chinese now keep these insects in cages for the sake of their song, so that it must be pleasing to the ears of some men. (23. These particulars are taken from Westwood's 'Modern Classification of Insects,' vol. ii. 1840, p. 422. See, also, on the Fulgoridae, Kirby and Spence, 'Introduct.' vol. ii. p. 401.) The Cicadidae usually sing during the day, whilst the Fulgoridae appear to be night-songsters. The sound, according to Landois (24. 'Zeitschrift fur wissenschaft Zoolog.' B. xvii. 1867, ss. 152-158.), is produced by the vibration of the lips of the spiracles, which are set into motion by a current of air emitted from the tracheae; but this view has lately been disputed. Dr. Powell appears to have proved (25. 'Transactions of the New Zealand Institute,' vol. v. 1873, p. 286.) that it is produced by the vibration of a membrane, set into action by a special muscle. In the living insect, whilst stridulating, this membrane can be seen to vibrate; and in the dead insect the proper sound is heard, if the muscle, when a little dried and hardened, is pulled with the point of a pin. In the female the whole complex musical apparatus is present, but is much less developed than in the male, and is never used for producing sound.

With respect to the object of the music, Dr. Hartman, in speaking of the Cicada septemdecim of the United States, says (26. I am indebted to Mr. Walsh for having sent me this extract from 'A Journal of the Doings of Cicada septemdecim,' by Dr. Hartman.), "the drums are now (June 6th and 7th, 1851) heard in all directions. This I believe to be the marital summons from the males. Standing in thick chestnut sprouts about as high as my head, where hundreds were around me, I observed the females coming around the drumming males." He adds, "this season (Aug. 1868) a dwarf pear-tree in my garden produced about fifty larvae of Cic. pruinosa; and I several times noticed the females to alight near a male while he was uttering his clanging notes." Fritz Muller writes to me from S. Brazil that he has often listened to a musical contest between two or three males of a species with a particularly loud voice, seated at a considerable distance from each other: as soon as one had finished his song, another immediately began, and then another. As there is so much rivalry between the males, it is probable that the females not only find them by their sounds, but that, like female birds, they are excited or allured by the male with the most attractive voice.

I have not heard of any well-marked cases of ornamental differences between the sexes of the Homoptera. Mr. Douglas informs me that there are three British species, in which the male is black or marked with black bands, whilst the females are pale-coloured or obscure.

ORDER, ORTHOPTERA (CRICKETS AND GRASSHOPPERS).

The males in the three saltatorial families in this Order are remarkable for their musical powers, namely the Achetidae or crickets, the Locustidae for which there is no equivalent English name, and the Acridiidae or grasshoppers. The stridulation produced by some of the Locustidae is so loud that it can be heard during the night at the distance of a mile (27. L. Guilding, 'Transactions of the Linnean Society,' vol. xv. p. 154.); and that made by certain species is not unmusical even to the human ear, so that the Indians on the Amazons keep them in wicker cages. All observers agree that the sounds serve either to call or excite the mute females. With respect to the migratory locusts of Russia, Korte has given (28. I state this on the authority of Koppen, 'Uber die Heuschrecken in Sudrussland,' 1866, p. 32, for I have in vain endeavoured to procure Korte's work.) an interesting case of selection by the female of a male. The males of this species (Pachytylus migratorius) whilst coupled with the female stridulate from anger or jealousy, if approached by other males. The house-cricket when surprised at night uses its voice to warn its fellows. (29. Gilbert White, 'Natural History of Selborne,' vol. ii. 1825, p. 262.) In North America the Katy-did (Platyphyllum concavum, one of the Locustidae) is described (30. Harris, 'Insects of New England,' 1842, p. 128.) as mounting on the upper branches of a tree, and in the evening beginning "his noisy babble, while rival notes issue from the neighbouring trees, and the groves resound with the call of Katy-did-she- did the live-long night." Mr. Bates, in speaking of the European field- cricket (one of the Achetidae), says "the male has been observed to place himself in the evening at the entrance of his burrow, and stridulate until a female approaches, when the louder notes are succeeded by a more subdued tone, whilst the successful musician caresses with his antennae the mate he has won." (31. 'The Naturalist on the Amazons,' vol. i. 1863, p. 252. Mr. Bates gives a very interesting discussion on the gradations in the musical apparatus of the three families. See also Westwood, 'Modern Classification of Insects,' vol. ii. pp. 445 and 453.) Dr. Scudder was able to excite one of these insects to answer him, by rubbing on a file with a quill. (32. 'Proceedings of the Boston Society of Natural History,' vol. xi. April 1868.) In both sexes a remarkable auditory apparatus has been discovered by Von Siebold, situated in the front legs. (33. 'Nouveau Manuel d'Anat. Comp.' (French translat.), tom. 1, 1850, p. 567.)

[Fig.11. Gryllus campestris (from Landois). Right-hand figure, under side of part of a wing-nervure, much magnified, showing the teeth, st. Left-hand figure, upper surface of wing-cover, with the projecting, smooth nervure, r, across which the teeth (st) are scraped.

Fig.12. Teeth of Nervure of Gryllus domesticus (from Landois).]

In the three Families the sounds are differently produced. In the males of the Achetidae both wing-covers have the same apparatus; and this in the field-cricket (see Gryllus campestris, Fig. 11) consists, as described by Landois (34. 'Zeitschrift fur wissenschaft. Zoolog.' B. xvii. 1867, s. 117.), of from 131 to 138 sharp, transverse ridges or teeth (st) on the under side of one of the nervures of the wing-cover. This toothed nervure is rapidly scraped across a projecting, smooth, hard nervure (r) on the upper surface of the opposite wing. First one wing is rubbed over the other, and then the movement is reversed. Both wings are raised a little at the same time, so as to increase the resonance. In some species the wing-covers of the males are furnished at the base with a talc-like plate. (35. Westwood, 'Modern Classification of Insects,' vol. i. p. 440.) I here give a drawing (Fig. 12) of the teeth on the under side of the nervure of another species of Gryllus, viz., G. domesticus. With respect to the formation of these teeth, Dr. Gruber has shewn (36. 'Ueber der Tonapparat der Locustiden, ein Beitrag zum Darwinismus,' 'Zeitschrift fur wissenschaft. Zoolog.' B. xxii. 1872, p. 100.) that they have been developed by the aid of selection, from the minute scales and hairs with which the wings and body are covered, and I came to the same conclusion with respect to those of the Coleoptera. But Dr. Gruber further shews that their development is in part directly due to the stimulus from the friction of one wing over the other.

[Fig.13. Chlorocoelus Tanana (from Bates). a,b. Lobes of opposite wing-covers.]

In the Locustidae the opposite wing-covers differ from each other in structure (Fig. 13), and the action cannot, as in the last family, be reversed. The left wing, which acts as the bow, lies over the right wing which serves as the fiddle. One of the nervures (a) on the under surface of the former is finely serrated, and is scraped across the prominent nervures on the upper surface of the opposite or right wing. In our British Phasgonura viridissima it appeared to me that the serrated nervure is rubbed against the rounded hind-corner of the opposite wing, the edge of which is thickened, coloured brown, and very sharp. In the right wing, but not in the left, there is a little plate, as transparent as talc, surrounded by nervures, and called the speculum. In Ephippiger vitium, a member of this same family, we have a curious subordinate modification; for the wing-covers are greatly reduced in size, but "the posterior part of the pro-thorax is elevated into a kind of dome over the wing-covers, and which has probably the effect of increasing the sound." (37. Westwood 'Modern Classification of Insects,' vol. i. p. 453.)

We thus see that the musical apparatus is more differentiated or specialised in the Locustidae (which include, I believe, the most powerful performers in the Order), than in the Achetidae, in which both wing-covers have the same structure and the same function. (38. Landois, 'Zeitschrift fur wissenschaft Zoolog.' B. xvii. 1867, ss. 121, 122.) Landois, however, detected in one of the Locustidae, namely in Decticus, a short and narrow row of small teeth, mere rudiments, on the inferior surface of the right wing-cover, which underlies the other and is never used as the bow. I observed the same rudimentary structure on the under side of the right wing-cover in Phasgonura viridissima. Hence we may infer with confidence that the Locustidae are descended from a form, in which, as in the existing Achetidae, both wing-covers had serrated nervures on the under surface, and could be indifferently used as the bow; but that in the Locustidae the two wing-covers gradually became differentiated and perfected, on the principle of the division of labour, the one to act exclusively as the bow, and the other as the fiddle. Dr. Gruber takes the same view, and has shewn that rudimentary teeth are commonly found on the inferior surface of the right wing. By what steps the more simple apparatus in the Achetidae originated, we do not know, but it is probable that the basal portions of the wing- covers originally overlapped each other as they do at present; and that the friction of the nervures produced a grating sound, as is now the case with the wing-covers of the females. (39. Mr. Walsh also informs me that he has noticed that the female of the Platyphyllum concavum, "when captured makes a feeble grating noise by shuffling her wing-covers together.") A grating sound thus occasionally and accidentally made by the males, if it served them ever so little as a love-call to the females, might readily have been intensified through sexual selection, by variations in the roughness of the nervures having been continually preserved.

[Fig.14. Hind-leg of Stenobothrus pratorum: r, the stridulating ridge; lower figure, the teeth forming the ridge, much magnified (from Landois).

Fig.15. Pneumora (from specimens in the British Museum). Upper figure, male; lower figure, female.]

In the last and third family, namely the Acridiidae or grasshoppers, the stridulation is produced in a very different manner, and according to Dr. Scudder, is not so shrill as in the preceding Families. The inner surface of the femur (Fig. 14, r) is furnished with a longitudinal row of minute, elegant, lancet-shaped, elastic teeth, from 85 to 93 in number (40. Landois, ibid. s. 113.); and these are scraped across the sharp, projecting nervures on the wing-covers, which are thus made to vibrate and resound. Harris (41. 'Insects of New England,' 1842, p. 133.) says that when one of the males begins to play, he first "bends the shank of the hind-leg beneath the thigh, where it is lodged in a furrow designed to receive it, and then draws the leg briskly up and down. He does not play both fiddles together, but alternately, first upon one and then on the other." In many species, the base of the abdomen is hollowed out into a great cavity which is believed to act as a resounding board. In Pneumora (Fig. 15), a S. African genus belonging to the same family, we meet with a new and remarkable modification; in the males a small notched ridge projects obliquely from each side of the abdomen, against which the hind femora are rubbed. (42. Westwood, 'Modern Classification,' vol i. p. 462.) As the male is furnished with wings (the female being wingless), it is remarkable that the thighs are not rubbed in the usual manner against the wing-covers; but this may perhaps be accounted for by the unusually small size of the hind-legs. I have not been able to examine the inner surface of the thighs, which, judging from analogy, would be finely serrated. The species of Pneumora have been more profoundly modified for the sake of stridulation than any other orthopterous insect; for in the male the whole body has been converted into a musical instrument, being distended with air, like a great pellucid bladder, so as to increase the resonance. Mr. Trimen informs me that at the Cape of Good Hope these insects make a wonderful noise during the night.

In the three foregoing families, the females are almost always destitute of an efficient musical apparatus. But there are a few exceptions to this rule, for Dr. Gruber has shewn that both sexes of Ephippiger vitium are thus provided; though the organs differ in the male and female to a certain extent. Hence we cannot suppose that they have been transferred from the male to the female, as appears to have been the case with the secondary sexual characters of many other animals. They must have been independently developed in the two sexes, which no doubt mutually call to each other during the season of love. In most other Locustidae (but not according to Landois in Decticus) the females have rudiments of the stridulatory organs proper to the male; from whom it is probable that these have been transferred. Landois also found such rudiments on the under surface of the wing-covers of the female Achetidae, and on the femora of the female Acridiidae. In the Homoptera, also, the females have the proper musical apparatus in a functionless state; and we shall hereafter meet in other divisions of the animal kingdom with many instances of structures proper to the male being present in a rudimentary condition of the female.

Landois has observed another important fact, namely, that in the females of the Acridiidae, the stridulating teeth on the femora remain throughout life in the same condition in which they first appear during the larval state in both sexes. In the males, on the other hand, they become further developed, and acquire their perfect structure at the last moult, when the insect is mature and ready to breed.

From the facts now given, we see that the means by which the males of the Orthoptera produce their sounds are extremely diversified, and are altogether different from those employed by the Homoptera. (43. Landois has recently found in certain Orthoptera rudimentary structures closely similar to the sound-producing organs in the Homoptera; and this is a surprising fact. See 'Zeitschrift fur wissenschaft Zoolog.' B. xxii. Heft 3, 1871, p. 348.) But throughout the animal kingdom we often find the same object gained by the most diversified means; this seems due to the whole organisation having undergone multifarious changes in the course of ages, and as part after part varied different variations were taken advantage of for the same general purpose. The diversity of means for producing sound in the three families of the Orthoptera and in the Homoptera, impresses the mind with the high importance of these structures to the males, for the sake of calling or alluring the females. We need feel no surprise at the amount of modification which the Orthoptera have undergone in this respect, as we now know, from Dr. Scudder's remarkable discovery (44. 'Transactions, Entomological Society,' 3rd series, vol. ii. ('Journal of Proceedings,' p. 117).), that there has been more than ample time. This naturalist has lately found a fossil insect in the Devonian formation of New Brunswick, which is furnished with "the well-known tympanum or stridulating apparatus of the male Locustidae." The insect, though in most respects related to the Neuroptera, appears, as is so often the case with very ancient forms, to connect the two related Orders of the Neuroptera and Orthoptera.

I have but little more to say on the Orthoptera. Some of the species are very pugnacious: when two male field-crickets (Gryllus campestris) are confined together, they fight till one kills the other; and the species of Mantis are described as manoeuvring with their sword-like front-limbs, like hussars with their sabres. The Chinese keep these insects in little bamboo cages, and match them like game-cocks. (45. Westwood, 'Modern Classification of Insects,' vol. i. p. 427; for crickets, p. 445.) With respect to colour, some exotic locusts are beautifully ornamented; the posterior wings being marked with red, blue, and black; but as throughout the Order the sexes rarely differ much in colour, it is not probable that they owe their bright tints to sexual selection. Conspicuous colours may be of use to these insects, by giving notice that they are unpalatable. Thus it has been observed (46. Mr. Ch. Horne, in 'Proceedings of the Entomological Society,' May 3, 1869, p. xii.) that a bright-coloured Indian locust was invariably rejected when offered to birds and lizards. Some cases, however, are known of sexual differences in colour in this Order. The male of an American cricket (47. The Oecanthus nivalis, Harris, 'Insects of New England,' 1842, p. 124. The two sexes of OE. pellucidus of Europe differ, as I hear from Victor Carus, in nearly the same manner.) is described as being as white as ivory, whilst the female varies from almost white to greenish-yellow or dusky. Mr. Walsh informs me that the adult male of Spectrum femoratum (one of the Phasmidae) "is of a shining brownish-yellow colour; the adult female being of a dull, opaque, cinereous brown; the young of both sexes being green." Lastly, I may mention that the male of one curious kind of cricket (48. Platyblemnus: Westwood, 'Modern Classification,' vol. i. p. 447.) is furnished with "a long membranous appendage, which falls over the face like a veil;" but what its use may be, is not known.

ORDER, NEUROPTERA.

Little need here be said, except as to colour. In the Ephemeridae the sexes often differ slightly in their obscure tints (49. B.D. Walsh, the 'Pseudo-neuroptera of Illinois,' in 'Proceedings of the Entomological Society of Philadelphia,' 1862, p. 361.); but it is not probable that the males are thus rendered attractive to the females. The Libellulidae, or dragon-flies, are ornamented with splendid green, blue, yellow, and vermilion metallic tints; and the sexes often differ. Thus, as Prof. Westwood remarks (50. 'Modern Classification,' vol. ii. p. 37.), the males of some of the Agrionidae, "are of a rich blue with black wings, whilst the females are fine green with colourless wings." But in Agrion Ramburii these colours are exactly reversed in the two sexes. (51. Walsh, ibid. p. 381. I am indebted to this naturalist for the following facts on Hetaerina, Anax, and Gomphus.) In the extensive N. American genus of Hetaerina, the males alone have a beautiful carmine spot at the base of each wing. In Anax junius the basal part of the abdomen in the male is a vivid ultramarine blue, and in the female grass-green. In the allied genus Gomphus, on the other hand, and in some other genera, the sexes differ but little in colour. In closely-allied forms throughout the animal kingdom, similar cases of the sexes differing greatly, or very little, or not at all, are of frequent occurrence. Although there is so wide a difference in colour between the sexes of many Libellulidae, it is often difficult to say which is the more brilliant; and the ordinary coloration of the two sexes is reversed, as we have just seen, in one species of Agrion. It is not probable that their colours in any case have been gained as a protection. Mr. MacLachlan, who has closely attended to this family, writes to me that dragon-flies—the tyrants of the insect-world—are the least liable of any insect to be attacked by birds or other enemies, and he believes that their bright colours serve as a sexual attraction. Certain dragon-flies apparently are attracted by particular colours: Mr. Patterson observed (52. 'Transactions, Ent. Soc.' vol. i. 1836, p. lxxxi.) that the Agrionidae, of which the males are blue, settled in numbers on the blue float of a fishing line; whilst two other species were attracted by shining white colours.