I have not found 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.—The males in the three saltatorial families belonging to this Order are remarkable for their musical powers, namely the Achetidæ or crickets, the Locustidæ for which there is no exact equivalent name in English, and the Acridiidæ or grasshoppers. The stridulation produced by some of the Locustidæ is so loud that it can be heard during the night at the distance of a mile;451 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. But it has been noticed452 that the male migratory locust of Russia (one of the Acridiidæ) whilst coupled with the female, stridulates from anger or jealousy when approached by another male. The house-cricket when surprised at night uses its voice to warn its fellows.453 In North America the Katy-did (Platyphyllum concavum, 353one of the Locustidæ) is described454 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 Achetidæ), says, “the male has been observed to place itself in the evening at the entrance of its 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 antennæ Fig. 10. Gryllus campestris (from Landois).
Right-hand figure, under side of part of the 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. the mate he has won.”455 Dr. Scudder was able to excite one of these insects to answer him, by rubbing on a file with a quill.456 In both sexes a remarkable auditory apparatus has been discovered by Von Siebold, situated in the front legs.457
In the three Families the sounds are differently produced. In the males Of the Achetidæ both wing-covers have the same structure; and this in the field-cricket (Gryllus campestris, fig. 10) consists, as de354scribed by Landois,458 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 Fig. 11. Teeth of Nervure of Gryllus domesticus (from Landois). 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.459 I have here given a drawing (fig. 11) of the teeth on the under side of the nervure of another species of Gryllus, viz. G. domesticus.
In the Locustidæ the opposite wing-covers differ in structure (fig. 12), and cannot, as in the last family, be indifferently used in a reversed manner. The left wing, which acts as the bow of the fiddle, lies over the right wing which serves as the fiddle itself. 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 355subordinate 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.”460
Fig. 12. Chlorocœlus Tanana (from Bates), a, b. Lobes of opposite wing-covers.
We thus see that the musical apparatus is more differentiated or specialised in the Locustidæ, which includes I believe the most powerful performers in the Order, than in the Achetidæ, in which both wing-covers have the same structure and the same function.461 Landois, however, detected in one of the Locustidæ, namely in Decticus, a short and narrow row of small 356teeth, 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 with confidence infer that the Locustidæ are descended from a form, in which, as in the existing Achetidæ, both wing-covers had serrated nervures on the under surface, and could be indifferently used as the bow; but that in the Locustidæ 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. By what steps the more simple apparatus in the Achetidæ originated, we do not know, but it is probable that the basal portions of the wing-covers overlapped each other formerly as at present, and that the friction of the nervures produced a grating sound, as I find is now the case with the wing-covers of the females.462 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 fitting variations in the roughness of the nervures having been continually preserved.
In the last and third Family, namely the Acridiidæ or grasshoppers, the stridulation is produced in a very different manner, and is not so shrill, according to Dr. Scudder, as in the preceding Families. The inner surface of the femur (fig. 13, r) is furnished with a longitudinal row of minute, elegant, lancet-shaped, elastic teeth, from 85 to 93 in number;463 and these are scraped 357across the sharp, projecting nervures on the wing-covers, which, are thus made to vibrate and resound. Harris464 Fig. 13, Hind-leg of Stenobothrus pratorum: r, the stridulating ridge; lower figure, the teeth, forming the ridge, much magnified (from Landois). 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. 14), a S. African genus belonging to this 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.465 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 358in 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 There is one exception to the rule that the females in these three Families are destitute of an efficient musical apparatus; for both sexes of Ephippiger (Locustidæ) are said466 to be thus provided. This case may 359be compared with that of the reindeer, in which species alone both sexes possess horns. Although the female orthoptera are thus almost invariably mute, yet Landois467 found rudiments of the stridulating organs on the femora of the female Acridiidæ, and similar rudiments on the under surface of the wing-covers of the female Achetidæ; but he failed to find any rudiments in the females of Decticus, one of the Locus
tidæ. In the Homoptera the mute females of Cicada, have the proper musical apparatus in an undeveloped state; and we shall hereafter meet in other divisions of the animal kingdom with innumerable instances of structures proper to the male being present in a rudimentary condition in the female. Such cases appear at first sight to indicate that both sexes were primordially constructed in the same manner, but that certain organs were subsequently lost by the females. It is, however, a more probable view, as previously explained, that the organs in question were acquired by the males and partially transferred to the females.
Fig. 14. Pneumora (from specimens in the British Museum). Upper figure, male; lower figure, female.
Landois has observed another interesting fact, namely that in the females of the Acridiidæ, the stridulating teeth on the femora remain throughout life in the same condition in which they first appear in both sexes during the larval state. In the males, on the other hand, they become fully 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 produce their sounds are extremely diversified in the Orthoptera, and are altogether different from those employed by the Homoptera. But throughout the animal kingdom we incessantly find the 360same object gained by the most diversified means; this being due to the whole organisation undergoing in the course of ages multifarious changes; and as part after part varies, different variations are taken advantage of for the same general purpose. The diversification of the 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,468 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 Locustidæ.” This insect, though in most respects related to the Neuroptera, appears to connect, as is so often the case with very ancient forms, the two Orders of the Neuroptera and Orthoptera which are now generally ranked as quite distinct.
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 manœuvring 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.469 With respect to colour, some exotic locusts are beautifully ornamented; the posterior wings being marked with red, 361blue, and black; but as throughout the Order the two sexes rarely differ much in colour, it is doubtful whether they owe these bright tints to sexual selection. Conspicuous colours may be of use to these insects as a protection, on the principle to be explained in the next chapter, by giving notice to their enemies that they are unpalatable. Thus it has been observed470 that an Indian brightly-coloured locust was invariably rejected when offered to birds and lizards. Some cases, however, of sexual differences in colour in this Order are known. The male of an American cricket471 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 Phasmidæ) “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 cricket472 is furnished with “a long membranous appendage, which falls over the face like a veil;” but whether this serves as an ornament is not known.
Order, Neuroptera.—Little need here be said, except in regard to colour. In the Ephemeridæ the sexes often differ slightly in their obscure tints;473 but it is not probable that the males are thus rendered attractive to the females. The Libellulidæ or dragon-flies are ornamented with splendid green, blue, yellow, and 362vermilion metallic tints; and the sexes often differ. Thus, the males of some of the Agrionidæ, as Prof. Westwood remarks474 “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.475 In the extensive N. American genus of Hetærina, 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 ultra-marine 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. Throughout the animal kingdom, similar cases of the sexes of closely-allied forms either differing greatly, or very little, or not at all, are of frequent occurrence. Although with many Libellulidæ there is so wide a difference in colour between the sexes, it is often difficult to say which is the most brilliant; and the ordinary coloration of the two sexes is exactly 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. As Mr. MacLachlan, who has closely attended to this family, writes to me, dragon-flies—the tyrants of the insect-world—are the least liable of any insect to be attacked by birds or other enemies. He believes that their bright colours serve as a sexual attraction. It deserves notice, as bearing on this subject, that certain dragon-flies appear to be attracted by particular colours: Mr. Patterson observed476 that the species of Agrionidæ, of which the males are blue, settled in numbers on the blue float of a fishing 363line; whilst two other species were attracted by shining white colours.
It is an interesting fact, first observed by Schelver, that the males, in several genera belonging to two sub-families, when they first emerge from the pupal state are coloured exactly like the females; but that their bodies in a short time assume a conspicuous milky-blue tint, owing to the exudation of a kind of oil, soluble in ether and alcohol. Mr. MacLachlan believes that in the male of Libellula depressa this change of colour does not occur until nearly a fortnight after the metamorphosis, when the sexes are ready to pair.
Certain species of Neurothemis present, according to Brauer477 a curious case of dimorphism, some of the females having their wings netted in the usual manner; whilst other females have them “very richly netted as in the males of the same species.” Brauer “explains the phenomenon on Darwinian principles by the supposition that the close netting of the veins is a secondary sexual character in the males.” This latter character is generally developed in the males alone, but being, like every other masculine character, latent in the female, is occasionally developed in them. We have here an illustration of the manner in which the two sexes of many animals have probably come to resemble each other, namely by variations first appearing in the males, being preserved in them, and then transmitted to and developed in the females; but in this particular genus a complete transference is occasionally and abruptly effected. Mr. MacLachlan informs me of another case of dimorphism occurring in several species of Agrion in which a certain number of individuals are found of an orange colour, and these are 364invariably females. This is probably a case of reversion, for in the true Libellulæ, when the sexes differ in colour, the females are always orange or yellow, so that supposing Agrion to be descended from some primordial form having the characteristic sexual colours of the typical Libellulæ, it would not be surprising that a tendency to vary in this manner should occur in the females alone.
Although many dragon-flies are such large, powerful, and fierce insects, the males have not been observed by Mr. MacLachlan to fight together, except, as he believes, in the case of some of the smaller species of Agrion. In another very distinct group in this Order, namely in the Termites or white ants, both sexes at the time of swarming may be seen running about, “the male after the female, sometimes two chasing one female, and contending with great eagerness who shall win the prize.”478
/> Order, Hymenoptera.—That inimitable observer, M. Fabre,479 in describing the habits of Cerceris, a wasp-like insect, remarks that “fights frequently ensue between the males for the possession of some particular female, who sits an apparently unconcerned beholder of the struggle for supremacy, and when the victory is decided, quietly flies away in company with the conqueror.” Westwood480 says that the males of one of the saw-flies (Tenthredinæ) “have been found fighting together, with their mandibles locked.” As M. Fabre speaks of the males of Cerceris striving to obtain a particular female, it may be well to bear in 365mind that insects belonging to this Order have the power of recognising each other after long intervals of time, and are deeply attached. For instance, Pierre Huber, whose accuracy no one doubts, separated some ants, and when after an interval of four months they met others which had formerly belonged to the same community, they mutually recognised and caressed each other with their antennæ. Had they been strangers they would have fought together. Again, when two communities engage in a battle, the ants on the same side in the general confusion sometimes attack each other, but they soon perceive their mistake, and the one ant soothes the other.481
In this Order slight differences in colour, according to sex, are common, but conspicuous differences are rare except in the family of Bees; yet both sexes of certain groups are so brilliantly coloured—for instance in Chrysis, in which vermilion and metallic greens prevail—that we are tempted to attribute the result to sexual selection. In the Ichneumonidæ, according to Mr. Walsh,482 the males are almost universally lighter coloured than the females. On the other hand, in the Tenthredinidæ the males are generally darker than the females. In the Siricidæ the sexes frequently differ; thus the male of Sirex juvencus is banded with orange, whilst the female is dark purple; but it is difficult to say which sex is the most ornamented. In Tremex columbæ the female is much brighter coloured than the male. With ants, as I am informed by Mr. F. Smith, the males of several species are black, the females being testaceous. In the family of Bees, especially in 366the solitary species, as I hear from the same distinguished entomologist, the sexes often differ in colour. The males are generally the brightest, and in Bombus as well as in Apathus, much more variable in colour than the females. In Anthophora retusa the male is of a rich fulvous-brown, whilst the female is quite black: so are the females of several species of Xylocopa, the males being bright yellow. In an Australian bee (Lestis bombylans), the female is of an extremely brilliant steel-blue, sometimes tinted with vivid green; the male being of a bright brassy colour clothed with rich fulvous pubescence. As in this group the females are provided with excellent defensive weapons in their stings, it is not probable that they have come to differ in colour from the males for the sake of protection.