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 colours--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 organs for locomotion, and often in their sense-organs, as in the pectinated and beautifully plumose antennae of the males of many species. In one of the Ephemerae, namely Chloeon, the male has great pillared eyes, of which the female is entirely destitute.[427] The ocelli are absent in the females of certain other insects, as in the Mutillidae, which are likewise destitute of wings. 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,[428] "it is astonishing," as Mr. B. D. Walsh[429] 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, &c.) has immense curved jaws, many times longer than those of the female; and they are smooth instead of being toothed, by which means he is enabled to seize her without injury.[430] 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;"[431] whilst the females use these organs for burrowing in sand-banks and making their nests.

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 object.[432] In the male of _Crabro cribrarius_ (fig. 8.), 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.[433] 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.[434]

In all the orders, the sexes of many species present differences, of which the meaning is not understood. One curious case is that of a beetle (fig. 9), the male of which has the left mandible much enlarged; so that the mouth is greatly distorted. In another Carabidous beetle, the Eurygnathus,[435] we have the unique case, 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,[436] and sometimes considerably in outline, as in the _Aricoris epitus_, which was shown 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.[437] In several British butterflies, the males alone, as shewn by Mr. Wonfor, are in parts clothed with peculiar scales.

The purpose of the luminosity in the female glow-worm is likewise not understood; for it is very doubtful whether the primary use of the light is to guide the male to the female. It is no serious objection to this latter belief that the males emit a feeble light; for secondary sexual characters proper to one sex are often developed in a slight degree in the other sex. It is a more valid objection that the larvae shine, and in some species brilliantly: Fritz Mueller informs me that the most luminous insect which he ever beheld in Brazil, was the larva of some beetle. Both sexes of certain luminous species of Elater emit light. Kirby and Spence suspect that the phosphorescence serves to frighten and drive away enemies.

_Difference in Size between the Sexes._--With insects of all kinds the males are commonly smaller than the females;[438] 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.[439] 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 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."[440] 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. E. Wallace has remarked,[441] 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 female; and in these cases the males, as with the stag-beetle (Lucanus), 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;[442] and they do not, as Mr. MacLachlan believes, 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 a 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 males require great strength and size in order to carry the females 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 Order are lowly organised for their class. They are wingless, dull-coloured, minute insects, with ugly, almost misshapen heads and bodies. The sexes do not differ; but they offer one interesting fact, by showing that the males pay sedulous court to their females even low down in the animal scale. Sir J. Lubbock[443] in describing the _Smynthurus luteus_, says: "it is very amusing to see these little creatures 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[444] 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 stags, being either branched or palmated. They equal in length the whole of the body in one of the species. They might be thought to serve 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 the horns serve as ornaments. That the males of some Diptera fight together is certain; for Prof. Westwood[445] has several times seen this with some species of Tipula or Harry-long-legs. Many observers believe that when gnats (Culicidae) dance in the air in a body, alternately rising and falling, the males are courting the females. The mental faculties of the Diptera are probably fairly well developed, for their nervous system is more highly developed than in most other Orders of insects.[446]

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 the body and elytra; in the second joints of their antennae and in their tarsi; but as the signification of these differences is quite 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 marked with vermilion and black. It is doubtful whether these colours serve as a protection. If in any species the males had differed from the females in an analogous manner, we might have been justified in attributing such conspicuous colours to sexual selection with transference to both sexes.

Some species of Reduvidae make a stridulating noise; and, in the case of _Pirates stridulus_, this is said[447] to be effected by the movement of the neck within the pro-thoracic cavity. According to Westring, _Reduvius personatus_ also stridulates. But I have not been able to learn any particulars about these insects; nor have I any reason to suppose that they differ sexually in this respect.

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.[448] The Cicadidae usually sing during the day; whilst the Fulgoridae appear to be night-songsters. The sound, according to Landois,[449] who has recently studied the subject, 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. It is increased by a wonderfully complex resounding apparatus, consisting of two cavities covered by scales. Hence the sound may truly be called a voice. In the female the musical apparatus is present, but very 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,[450] "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 Mueller writes to me from S. Brazil that he has often listened to a musical contest between two or three males of a Cicada, having a particularly loud voice, and seated at a considerable distance from each other. As soon as the first had finished his song, a second immediately began; and after he had concluded, another began, and so on. As there is so much rivalry between the males, it is probable that the females not only discover them by the sounds emitted, but that, like female birds, they are excited or allured by the male with the most attractive voice.

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 Achetidae or crickets, the Locustidae for which there is no exact equivalent name in English, 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;[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 noticed[452] that the male migratory locust of Russia (one of the Acridiidae) 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_, one of the Locustidae) is described[454] 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 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 antennae 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 Achetidae both wing-covers have the same structure; and this in the field-cricket (_Gryllus campestris_, fig. 10) consists, as described 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 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 Locustidae 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 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."[460]

We thus see that the musical apparatus is more differentiated or specialised in the Locustidae, which includes 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.[461] 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 with confidence infer 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. 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 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 Acridiidae 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 across the sharp, projecting nervures on the wing-covers, which, are thus made to vibrate and resound. Harris[464] 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 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 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 (Locustidae) are said[466] to be thus provided. This case may be compared with that of the reindeer, in which species alone both sexes possess horns. Although the female orthoptera are thus almost invariably mute, yet Landois[467] found rudiments of the stridulating organs on the femora of the female Acridiidae, and similar rudiments on the under surface of the wing-covers of the female Achetidae; but he failed to find any rudiments in the females of Decticus, one of the Locustidae. 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.

Landois has observed another interesting 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 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 same 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 Locustidae." 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 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.[469] 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 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 observed[470] 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 cricket[471] 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[472] 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 Ephemeridae 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 Libellulidae or dragon-flies are ornamented with splendid green, blue, yellow, and vermilion metallic tints; and the sexes often differ. Thus, the males of some of the Agrionidae, as Prof. Westwood remarks[474] "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 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 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 Libellulidae 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 observed[476] that the species of 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.

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 Brauer[477] 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 invariably females. This is probably a case of reversion, for in the true Libellulae, 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 Libellulae, 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." Westwood[480] says that the males of one of the saw-flies (Tenthredinae) "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 mind 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 antennae. 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 Ichneumonidae, according to Mr. Walsh,[482] the males are almost universally lighter coloured than the females. On the other hand, in the Tenthredinidae the males are generally darker than the females. In the Siricidae 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 columbae_ 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 the 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.

_Mutilla Europaea_ emits a stridulating noise; and according to Goureau[483] both sexes have this power. He attributes the sound to the friction of the third and preceding abdominal segments; and I find that these surfaces are marked with very fine concentric ridges, but so is the projecting thoracic collar, on which the head articulates; and this collar, when scratched with the point of a needle, emits the proper sound. It is rather surprising that both sexes should have the power of stridulating, as the male is winged and the female wingless. It is notorious that Bees express certain emotions, as of anger, by the tone of their humming, as do some dipterous insects; but I have not referred to these sounds, as they are not known to be in any way connected with the act of courtship.

Order, _Coleoptera_ (Beetles).--Many beetles are coloured so as to resemble the surfaces which they habitually frequent. Other species are ornamented with gorgeous metallic tints,--for instance, many Carabidae, which live on the ground and have the power of defending themselves by an intensely acrid secretion,--the splendid diamond-beetles which are protected by an extremely hard covering,--many species of Chrysomela, such as _C. cerealis_, a large species beautifully striped with various colours, and in Britain confined to the bare summit of Snowdon,--and a host of other species. These splendid colours, which are often arranged in stripes, spots, crosses and other elegant patterns, can hardly be beneficial, as a protection, except in the case of some flower-feeding species; and we cannot believe that they are purposeless. Hence the suspicion arises, that they serve as a sexual attraction; but we have no evidence on this head, for the sexes rarely differ in colour. Blind beetles, which cannot of course behold each other's beauty, never exhibit, as I hear from Mr. Waterhouse, jun., bright colours, though they often have polished coats: but the explanation of their obscurity may be that blind insects inhabit caves and other obscure stations.

Some Longicorns, however, especially certain Prionidae, offer an exception to the common rule that the sexes of beetles do not differ in colour. Most of these insects are large and splendidly coloured. The males in the genus Pyrodes,[484] as I saw in Mr. Bates' collection, are generally redder but rather duller than the females, the latter being coloured of a more or less splendid golden green. On the other hand, in one species the male is golden-green, the female being richly tinted with red and purple. In the genus Esmeralda the sexes differ so greatly in colour that they have been ranked as distinct species: in one species both are of a beautiful shining green, but the male has a red thorax. On the whole, as far as I could judge, the females of those Prionidae, in which the sexes differ, are coloured more richly than the males; and this does not accord with the common rule in regard to colour when acquired through sexual selection.

A most remarkable distinction between the sexes of many beetles is presented by the great horns which rise from the head, thorax, or clypeus of the males; and in some few cases from the under surface of the body. These horns, in the great family of the Lamellicorns, resemble those of various quadrupeds, such as stags, rhinoceroses, &c., and are wonderful both from their size and diversified shapes. Instead of describing them, I have given figures of the males and females of some of the more remarkable forms. (Figs. 15 to 19.) The females generally exhibit rudiments of the horns in the form of small knobs or ridges; but some are destitute of even a rudiment. On the other hand, the horns are nearly as well developed in the female as in the male of _Phanaeus lancifer_; and only a little less well developed in the females of some other species of the same genus and of Copris. In the several subdivisions of the family, the differences in structure of the horns do not run parallel, as I am informed by Mr. Bates, with their more important and characteristic differences; thus within the same natural section of the genus Onthophagus, there are species which have either a single cephalic horn, or two distinct horns.

In almost all cases, the horns are remarkable from their excessive variability; so that a graduated series can be formed, from the most highly developed males to others so degenerate that they can barely be distinguished from the females. Mr. Walsh[485] found that in _Phanaeus carnifex_ the horns were thrice as long in some males as in others. Mr. Bates, after examining above a hundred males of _Onthophagus rangifer_ (fig. 19), thought that he had at last discovered a species in which the horns did not vary; but further research proved the contrary.

The extraordinary size of the horns, and their widely different structure in closely-allied forms, indicate that they have been formed for some important purpose; but their excessive variability in the males of the same species leads to the inference that this purpose cannot be of a definite nature. The horns do not show marks of friction, as if used for any ordinary work. Some authors suppose[486] that as the males wander much more than the females, they require horns as a defence against their enemies; but in many cases the horns do not seem well adapted for defence, as they are not sharp. The most obvious conjecture is that they are used by the males for fighting together; but they have never been observed to fight; nor could Mr. Bates, after a careful examination of numerous species, find any sufficient evidence in their mutilated or broken condition of their having been thus used. If the males had been habitual fighters, their size would probably have been increased through sexual selection, so as to have exceeded that of the female; but Mr. Bates, after comparing the two sexes in above a hundred species of the Copridae, does not find in well-developed individuals any marked difference in this respect. There is, moreover, one beetle, belonging to the same great division of the Lamellicorns, namely Lethrus, the males of which are known to fight, but they are not provided with horns, though their mandibles are much larger than those of the female.

The conclusion, which best agrees with the fact of the horns having been so immensely yet not fixedly developed,--as shewn by their extreme variability in the same species and by their extreme diversity in closely-allied species--is that they have been acquired as ornaments. This view will at first appear extremely improbable; but we shall hereafter find with many animals, standing much higher in the scale, namely fishes, amphibians, reptiles and birds, that various kinds of crests, knobs, horns and combs have been developed apparently for this sole purpose.

The males of _Onitis furcifer_ (fig. 20) are furnished with singular projections on their anterior femora, and with a great fork or pair of horns on the lower surface of the thorax. This situation seems extremely ill adapted for the display of these projections, and they may be of some real service; but no use can at present be assigned to them. It is a highly remarkable fact, that although the males do not exhibit even a trace of horns on the upper surface of the body, yet in the females a rudiment of a single horn on the head (fig. 21, _a_), and of a crest (_b_) on the thorax, are plainly visible. That the slight thoracic crest in the female is a rudiment of a projection proper to the male, though entirely absent in the male of this particular species, is clear: for the female of _Bubas bison_ (a form which comes next to _Onitis_) has a similar slight crest on the thorax, and the male has in the same situation a great projection. So again there can be no doubt that the little point (_a_) on the head of the female _Onitis furcifer_, as well of the females of two or three allied species, is a rudimentary representative of the cephalic horn, which is common to the males of so many lamellicorn beetles, as in Phanaeus, fig. 17. The males indeed of some unnamed beetles in the British Museum, which are believed actually to belong to the genus Onitis, are furnished with a similar horn. The remarkable nature of this case will be best perceived by an illustration: the Ruminant quadrupeds run parallel with the lamellicorn beetles, in some females possessing horns as large as those of the male, in others having them much smaller, or existing as mere rudiments (though this is as rare with ruminants as it is common with Lamellicorns), or in having none at all. Now if a new species of deer or sheep were discovered with the female bearing distinct rudiments of horns, whilst the head of the male was absolutely smooth, we should have a case like that of _Onitis furcifer_.

In this case the old belief of rudiments having been created to complete the scheme of nature is so far from holding good, that all ordinary rules are completely broken through. The view which seems the most probable is that some early progenitor of Onitis acquired, like other Lamellicorns, horns on the head and thorax, and then transferred them, in a rudimentary condition, as with so many existing species, to the female, by whom they have ever since been retained. The subsequent loss of the horns by the male may have resulted through the principle of compensation from the development of the projections on the lower surface, whilst the female has not been thus affected, as she is not furnished with these projections, and consequently has retained the rudiments of the horns on the upper surface. Although this view is supported by the case of Bledius immediately to be given, yet the projections on the lower surface differ greatly in structure and development in the males of the several species of Onitis, and are even rudimentary in some; nevertheless the upper surface in all these species is quite destitute of horns. As secondary sexual characters are so eminently variable, it is possible that the projections on the lower surface may have been first acquired by some progenitor of Onitis and produced their effect through compensation, and then have been in certain cases almost completely lost.

All the cases hitherto given refer to the Lamellicorns, but the males of some few other beetles, belonging to two widely distinct groups, namely, the Curculionidae and Staphylinidae, are furnished with horns,--in the former on the lower surface of the body,[487] in the latter on the upper surface of the head and thorax. In the Staphylinidae the horns of the males in the same species are extraordinarily variable, just as we have seen with the Lamellicorns. In Siagonium we have a case of dimorphism, for the males can be divided into two sets, differing greatly in the size of their bodies, and in the development of their horns, without any intermediate gradations. In a species of Bledius (fig. 22), also belonging to the Staphylinidae, male specimens can be found in the same locality, as Professor Westwood states, "in which the central horn of the thorax is very large, but the horns of the head quite rudimental; and others, in which the thoracic horn is much shorter, whilst the protuberances on the head are long."[488] Here, then, we apparently have an instance of compensation of growth, which throws light on the curious case just given of the loss of the upper horns by the males of _Onitis furcifer_.

_Law of Battle._--Some male beetles, which seem ill fitted for fighting, nevertheless engage in conflicts for the possession of the females. Mr. Wallace[489] saw two males of _Leptorhynchus angustatus_, a linear beetle with a much elongated rostrum, "fighting for a female, who stood close by busy at her boring. They pushed at each other with their rostra, and clawed and thumped, apparently in the greatest rage." The smaller male, however, "soon ran away, acknowledging himself vanquished." In some few cases the males are well adapted for fighting, by possessing great toothed mandibles, much larger than those of the females. This is the case with the common stag-beetle (_Lucanus cervus_), the males of which emerge from the pupal state about a week before the other sex, so that several may often be seen pursuing the same female. At this period they engage in fierce conflicts. When Mr. A. H. Davis[490] enclosed two males with one female in a box, the larger male severely pinched the smaller one, until he resigned his pretensions. A friend informs me that when a boy he often put the males together to see them fight, and he noticed that they were much bolder and fiercer than the females, as is well known to be the case with the higher animals. The males would seize hold of his finger, if held in front, but not so the females. With many of the Lucanidae, as well as with the above-mentioned Leptorhynchus, the males are larger and more powerful insects than the females. The two sexes of _Lethrus cephalotes_ (one of the Lamellicorns) inhabit the same burrow; and the male has larger mandibles than the female. If, during the breeding-season, a strange male attempts to enter the burrow, he is attacked; the female does not remain passive, but closes the mouth of the burrow, and encourages her mate by continually pushing him on from behind. The action does not cease until the aggressor is killed or runs away.[491] The two sexes of another lamellicorn beetle, the _Ateuchus cicatricosus_ live in pairs, and seem much attached to each other; the male excites the female to roll the balls of dung in which the ova are deposited; and if she is removed, he becomes much agitated. If the male is removed, the female ceases all work, and as M. Brulerie[492] believes, would remain on the spot until she died.

The great mandibles of the male Lucanidae are extremely variable both in size and structure, and in this respect resemble the horns on the head and thorax of many male Lamellicorns and Staphylinidae. A perfect series can be formed from the best-provided to the worst-provided or degenerate males. Although the mandibles of the common stag-beetle, and probably of many other species, are used as efficient weapons for fighting, it is doubtful whether their great size can thus be accounted for. We have seen that with the _Lucanus elaphus_ of N. America they are used for seizing the female. As they are so conspicuous and so elegantly branched, the suspicion has sometimes crossed my mind that they may be serviceable to the males as an ornament, in the same manner as the horns on the head and thorax of the various above described species. The male _Chiasognathus grantii_ of S. Chile--a splendid beetle belonging to the same family--has enormously-developed mandibles (fig. 23); he is bold and pugnacious; when threatened on any side he faces round, opening his great jaws, and at the same time stridulating loudly; but the mandibles were not strong enough to pinch my finger so as to cause actual pain.

Sexual selection, which implies the possession of considerable perceptive powers and of strong passions, seems to have been more effective with the Lamellicorns than with any other family of the Coleoptera or beetles. With some species the males are provided with weapons for fighting; some live in pairs and show mutual affection; many have the power of stridulating when excited; many are furnished with the most extraordinary horns, apparently for the sake of ornament; some which are diurnal in their habits are gorgeously coloured; and, lastly, several of the largest beetles in the world belong to this family, which was placed by Linnaeus and Fabricius at the head of the Order of the Coleoptera.[493]

_Stridulating organs._--Beetles belonging to many and widely distinct families possess these organs. The sound can sometimes be heard at the distance of several feet or even yards,[494] but is not comparable with that produced by the Orthoptera. The part which may be called the rasp generally consists of a narrow slightly-raised surface, crossed by very fine, parallel ribs, sometimes so fine as to cause iridescent colours, and having a very elegant appearance under the microscope. In some cases, for instance, with Typhaeus, it could be plainly seen that extremely minute, bristly, scale-like prominences, which cover the whole surrounding surface in approximately parallel lines, give rise to the ribs of the rasp by becoming confluent and straight, and at the same time more prominent and smooth. A hard ridge on any adjoining part of the body, which in some cases is specially modified for the purpose, serves as the scraper for the rasp. The scraper is rapidly moved across the rasp, or conversely the rasp across the scraper.

These organs are situated in widely different positions. In the carrion-beetles (Necrophorus) two parallel rasps (_r_, fig. 24) stand on the dorsal surface of the fifth abdominal segment, each rasp being crossed, as described by Landois,[495] by from 126 to 140 fine ribs. These ribs are scraped by the posterior margins of the elytra, a small portion of which projects beyond the general outline. In many Crioceridae, and in _Clythra 4-punctata_ (one of the Chrysomelidae), and in some Tenebrionidae, &c.,[496] the rasp is seated on the dorsal apex of the abdomen, on the pygidium or pro-pygidium, and is scraped as above by the elytra. In Heterocerus, which belongs to another family, the rasps are placed on the sides of the first abdominal segment, and are scraped by ridges on the femora.[497] In certain Curculionidae and Carabidae,[498] the parts are completely reversed in position, for the rasps are seated on the inferior surface of the elytra, near their apices, or along their outer margins, and the edges of the abdominal segments serve as the scrapers. In _Pelobius hermanni_ (one of Dytiscidae or water-beetles) a strong ridge runs parallel and near to the sutural margin of the elytra, and is crossed by ribs, coarse in the middle part, but becoming gradually finer at both ends, especially at the upper end; when this insect is held under water or in the air, a stridulating noise is produced by scraping the extreme horny margin of the abdomen against the rasp. In a great number of long-horned beetles (Longicornia) the organs are altogether differently situated, the rasp being on the meso-thorax, which is rubbed against the pro-thorax; Landois counted 238 very fine ribs on the rasp of _Cerambyx heros_.

Many Lamellicorns have the power of stridulating, and the organs differ greatly in position. Some species stridulate very loudly, so that when Mr. F. Smith caught a _Trox sabulosus_, a gamekeeper who stood by thought that he had caught a mouse; but I failed to discover the proper organs in this beetle. In Geotrupes and Typhaeus a narrow ridge runs obliquely across (_r_, fig. 25) the coxa of each hind-leg, having in _G. stercorarius_ 84 ribs, which are scraped by a specially-projecting part of one of the abdominal segments. In the nearly allied _Copris lunaris_, an excessively narrow fine rasp runs along the sutural margin of the elytra, with another short rasp near the basal outer margin; but in some other Coprini the rasp is seated, according to Leconte,[499] on the dorsal surface of the abdomen. In Oryctes it is seated on the pro-pygidium, and in some other Dynastini, according to the same entomologist, on the under surface of the elytra. Lastly, Westring states that in _Omaloplia brunnea_ the rasp is placed on the pro-sternum, and the scraper on the meta-sternum, the parts thus occupying the under surface of the body, instead of the upper surface as in the Longicorns.

We thus see that the stridulating organs in the different coleopterous families are wonderfully diversified in position, but not much in structure. Within the same family some species are provided with these organs, and some are quite destitute of them. This diversity is intelligible, if we suppose that originally various species made a shuffling or hissing noise by the rubbing together of the hard and rough parts of their bodies which were in contact; and that from the noise thus produced being in some way useful, the rough surfaces were gradually developed into regular stridulating organs. Some beetles as they move, now produce, either intentionally or unintentionally, a shuffling noise, without possessing any proper organs for the purpose. Mr. Wallace informs me that the _Euchirus longimanus_ (a Lamellicorn, with the anterior legs wonderfully elongated in the male) "makes, whilst moving, a low hissing sound by the protrusion and contraction of the abdomen; and when seized it produces a grating sound by rubbing its hind-legs against the edges of the elytra." The hissing sound is clearly due to a narrow rasp running along the sutural margin of each elytron; and I could likewise make the grating sound by rubbing the shagreened surface of the femur against the granulated margin of the corresponding elytron; but I could not here detect any proper rasp; nor is it likely that I could have overlooked it in so large an insect. After examining Cychrus and reading what Westring has written in his two papers about this beetle, it seems very doubtful whether it possesses any true rasp, though it has the power of emitting a sound.

From the analogy of the Orthoptera and Homoptera, I expected to find that the stridulating organs in the Coleoptera differed according to sex; but Landois, who has carefully examined several species, observed no such difference; nor did Westring; nor did Mr. G. R. Crotch in preparing the numerous specimens which he had the kindness to send me for examination. Any slight sexual difference, however, would be difficult to detect, on account of the great variability of these organs. Thus in the first pair of the _Necrophorus humator_ and of the _Pelobius_ which I examined, the rasp was considerably larger in the male than in the female; but not so with succeeding specimens. In _Geotrupes stercorarius_ the rasp appeared to me thicker, opaquer, and more prominent in three males than in the same number of females; consequently my son, Mr. F. Darwin, in order to discover whether the sexes differed in their power of stridulating, collected 57 living specimens, which he separated into two lots, according as they made, when held in the same manner, a greater or lesser noise. He then examined their sexes, but found that the males were very nearly in the same proportion to the females in both lots. Mr. F. Smith has kept alive numerous specimens of _Mononychus pseudacori_ (Curculionidae), and is satisfied that both sexes stridulate, and apparently in an equal degree.

Nevertheless the power of stridulating is certainly a sexual character in some few Coleoptera. Mr. Crotch has discovered that the males alone of two species of Heliopathes (Tenebrionidae) possess stridulating organs. I examined five males of _H. gibbus_, and in all these there was a well-developed rasp, partially divided into two, on the dorsal surface of the terminal abdominal segment; whilst in the same number of females there was not even a rudiment of the rasp, the membrane of this segment being transparent and much thinner than in the male. In _H. cribratostriatus_ the male has a similar rasp, excepting that it is not partially divided into two portions, and the female is completely destitute of this organ; but in addition the male has on the apical margins of the elytra, on each side of the suture, three or four short longitudinal ridges, which are crossed by extremely fine ribs, parallel to and resembling those on the abdominal rasp; whether these ridges serve as an independent rasp, or as a scraper for the abdominal rasp, I could not decide: the female exhibits no trace of this latter structure.

Again, in three species of the Lamellicorn genus Oryctes, we have a nearly parallel case. In the females of _O. gryphus_ and _nasicornis_ the ribs on the rasp of the pro-pygidium are less continuous and less distinct than in the males; but the chief difference is that the whole upper surface of this segment, when held in the proper light, is seen to be clothed with hairs, which are absent or are represented by excessively fine down in the males. It should be noticed that in all Coleoptera the effective part of the rasp is destitute of hairs. In _O. senegalensis_ the difference between the sexes is more strongly marked, and this is best seen when the proper segment is cleaned and viewed as a transparent object. In the female the whole surface is covered with little separate crests, bearing spines; whilst in the male these crests become, in proceeding towards the apex, more and more confluent, regular, and naked; so that three-fourths of the segment is covered with extremely fine parallel ribs, which are quite absent in the female. In the females, however, of all three species of Oryctes, when the abdomen of a softened specimen is pushed backwards and forwards, a slight grating or stridulating sound can be produced.

In the case of the Heliopathes and Oryctes there can hardly be a doubt that the males stridulate in order to call or to excite the females; but with most beetles the stridulation apparently serves both sexes as a mutual call. This view is not rendered improbable from beetles stridulating under various emotions; we know that birds use their voices for many purposes besides singing to their mates. The great Chiasognathus stridulates in anger or defiance; many species do the same from distress or fear, when held so that they cannot escape; Messrs. Wollaston and Crotch were able, by striking the hollow stems of trees in the Canary Islands, to discover the presence of beetles belonging to the genus Acalles by their stridulation. Lastly the male Ateuchus stridulates to encourage the female in her work, and from distress when she is removed.[500] Some naturalists believe that beetles make this noise to frighten away their enemies; but I cannot think that the quadrupeds and birds which are able to devour the larger beetles with their extremely hard coats, would be frightened by so slight a grating sound. The belief that the stridulation serves as a sexual call is supported by the fact that death-ticks (_Anobium tessellatum_) are well known to answer each other's ticking, or, as I have myself observed, a tapping noise artificially made; and Mr. Doubleday informs me that he has twice or thrice observed a female ticking,[501] and in the course of an hour or two has found her united with a male, and on one occasion surrounded by several males. Finally, it seems probable that the two sexes of many kinds of beetles were at first enabled to find each other by the slight shuffling noise produced by the rubbing together of the adjoining parts of their hard bodies; and that as the males or females which made the greatest noise succeeded best in finding partners, the rugosities on various parts of their bodies were gradually developed by means of sexual selection into true stridulating organs.