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 in 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 für 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.
It is an interesting fact, first noticed by Schelver, that, in several genera belonging to two sub-families, the males on first emergence 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 (53. See abstract in the ‘Zoological Record’ for 1867, p. 450.), a curious case of dimorphism, some of the females having ordinary wings, whilst others 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, which has been abruptly transferred to some of the females, instead of, as generally occurs, to all of them.” Mr. MacLachlan informs me of another instance of dimorphism in several species of Agrion, in which some individuals are 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 orange or yellow; so that supposing Agrion to be descended from some primordial form which resembled the typical Libellulae in its sexual characters, it would not be surprising that a tendency to vary in this manner should occur in the females alone.