The males of Onitis furcifer (Fig. 21), and of some other species of the genus, 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. Judging from other insects, these may aid the male in clinging to the female. Although the males have not even a trace of a horn on the upper surface of the body, yet the females plainly exhibit a rudiment of a single horn on the head (Fig. 22, a), and of a crest (b) on the thorax. 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 genus which comes next to Onitis) has a similar slight crest on the thorax, and the male bears a great projection in the same situation. So, again, there can hardly be a doubt that the little point (a) on the head of the female Onitis furcifer, as well as on the head 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. 18).
The old belief that rudiments have been created to complete the scheme of nature is here so far from holding good, that we have a complete inversion of the ordinary state of things in the family. We may reasonably suspect that the males originally bore horns and transferred them to the females in a rudimentary condition, as in so many other Lamellicorns. Why the males subsequently lost their horns, we know not; but this may have been caused through the principle of compensation, owing to the development of the large horns and projections on the lower surface; and as these are confined to the males, the rudiments of the upper horns on the females would not have been thus obliterated.
[Fig. 23. Bledius taurus, magnified. Left-hand figure, male; right-hand figure, female.]
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 (66. Kirby and Spence, ‘Introduction to Entomology,’ vol. iii. p. 329.), in the latter on the upper surface of the head and thorax. In the Staphylinidae, the horns of the males are extraordinarily variable in the same species, 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 intermediate gradations. In a species of Bledius (Fig. 23), also belonging to the Staphylinidae, Professor Westwood states that, “male specimens can be found in the same locality 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.” (67. ‘Modern Classification of Insects,’ vol. i. p. 172: Siagonium, p. 172. In the British Museum I noticed one male specimen of Siagonium in an intermediate condition, so that the dimorphism is not strict.) Here we apparently have a case of compensation, which throws light on that just given, of the supposed loss of the upper horns by the males of Onitis.
LAW OF BATTLE.
Some male beetles, which seem ill-fitted for fighting, nevertheless engage in conflicts for the possession of the females. Mr. Wallace (68. ‘The Malay Archipelago,’ vol. ii. 1869, p. 276. Riley, Sixth ‘Report on Insects of Missouri,’ 1874, p. 115.) 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 male beetles 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 season they engage in fierce conflicts. When Mr. A.H. Davis (69. ‘Entomological Magazine,’ vol. i. 1833, p. 82. See also on the conflicts of this species, Kirby and Spence, ibid. vol. iii. p. 314; and Westwood, ibid. vol. i. p. 187.) 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 with the higher animals. The males would seize hold of his finger, if held in front of them, but not so the females, although they have stronger jaws. The males of many of the Lucanidae, as well as of the above-mentioned Leptorhynchus, 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; and the battle lasts until the aggressor is killed or runs away. (70. Quoted from Fischer, in ‘Dict. Class. d’Hist. Nat.’ tom. x. p. 324.) The two sexes of another Lamellicorn beetle, the Ateuchus cicatricosus, live in pairs, and seem much attached to each other; the male excites the females 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 believes, would remain on the same spot until she died. (71. ‘Ann. Soc. Entomolog. France,’ 1866, as quoted in ‘Journal of Travel,’ by A. Murray, 1868, p. 135.)
[Fig. 24. Chiasognathus Grantii, reduced. Upper figure, male; lower figure, female.]
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 they are used by the Lucanus elaphus of N. America for seizing the female. As they are so conspicuous and so elegantly branched, and as owing to their great length they are not well adapted for pinching, the suspicion has crossed my mind that they may in addition serve as an ornament, like the horns on the head and thorax of the various species above described. The male Chiasognathus grantii of S. Chile—a splendid beetle belonging to the same family—has enormously developed mandibles (Fig. 24); he is bold and pugnacious; when threatened he faces round, opens his great jaws, and at the same time stridulates 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 beetles. With some species the males are provided with weapons for fighting; some live in pairs and shew mutual affection; many have the power of stridulating when excited; many are furnished with the most extraordinary horns, apparently for the sake of ornament; and some, which are diurnal in their habits, are gorgeously coloured. Lastly, several of the largest beetles in the world belong to this family, which was placed by Linnaeus and Fabricius as the head of the Order. (72. Westwood, ‘Modern Classification,’ vol. i. p. 184.)