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 Curculionidæ and Staphylinidæ, 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 Staphylinidæ 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 Staphylinidæ, 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 Lucanidæ, 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 Lucanidæ 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 Staphylinidæ. 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
Fig. 23. Chiasognathus grantii, reduced. Upper figure, male; lower figure, female. 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 Linnæus 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 Typhæus, 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.
Fig. 24. Necrophorus (from Landois). r. The two rasps. Left-hand figure, part of the rasp highly magnified.