Fig. 18. Dipelicus cantori.
Fig. 19. Onthophagus rangifer, enlarged.
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 Phanæus 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 Phanæus 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 Copridæ, 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.