I then applied to Mr. Wallace, who has an innate genius for solving difficulties. After some consideration he replied: “Most caterpillars require protection, as may be inferred from some kinds being furnished with spines or irritating hairs, and from many being coloured green like the leaves on which they feed, or curiously like the twigs of the trees on which they live.” I may add as another instance of protection, that there is a caterpillar of a moth, as I am informed by Mr. J. Mansel Weale, which lives on the mimosas in South Africa, and fabricates for itself a case, quite undistinguishable from the surrounding thorns. From such considerations Mr. Wallace thought it probable that conspicuously-coloured caterpillars were protected by having a nauseous taste; but as their skin is extremely tender, and as their intestines readily protrude from a wound, a slight peck from the beak of a bird would be as fatal to them as if they had been devoured. Hence, as Mr. Wallace remarks, “distastefulness alone would be insufficient to protect a caterpillar unless some outward sign indicated to its would-be destroyer that its prey was a disgusting morsel.” Under these circumstances it would be highly advantageous to a caterpillar to be instantaneously and certainly recognised as unpalatable by all birds and other animals.417 Thus the most gaudy colours would be serviceable, and might have been gained by variation and the survival of the most easily-recognised individuals.
This hypothesis appears at first sight very bold; but when it was brought before the Entomological Society533 it was supported by various statements; and Mr. J. Jenner Weir, who keeps a large number of birds in an aviary, has made, as he informs me, numerous trials, and finds no exception to the rule, that all caterpillars of nocturnal and retiring habits with smooth skins, all of a green colour, and all which imitate twigs, are greedily devoured by his birds. The hairy and spinose kinds are invariably rejected, as were four conspicuously-coloured species. When the birds rejected a caterpillar, they plainly shewed, by shaking their heads and cleansing their beaks, that they were disgusted by the taste.534 Three conspicuous kinds of caterpillars and moths were also given by Mr. A. Butler to some lizards and frogs, and were rejected; though other kinds were eagerly eaten. Thus the probable truth of Mr. Wallace’s view is confirmed, namely, that certain caterpillars have been made conspicuous for their own good, so as to be easily recognised by their enemies, on nearly the same principle that certain poisons are coloured by druggists for the good of man. This view will, it is probable, be hereafter extended to many animals, which are coloured in a conspicuous manner.
Summary and Concluding Remarks on Insects.—Looking back to the several Orders, we have seen that the sexes often differ in various characters, the meaning 418of which is not understood. The sexes, also, often differ in their organs of sense or locomotion, so that the males may quickly discover or reach the females, and still oftener in the males possessing diversified contrivances for retaining the females when found. But we are not here much concerned with sexual differences of these kinds.
In almost all the Orders, the males of some species, even of weak and delicate kinds, are known to be highly pugnacious; and some few are furnished with special weapons for fighting with their rivals. But the law of battle does not prevail nearly so widely with insects as with the higher animals. Hence probably it is that the males have not often been rendered larger and stronger than the females. On the contrary they are usually smaller, in order that they may be developed within a shorter time, so as to be ready in large numbers for the emergence of the females.
In two families of the Homoptera the males alone possess, in an efficient state, organs which may be called vocal; and in three families of the Orthoptera the males alone possess stridulating organs. In both cases these organs are incessantly used during the breeding-season, not only for calling the females, but for charming or exciting them in rivalry with other males. No one who admits the agency of natural selection, will dispute that these musical instruments have been acquired through sexual selection. In four other Orders the members of one sex, or more commonly of both sexes, are provided with organs for producing various sounds, which apparently serve merely as call-notes. Even when both sexes are thus provided, the individuals which were able to make the loudest or most continuous noise would gain partners before those which were less noisy, so that their organs have probably been gained419 through sexual selection. It is instructive to reflect on the wonderful diversity of the means for producing sound, possessed by the males alone or by both sexes in no less than six Orders, and which were possessed by at least one insect at an extremely remote geological epoch. We thus learn how effectual sexual selection has been in leading to modifications of structure, which sometimes, as with the Homoptera, are of an important nature.
From the reasons assigned in the last chapter, it is probable that the great horns of the males of many lamellicorn, and some other beetles, have been acquired as ornaments. So perhaps it may be with certain other peculiarities confined to the male sex. From the small size of insects, we are apt to undervalue their appearance. If we could imagine a male Chalcosoma (fig. 15) with its polished, bronzed coat of mail, and vast complex horns, magnified to the size of a horse or even of a dog, it would be one of the most imposing animals in the world.
The colouring of insects is a complex and obscure subject. When the male differs slightly from the female, and neither are brilliantly coloured, it is probable that the two sexes have varied in a slightly different manner, with the variations transmitted to the same sex, without any benefit having been thus derived or evil suffered. When the male is brilliantly coloured and differs conspicuously from the female, as with some dragon-flies and many butterflies, it is probable that he alone has been modified, and that he owes his colours to sexual selection; whilst the female has retained a primordial or very ancient type of colouring, slightly modified by the agencies before explained, and has therefore not been rendered obscure, at least in most cases, for the sake of protection. But the female alone has some420times been coloured brilliantly so as to imitate other protected species inhabiting the same district. When the sexes resemble each other and both are obscurely coloured, there is no doubt that they have been in a multitude of cases coloured for the sake of protection. So it is in some instances when both are brightly coloured, causing them to resemble surrounding objects such as flowers, or other protected species, or indirectly by giving notice to their enemies that they are of an unpalatable nature. In many other cases in which the sexes resemble each other and are brilliantly coloured, especially when the colours are arranged for display, we may conclude that they have been gained by the male sex as an attraction, and have been transferred to both sexes. We are more especially led to this conclusion whenever the same type of coloration prevails throughout a group, and we find that the males of some species differ widely in colour from the females, whilst both sexes of other species are quite alike, with intermediate gradations connecting these extreme states.
In the same manner as bright colours have often been partially transferred from the males to the females, so it has been with the extraordinary horns of many lamellicorn and some other beetles. So, again, the vocal or instrumental organs proper to the males of the Homoptera and Orthoptera have generally been transferred in a rudimentary, or even in a nearly perfect condition to the females; yet not sufficiently perfect to be used for producing sound. It is also an interesting fact, as bearing on sexual selection, that the stridulating organs of certain male Orthoptera are not fully developed until the last moult; and that the colours of certain male dragon-flies are not fully developed until some little time after their emergence from the pupal state, and when they are ready to breed.
421Sexual selection implies that the more attractive individuals are preferred by the opposite sex; and as with insects, when the sexes differ, it is the male which, with rare exceptions, is the most ornamented and departs most from the type to which the species belongs;—and as it is the male which searches eagerly for the female, we must suppose that the females habitu
ally or occasionally prefer the more beautiful males, and that these have thus acquired their beauty. That in most or all the orders the females have the power of rejecting any particular male, we may safely infer from the many singular contrivances possessed by the males, such as great jaws, adhesive cushions, spines, elongated legs, &c., for seizing the female; for these contrivances shew that there is some difficulty in the act. In the case of unions between distinct species, of which many instances have been recorded, the female must have been a consenting party. Judging from what we know of the perceptive powers and affections of various insects, there is no antecedent improbability in sexual selection having come largely into action; but we have as yet no direct evidence on this head, and some facts are opposed to the belief. Nevertheless, when we see many males pursuing the same female, we can hardly believe that the pairing is left to blind chance—that the female exerts no choice, and is not influenced by the gorgeous colours or other ornaments, with which the male alone is decorated.
If we admit that the females of the Homoptera and Orthoptera appreciate the musical tones emitted by their male partners, and that the various instruments for this purpose have been perfected through sexual selection, there is little improbability in the females of other insects appreciating beauty in form or colour, and consequently in such characters having been thus gained422 by the males. But from the circumstance of colour being so variable, and from its having been so often modified for the sake of protection, it is extremely difficult to decide in how large a proportion of cases sexual selection has come into play. This is more especially difficult in those Orders, such as the Orthoptera, Hymenoptera, and Coleoptera, in which the two sexes rarely differ much in colour; for we are thus cut off from our best evidence of some relation between the reproduction of the species and colour. With the Coleoptera, however, as before remarked, it is in the great lamellicorn group, placed by some authors at the head of the Order, and in which we sometimes see a mutual attachment between the sexes, that we find the males of some species possessing weapons for sexual strife, others furnished with wonderful horns, many with stridulating organs, and others ornamented with splendid metallic tints. Hence it seems probable that all these characters have been gained through the same means, namely sexual selection.
When we treat of Birds, we shall see that they present in their secondary sexual characters the closest analogy with insects. Thus, many male birds are highly pugnacious, and some are furnished with special weapons for fighting with their rivals. They possess organs which are used during the breeding-season for producing vocal and instrumental music. They are frequently ornamented with combs, horns, wattles and plumes of the most diversified kinds, and are decorated with beautiful colours, all evidently for the sake of display. We shall find that, as with insects, both sexes, in certain groups, are equally beautiful, and are equally provided with ornaments which are usually confined to the male sex. In other groups both sexes are equally plain-coloured and unornamented. Lastly, in some few423 anomalous cases, the females are more beautiful than the males. We shall often find, in the same group of birds, every gradation from no difference between the sexes, to an extreme difference. In the latter case we shall see that the females, like female insects, often possess more or less plain traces of the characters which properly belong to the males. The analogy, indeed, in all these respects between birds and insects, is curiously close. Whatever explanation applies to the one class probably applies to the other; and this explanation, as we shall hereafter attempt to shew, is almost certainly sexual selection.
FOOTNOTES:
1 As the works of the first-named authors are so well known, I need not give the titles; but as those of the latter are less well known in England, I will give them:—‘Sechs Vorlesungen über die Darwin’sche Theorie:’ zweite Auflage, 1868, von Dr. L. Büchner; translated into French under the title ‘Conférences sur la Théorie Darwinienne,’ 1869. ‘Der Mensch, im Lichte der Darwin’sche Lehre,’ 1865, von Dr. F. Rolle. I will not attempt to give references to all the authors who have taken the same side of the question. Thus G. Canestrini has published (‘Annuario della Soc. d. Nat.,’ Modena, 1867, p. 81) a very curious paper on rudimentary characters, as bearing on the origin of man. Another work has (1869) been published by Dr. Barrago Francesco, bearing in Italian the title of “Man, made in the image of God, was also made in the image of the ape.”
2 Prof. Häckel is the sole author who, since the publication of the ‘Origin,’ has discussed, in his various works, in a very able manner, the subject of sexual selection, and has seen its full importance.
3 ‘Grosshirnwindungen des Menschen,’ 1868, s. 96.
4 ‘Leç. sur la Phys.’ 1866, p. 890, as quoted by M. Dally, ‘L’Ordre des Primates et le Transformisme,’ 1868, p. 29.
5 ‘Naturgeschichte der Säugethiere von Paraguay,’ 1830, s. 50.
6 Brehm, ‘Thierleben,’ B. i. 1864, s. 75, 86. On the Ateles, s. 105. For other analogous statements, see s. 25, 107.
7 With respect to insects see Dr. Laycock ‘On a General Law of Vital Periodicity,’ British Association, 1842. Dr. Macculloch, ‘Silliman’s North American Journal of Science,’ vol. xvii. p. 305, has seen a dog suffering from tertian ague.
8 I have given the evidence on this head in my ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 15.
9 “Mares e diversis generibus Quadrumanorum sine dubio dignoscunt feminas humanas a maribus. Primum, credo, odoratu, postea aspectu. Mr. Youatt, qui diu in Hortis Zoologicis (Bestiariis) medicus animalium erat, vir in rebus observandis cautus et sagax, hoc mihi certissime probavit, et curatores ejusdem loci et alii e ministris confirmaverunt. Sir Andrew Smith et Brehm notabant idem in Cynocephalo. Illustrissimus Cuvier etiam narrat multa de hac re quâ ut opinor nihil turpius potest indicari inter omnia hominibus et Quadrumanis communia. Narrat enim Cynocephalum quendam in furorem incidere aspectu feminarum aliquarum, sed nequaquam accendi tanto furore ab omnibus. Semper eligebat juniores, et dignoscebat in turba, et advocabat voce gestuque.”
10 This remark is made with respect to Cynocephalus and the anthropomorphous apes by Geoffroy Saint-Hilaire and F. Cuvier, ‘Hist. Nat. des Mammifères,’ tom. i. 1824.
11 Huxley, ‘Man’s Place in Nature,’ 1863, p. 34.
12 ‘Man’s Place in Nature,’ 1863, p. 67.
13 The human embryo (upper fig.) is from Ecker, ‘Icones Phys.,’ 1851-1859, tab. xxx. fig. 2. This embryo was ten lines in length, so that the drawing is much magnified. The embryo of the dog is from Bischoff, ‘Entwicklungsgeschichte des Hunde-Eies,’ 1845, tab. xi. fig. 42 B. This drawing is five times magnified, the embryo being 25 days old. The internal viscera have been omitted, and the uterine appendages in both drawings removed. I was directed to these figures by Prof. Huxley, from whose work, ‘Man’s Place in Nature.’ the idea of giving them was taken. Häckel has also given analogous drawings in his ‘Schöpfungsgeschichte.’
14 Prof. Wyman in ‘Proc. of American Acad. of Sciences,’ vol. iv. 1860, p. 17.
15 Owen, ‘Anatomy of Vertebrates,’ vol. i. p. 533.
16 ‘Die Grosshirnwindungen des Menschen,’ 1868, s. 95.
17 ‘Anatomy of Vertebrates,’ vol. ii. p. 553.
18 ‘Proc. Soc. Nat. Hist.’ Boston, 1863, vol. ix. p. 185.
19 ‘Man’s Place in Nature,’ p. 65.
20 I had written a rough copy of this chapter before reading a valuable paper, “Caratteri rudimentali in ordine all’origine del uomo” (‘Annuario della Soc. d. Nat.,’ Modena, 1867, p. 81), by G. Canestrini, to which paper I am considerably indebted. Häckel has given admirable discussions on this whole subject, under the title of Dysteleology, in his ‘Generelle Morphologie’ and ‘Schöpfungsgeschichte.’
21 Some good criticisms on this subject have been given by Messrs. Murie and Mivart, in ‘Transact. Zoolog. Soc.’ 1869, vol. vii. p. 92.
22 ‘Variation of Animals and Plants under Domestication,’ vol. ii. pp. 317 and 397. See also ‘Origin of Species,
’ 5th edit. p. 535.
23 For instance M. Richard (‘Annales des Sciences Nat.’ 3rd series, Zoolog. 1852, tom. xviii. p. 13) describes and figures rudiments of what he calls the “muscle pédieux de la main,” which he says is sometimes “infiniment petit.” Another muscle, called “le tibial postérieur,” is generally quite absent in the hand, but appears from time to time in a more or less rudimentary condition.
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