CHAPTER XI.

INSECTS, _continued_.--ORDER LEPIDOPTERA.

Courtship of butterflies--Battles--Ticking noise--Colours common to both sexes, or more brilliant in the males--Examples--Not due to the direct action of the conditions of life--Colours adapted for protection--Colours of moths--Display--Perceptive powers of the Lepidoptera--Variability--Causes of the difference in colour between the males and females--Mimickry, female butterflies more brilliantly coloured than the males--Bright colours of caterpillars--Summary and concluding remarks on the secondary sexual characters of insects--Birds and insects compared.

In this great Order the most interesting point for us is the difference in colour between the sexes of the same species, and between the distinct species of the same genus. Nearly the whole of the following chapter will be devoted to this subject; but I will first make a few remarks on one or two other points. Several males may often be seen pursuing and crowding round the same female. Their courtship appears to be a prolonged affair, for I have frequently watched one or more males pirouetting round a female until I became tired, without seeing the end of the courtship. Although butterflies are such weak and fragile creatures, they are pugnacious, and an Emperor butterfly[502] has been captured with the tips of its wings broken from a conflict with another male. Mr. Collingwood in speaking of the frequent battles between the butterflies of Borneo says, "They whirl round each other with the greatest rapidity, and appear to be incited by the greatest ferocity." One case is known of a butterfly, namely the _Ageronia feronia_, which makes a noise like that produced by a toothed wheel passing under a spring catch, and which could be heard at the distance of several yards. At Rio de Janeiro this sound was noticed by me, only when two were chasing each other in an irregular course, so that it is probably made during the courtship of the sexes; but I neglected to attend to this point.[503]

Every one has admired the extreme beauty of many butterflies and of some moths; and we are led to ask, how has this beauty been acquired? Have their colours and diversified patterns simply resulted from the direct action of the physical conditions to which these insects have been exposed, without any benefit being thus derived? Or have successive variations been accumulated and determined either as a protection or for some unknown purpose, or that one sex might be rendered attractive to the other? And, again, what is the meaning of the colours being widely different in the males and females of certain species, and alike in the two sexes of other species? Before attempting to answer these questions a body of facts must be given.

With most of our English butterflies, both those which are beautiful, such as the admiral, peacock, and painted lady (Vanessae), and those which are plain-coloured, such as the meadow-browns (Hipparchiae), the sexes are alike. This is also the case with the magnificent Heliconidae and Danaidae of the tropics. But in certain other tropical groups, and with some of our English butterflies, as the purple emperor, orange-tip, &c. (_Apatura Iris_ and _Anthocharis cardamines_), the sexes differ either greatly or slightly in colour. No language suffices to describe the splendour of the males of some tropical species. Even within the same genus we often find species presenting an extraordinary difference between the sexes, whilst others have their sexes closely alike. Thus in the South American genus Epicalia, Mr. Bates, to whom I am much indebted for most of the following facts and for looking over this whole discussion, informs me that he knows twelve species, the two sexes of which haunt the same stations (and this is not always the case with butterflies), and therefore cannot have been differently affected by external conditions[504]. In nine of these species the males rank amongst the most brilliant of all butterflies, and differ so greatly from the comparatively plain females that they were formerly placed in distinct genera. The females of these nine species resemble each other in their general type of coloration, and likewise resemble both sexes in several allied genera, found in various parts of the world. Hence in accordance with the descent-theory we may infer that these nine species, and probably all the others of the genus, are descended from an ancestral form which was coloured in nearly the same manner. In the tenth species the female still retains the same general colouring, but the male resembles her, so that he is coloured in a much less gaudy and contrasted manner than the males of the previous species. In the eleventh and twelfth species, the females depart from the type of colouring which is usual with their sex in this genus, for they are gaily decorated in nearly the same manner as the males, but in a somewhat less degree. Hence in these two species the bright colours of the males seem to have been transferred to the females; whilst the male of the tenth species has either retained or recovered the plain colours of the female as well as of the parent-form of the genus; the two sexes being thus rendered in both cases, though in an opposite manner, nearly alike. In the allied genus Eubagis, both sexes of some of the species are plain-coloured and nearly alike; whilst with the greater number the males are decorated with beautiful metallic tints, in a diversified manner, and differ much from their females. The females throughout the genus retain the same general style of colouring, so that they commonly resemble each other much more closely than they resemble their own proper males.

In the genus Papilio, all the species of the AEneas group are remarkable for their conspicuous and strongly contrasted colours, and they illustrate the frequent tendency to gradation in the amount of difference between the sexes. In a few species, for instance in _P. ascanius_, the males and females are alike; in others the males are a little or very much more superbly coloured than the females. The genus Junonia allied to our Vanessae offers a nearly parallel case, for although the sexes of most of the species resemble each other and are destitute of rich colours, yet in certain species, as in _J. oenone_, the male is rather more brightly coloured than the female, and in a few (for instance _J. andremiaja_) the male is so different from the female that he might be mistaken for an entirely distinct species.

Another striking case was pointed out to me in the British museum by Mr. A. Butler, namely one of the Tropical American Theclae, in which both sexes are nearly alike and wonderfully splendid; in another, the male is coloured in a similarly gorgeous manner, whilst the whole upper surface of the female is of a dull uniform brown. Our common little English blue butterflies of the genus Lycaena, illustrate the various differences in colour between the sexes, almost as well, though not in so striking a manner, as the above exotic genera. In _Lycaena agestis_ both sexes have wings of a brown colour, bordered with small ocellated orange spots, and are consequently alike. In _L. oegon_ the wings of the male are of a fine blue, bordered with black; whilst the wings of the female are brown, with a similar border, and closely resemble those of _L. agestis_. Lastly, in _L. arion_ both sexes are of a blue colour and nearly alike, though in the female the edges of the wings are rather duskier, with the black spots plainer; and in a bright blue Indian species both sexes are still more closely alike.

I have given the foregoing cases in some detail in order to shew, in the first place, that when the sexes of butterflies differ, the male as a general rule is the most beautiful, and departs most from the usual type of colouring of the group to which the species belongs. Hence in most groups the females of the several species resemble each other much more closely than do the males. In some exceptional cases, however, to which I shall hereafter allude, the females are coloured more splendidly than the males. In the second place these cases have been given to bring clearly before the mind that within the same genus, the two sexes frequently present every gradation from no difference in colour to so great a difference that it was long before the two were placed by entomologists in the same genus. In the third place, we have seen that when the sexes nearly resemble each other, this apparently may be due either to the male having transferred his colours to the female, or to the male having retained, or perhaps recovered, the primordial colours of the genus to which the species belongs. It also deserves notice that in those groups in which the sexes present any difference of colour, the females usually resemble the males to a certain extent, so that when the males are beautiful to an extraordinary degree, the females almost invariably exhibit some degree of beauty. From the numerous cases of gradation in the amount of difference between the sexes, and from the prevalence of the same general type of coloration throughout the whole of the same group, we may conclude that the causes, whatever they may be, which have determined the brilliant colouring of the males alone of some species, and of both sexes in a more or less equal degree of other species, have generally been the same.

As so many gorgeous butterflies inhabit the tropics, it has often been supposed that they owe their colours to the great heat and moisture of these zones; but Mr. Bates[505] has shewn by the comparison of various closely-allied groups of insects from the temperate and tropical regions, that this view cannot be maintained; and the evidence becomes conclusive when brilliantly-coloured males and plain-coloured females of the same species inhabit the same district, feed on the same food, and follow exactly the same habits of life. Even when the sexes resemble each other, we can hardly believe that their brilliant and beautifully-arranged colours are the purposeless result of the nature of the tissues, and the action of the surrounding conditions.

With animals of all kinds, whenever colour has been modified for some special purpose, this has been, as far as we can judge, either for protection or as an attraction between the sexes. With many species of butterflies the upper surfaces of the wings are obscurely coloured, and this in all probability leads to their escaping observation and danger. But butterflies when at rest would be particularly liable to be attacked by their enemies; and almost all the kinds when resting raise their wings vertically over their backs, so that the lower sides alone are exposed to view. Hence it is this side which in many cases is obviously coloured so as to imitate the surfaces on which these insects commonly rest. Dr. Roessler, I believe, first noticed the similarity of the closed wings of certain Vanessae and other butterflies to the bark of trees. Many analogous and striking facts could be given. The most interesting one is that recorded by Mr. Wallace[506] of a common Indian and Sumatran butterfly (Kallima), which disappears like magic when it settles in a bush; for it hides its head and antennae between its closed wings, and these in form, colour, and veining cannot be distinguished from a withered leaf together with the footstalk. In some other cases the lower surfaces of the wings are brilliantly coloured, and yet are protective; thus in _Thecla rubi_ the wings when closed are of an emerald green and resemble the young leaves of the bramble, on which this butterfly in the spring may often be seen seated.

Although the obscure tints of the upper or under surface of many butterflies no doubt serve to conceal them, yet we cannot possibly extend this view to the brilliant and conspicuous colours of many kinds, such as our admiral and peacock Vanessae, our white cabbage-butterflies (Pieris), or the great swallow-tail Papilio which haunts the open fens--for these butterflies are thus rendered visible to every living creature. With these species both sexes are alike; but in the common brimstone butterfly (_Gonepteryx rhamni_), the male is of an intense yellow, whilst the female is much paler; and in the orange-tip (_Anthocharis cardamines_) the males alone have the bright orange tips to their wings. In these cases the males and females are equally conspicuous, and it is not credible that their difference in colour stands in any relation to ordinary protection. Nevertheless it is possible that the conspicuous colours of many species may be in an indirect manner beneficial, as will hereafter be explained, by leading their enemies at once to recognise them as unpalatable. Even in this case it does not certainly follow that their bright colours and beautiful patterns were acquired for this special purpose. In some other remarkable cases, beauty has been gained for the sake of protection, through the imitation of other beautiful species, which inhabit the same district and enjoy an immunity from attack by being in some way offensive to their enemies.

The female of our orange-tip butterfly, above referred to, and of an American species (_Anth. genutia_) probably shew us, as Mr. Walsh has remarked to me, the primordial colours of the parent-species of the genus; for both sexes of four or five widely-distributed species are coloured in nearly the same manner. We may infer here, as in several previous cases, that it is the males of _Anth. cardamines_ and _genutia_ which have departed from the usual type of colouring of their genus. In the _Anth. sara_ from California, the orange-tips have become partially developed in the female; for her wings are tipped with reddish-orange, but paler than in the male, and slightly different in some other respects. In an allied Indian form, the _Iphias glaucippe_, the orange-tips are fully developed in both sexes. In this Iphias the under surface of the wings marvellously resembles, as pointed out to me by Mr. A. Butler, a pale-coloured leaf; and in our English orange-tip, the under surface resembles the flower-head of the wild parsley, on which it may be seen going to rest at night.[507] The same reasoning power which compels us to believe that the lower surfaces have here been coloured for the sake of protection, leads us to deny that the wings have been tipped, especially when this character is confined to the males, with bright orange for the same purpose.

Turning now to Moths: most of these rest motionless with their wings depressed during the whole or greater part of the day; and the upper surfaces of their wings are often shaded and coloured in an admirable manner, as Mr. Wallace has remarked, for escaping detection. With most of the Bombycidae and Noctuidae,[508] when at rest, the front-wings overlap and conceal the hind-wings; so that the latter might be brightly coloured without much risk; and they are thus coloured in many species of both families. During the act of flight, moths would often be able to escape from their enemies; nevertheless, as the hind-wings are then fully exposed to view, their bright colours must generally have been acquired at the cost of some little risk. But the following fact shews us how cautious we ought to be in drawing conclusions on this head. The common yellow under-wings (Triphaena) often fly about during the day or early evening, and are then conspicuous from the colour of their hind-wings. It would naturally be thought that this would be a source of danger; but Mr. J. Jenner Weir believes that it actually serves them as a means of escape, for birds strike at these brightly coloured and fragile surfaces, instead of at the body. For instance, Mr. Weir turned into his aviary a vigorous specimen of _Triphaena pronuba_, which was instantly pursued by a robin; but the bird's attention being caught by the coloured wings, the moth was not captured until after about fifty attempts, and small portions of the wings were repeatedly broken off. He tried the same experiment, in the open air, with a _T. fimbria_ and swallow; but the large size of this moth probably interfered with its capture.[509] We are thus reminded of a statement made by Mr. Wallace,[510] namely, that in the Brazilian forests and Malayan islands, many common and highly-decorated butterflies are weak flyers, though furnished with a broad expanse of wings; and they "are often captured with pierced and broken wings, as if they had been seized by birds, from which they had escaped: if the wings had been much smaller in proportion to the body, it seems probable that the insect would more frequently have been struck or pierced in a vital part, and thus the increased expanse of the wings may have been indirectly beneficial."

_Display._--The bright colours of butterflies and of some moths are specially arranged for display, whether or not they serve in addition as a protection. Bright colours would not be visible during the night; and there can be no doubt that moths, taken as a body, are much less gaily decorated than butterflies, all of which are diurnal in their habits. But the moths in certain families, such as the Zygaenidae, various Sphingidae, Uraniidae, some Arctiidae and Saturniidae, fly about during the day or early evening, and many of these are extremely beautiful, being far more brightly coloured than the strictly nocturnal kinds. A few exceptional cases, however, of brightly-coloured nocturnal species have been recorded.[511]

There is evidence of another kind in regard to display. Butterflies, as before remarked, elevate their wings when at rest, and whilst basking in the sunshine often alternately raise and depress them, thus exposing to full view both surfaces; and although the lower surface is often coloured in an obscure manner as a protection, yet in many species it is as highly coloured as the upper surface, and sometimes in a very different manner. In some tropical species the lower surface is even more brilliantly coloured than the upper.[512] In one English fritillary, the _Argynnis aglaia_, the lower surface alone is ornamented with shining silver discs. Nevertheless, as a general rule, the upper surface, which is probably the most fully exposed, is coloured more brightly and in a more diversified manner than the lower. Hence the lower surface generally affords to entomologists the most useful character for detecting the affinities of the various species.

Now if we turn to the enormous group of moths, which do not habitually expose to full view the under surface of their wings, this side is very rarely, as I hear from Mr. Stainton, coloured more brightly than the upper side, or even with equal brightness. Some exceptions to the rule, either real or apparent, must be noticed, as that of Hypopyra, specified by Mr. Wormald.[513] Mr. R. Trimen informs me that in Guenee's great work, three moths are figured, in which the under surface is much the most brilliant. For instance, in the Australian Gastrophora the upper surface of the fore-wing is pale greyish-ochreous, while the lower surface is magnificently ornamented by an ocellus of cobalt-blue, placed in the midst of a black mark, surrounded by orange-yellow, and this by bluish-white. But the habits of these three moths are unknown; so that no explanation can be given of their unusual style of colouring. Mr. Trimen also informs me that the lower surface of the wings in certain other Geometrae[514] and quadrifid Noctuae are either more variegated or more brightly-coloured than the upper surface; but some of these species have the habit of "holding their wings quite erect over their backs, retaining them in this position for a considerable time," and thus exposing to view the under surface. Other species when settled on the ground or herbage have the habit of now and then suddenly and slightly lifting up their wings. Hence the lower surface of the wings being more brightly-coloured than the upper surface in certain moths is not so anomalous a circumstance as it at first appears. The Saturniidae include some of the most beautiful of all moths, their wings being decorated, as in our British Emperor moth, with fine ocelli; and Mr. T. W. Wood[515] observes that they resemble butterflies in some of their movements; "for instance, in the gentle waving up and down of the wings, as if for display, which is more characteristic of diurnal than of nocturnal Lepidoptera."

It is a singular fact that no British moths, nor as far as I can discover hardly any foreign species, which are brilliantly coloured, differ much in colour according to sex; though this is the case with many brilliant butterflies. The male, however, of one American moth, the _Saturnia Io_, is described as having its fore-wings deep yellow, curiously marked with purplish-red spots; whilst the wings of the female are purple-brown, marked with grey lines.[516] The British moths which differ sexually in colour are all brown, or various tints of dull yellow, or nearly white. In several species the males are much darker than the females,[517] and these belong to groups which generally fly about during the afternoon. On the other hand, in many genera, as Mr. Stainton informs me, the males have the hind-wings whiter than those of the female--of which, fact _Agrotis exclamationis_ offers a good instance. The males are thus rendered more conspicuous than the females, whilst flying about in the dusk. In the Ghost Moth (_Hepialus humuli_) the difference is more strongly marked; the males being white and the females yellow with darker markings. It is difficult to conjecture what the meaning can be of these differences between the sexes in the shades of darkness or lightness; but we can hardly suppose that they are the result of mere variability with sexually-limited inheritance, independently of any benefit thus derived.

From the foregoing statements it is impossible to admit that the brilliant colours of butterflies and of some few moths, have commonly been acquired for the sake of protection. We have seen that their colours and elegant patterns are arranged and exhibited as if for display. Hence I am led to suppose that the females generally prefer, or are most excited by the more brilliant males; for on any other supposition the males would be ornamented, as far as we can see, for no purpose. We know that ants and certain lamellicorn beetles are capable of feeling an attachment for each other, and that ants recognise their fellows after an interval of several months. Hence there is no abstract improbability in the Lepidoptera, which probably stand nearly or quite as high in the scale as these insects, having sufficient mental capacity to admire bright colours. They certainly discover flowers by colour, and, as I have elsewhere shewn, the plants which are fertilised exclusively by the wind never have a conspicuously-coloured corolla. The Humming-bird Sphinx may often be seen to swoop down from a distance on a bunch of flowers in the midst of green foliage; and I have been assured by a friend, that these moths repeatedly visited flowers painted on the walls of a room in the South of France. The common white butterfly, as I hear from Mr. Doubleday, often flies down to a bit of paper on the ground, no doubt mistaking it for one of its own species. Mr. Collingwood[518] in speaking of the difficulty of collecting certain butterflies in the Malay Archipelago, states that "a dead specimen pinned upon a conspicuous twig will often arrest an insect of the same species in its headlong flight, and bring it down within easy reach of the net, especially if it be of the opposite sex."

The courtship of butterflies is a prolonged affair. The males sometimes fight together in rivalry; and many may be seen pursuing or crowding round the same female. If, then, the females do not prefer one male to another, the pairing must be left to mere chance, and this does not appear to me a probable event. If, on the other hand, the females habitually, or even occasionally, prefer the more beautiful males, the colours of the latter will have been rendered brighter by degrees, and will have been transmitted to both sexes or to one sex, according to which law of inheritance prevailed. The process of sexual selection will have been much facilitated, if the conclusions arrived at from various kinds of evidence in the supplement to the ninth chapter can be trusted; namely that the males of many Lepidoptera, at least in the imago state, greatly exceed in number the females.

Some facts, however, are opposed to the belief that female butterflies prefer the more beautiful males; thus, as I have been assured by several observers, fresh females may frequently be seen paired with battered, faded or dingy males; but this is a circumstance which could hardly fail often to follow from the males emerging from their cocoons earlier than the females. With moths of the family of the Bombycidae, the sexes pair immediately after assuming the imago state; for they cannot feed, owing to the rudimentary condition of their mouths. The females, as several entomologists have remarked to me, lie in an almost torpid state, and appear not to evince the least choice in regard to their partners, This is the case with the common silk-moth (_B. mori_), as I have been told by some continental and English breeders. Dr. Wallace, who has had such immense experience in breeding _Bombyx cynthia_, is convinced that the females evince no choice or preference. He has kept above 300 of these moths living together, and has often found the most vigorous females mated with stunted males. The reverse apparently seldom occurs; for, as he believes, the more vigorous males pass over the weakly females, being attracted by those endowed with most vitality. Although we have been indirectly induced to believe that the females of many species prefer the more beautiful males, I have no reason to suspect, either with moths or butterflies, that the males are attracted by the beauty of the females. If the more beautiful females had been continually preferred, it is almost certain, from the colours of butterflies being so frequently transmitted to one sex alone, that the females would often have been rendered more beautiful than their male partners. But this does not occur except in a few instances; and these can be explained, as we shall presently see, on the principle of mimickry and protection.

As sexual selection primarily depends on variability, a few words must be added on this subject. In respect to colour there is no difficulty, as any number of highly variable Lepidoptera could be named. One good instance will suffice. Mr. Bates shewed me a whole series of specimens of _Papilio sesostris_ and _childrenae_; in the latter the males varied much in the extent of the beautifully enamelled green patch on the fore-wings, and in the size of the white mark, as well as of the splendid crimson stripe on the hind-wings; so that there was a great contrast between the most and least gaudy males. The male of _Papilio sesostris_, though a beautiful insect, is much less so than _P. childrenae_. It likewise varies a little in the size of the green patch on the fore-wings, and in the occasional appearance of a small crimson stripe on the hind-wings, borrowed, as it would seem, from its own female; for the females of this and of many other species in the AEneas group possess this crimson stripe. Hence between the brightest specimens of _P. sesostris_ and the least bright of _P. childrenae_, there was but a small interval; and it was evident that as far as mere variability is concerned, there would be no difficulty in permanently increasing by means of selection the beauty of either species. The variability is here almost confined to the male sex; but Mr. Wallace and Mr. Bates have shewn[519] that the females of some other species are extremely variable, the males being nearly constant. As I have before mentioned the Ghost Moth (_Hepialus humuli_) as one of the best instances in Britain of a difference in colour between the sexes of moths, it may be worth adding[520] that in the Shetland Islands, males are frequently found which closely resemble the females. In a future chapter I shall have occasion to shew that the beautiful eye-like spots or ocelli, so common on the wings of many Lepidoptera, are eminently variable.

On the whole, although many serious objections may be urged, it seems probable that most of the species of Lepidoptera which are brilliantly coloured, owe their colours to sexual selection, excepting in certain cases, presently to be mentioned, in which conspicuous colours are beneficial as a protection. From the ardour of the male throughout the animal kingdom, he is generally willing to accept any female; and it is the female which usually exerts a choice. Hence if sexual selection has here acted, the male, when the sexes differ, ought to be the most brilliantly coloured; and this undoubtedly is the ordinary rule. When the sexes are brilliantly coloured and resemble each other, the characters acquired by the males appear to have been transmitted to both sexes. But will this explanation of the similarity and dissimilarity in colour between the sexes suffice?

The males and females of the same species of butterfly are known[521] in several cases to inhabit different stations, the former commonly basking in the sunshine, the latter haunting gloomy forests. It is therefore possible that different conditions of life may have acted directly on the two sexes; but this is not probable,[522] as in the adult state they are exposed during a very short period to different conditions; and the larvae of both are exposed to the same conditions. Mr. Wallace believes that the less brilliant colours of the female have been specially gained in all or almost all cases for the sake of protection. On the contrary it seems to me more probable that the males alone, in the large majority of cases, have acquired their bright colours through sexual selection, the females having been but little modified. Consequently the females of distinct but allied species ought to resemble each other much more closely than do the males of the same species; and this is the general rule. The females thus approximately show us the primordial colouring of the parent-species of the group to which they belong. They have, however, almost always been modified to a certain extent by some of the successive steps of variation, through the accumulation of which the males were rendered beautiful, having been transferred to them. The males and females of allied though distinct species will also generally have been exposed during their prolonged larval state to different conditions, and may have been thus indirectly affected; though with the males any slight change of colour thus caused will often have been completely masked by the brilliant tints gained through sexual selection. When we treat of Birds, I shall have to discuss the whole question whether the differences in colour between the males and females have been in part specially gained by the latter as a protection; so that I will here only give unavoidable details.

In all cases when the more common form of equal inheritance by both sexes has prevailed, the selection of bright-coloured males would tend to make the females bright-coloured; and the selection of dull-coloured females would tend to make the males dull. If both processes were carried on simultaneously, they would tend to neutralise each other. As far as I can see, it would be extremely difficult to change through selection the one form of inheritance into the other. But by the selection of successive variations, which were from the first sexually limited in their transmission, there would not be the slightest difficulty in giving bright colours to the males alone, and at the same time or subsequently, dull colours to the females alone. In this latter manner female butterflies and moths may, as I fully admit, have been rendered inconspicuous for the sake of protection, and widely different from their males.

Mr. Wallace[523] has argued with much force in favour of his view that when the sexes differ, the female has been specially modified for the sake of protection; and that this has been effected by one form of inheritance, namely, the transmission of characters to both sexes, having been changed through the agency of natural selection into the other form, namely, transmission to one sex. I was at first strongly inclined to accept this view; but the more I have studied the various classes throughout the animal kingdom, the less probable it has appeared. Mr. Wallace urges that both sexes of the _Heliconidae_, _Danaidae_, _Acroeidae_ are equally brilliant because both are protected from the attacks of birds and other enemies, by their offensive odour; but that in other groups, which do not possess this immunity, the females have been rendered inconspicuous, from having more need of protection than the males. This supposed difference in the "need of protection by the two sexes" is rather deceptive, and requires some discussion. It is obvious that brightly-coloured individuals, whether males or females, would equally attract, and obscurely-coloured individuals equally escape, the attention of their enemies. But we are concerned with the effects of the destruction or preservation of certain individuals of either sex, on the character of the race. With insects, after the male has fertilised the female, and after the latter has laid her eggs, the greater or less immunity from danger of either sex could not possibly have any effect on the offspring. Before the sexes have performed their proper functions, if they existed in equal numbers and if they strictly paired (all other circumstances being the same), the preservation of the males and females would be equally important for the existence of the species and for the character of the offspring. But with most animals, as is known to be the case with the domestic silk-moth, the male can fertilise two or three females; so that the destruction of the males would not be so injurious to the species as that of the females. On the other hand, Dr. Wallace believes that with moths the progeny from a second or third fertilisation is apt to be weakly, and therefore would not have so good chance of surviving. When the males exist in much greater numbers than the females, no doubt many males might be destroyed with impunity to the species; but I cannot see that the results of ordinary selection for the sake of protection would be influenced by the sexes existing in unequal numbers; for the same proportion of the more conspicuous individuals, whether males or females, would probably be destroyed. If indeed the males presented a greater range of variation in colour, the result would be different; but we need not here follow out such complex details. On the whole I cannot perceive that an inequality in the numbers of the two sexes would influence in any marked manner the effects of ordinary selection on the character of the offspring.

Female Lepidoptera require, as Mr. Wallace insists, some days to deposit their fertilised ova and to search for a proper place; during this period (whilst the life of the male was of no importance) the brighter-coloured females would be exposed to danger and would be liable to be destroyed. The duller-coloured females on the other hand would survive, and thus would influence, it might be thought, in a marked manner the character of the species,--either of both sexes or of one sex, according to which form of inheritance prevailed. But it must not be forgotten that the males emerge from the cocoon-state some days before the females, and during this period, whilst the unborn females were safe, the brighter-coloured males would be exposed to danger; so that ultimately both sexes would probably be exposed during a nearly equal length of time to danger, and the elimination of conspicuous colours would not be much more effective in the one than the other sex.

It is a more important consideration that female Lepidoptera, as Mr. Wallace remarks, and as is known to every collector, are generally slower flyers than the males. Consequently the latter, if exposed to greater danger from being conspicuously coloured, might be able to escape from their enemies, whilst the similarly-coloured females would be destroyed; and thus the females would have the most influence in modifying the colour of their progeny.

There is one other consideration: bright colours, as far as sexual selection is concerned, are commonly of no service to the females; so that if the latter varied in brightness, and the variations were sexually limited in their transmission, it would depend on mere chance whether the females had their bright colours increased; and this would tend throughout the Order to diminish the number of species with brightly-coloured females in comparison with the species having brightly-coloured males. On the other hand, as bright colours are supposed to be highly serviceable to the males in their love-struggles, the brighter males (as we shall see in the chapter on Birds) although exposed to rather greater danger, would on an average procreate a greater number of offspring than the duller males. In this case, if the variations were limited in their transmission to the male sex, the males alone would be rendered more brilliantly coloured; but if the variations were not thus limited, the preservation and augmentation of such variations would depend on whether more evil was caused to the species by the females being rendered conspicuous, than good to the males by certain individuals being successful over their rivals.

As there can hardly be a doubt that both sexes of many butterflies and moths have been rendered dull-coloured for the sake of protection, so it may have been with the females alone of some species in which successive variations towards dullness first appeared in the female sex and were from the first limited in their transmission to the same sex. If not thus limited, both sexes would become dull-coloured. We shall immediately see, when we treat of mimickry, that the females alone of certain butterflies have been rendered extremely beautiful for the sake of protection, without any of the successive protective variations having been transferred to the male, to whom they could not possibly have been in the least degree injurious, and therefore could not have been eliminated through natural selection. Whether in each particular species, in which the sexes differ in colour, it is the female which has been specially modified for the sake of protection; or whether it is the male which has been specially modified for the sake of sexual attraction, the female having retained her primordial colouring only slightly changed through the agencies before alluded to; or whether again both sexes have been modified, the female for protection and the male for sexual attraction, can only be definitely decided when we know the life-history of each species.

Without distinct evidence, I am unwilling to admit that a double process of selection has long been going on with a multitude of species,--the males having been rendered more brilliant by beating their rivals; and the females more dull-coloured by having escaped from their enemies. We may take as an instance the common brimstone butterfly (Gonepteryx), which appears early in the spring before any other kind. The male of this species is of a far more intense yellow than the female, though she is almost equally conspicuous; and in this case it does not seem probable that she specially acquired her pale tints as a protection, though it is probable that the male acquired his bright colours as a sexual attraction. The female of _Anthocharis cardamines_ does not possess the beautiful orange tips to her wings with which the male is ornamented; consequently she closely resembles the white butterflies (Pieris) so common in our gardens; but we have no evidence that this resemblance is beneficial. On the contrary, as she resembles both sexes of several species of the same genus inhabiting various quarters of the world, it is more probable that she has simply retained to a large extent her primordial colours.

Various facts support the conclusion that with the greater number of brilliantly-coloured Lepidoptera, it is the male which has been modified; the two sexes having come to differ from each other, or to resemble each other, according to which form of inheritance has prevailed. Inheritance is governed by so many unknown laws or conditions, that they seem to us to be most capricious in their action;[524] and we can so far understand how it is that with closely-allied species the sexes of some differ to an astonishing degree, whilst the sexes of others are identical in colour. As the successive steps in the process of variation are necessarily all transmitted through the female, a greater or less number of such steps might readily become developed in her; and thus we can understand the frequent gradations from an extreme difference to no difference at all between the sexes of the species within the same group. These cases of gradation are much too common to favour the supposition that we here see females actually undergoing the process of transition and losing their brightness for the sake of protection; for we have every reason to conclude that at any one time the greater number of species are in a fixed condition. With respect to the differences between the females of the species in the same genus or family, we can perceive that they depend, at least in part, on the females partaking of the colours of their respective males. This is well illustrated in those groups in which the males are ornamented to an extraordinary degree; for the females in these groups generally partake to a certain extent of the splendour of their male partners. Lastly, we continually find, as already remarked, that the females of almost all the species in the same genus, or even family, resemble each other much more closely in colour than do the males; and this indicates that the males have undergone a greater amount of modification than the females.

_Mimickry._--This principle was first made clear in an admirable paper by Mr. Bates,[525] who thus threw a flood of light on many obscure problems. It had previously been observed that certain butterflies in S. America belonging to quite distinct families, resembled the Heliconidae so closely in every stripe and shade of colour that they could not be distinguished except by an experienced entomologist. As the Heliconidae are coloured in their usual manner, whilst the others depart from the usual colouring of the groups to which they belong, it is clear that the latter are the imitators, and the Heliconidae the imitated. Mr. Bates further observed that the imitating species are comparatively rare, whilst the imitated swarm in large numbers; the two sets living mingled together. From the fact of the Heliconidae being conspicuous and beautiful insects, yet so numerous in individuals and species, he concluded that they must be protected from the attacks of birds by some secretion or odour; and this hypothesis has now been confirmed by a considerable body of curious evidence.[526] From these considerations Mr. Bates inferred that the butterflies which imitate the protected species had acquired their present marvellously deceptive appearance, through variation and natural selection, in order to be mistaken for the protected kinds and thus to escape being devoured. No explanation is here attempted of the brilliant colours of the imitated, but only of the imitating butterflies. We must account for the colours of the former in the same general manner, as in the cases previously discussed in this chapter. Since the publication of Mr. Bates' paper, similar and equally striking facts have been observed by Mr. Wallace[527] in the Malayan region, and by Mr. Trimen in South Africa.

As some writers[528] have felt much difficulty in understanding how the first steps in the process of mimickry could have been effected through natural selection, it may be well to remark that the process probably has never commenced with forms widely dissimilar in colour. But with two species moderately like each other, the closest resemblance if beneficial to either form could readily be thus gained; and if the imitated form was subsequently and gradually modified through sexual selection or any other means, the imitating form would be led along the same track, and thus be modified to almost any extent, so that it might ultimately assume an appearance or colouring wholly unlike that of the other members of the group to which it belonged. As extremely slight variations in colour would not in many cases suffice to render a species so like another protected species as to lead to its preservation, it should be remembered that many species of Lepidoptera are liable to considerable and abrupt variations in colour. A few instances have been given in this chapter; but under this point of view Mr. Bates' original paper on mimickry, as well as Mr. Wallace's papers, should be consulted.

In the foregoing cases both sexes of the imitating species resemble the imitated; but occasionally the female alone mocks a brilliantly-coloured and protected species inhabiting the same district. Consequently the female differs in colour from her own male, and, which is a rare and anomalous circumstance, is the more brightly-coloured of the two. In all the few species of Pieridae, in which the female is more conspicuously coloured than the male, she imitates, as I am informed by Mr. Wallace, some protected species inhabiting the same region. The female of _Diadema anomala_ is rich purple-brown with almost the whole surface glossed with satiny blue, and she closely imitates the _Euploea midamus_, "one of the commonest butterflies of the East;" whilst the male is bronzy or olive-brown, with only a slight blue gloss on the outer parts of the wings.[529] Both sexes of this Diadema and of _D. bolina_ follow the same habits of life, so that the differences in colour between the sexes cannot be accounted for by exposure to different conditions;[530] even if this explanation were admissible in other instances.[531]

The above cases of female butterflies which are more brightly-coloured than the males, shew us, firstly, that variations have arisen in a state of nature in the female sex, and have been transmitted exclusively, or almost exclusively, to the same sex; and, secondly, that this form of inheritance has not been determined through natural selection. For if we assume that the females, before they became brightly coloured in imitation of some protected kind, were exposed during each season for a longer period to danger than the males; or if we assume that they could not escape so swiftly from their enemies, we can understand how they alone might originally have acquired through natural selection and sexually-limited inheritance their present protective colours. But except on the principle of these variations having been transmitted exclusively to the female offspring, we cannot understand why the males should have remained dull-coloured; for it would surely not have been in any way injurious to each individual male to have partaken by inheritance of the protective colours of the female, and thus to have had a better chance of escaping destruction. In a group in which brilliant colours are so common as with butterflies, it cannot be supposed that the males have been kept dull-coloured through sexual selection by the females rejecting the individuals which were rendered as beautiful as themselves. We may, therefore, conclude that in these cases inheritance by one sex is not due to the modification through natural selection of a tendency to equal inheritance by both sexes.

It may be well here to give an analogous case in another Order, of characters acquired only by the female, though not in the least injurious, as far as we can judge, to the male. Amongst the Phasmidae, or spectre-insects, Mr. Wallace states that "it is often the females alone that so strikingly resemble leaves, while the males show only a rude approximation." Now, whatever may be the habits of these insects, it is highly improbable that it could be disadvantageous to the males to escape detection by resembling leaves.[532] Hence we may conclude that the females alone in this latter as in the previous cases originally varied in certain characters; these characters having been preserved and augmented through ordinary selection for the sake of protection and from the first transmitted to the female offspring alone.

_Bright Colours of Caterpillars._--Whilst reflecting on the beauty of many butterflies, it occurred to me that some caterpillars were splendidly coloured, and as sexual selection could not possibly have here acted, it appeared rash to attribute the beauty of the mature insect to this agency, unless the bright colours of their larvae could be in some manner explained. In the first place it may be observed that the colours of caterpillars do not stand in any close correlation with those of the mature insect. Secondly, their bright colours do not serve in any ordinary manner as a protection. As an instance of this, Mr. Bates informs me that the most conspicuous caterpillar which he ever beheld (that of a Sphinx) lived on the large green leaves of a tree on the open llanos of South America; it was about four inches in length, transversely banded with black and yellow, and with its head, legs, and tail of a bright red. Hence it caught the eye of any man who passed by at the distance of many yards, and no doubt of every passing bird.

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. 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 Society[533] 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 of 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 gained 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 sometimes 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.

Sexual 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 habitually 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 gained 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 few 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 ueber die Darwin'sche Theorie:' zweite Auflage, 1868, von Dr. L. Buechner; translated into French under the title 'Conferences sur la Theorie 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. Haeckel 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] 'Lec. sur la Phys.' 1866, p. 890, as quoted by M. Dally, 'L'Ordre des Primates et le Transformisme,' 1868, p. 29.

[5] 'Naturgeschichte der Saeugethiere 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 qua 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 Mammiferes,' 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. Haeckel has also given analogous drawings in his 'Schoepfungsgeschichte.'

[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. Haeckel has given admirable discussions on this whole subject, under the title of Dysteleology, in his 'Generelle Morphologie' and 'Schoepfungsgeschichte.'

[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 pedieux de la main," which he says is sometimes "infiniment petit." Another muscle, called "le tibial posterieur," is generally quite absent in the hand, but appears from time to time in a more or less rudimentary condition.

[24] Prof. W. Turner, 'Proc. Royal Soc. Edinburgh,' 1866-67, p. 65.

[25] Canestrini quotes Hyrt. ('Annuario della Soc. dei Naturalisti,' Modena, 1867, p. 97) to the same effect.

[26] 'The Diseases of the Ear,' by J. Toynbee, F.R.S., 1860, p. 12.

[27] See also some remarks, and the drawings of the ears of the Lemuroidea, in Messrs. Murie and Mivart's excellent paper in 'Transact. Zoolog. Soc.' vol. vii. 1869, pp. 6 and 90.

[28] Mueller's 'Elements of Physiology,' Eng. translat., 1842, vol. ii. p. 1117. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 260; ibid. on the Walrus, 'Proc. Zoolog. Soc.' November 8th, 1854. See also R. Knox, 'Great Artists and Anatomists,' p. 106. This rudiment apparently is somewhat larger in Negroes and Australians than in Europeans, see Carl Vogt, 'Lectures on Man,' Eng. translat. p. 129.

[29] 'The Physiology and Pathology of Mind,' 2nd edit. 1868, p. 134.

[30] Eschricht, Ueber die Richtung der Haare am menschlichen Koerper 'Muellers Archiv fuer Anat. und Phys.' 1837, s. 47. I shall often have to refer to this very curious paper.

[31] Paget, 'Lectures on Surgical Pathology,' 1853, vol. i. p. 71.

[32] Eschricht, ibid. s. 40, 47.

[33] Dr. Webb, 'Teeth in Man and the Anthropoid Apes,' as quoted by Dr. C. Carter Blake in 'Anthropological Review,' July, 1867, p. 299.

[34] Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 320, 321, and 325.

[35] 'On the Primitive Form of the Skull,' Eng. translat. in 'Anthropological Review,' Oct. 1868, p. 426.

[36] Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 416, 434, 441.

[37] 'Annuario della Soc. d. Nat.' Modena, 1867, p. 94.

[38] M. C. Martins ("De l'Unite Organique," in 'Revue des Deux Mondes,' June 15, 1862, p. 16), and Haeckel ('Generelle Morphologie,' B. ii. s. 278), have both remarked on the singular fact of this rudiment sometimes causing death.

[39] 'The Lancet,' Jan. 24, 1863, p. 83. Dr. Knox, 'Great Artists and Anatomists,' p. 63. See also an important memoir on this process by Dr. Grube, in the 'Bulletin de l'Acad. Imp. de St. Petersbourg,' tom. xii. 1867, p. 448.

[40] "On the Caves of Gibraltar," 'Transact. Internat. Congress of Prehist. Arch.' Third Session, 1869, p. 54.

[41] Quatrefages has lately collected the evidence on this subject. 'Revue des Cours Scientifiques,' 1867-1868, p. 625.

[42] Owen, 'On the Nature of Limbs,' 1849, p. 114.

[43] Leuckart, in Todd's 'Cyclop. of Anat.' 1849-52, vol. iv. p. 1415. In man this organ is only from three to six lines in length, but, like so many other rudimentary parts, it is variable in development as well as in other characters.

[44] See, on this subject, Owen, 'Anatomy of Vertebrates,' vol. iii. pp. 675, 676, 706.

[45] See the evidence on these points, as given by Lubbock, 'Prehistoric Times,' p. 354, &c.

[46] 'L'Instinct chez les Insectes.' 'Revue des Deux Mondes,' Feb. 1870, p. 690.

[47] 'The American Beaver and his Works,' 1868.

[48] 'The Principles of Psychology,' 2nd edit. 1870, pp. 418-443.

[49] 'Contributions to the Theory of Natural Selection,' 1870, p. 212

[50] 'Recherches sur les Moeurs des Fourmis,' 1810, p. 173.

[51] All the following statements, given on the authority of these two naturalists, are taken from Rengger's 'Naturges. der Saeugethiere von Paraguay,' 1830, s. 41-57, and from Brehm's 'Thierleben,' B. i. s. 10-87.

[52] 'Bridgewater Treatise,' p. 263.

[53] W. C. L. Martin, 'Nat. Hist. of Mammalia,' 1841, p. 405.

[54] Quoted by Vogt, 'Memoire sur les Microcephales,' 1867, p. 168.

[55] 'The Variation of Animals and Plants under Domestication,' vol. i. p. 27.

[56] 'Les Moeurs des Fourmis,' 1810, p. 150.

[57] Quoted in Dr. Maudsley's 'Physiology and Pathology of Mind,' 1868, pp. 19, 220.

[58] Dr. Jerdon, 'Birds of India,' vol. i. 1862, p. xxi.

[59] Mr. L. H. Morgan's work on 'The American Beaver,' 1868, offers a good illustration of this remark. I cannot, however, avoid thinking that he goes too far in underrating the power of Instinct.

[60] 'The Moor and the Loch,' p. 45. Col. Hutchinson on 'Dog Breaking,' 1850, p. 46.

[61] 'Personal Narrative,' Eng. translat., vol. iii. p. 106.

[62] Quoted by Sir C. Lyell, 'Antiquity of Man,' p. 497.

[63] 'Journal of Researches during the Voyage of the "Beagle,"' 1845, p. 398. 'Origin of Species,' 5th edit. p. 260.

[64] 'Lettres Phil. sur l'Intelligence des Animaux,' nouvelle edit. 1802, p. 86.

[65] See the evidence on this head in chap. i. vol. i. 'On the Variation of Animals and Plants under Domestication.'

[66] 'Proc. Zoolog. Soc.' 1864, p. 186.

[67] Savage and Wyman in 'Boston Journal of Nat. Hist.' vol. iv. 1843-44, p. 383.

[68] 'Saeugethiere von Paraguay,' 1830, s. 51-56.

[69] 'Thierleben,' B. i. s. 79, 82.

[70] 'The Malay Archipelago,' vol. i. 1869, p. 87.

[71] 'Primeval Man,' 1869, pp. 145, 147.

[72] 'Prehistoric Times,' 1865, p. 473, &c.

[73] Quoted in 'Anthropological Review,' 1864, p. 158.

[74] Rengger, ibid. s. 45.

[75] See my 'Variation of Animals and Plants under Domestication,' vol. i. p. 27.

[76] See a discussion on this subject in Mr. E. B. Tylor's very interesting work, 'Researches into the Early History of Mankind,' 1865, chaps. ii. to iv.

[77] Hon. Daines Barrington in 'Philosoph. Transactions,' 1773, p. 262. See also Dureau de la Malle, in 'Ann. des Sc. Nat.' 3rd series, Zoolog. tom. x. p. 119.

[78] 'On the Origin of Language,' by H. Wedgwood, 1866. 'Chapters on Language,' by the Rev. F. W. Farrar, 1865. These works are most interesting. See also 'De la Phys. et de Parole,' par Albert Lemoine, 1865, p. 190. The work on this subject, by the late Prof. Aug. Schleicher, has been translated by Dr. Bikkers into English, under the title of 'Darwinism tested by the Science of Language,' 1869.

[79] Vogt, 'Memoire sur les Microcephales,' 1867, p. 169. With respect to savages, I have given some facts in my 'Journal of Researches,' &c., 1845, p. 206.

[80] See clear evidence on this head in the two works so often quoted, by Brehm and Rengger.

[81] See remarks on this head by Dr. Maudsley, 'The Physiology and Pathology of Mind,' 2nd edit. 1868, p. 199.

[82] Many curious cases have been recorded. See, for instance, 'Inquiries Concerning the Intellectual Powers,' by Dr. Abercrombie, 1838, p. 150.

[83] 'The Variation of Animals and Plants under Domestication,' vol. ii. p. 6.

[84] See some good remarks to this effect by Dr. Maudsley, 'The Physiology and Pathology of Mind,' 1808, p. 199.

[85] Macgillivray, 'Hist. of British Birds,' vol. ii. 1839, p. 29. An excellent observer, Mr. Blackwall, remarks that the magpie learns to pronounce single words, and even short sentences, more readily than almost any other British bird; yet, as he adds, after long and closely investigating its habits, he has never known it, in a state of nature, display any unusual capacity for imitation. 'Researches in Zoology,' 1834, p. 158.

[86] See the very interesting parallelism between the development of species and languages, given by Sir C. Lyell in 'The Geolog. Evidences of the Antiquity of Man,' 1863, chap. xxiii.

[87] See remarks to this effect by the Rev. F. W. Farrar, in an interesting article, entitled "Philology and Darwinism" in 'Nature,' March 24th, 1870, p. 528.

[88] 'Nature,' Jan. 6th, 1870, p. 257.

[89] Quoted by C. S. Wake, 'Chapters on Man,' 1868, p. 101.

[90] Buckland, 'Bridgewater Treatise,' p. 411.

[91] See some good remarks on the simplification of languages, by Sir J. Lubbock, 'Origin of Civilisation,' 1870, p. 278.

[92] 'Conferences sur la Theorie Darwinienne,' French translat., 1869, p. 132.

[93] The Rev. Dr. J. M'Cann, 'Anti-Darwinism,' 1869, p. 13.

[94] 'The Spectator,' Dec. 4th, 1869, p. 1430.

[95] See an excellent article on this subject by the Rev. F. W. Farrar, in the 'Anthropological Review,' Aug. 1864, p. ccxvii. For further facts see Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. 1869. p. 564; and especially the chapters on Religion in his 'Origin of Civilisation,' 1870.

[96] The Worship of Animals and Plants, in the 'Fortnightly Review,' Oct. 1, 1869, p. 422.

[97] Tylor, 'Early History of Mankind,' 1865, p. 6. See also the three striking chapters on the Development of Religion, in Lubbock's 'Origin of Civilisation,' 1870. In a like manner Mr. Herbert Spencer, in his ingenious essay in the 'Fortnightly Review' (May 1st, 1870, p. 535), accounts for the earliest forms of religious belief throughout the world, by man being led through dreams, shadows, and other causes, to look at himself as a double essence, corporeal and spiritual. As the spiritual being is supposed to exist after death and to be powerful, it is propitiated by various gifts and ceremonies, and its aid invoked. He then further shews that names or nicknames given from some animal or other object to the early progenitors or founders of a tribe, are supposed after a long interval to represent the real progenitor of the tribe; and such animal or object is then naturally believed still to exist as a spirit, is held sacred, and worshipped as a god. Nevertheless I cannot but suspect that there is a still earlier and ruder stage, when anything which manifests power or movement is thought to be endowed with some form of life, and with mental faculties analogous to our own.

[98] See an able article on the Psychical Elements of Religion, by Mr. L. Owen Pike, in 'Anthropolog. Review,' April, 1870, p. lxiii.

[99] 'Religion, Moral, &c., der Darwin'schen Art-Lehre,' 1869, s. 53.

[100] 'Prehistoric Times,' 2nd edit. p. 571. In this work (at p. 553) there will be found an excellent account of the many strange and capricious customs of savages.

[101] See, for instance, on this subject, Quatrefages, 'Unite de l'Espece Humaine,' 1861, p. 21, &c.

[102] 'Dissertation on Ethical Philosophy,' 1837, p. 231, &c.

[103] 'Metaphysics of Ethics,' translated by J. W. Semple, Edinburgh, 1836, p. 136.

[104] Mr. Bain gives a list ('Mental and Moral Science,' 1868, p. 543-725) of twenty-six British authors who have written on this subject, and whose names are familiar to every reader; to these, Mr. Bain's own name, and those of Mr. Lecky, Mr. Shadworth Hodgson, and Sir J. Lubbock, as well as of others, may be added.

[105] Sir B. Brodie, after observing that man is a social animal ('Psychological Enquiries,' 1854, p. 192), asks the pregnant question, "ought not this to settle the disputed question as to the existence of a moral sense?" Similar ideas have probably occurred to many persons, as they did long ago to Marcus Aurelius. Mr. J. S. Mill speaks, in his celebrated work, 'Utilitarianism,' (1864, p. 46), of the social feelings as a "powerful natural sentiment," and as "the natural basis of sentiment for utilitarian morality;" but on the previous page he says, "if, as is my own belief, the moral feelings are not innate, but acquired, they are not for that reason less natural." It is with hesitation that I venture to differ from so profound a thinker, but it can hardly be disputed that the social feelings are instinctive or innate in the lower animals; and why should they not be so in man? Mr. Bain (see, for instance, 'The Emotions and the Will,' 1865, p. 481) and others believe that the moral sense is acquired by each individual during his lifetime. On the general theory of evolution this is at least extremely improbable.

[106] 'Die Darwin'sche Theorie,' s. 101.

[107] Mr. R. Brown in 'Proc. Zoolog. Soc.' 1868, p. 409.

[108] Brehm, 'Thierleben,' B. i. 1864, s. 52, 79. For the case of the monkeys extracting thorns from each other, see s. 54. With respect to the Hamadryas turning over stones, the fact is given (s. 76) on the evidence of Alvarez, whose observations Brehm thinks quite trustworthy. For the cases of the old male baboons attacking the dogs, see s. 79; and with respect to the eagle, s. 56.

[109] 'Annals and Mag. of Nat. Hist.' November, 1868, p. 382.

[110] Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. p. 446.

[111] As quoted by Mr. L. H. Morgan, 'The American Beaver,' 1868, p. 272. Capt. Stansbury also gives an interesting account of the manner in which a very young pelican, carried away by a strong stream, was guided and encouraged in its attempts to reach the shore by half a dozen old birds.

[112] As Mr. Bain states, "effective aid to a sufferer springs from sympathy proper:" 'Mental and Moral Science,' 1868, p. 245.

[113] 'Thierleben,' B. i. s. 85.

[114] 'De l'Espece et de la Class.' 1869, p. 97.

[115] 'Der Darwin'schen Art-Lehre,' 1869, s. 54.

[116] Brehm, 'Thierleben,' B. i. s. 76.

[117] See the first and striking chapter in Adam Smith's 'Theory of Moral Sentiments.' Also Mr. Bain's 'Mental and Moral Science,' 1868, p. 244, and 275-282. Mr. Bain states, that "sympathy is, indirectly, a source of pleasure to the sympathiser;" and he accounts for this through reciprocity. He remarks that "the person benefited, or others in his stead, may make up, by sympathy and good offices returned, for all the sacrifice." But if, as appears to be the case, sympathy is strictly an instinct, its exercise would give direct pleasure, in the same manner as the exercise, as before remarked, of almost every other instinct.

[118] This fact, the Rev. L. Jenyns states (see his edition of 'White's Nat. Hist. of Selborne,' 1853, p. 204) was first recorded by the illustrious Jenner, in 'Phil. Transact.' 1824, and has since been confirmed by several observers, especially by Mr. Blackwall. This latter careful observer examined, late in the autumn, during two years, thirty-six nests; he found that twelve contained young dead birds, five contained eggs on the point of being hatched, and three eggs not nearly hatched. Many birds not yet old enough for a prolonged flight are likewise deserted and left behind. See Blackwall, 'Researches in Zoology,' 1834, pp. 108, 118. For some additional evidence, although this is not wanted, see Leroy, 'Lettres Phil.' 1802, p. 217.

[119] Hume remarks ('An Enquiry Concerning the Principles of Morals,' edit. of 1751, p. 132), "there seems a necessity for confessing that the happiness and misery of others are not spectacles altogether indifferent to us, but that the view of the former ... communicates a secret joy; the appearance of the latter ... throws a melancholy damp over the imagination."

[120] 'Mental and Moral Science,' 1868, p. 254.

[121] I have given one such case, namely of three Patagonian Indians who preferred being shot, one after the other, to betraying the plans of their companions in war ('Journal of Researches,' 1845, p. 103).

[122] Dr. Prosper Despine, in his 'Psychologie Naturelle,' 1868 (tom. i. p. 243; tom ii. p. 169) gives many curious cases of the worst criminals, who apparently have been entirely destitute of conscience.

[123] See an able article in the 'North British Review,' 1867, p. 395. See also Mr. W. Bagehot's articles on the Importance of Obedience and Coherence to Primitive Man, in the 'Fortnightly Review,' 1867, p. 529, and 1868, p. 457, &c.

[124] The fullest account which I have met with is by Dr. Gerland, in his 'Ueber das Aussterben der Naturvoelker,' 1868; but I shall have to recur to the subject of infanticide in a future chapter.

[125] See the very interesting discussion on Suicide in Lecky's 'History of European Morals,' vol. i. 1869, p. 223.

[126] See, for instance, Mr. Hamilton's account of the Kaffirs, 'Anthropological Review,' 1870, p. xv.

[127] Mr. M'Lennan has given 'Primitive Marriage,' 1865, p. 176, a good collection of facts on this head.

[128] Lecky, 'History of European Morals,' vol. i. 1869, p. 109.

[129] 'Embassy to China,' vol. ii. p. 348.

[130] See on this subject copious evidence in Chap. vii. of Sir J. Lubbock, 'Origin of Civilisation,' 1870.

[131] For instance Lecky, 'Hist. European Morals,' vol. i. p. 124.

[132] This term is used in an able article in the 'Westminster Review,' Oct. 1869, p. 498. For the Greatest Happiness principle, see J. S. Mill, 'Utilitarianism,' p. 17.

[133] Good instances are given by Mr. Wallace in 'Scientific Opinion,' Sept. 15, 1869; and more fully in his 'Contributions to the Theory of Natural Selection,' 1870, p. 353.

[134] Tennyson, 'Idylls of the King,' p. 244.

[135] 'The Thoughts of the Emperor M. Aurelius Antoninus,' Eng. translat., 2nd edit., 1869, p. 112. Marcus Aurelius was born A.D. 121.

[136] Letter to Mr. Mill in Bain's 'Mental and Moral Science,' 1868, p. 722.

[137] A writer in the 'North British Review' (July, 1869, p. 531), well capable of forming a sound judgment, expresses himself strongly to this effect. Mr. Lecky ('Hist. of Morals,' vol. i. p. 143) seems to a certain extent to coincide.

[138] See his remarkable work on 'Hereditary Genius,' 1869, p. 349. The Duke of Argyll ('Primeval Man,' 1869, p. 188) has some good remarks on the contest in man's nature between right and wrong.

[139] 'The Thoughts of Marcus Aurelius,' &c., p. 139.

[140] 'Investigations in Military and Anthropolog. Statistics of American Soldiers,' by B. A. Gould, 1869, p. 256.

[141] With respect to the "Cranial forms of the American aborigines," see Dr. Aitken Meigs in 'Proc. Acad. Nat. Sci.' Philadelphia, May, 1866. On the Australians, see Huxley, in Lyell's 'Antiquity of Man,' 1863, p. 87. On the Sandwich Islanders, Prof. J. Wyman, 'Observations on Crania,' Boston, 1868, p. 18.

[142] 'Anatomy of the Arteries,' by R. Quain.

[143] 'Transact. Royal Soc.' Edinburgh, vol. xxiv. p. 175, 189.

[144] 'Proc. Royal Soc.' 1867, p. 544; also 1868, p. 483, 524. There is a previous paper, 1866, p. 229.

[145] 'Proc. R. Irish Academy,' vol. x. 1868, p. 141.

[146] 'Act. Acad.,' St. Petersburg, 1778, part ii. p. 217.

[147] Brehm, 'Thierleben,' B. i. s. 58, 87. Rengger, 'Saeugethiere von Paraguay,' s. 57.

[148] 'Variation of Animals and Plants under Domestication,' vol. ii. chap. xii.

[149] 'Hereditary Genius: an Inquiry into its Laws and Consequences,' 1869.

[150] Mr. Bates remarks ('The Naturalist on the Amazons,' 1863, vol. ii. p. 159), with respect to the Indians of the same S. American tribe, "no two of them were at all similar in the shape of the head; one man had an oval visage with fine features, and another was quite Mongolian in breadth and prominence of cheek, spread of nostrils, and obliquity of eyes."

[151] Blumenbach, 'Treatises on Anthropolog.' Eng. translat., 1865, p. 205.

[152] Godron, 'De l'Espece,' 1859, tom. ii. livre 3. Quatrefages, 'Unite de l'Espece Humaine,' 1861. Also Lectures on Anthropology, given in the 'Revue des Cours Scientifiques,' 1866-1868.

[153] 'Hist. Gen. et Part. des Anomalies de l'Organisation,' in three volumes, tom. i. 1832.

[154] I have fully discussed these laws in my 'Variation of Animals and Plants under Domestication,' vol. ii. chap. xxii. and xxiii. M. J. P. Durand has lately 1868; published a valuable essay 'De l'Influence des Milieux, &c.' He lays much stress on the nature of the soil.

[155] 'Investigations in Military and Anthrop. Statistics,' &c. 1869, by B. A. Gould, p. 93, 107, 126, 131, 134.

[156] For the Polynesians, see Prichard's 'Physical Hist. of Mankind,' vol. v. 1847, p. 145, 283. Also Godron, 'De l'Espece,' tom. ii. p. 289. There is also a remarkable difference in appearance between the closely-allied Hindoos inhabiting the Upper Ganges and Bengal; see Elphinstone's 'History of India,' vol. i. p. 324.

[157] 'Memoirs, Anthropolog. Soc.' vol. iii. 1867-69, p. 561, 565, 567.

[158] Dr. Brakenridge, 'Theory of Diathesis,' 'Medical Times,' June 19 and July 17, 1869.

[159] I have given authorities for these several statements in my 'Variation of Animals under Domestication,' vol. ii. p. 297-300. Dr. Jaeger, "Ueber das Laengenwachsthum der Knochen," 'Jenaischen Zeitschrift,' B. v. Heft i.

[160] 'Investigations,' &c. By B. A. Gould, 1869, p. 288.

[161] 'Saeugethiere von Paraguay,' 1830, s. 4.

[162] 'History of Greenland,' Eng. translat. 1767, vol. i. p. 230.

[163] 'Intermarriage.' By Alex. Walker, 1838. p. 377.

[164] 'The Variation of Animals under Domestication,' vol. i. p. 173.

[165] 'Principles of Biology,' vol. i. p. 455.

[166] Paget, 'Lectures on Surgical Pathology,' vol. i. 1853, p. 209.

[167] 'The Variation of Animals under Domestication,' vol. ii. p. 8.

[168] 'Saeugethiere von Paraguay,' s. 8, 10. I have had good opportunities for observing the extraordinary power of eyesight in the Fuegians.' See also Lawrence ('Lectures on Physiology,' &c., 1822, p. 404) on this same subject. M. Giraud-Teulon has recently collected ('Revue des Cours Scientifiques,' 1870, p. 625) a large and valuable body of evidence proving that the cause of short-sight, "_C'est le travail assidu, de pres._"

[169] Prichard, 'Phys. Hist. of Mankind,' on the authority of Blumenbach, vol. i. 1851, p. 311; for the statement by Pallas, vol. iv. 1844, p. 407.

[170] Quoted by Prichard, 'Researches into the Phys. Hist. of Mankind,' vol. v. p. 463.

[171] Mr. Forbes' valuable paper is now published in the 'Journal of the Ethnological Soc. of London,' new series, vol. ii. 1870, p. 193.

[172] Dr. Wilckens ('Landwirthschaft. Wochenblatt,' No. 10, 1869) has lately published an interesting essay shewing how domestic animals, which live in mountainous regions, have their frames modified.

[173] 'Memoire sur les Microcephales,' 1867, p. 50, 125, 169, 171, 184-198.

[174] See Dr. A. Farre's well-known article in the 'Cyclop. of Anat. and Phys.' vol. v. 1859, p. 642. Owen 'Anatomy of Vertebrates,' vol. iii. 1868, p. 687. Prof. Turner in 'Edinburgh Medical Journal,' Feb. 1865.

[175] 'Annuario della Soc. dei Naturalisti in Modena,' 1867, p. 83. Prof. Canestrini gives extracts on this subject from various authorities. Laurillard remarks, that as he has found a complete similarity in the form, proportions, and connexion of the two malar bones in several human subjects and in certain apes, he cannot consider this disposition of the parts as simply accidental.

[176] A whole series of cases is given by Isid. Geoffroy St.-Hilaire, 'Hist. des Anomalies,' tom. iii. p. 437.

[177] In my 'Variation of Animals under Domestication' (vol. ii. p. 57) I attributed the not very rare cases of supernumerary mammae in women to reversion. I was led to this as a _probable_ conclusion, by the additional mammae being generally placed symmetrically on the breast, and more especially from one case, in which a single efficient mamma occurred in the inguinal region of a woman, the daughter of another woman with supernumerary mammae. But Prof. Preyer ('Der Kampf um das Dasein,' 1869, s. 45) states that _mammae erraticae_ have been known to occur in other situations, even on the back; so that the force of my argument is greatly weakened or perhaps quite destroyed.

With much hesitation I, in the same work (vol. ii. p. 12), attributed the frequent cases of polydactylism in men to reversion. I was partly led to this through Prof. Owen's statement, that some of the Ichthyopterygia possess more than five digits, and therefore, as I supposed, had retained a primordial condition; but after reading Prof. Gegenbaur's paper ('Jenaischen Zeitschrift,' B. v. Heft 3, s. 341), who is the highest authority in Europe on such a point, and who disputes Owen's conclusion, I see that it is extremely doubtful whether supernumerary digits can thus be accounted for. It was the fact that such digits not only frequently occur and are strongly inherited, but have the power of regrowth after amputation, like the normal digits of the lower vertebrata, that chiefly led me to the above conclusion. This extraordinary fact of their regrowth remains inexplicable, if the belief in reversion to some extremely remote progenitor must be rejected. I cannot, however, follow Prof. Gegenbaur in supposing that additional digits could not reappear through reversion, without at the same time other parts of the skeleton being simultaneously and similarly modified; for single characters often reappear through reversion.

[178] 'Anatomy of Vertebrates,' vol. iii. 1868, p. 323.

[179] 'Generelle Morphologie,' 1866, B. ii. s. clv.

[180] Carl Vogt's 'Lectures on Man,' Eng. translat. 1864, p. 151.

[181] C. Carter Blake, on a jaw from La Naulette, 'Anthropolog. Review,' 1867, p. 295. Schaaffhausen, ibid. 1868, p. 426.

[182] 'The Anatomy of Expression,' 1844, p. 110, 131.

[183] Quoted by Prof. Canestrini in the 'Annuario,' &c., 1867, p. 90.

[184] These papers deserve careful study by any one who desires to learn how frequently our muscles vary, and in varying come to resemble those of the Quadrumana. The following references relate to the few points touched on in my text: Proc. Royal Soc. vol. xiv. 1865, p. 379-384; vol. xv. 1866, p. 241, 242; vol. xv. 1867, p. 544; vol. xvi. 1868, p. 524. I may here add that Dr. Murie and Mr. St. George Mivart have shewn in their Memoir on the Lemuroidea ('Transact. Zoolog. Soc.' vol. vii. 1869, p. 96), how extraordinarily variable some of the muscles are in these animals, the lowest members of the Primates. Gradations, also, in the muscles leading to structures found in animals still lower in the scale, are numerous in the Lemuroidea.

[185] Prof. Macalister in 'Proc. R. Irish Academy,' vol. x. 1868, p. 124.

[186] Prof. Macalister (ibid. p. 121) has tabulated his observations, and finds that muscular abnormalities are most frequent in the fore-arms, secondly in the face, thirdly in the foot, &c.

[187] The Rev. Dr. Haughton, after giving ('Proc. R. Irish Academy,' June 27, 1864, p. 715) a remarkable case of variation in the human _flexor pollicis longus_, adds, "This remarkable example shews that man may sometimes possess the arrangement of tendons of thumb and fingers characteristic of the macaque; but whether such a case should be regarded as a macaque passing upwards into a man, or a man passing downwards into a macaque, or as a congenital freak of nature, I cannot undertake to say." It is satisfactory to hear so capable an anatomist, and so embittered an opponent of evolutionism, admitting even the possibility of either of his first propositions. Prof. Macalister has also described ('Proc. R. Irish Acad.' vol. x. 1864, p. 138) variations in the _flexor pollicis longus_, remarkable from their relations to the same muscle in the Quadrumana.

[188] The authorities for these several statements are given in my 'Variation of Animals under Domestication,' vol. ii. p. 320-335.

[189] This whole subject has been discussed in chap. xxiii. vol. ii. of my 'Variation of Animals and Plants under Domestication.'

[190] See the ever memorable 'Essay on the Principle of Population,' by the Rev. T. Malthus, vol. i. 1826, p. 6, 517.

[191] 'Variation of Animals and Plants under Domestication,' vol. ii. p. 111-113, 163.

[192] Mr. Sedgwick, 'British and Foreign Medico-Chirurg. Review,' July, 1863, p. 170.

[193] 'The Annals of Rural Bengal,' by W. W. Hunter, 1868, p. 259.

[194] 'Primitive Marriage,' 1865.

[195] See some good remarks to this effect by W. Stanley Jevons, "A Deduction from Darwin's Theory," 'Nature,' 1869, p. 231.

[196] Latham, 'Man and his Migrations,' 1851, p. 135.

[197] Messrs. Murie and Mivart in their "Anatomy of the Lemuroidea" ('Transact. Zoolog. Soc.' vol. vii. 1869, p. 96-98) say, "some muscles are so irregular in their distribution that they cannot be well classed in any of the above groups." These muscles differ even on the opposite sides of the same individual.

[198] 'Quarterly Review,' April, 1869, p. 392. This subject is more fully discussed in Mr. Wallace's 'Contributions to the Theory of Natural Selection,' 1870, in which all the essays referred to in this work are republished. The 'Essay on Man' has been ably criticised by Prof. Claparede, one of the most distinguished zoologists in Europe, in an article published in the 'Bibliotheque Universelle,' June, 1870. The remark quoted in my text will surprise every one who has read Mr. Wallace's celebrated paper on 'The Origin of Human Races deduced from the Theory of Natural Selection,' originally published in the 'Anthropological Review,' May, 1864, p. clviii. I cannot here resist quoting a most just remark by Sir J. Lubbock ('Prehistoric Times,' 1865, p. 479) in reference to this paper, namely, that Mr. Wallace, "with characteristic unselfishness, ascribes it (_i.e._ the idea of natural selection) unreservedly to Mr. Darwin, although, as is well known, he struck out the idea independently, and published it, though not with the same elaboration, at the same time."

[199] Quoted by Mr. Lawson Tait in his "Law of Natural Selection,"--'Dublin Quarterly Journal of Medical Science,' Feb. 1869. Dr. Keller is likewise quoted to the same effect.

[200] Owen, 'Anatomy of Vertebrates,' vol. iii. p. 71.

[201] 'Quarterly Review,' April, 1869, p. 392.

[202] In _Hylobates syndactylus_, as the name expresses, two of the digits regularly cohere; and this, as Mr. Blyth informs me, is occasionally the case with the digits of _H. agilis_, _lar_, and _leuciscus_.

[203] Brehm, 'Thierleben,' B. i. s. 80.

[204] "The Hand, its mechanism," &c. 'Bridgewater Treatise,' 1833, p. 38.

[205] Haeckel has an excellent discussion on the steps by which man became a biped: 'Natuerliche Schoepfungsgeschichte,' 1868, s. 507. Dr. Buechner ('Conferences sur la Theorie Darwinienne,' 1869, p. 135) has given good cases of the use of the foot as a prehensile organ by man; also on the manner of progression of the higher apes to which I allude in the following paragraph: see also Owen ('Anatomy of Vertebrates,' vol. iii. p. 71) on this latter subject.

[206] "On the Primitive Form of the Skull," translated in 'Anthropological Review,' Oct. 1868, p. 428. Owen ('Anatomy of Vertebrates,' vol. ii. 1866, p. 551) on the mastoid processes in the higher apes.

[207] 'Die Grenzen der Thierwelt, eine Betrachtung zu Darwin's Lehre,' 1868, s. 51.

[208] Dujardin, 'Annales des Sc. Nat.' 3rd series, Zoolog. tom. xiv. 1850, p. 203. See also Mr. Lowne, 'Anatomy and Phys. of the _Musca vomitoria_,' 1870, p. 14. My son, Mr. F. Darwin, dissected for me the cerebral ganglia of the _Formica rufa_.

[209] 'Philosophical Transactions,' 1869, p. 513.

[210] Quoted in C. Vogt's 'Lectures on Man,' Eng. translat. 1864, p. 88, 90. Prichard, 'Phys. Hist. of Mankind,' vol. i. 1838, p. 305.

[211] 'Comptes Rendus des Seances,' &c. June 1, 1868.

[212] 'The Variation of Animals and Plants under Domestication,' vol. ii. p. 124-129.

[213] Schaaffhausen gives from Blumenbach and Busch, the cases of the spasms and cicatrix, in 'Anthropolog. Review,' Oct. 1868, p. 420. Dr. Jarrold ('Anthropologia,' 1808, p. 115, 116) adduces from Camper and from his own observations, cases of the modification of the skull from the head being fixed in an unnatural position. He believes that certain trades, such as that of a shoemaker, by causing the head to be habitually held forward, makes the forehead more rounded and prominent.

[214] 'Variation of Animals,' &c., vol. i. p. 117 on the elongation of the skull; p. 119, on the effect of the lopping of one ear.

[215] Quoted by Schaaffhausen, in 'Anthropolog. Review,' Oct. 1868, p. 419.

[216] Owen, 'Anatomy of Vertebrates,' vol. iii. p. 619.

[217] Isidore Geoffroy St.-Hilaire remarks ('Hist. Nat. Generale,' tom. ii. 1859, p. 215-217) on the head of man being covered with long hair; also on the upper surfaces of monkeys and of other mammals being more thickly clothed than the lower surfaces. This has likewise been observed by various authors. Prof. P. Gervais ('Hist. Nat. des Mammiferes,' tom. i. 1854, p. 28), however, states that in the Gorilla the hair is thinner on the back, where it is partly rubbed off, than on the lower surface.

[218] Mr. St. George Mivart, 'Proc. Zoolog. Soc.' 1865, p. 562, 583. Dr. J. E. Gray, 'Cat. Brit. Mus.: Skeletons.' Owen, 'Anatomy of Vertebrates,' vol. ii. p. 517. Isidore Geoffroy, 'Hist. Nat. Gen.' tom. ii. p. 244.

[219] 'The Variation of Animals and Plants under Domestication,' vol. ii. p. 280, 282.

[220] 'Primeval Man,' 1869, p. 66.

[221] 'Anthropological Review,' May, 1864, p. clviii.

[222] After a time the members or tribes which are absorbed into another tribe assume, as Mr. Maine remarks ('Ancient Law,' 1861, p. 131), that they are the co-descendants of the same ancestors.

[223] Morlot, 'Soc. Vaud. Sc. Nat.' 1860, p. 294.

[224] I have given instances in my 'Variation of Animals under Domestication,' vol. ii. p. 196.

[225] See a remarkable series of articles on Physics and Politics in the 'Fortnightly Review,' Nov. 1867; April 1, 1868; July 1, 1869.

[226] 'Origin of Civilisation,' 1870, p. 265.

[227] Mr. Wallace gives cases in his 'Contributions to the Theory of Natural Selection,' 1870, p. 354.

[228] 'Ancient Law,' 1861, p. 22. For Mr. Bagehot's remarks, 'Fortnightly Review,' April 1, 1868, p. 452.

[229] 'The Variation of Animals and Plants under Domestication,' vol. i. p. 309.

[230] 'Fraser's Magazine,' Sept. 1868, p. 353. This article seems to have struck many persons, and has given rise to two remarkable essays and a rejoinder in the 'Spectator,' Oct. 3rd and 17th 1868. It has also been discussed in the 'Q. Journal of Science,' 1869, p. 152, and by Mr. Lawson Tait in the 'Dublin Q. Journal of Medical Science,' Feb. 1869, and by Mr. E. Ray Lankester in his 'Comparative Longevity,' 1870, p. 128. Similar views appeared previously in the 'Australasian,' July 13, 1867. I have borrowed ideas from several of these writers.

[231] For Mr. Wallace, see 'Anthropolog. Review,' as before cited. Mr. Galton in 'Macmillan's Magazine,' Aug. 1865, p. 318; also his great work, 'Hereditary Genius,' 1870.

[232] 'Hereditary Genius,' 1870, p. 132-140.

[233] See the fifth and sixth columns, compiled from good authorities, in the table given in Mr. E. R. Lankester's 'Comparative Longevity,' 1870, p. 115.

[234] 'Hereditary Genius,' 1870, p. 330.

[235] 'Origin of Species' (fifth edition, 1869), p. 104.

[236] 'Hereditary Genius,' 1870, p. 347.

[237] E. Ray Lankester, 'Comparative Longevity,' 1870, p. 115. The table of the intemperate is from Nelson's 'Vital Statistics.' In regard to profligacy, see Dr. Farr, "Influence of Marriage on Mortality," 'Nat. Assoc. for the Promotion of Social Science,' 1858.

[238] 'Fraser's Magazine,' Sept. 1868, p. 353. 'Macmillan's Magazine,' Aug. 1865, p. 318. The Rev. F. W. Farrar ('Fraser's Mag.,' Aug. 1870, p. 264) takes a different view.

[239] "On the Laws of the Fertility of Women," in 'Transact. Royal Soc.' Edinburgh, vol. xxiv. p. 287. See, also, Mr. Galton, 'Hereditary Genius,' p. 352-357, for observations to the above effect.

[240] 'Tenth Annual Report of Births, Deaths, &c., in Scotland,' 1867, p. xxix.

[241] These quotations are taken from our highest authority on such questions, namely, Dr. Farr, in his paper "On the Influence of Marriage on the Mortality of the French People," read before the Nat. Assoc. for the Promotion of Social Science, 1858.

[242] Dr. Farr, ibid. The quotations given below are extracted from the same striking paper.

[243] I have taken the mean of the quinquennial means, given in 'The Tenth Annual Report of Births, Deaths, &c., in Scotland,' 1867. The quotation from Dr. Stark is copied from an article in the 'Daily News,' Oct. 17th, 1868, which Dr. Farr considers very carefully written.

[244] See the ingenious and original argument on this subject by Mr. Galton, 'Hereditary Genius,' p. 340-342.

[245] Mr. Greg, 'Fraser's Magazine,' Sept. 1868, p. 357.

[246] 'Hereditary Genius,' 1870, p. 357-359. The Rev. F. H. Farrar ('Fraser's Mag.' , Aug. 1870, p. 257) advances arguments on the other side. Sir C. Lyell had already ('Principles of Geology,' vol. ii. 1868, p. 489) called attention, in a striking passage, to the evil influence of the Holy Inquisition in having lowered, through selection, the general standard of intelligence in Europe.

[247] Mr. Galton, 'Macmillan's Magazine,' August, 1865, p. 325. See, also, 'Nature,' "On Darwinism and National Life," Dec. 1869, p. 184.

[248] 'Last Winter in the United States,' 1868, p. 29.

[249] 'On the Origin of Civilisation,' 'Proc. Ethnological Soc.' Nov. 26, 1867.

[250] 'Primeval Man,' 1869.

[251] 'Royal Institution of Great Britain,' March 15, 1867. Also, 'Researches into the Early History of Mankind,' 1865.

[252] 'Primitive Marriage,' 1865. See, likewise, an excellent article, evidently by the same author, in the 'North British Review,' July, 1869. Also, Mr. L. H. Morgan, "A Conjectural Solution of the Origin of the Class. System of Relationship," in 'Proc. American Acad. of Sciences,' vol. vii. Feb. 1868. Prof. Schaaffhausen ('Anthropolog. Review,' Oct. 1869, p. 373) remarks on "the vestiges of human sacrifices found both in Homer and the Old Testament."

[253] Sir J. Lubbock, 'Prehistoric Times,' 2nd edit. 1869, chap. xv. and xvi. _et passim_.

[254] Dr. F. Mueller has made some good remarks to this effect in the 'Reise der Novara: Anthropolog. Theil,' Abtheil. iii. 1868, s. 127.

[255] Isidore Geoffroy St.-Hilaire gives a detailed account of the position assigned to man by various naturalists in their classifications: 'Hist. Nat. Gen.' tom. ii. 1859, p. 170-189.

[256] See the very interesting article, "L'Instinct chez les Insectes," by M. George Pouchet, 'Revue des Deux Mondes,' Feb. 1870, p. 682.

[257] Westwood, 'Modern Class. of Insects,' vol. ii. 1840, p. 87.

[258] 'Proc. Zoolog. Soc.' 1869, p. 4.

[259] 'Evidence as to Man's Place in Nature,' 1863, p. 70, _et passim_.

[260] Isid. Geoffroy, 'Hist. Nat. Gen.' tom. ii. 1859, p. 217.

[261] "Ueber die Richtung der Haare," &c., Mueller's 'Archiv fuer Anat. und Phys.' 1837, s. 51.

[262] On the hair in Hylobates, see 'Nat. Hist. of Mammals,' by C. L. Martin, 1841, p. 415. Also, Isid. Geoffroy on the American monkeys and other kinds, 'Hist. Nat. Gen.' vol. ii. 1859, p. 216, 243. Eschricht, ibid., s. 46, 55, 61. Owen, 'Anat. of Vertebrates,' vol. iii. p. 619. Wallace, 'Contributions to the Theory of Natural Selection,' 1870. p. 344.

[263] 'Origin of Species,' 5th edit. 1869, p. 194. 'The Variation of Animals and Plants under Domestication,' vol. ii. 1868, p. 348.

[264] 'An Introduction to the Classification of Animals,' 1869, p. 99.

[265] This is nearly the same classification as that provisionally adopted by Mr. St. George Mivart ('Transact. Philosoph. Soc.' 1867, p. 300), who, after separating the Lemuridae, divides the remainder of the Primates into the Hominidae, the Simiadae answering to the Catarhines, the Cebidae, and the Hapalidae,--these two latter groups answering to the Platyrhines.

[266] 'Transact. Zoolog. Soc.' vol. vi. 1867, p. 214.

[267] Mr. St. G. Mivart, 'Transact. Phil. Soc.' 1867, p. 410.

[268] Messrs. Murie and Mivart on the Lemuroidea. 'Transact. Zoolog. Soc.' vol. vii. 1869, p. 5.

[269] Haeckel has come to this same conclusion. See 'Ueber die Entstehung des Menschengeschlechts,' in Virchow's 'Sammlung. gemein. wissen. Vortraege,' 1868, s. 61. Also his 'Natuerliche Schoepfungsgeschichte,' 1868, in which he gives in detail his views on the genealogy of man.

[270] 'Anthropological Review,' April, 1867, p. 236.

[271] 'Elements of Geology,' 1865, p. 583-585. 'Antiquity of Man', 1863; p. 145.

[272] 'Man's Place in Nature,' p. 105.

[273] Elaborate tables are given in his 'Generelle Morphologie' (B. ii. s. cliii. and s. 425); and with more especial reference to man in his 'Natuerliche Schoepfungsgeschichte,' 1868. Prof. Huxley, in reviewing this latter work ('The Academy,' 1869, p. 42) says, that he considers the phylum or lines of descent of the Vertebrata to be admirably discussed by Haeckel, although he differs on some points. He expresses, also, his high estimate of the value of the general tenor and spirit of the whole work.

[274] 'Palaeontology,' 1860, p. 199.

[275] I had the satisfaction of seeing, at the Falkland Islands, in April, 1833, and therefore some years before any other naturalist, the locomotive larvae of a compound Ascidian, closely allied to, but apparently generically distinct from, Synoicum. The tail was about five times as long as the oblong head, and terminated in a very fine filament. It was plainly divided, as sketched by me under a simple microscope, by transverse opaque partitions, which I presume represent the great cells figured by Kowalevsky. At an early stage of development the tail was closely coiled round the head of the larva.

[276] 'Memoires de l'Acad. des Sciences de St. Petersbourg,' tom. x. No. 15, 1866.

[277] This is the conclusion of one of the highest authorities in comparative anatomy, namely, Prof. Gegenbaur: 'Grundzuege der vergleich. Anat.' 1870, s. 876. The result has been arrived at chiefly from the study of the Amphibia; but it appears from the researches of Waldeyer (as quoted in Humphry's 'Journal of Anat. and Phys.' 1869, p. 161), that the sexual organs of even "the higher vertebrata are, in their early condition, hermaphrodite." Similar views have long been held by some authors, though until recently not well based.

[278] The male Thylacinus offers the best instance. Owen, 'Anatomy of Vertebrates,' vol. iii. p. 771.

[279] Serranus is well known often to be in an hermaphrodite condition; but Dr. Guenther informs me that he is convinced that this is not its normal state. Descent from an ancient androgynous prototype would, however, naturally favour and explain, to a certain extent, the recurrence of this condition in these fishes.

[280] Mr. Lockwood believes (as quoted in 'Quart. Journal of Science,' April, 1868, p. 269), from what he has observed of the development of Hippocampus, that the walls of the abdominal pouch of the male in some way afford nourishment. On male fishes hatching the ova in their mouths, see a very interesting paper by Prof. Wyman, in 'Proc. Boston Soc. of Nat. Hist.' Sept. 15, 1857; also Prof. Turner, in 'Journal of Anat. and Phys.' Nov. 1, 1866, p. 78. Dr. Guenther has likewise described similar cases.

[281] All vital functions tend to run their course in fixed and recurrent periods, and with tidal animals the periods would probably be lunar; for such animals must have been left dry or covered deep with water,--supplied with copious food or stinted,--during endless generations, at regular lunar intervals. If then the Vertebrata are descended from an animal allied to the existing tidal Ascidians, the mysterious fact, that with the higher and now terrestrial Vertebrata, not to mention other classes, many normal and abnormal vital processes run their course according to lunar periods, is rendered intelligible. A recurrent period, if approximately of the right duration, when once gained, would not, as far as we can judge, be liable to be changed; consequently it might be thus transmitted during almost any number of generations. This conclusion, if it could be proved sound, would be curious; for we should then see that the period of gestation in each mammal, and the hatching of each bird's eggs, and many other vital processes, still betrayed the primordial birthplace of these animals.

[282] 'History of India,' 1841, vol. i. p. 323. Father Ripa makes exactly the same remark with respect to the Chinese.

[283] A vast number of measurements of Whites, Blacks, and Indians, are given in the 'Investigations in the Military and Anthropolog. Statistics of American Soldiers,' by B. A. Gould, 1869, p. 298-358; on the capacity of the lungs, p. 471. See also the numerous and valuable tables, by Dr. Weisbach, from the observations of Dr. Scherzer and Dr. Schwarz, in the 'Reise der Novara: Anthropolog. Theil,' 1867.

[284] See, for instance, Mr. Marshall's account of the brain of a Bush-woman, in 'Phil. Transact.' 1864, p. 519.

[285] Wallace, 'The Malay Archipelago,' vol. ii. 1869, p. 178.

[286] With respect to the figures in the famous Egyptian caves of Abou-Simbel, M. Pouchet says ('The Plurality of the Human Races,' Eng. translat. 1864, p. 50), that he was far from finding recognisable representations of the dozen or more nations which some authors believe that they can recognise. Even some of the most strongly-marked races cannot be identified with that degree of unanimity which might have been expected from what has been written on the subject. Thus Messrs. Nott and Gliddon ('Types of Mankind,' p. 148) state that Rameses II., or the Great, has features superbly European; whereas Knox, another firm believer in the specific distinction of the races of man ('Races of Man,' 1850, p. 201), speaking of young Memnon (the same person with Rameses II., as I am informed by Mr. Birch) insists in the strongest manner that he is identical in character with the Jews of Antwerp. Again, whilst looking in the British Museum with two competent judges, officers of the establishment, at the statue of Amunoph III., we agreed that he had a strongly negro cast of features; but Messrs. Nott and Gliddon (ibid. p. 146, fig. 53) describe him as "a hybrid, but not of negro intermixture."

[287] As quoted by Nott and Gliddon, 'Types of Mankind,' 1854, p. 439. They give also corroborative evidence; but C. Vogt thinks that the subject requires further investigation.

[288] "Diversity of Origin of the Human Races," in the 'Christian Examiner,' July, 1850.

[289] 'Transact. B. Soc. of Edinburgh,' vol. xxii. 1861, p. 567.

[290] 'On the Phenomena of Hybridity in the Genus Homo,' Eng. translat. 1864.

[291] See the interesting letter by Mr. T. A. Murray, in the 'Anthropolog. Review,' April, 1868, p. liii. In this letter Count Strzelecki's statement, that Australian women who have borne children to a white man are afterwards sterile with their own race, is disproved. M. A. de Quatrefages has also collected ('Revue des Cours Scientifiques,' March, 1869, p. 239) much evidence that Australians and Europeans are not sterile when crossed.

[292] 'An Examination of Prof. Agassiz's Sketch of the Nat. Provinces of the Animal World,' Charleston, 1855, p. 44.

[293] 'Military and Anthropolog. Statistics of American Soldiers,' by B. A. Gould, 1869, p. 319.

[294] 'The Variation of Animals and Plants under Domestication,' vol. ii. p. 109. I may here remind the reader that the sterility of species when crossed is not a specially-acquired quality; but, like the incapacity of certain trees to be grafted together, is incidental on other acquired differences. The nature of these differences is unknown, but they relate more especially to the reproductive system, and much less to external structure or to ordinary differences in constitution. One important element in the sterility of crossed species apparently lies in one or both having been long habituated to fixed conditions; for we know that changed conditions have a special influence on the reproductive system, and we have good reason to believe (as before remarked) that the fluctuating conditions of domestication tend to eliminate that sterility which is so general with species in a natural state when crossed. It has elsewhere been shewn by me (ibid. vol. ii. p. 185, and 'Origin of Species,' 5th edit. p. 317) that the sterility of crossed species has not been acquired through natural selection: we can see that when two forms have already been rendered very sterile, it is scarcely possible that their sterility should be augmented by the preservation or survival of the more and more sterile individuals; for as the sterility increases fewer and fewer offspring will be produced from which to breed, and at last only single individuals will be produced, at the rarest intervals. But there is even a higher grade of sterility than this. Both Gaertner and Koelreuter have proved that in genera of plants including numerous species, a series can be formed from species which when crossed yield fewer and fewer seeds, to species which never produce a single seed, but yet are affected by the pollen of the other species, for the germen swells. It is here manifestly impossible to select the more sterile individuals, which have already ceased to yield seeds; so that the acme of sterility, when the germen alone is affected, cannot be gained through selection. This acme, and no doubt the other grades of sterility, are the incidental results of certain unknown differences in the constitution of the reproductive system of the species which are crossed.

[295] 'The Variation of Animals,' &c., vol. ii. p. 92.

[296] M. de Quatrefages has given ('Anthropolog. Review,' Jan. 1869, p. 22) an interesting account of the success and energy of the Paulistas in Brazil, who are a much crossed race of Portuguese and Indians, with a mixture of the blood of other races.

[297] For instance with the aborigines of America and Australia. Prof. Huxley says ('Transact. Internat. Congress of Prehist. Arch.' 1868. p. 105) that the skulls of many South Germans and Swiss are "as short and as broad as those of the Tartars," &c.

[298] See a good discussion on this subject in Waitz, 'Introduct. to Anthropology,' Eng. translat. 1863, p. 198-208, 227. I have taken some of the above statements from H. Tuttle's 'Origin and Antiquity of Physical Man,' Boston, 1866, p. 35.

[299] Prof. Naegeli has carefully described several striking cases in his 'Botanische Mittheilungen,' B. ii. 1866, s. 294-369. Prof. Asa Gray has made analogous remarks on some intermediate forms in the Compositae of N. America.

[300] 'Origin of Species,' 5th edit. p. 68.

[301] See Prof. Huxley to this effect in the 'Fortnightly Review,' 1865, p. 275.

[302] 'Lectures on Man,' Eng. translat. 1864, p. 468.

[303] 'Die Racen des Schweines,' 1860, s. 46. 'Vorstudien fuer Geschichte, &c., Schweineschaedel,' 1864, s. 104. With respect to cattle, see M. de Quatrefages, 'Unite de l'Espece Humaine,' 1861, p. 119.

[304] Tylor's 'Early History of Mankind,' 1865; for the evidence with respect to gesture-language, see p. 54. Lubbock's 'Prehistoric Times,' 2nd edit. 1869.

[305] 'The Primitive Inhabitants of Scandinavia,' Eng. translat. edited by Sir J. Lubbock, 1868, p. 104.

[306] Hodder M. Westropp, on Cromlechs, &c., 'Journal of Ethnological Soc.' as given in 'Scientific Opinion,' June 2nd. 1869, p. 3.

[307] 'Journal of Researches: Voyage of the "Beagle,"' p. 46.

[308] 'Prehistoric Times,' 1869, p. 574.

[309] Translation in 'Anthropological Review,' Oct. 1868, p. 431.

[310] 'Transact. Internat. Congress of Prehistoric Arch.' 1868, p. 172-175. See also Broca (translation) in 'Anthropological Review,' Oct. 1868, p. 410.

[311] Dr. Gerland, 'Ueber das Aussterben der Naturvoelker,' 1868, s. 82.

[312] Gerland (ibid. s. 12) gives facts in support of this statement.

[313] See remarks to this effect in Sir H. Holland's 'Medical Notes and Reflections,' 1839, p. 390.

[314] I have collected ('Journal of Researches, Voyage of the "Beagle,"' p. 435) a good many cases bearing on this subject: see also Gerland, ibid. s. 8. Poeppig speaks of the "breath of civilisation as poisonous to savages."

[315] Sproat, 'Scenes and Studies of Savage Life,' 1868, p. 284.

[316] Bagehot, "Physics and Politics," 'Fortnightly Review,' April 1, 1868, p. 455.

[317] "On Anthropology," translation, 'Anthropolog. Review,' Jan. 1868, p. 38.

[318] 'The Annals of Rural Bengal,' 1868, p. 134.

[319] 'The Variation of Animals and Plants under Domestication,' vol. ii. p. 95.

[320] Pallas, 'Act. Acad. St. Petersburgh,' 1780, part ii. p. 69. He was followed by Rudolphi, in his 'Beytraege zur Anthropologie,' 1812. An excellent summary of the evidence is given by Godron, 'De l'Espece,' 1859, vol. ii. p. 246, &c.

[321] Sir Andrew Smith, as quoted by Knox, 'Races of Man,' 1850, p. 473.

[322] See De Quatrefages on this head, 'Revue des Cours Scientifiques,' Oct. 17, 1868, p. 731.

[323] Livingstone's 'Travels and Researches in S. Africa,' 1857, p. 338, 329. D'Orbigny, as quoted by Godron, 'De l'Espece,' vol. ii. p. 266.

[324] See a paper read before the Royal Soc. in 1813, and published in his Essays in 1818. I have given an account of Dr. Wells' views in the Historical Sketch (p. xvi) to my 'Origin of Species.' Various cases of colour correlated with constitutional peculiarities are given in my 'Variation of Animals under Domestication,' vol. ii. p. 227, 335.

[325] See, for instance, Nott and Gliddon, 'Types of Mankind,' p. 68.

[326] Major Tulloch, in a paper read before the Statistical Society, April 20th, 1840, and given in the 'Athenaeum,' 1840, p. 353.

[327] 'The Plurality of the Human Race' (translat.), 1864, p. 60.

[328] Quatrefages, 'Unite de l'Espece Humaine,' 1861, p. 205. Waitz, 'Introduct. to Anthropology,' translat. vol. i. 1863, p. 124. Livingstone gives analogous cases in his 'Travels.'

[329] In the spring of 1862 I obtained permission from the Director-General of the Medical department of the Army, to transmit to the surgeons of the various regiments on foreign service a blank table, with the following appended remarks, but I have received no returns. "As several well-marked cases have been recorded with our domestic animals of a relation between the colour of the dermal appendages and the constitution; and it being notorious that there is some limited degree of relation between the colour of the races of man and the climate inhabited by them; the following investigation seems worth consideration. Namely, whether there is any relation in Europeans between the colour of their hair, and their liability to the diseases of tropical countries. If the surgeons of the several regiments, when stationed in unhealthy tropical districts, would be so good as first to count, as a standard of comparison, how many men, in the force whence the sick are drawn, have dark and light-coloured hair, and hair of intermediate or doubtful tints; and if a similar account were kept by the same medical gentlemen, of all the men who suffered from malarious and yellow fevers, or from dysentery, it would soon be apparent, after some thousand cases had been tabulated, whether there exists any relation between the colour of the hair and constitutional liability to tropical diseases. Perhaps no such relation would be discovered, but the investigation is well worth making. In case any positive result were obtained, it might be of some practical use in selecting men for any particular service. Theoretically the result would be of high interest, as indicating one means by which a race of men inhabiting from a remote period an unhealthy tropical climate, might have become dark-coloured by the better preservation of dark-haired or dark-complexioned individuals during a long succession of generations."

[330] 'Anthropological Review,' Jan. 1866, p. xxi.

[331] See, for instance, Quatrefages ('Revue des Cours Scientifiques,' Oct. 10, 1868, p. 724) on the effects of residence in Abyssinia and Arabia, and other analogous cases. Dr. Rolle ('Der Mensch, seine Abstammung,' &c., 1865, s. 99) states, on the authority of Khanikof, that the greater number of German families settled in Georgia, have acquired in the course of two generations dark hair and eyes. Mr. D. Forbes informs me that the Quichuas in the Andes vary greatly in colour, according to the position of the valleys inhabited by them.

[332] Harlan, 'Medical Researches,' p. 532. Quatrefages ('Unite de l'Espece Humaine,' 1861, p. 128) has collected much evidence on this head.

[333] See Prof. Schaaffhausen, translat. in 'Anthropological Review,' Oct. 1868, p. 429.

[334] Mr. Catlin states ('N. American Indians,' 3rd edit. 1842, vol. i. p. 49) that in the whole tribe of the Mandans, about one in ten or twelve of the members of all ages and both sexes have bright silvery grey hair, which is hereditary. Now this hair is as coarse and harsh as that of a horse's mane, whilst the hair of other colours is fine and soft.

[335] On the odour of the skin, Godron, 'Sur l'Espece,' tom. ii. p. 217. On the pores in the skin, Dr. Wilckens, 'Die Aufgaben der landwirth. Zootechnik,' 1869, s. 7.

[336] Westwood, 'Modern Class. of Insects,' vol. ii. 1810, p. 541. In regard to the statement about Tanais, mentioned below, I am indebted to Fritz Mueller.

[337] Kirby and Spence, 'Introduction to Entomology,' vol. iii. 1826, p. 309.

[338] Even with those of plants in which the sexes are separate, the male flowers are generally mature before the female. Many hermaphrodite plants are, as first shewn by C. K. Sprengel, dichogamous; that is, their male and female organs are not ready at the same time, so that they cannot be self-fertilised. Now with such plants the pollen is generally mature in the same flower before the stigma, though there are some exceptional species in which the female organs are mature before the male.

[339] I have received information, hereafter to be given, to this effect with respect to poultry. Even with birds, such as pigeons, which pair for life, the female, as I hear from Mr. Jenner Weir, will desert her mate if he is injured or grows weak.

[340] On the Gorilla, Savage and Wyman, 'Boston Journal of Nat. Hist.' vol. v. 1845-47, p. 423. On Cynocephalus, Brehm, 'Illust. Thierleben,' B. i. 1864, s. 77. On Mycetes, Rengger, 'Naturgesch.: Saeugethiere von Paraguay,' 1830, s. 14, 20. On Cebus, Brehm, ibid. s. 108.

[341] Pallas, 'Spicilegia Zoolog.' Fasc. xii. 1777, p. 29. Sir Andrew Smith, 'Illustrations of the Zoology of S. Africa,' 1849, pl. 29, on the Kobus. Owen, in his 'Anatomy of Vertebrates' (vol. iii. 1868, p. 633) gives a table incidentally showing which species of Antelopes pair and which are gregarious.

[342] Dr. Campbell, in 'Proc. Zoolog. Soc.' 1869, p. 138. See also an interesting paper, by Lieut. Johnstone, in 'Proc. Asiatic Soc. of Bengal,' May, 1868.

[343] 'The Ibis,' vol. iii. 1861, p. 133, on the Progne Widow-bird. See also on the Vidua axillaris, ibid. vol. ii. 1860, p. 211. On the polygamy of the Capercailzie and Great Bustard, see L. Lloyd, 'Game Birds of Sweden,' 1867, p. 19, and 182. Montagu and Selby speak of the Black Grouse as polygamous and of the Red Grouse as monogamous.

[344] The Rev. E. S. Dixon, however, speaks positively ('Ornamental Poultry,' 1848, p. 76) about the eggs of the guinea-fowl being infertile when more than one female is kept with the same male.

[345] Noel Humphreys, 'River Gardens,' 1857.

[346] Kirby and Spence, 'Introduction to Entomology,' vol. iii. 1826, p. 342.

[347] One parasitic Hymenopterous insect (Westwood, 'Modern Class. of Insects,' vol. ii, p. 160) forms an exception to the rule, as the male has rudimentary wings, and never quits the cell in which it is born, whilst the female has well-developed wings. Audouin believes that the females are impregnated by the males which are born in the same cells with them; but it is much more probable that the females visit other cells, and thus avoid close inter-breeding. We shall hereafter meet with a few exceptional cases, in various classes, in which the female, instead of the male, is the seeker and wooer.

[348] 'Essays and Observations,' edited by Owen, vol. i. 1861, p. 194.

[349] Prof. Sachs ('Lehrbuch der Botanik,' 1870, s. 633) in speaking of the male and female reproductive cells, remarks, "verhaelt sich die eine bei der Vereinigung activ, ... die andere erscheint bei der Vereinigung passiv."

[350] 'Reise der Novara: Anthropolog. Theil,' 1867, s. 216-269. The results were calculated by Dr. Weisbach from measurements made by Drs. K. Scherzer and Schwarz. On the greater variability of the males of domesticated animals, see my 'Variation of Animals and Plants under Domestication,' vol. ii. 1868, p. 75.

[351] 'Proceedings Royal Soc.' vol. xvi. July, 1868, p. 519 and 524.

[352] 'Proc. Royal Irish Academy,' vol. x. 1868, p. 123.

[353] 'Massachusetts Medical Soc.' vol. ii. No. 3, 1808, p. 9.

[354] 'The Variation of Animals and Plants under Domestication,' vol. ii. 1868, p. 75. In the last chapter but one, the provisional hypothesis of pangenesis, above alluded to, is fully explained.

[355] These facts are given on the high authority of a great breeder, Mr. Teebay, in Tegetmeier's 'Poultry Book,' 1868, p. 158. On the characters of chickens of different breeds, and on the breeds of the pigeon, alluded to in the above paragraph, see 'Variation of Animals,' &c., vol. i. p. 160, 249; vol. ii. p. 77.

[356] 'Novae species Quadrupedum e Glirium ordine,' 1778, p. 7. On the transmission of colour by the horse, see 'Variation of Animals, &c. under Domestication,' vol. i. p. 21. Also vol. ii. p. 71, for a general discussion on Inheritance as limited by Sex.

[357] Dr. Chapuis, 'Le Pigeon Voyageur Belge,' 1865, p. 87. Boitard et Corbie, 'Les Pigeons de Voliere,' &c., 1824, p. 173.

[358] References are given in my 'Variation of Animals under Domestication,' vol. ii. p. 72.

[359] I am much obliged to Mr. Cupples for having made enquiries for me in regard to the Roebuck and Red Deer of Scotland from Mr. Robertson, the experienced head-forester to the Marquis of Breadalbane. In regard to Fallow-deer, I am obliged to Mr. Eyton and others for information. For the _Cervus alces_ of N. America, see 'Land and Water,' 1868, p. 221 and 254; and for the _C. Virginianus_ and _strongyloceros_ of the same continent, see J. D. Caton, in 'Ottawa Acad. of Nat. Sc.' 1868, p. 13. For _Cervus Eldi_ of Pegu, see Lieut. Beavan, 'Proc. Zoolog. Soc.' 1867, p. 762.

[360] _Antilocapra Americana._ Owen, 'Anatomy of Vertebrates,' vol. iii. p. 627.

[361] I have been assured that the horns of the sheep in North Wales can always be felt, and are sometimes even an inch in length, at birth. With cattle Youatt says ('Cattle,' 1834, p. 277) that the prominence of the frontal bone penetrates the cutis at birth, and that the horny matter is soon formed over it.

[362] I am greatly indebted to Prof. Victor Carus for having made inquiries for me, from the highest authorities, with respect to the merino sheep of Saxony. On the Guinea coast of Africa there is a breed of sheep in which, as with merinos, the rams alone bear horns; and Mr. Winwood Reade informs me that in the one case observed, a young ram born on Feb. 10th first showed horns on March 6th, so that in this instance the development of the horns occurred at a later period of life, conformably with our rule, than in the Welsh sheep, in which both sexes are horned.

[363] In the common peacock (_Pavo cristatus_) the male alone possesses spurs, whilst both sexes of the Java peacock (_P. muticus_) offer the unusual case of being furnished with spurs. Hence I fully expected that in the latter species they would have been developed earlier in life than in the common peacock; but M. Hegt of Amsterdam informs me, that with young birds of the previous year, belonging to both species, compared on April 23rd, 1869, there was no difference in the development of the spurs. The spurs, however, were as yet represented merely by slight knobs or elevations. I presume that I should have been informed if any difference in the rate of development had subsequently been observed.

[364] In some other species of the Duck Family the speculum in the two sexes differs in a greater degree; but I have not been able to discover whether its full development occurs later in life in the males of such species, than in the male of the common duck, as ought to be the case according to our rule. With the allied _Mergus cucullatus_ we have, however, a case of this kind: the two sexes differ conspicuously in general plumage, and to a considerable degree in the speculum, which is pure white in the male and greyish-white in the female. Now the young males at first resemble, in all respects, the female, and have a greyish-white speculum, but this becomes pure white at an earlier age than that at which the adult male acquires his other more strongly-marked sexual differences in plumage: see Audubon, 'Ornithological Biography,' vol. iii. 1835, p. 249-250.

[365] 'Das Ganze der Taubenzucht,' 1837, s. 21, 24. For the case of the streaked pigeons, see Dr. Chapuis, 'Le Pigeon Voyageur Belge.' 1865, p. 87.

[366] For full particulars and references on all these points respecting the several breeds of the Fowl, see 'Variation of Animals and Plants under Domestication,' vol. i. p. 250, 256. In regard to the higher animals, the sexual differences which have arisen under domestication are described in the same work under the head of each species.

[367] 'Twenty-ninth Annual Report of the Registrar-General for 1866.' In this report (p. xii) a special decennial table is given.

[368] For Norway and Russia, see abstract of Prof. Faye's researches, in 'British and Foreign Medico-Chirurg. Review,' April, 1867, p. 343, 345. For France, the 'Annuaire pour l'An 1867.' p. 213.

[369] In regard to the Jews, see M. Thury, 'La Loi de Production des Sexes,' 1863, p. 25.

[370] Babbage, 'Edinburgh Journal of Science,' 1829, vol. i. p. 88; also p. 90, on still-born children. On illegitimate children in England, see 'Report of Registrar-General for 1866,' p. xv.

[371] 'British and Foreign Medico-Chirurg. Review,' April, 1867, p. 343. Dr. Stark also remarks ('Tenth Annual Report of Births, Deaths, &c., in Scotland,' 1867, p. xxviii) that "These examples may suffice to shew that, at almost every stage of life, the males in Scotland have a greater liability to death and a higher death-rate than the females. The fact, however, of this peculiarity being most strongly developed at that infantile period of life when the dress, food, and general treatment of both sexes are alike, seems to prove that the higher male death-rate is an impressed, natural, and constitutional peculiarity due to sex alone."

[372] With the savage Guaranys of Paraguay, according to the accurate Azara ('Voyages dans l'Amerique merid.' tom. ii. 1809, p. 60, 179), the women in proportion to the men are as 14 to 13.

[373] Leuckart in Wagner, 'Handwoerterbuch der Phys.' B. iv. 1853, s. 774.

[374] Anthropological Review, April, 1870, p. cviii.

[375] During the last eleven years a record has been kept of the number of mares which have proved barren or prematurely slipped their foals; and it deserves notice, as shewing how infertile these highly-nurtured and rather closely-interbred animals have become, that not far from one-third of the mares failed to produce living foals. Thus during 1866, 809 male colts and 816 female colts were born, and 743 mares failed to produce offspring. During 1867, 836 males and 902 females were born, and 794 mares failed.

[376] I am much indebted to Mr. Cupples for having procured for me the above returns from Scotland, as well as some of the following returns on cattle. Mr. R. Elliot, of Laighwood, first called my attention to the premature deaths of the males,--a statement subsequently confirmed by Mr. Aitchison and others. To this latter gentleman, and to Mr. Payan, I owe my thanks for the larger returns on sheep.

[377] Bell, 'History of British Quadrupeds,' p. 100.

[378] 'Illustrations of the Zoology of S. Africa,' 1849, pl. 29.

[379] Brehm ('Illust. Thierleben,' B. iv. s. 990) comes to the same conclusion.

[380] On the authority of L. Lloyd, 'Game Birds of Sweden,' 1867, p. 12, 132.

[381] 'Nat. Hist. of Selbourne,' letter xxix. edit. of 1825, vol. i. p. 139.

[382] Mr. Jenner Weir received similar information, on making enquiries during the following year. To shew the number of chaffinches caught, I may mention that in 1869 there was a match between two experts; and one man caught in a day 62, and another 40, male chaffinches. The greatest number ever caught by one man in a single day was 70.

[383] 'Ibis,' vol. ii. p. 260, as quoted in Gould's 'Trochilidae,' 1861, p. 52. For the foregoing proportions, I am indebted to Mr. Salvin for a table of his results.

[384] 'Ibis,' 1860, p. 137; and 1867, p. 369.

[385] 'Ibis,' 1862, p. 137.

[386] Leuckart quotes Bloch (Wagner, 'Handwoerterbuch der Phys.' B. iv. 1853, s. 775), that with fish there are twice as many males as females.

[387] Quoted in the 'Farmer,' March 18, 1869, p. 369.

[388] 'The Stormontfield Piscicultural Experiments,' 1866, p. 23. The 'Field' newspaper, June 29th, 1867.

[389] 'Land and Water,' 1868, p. 41.

[390] Yarrell, 'Hist. British Fishes,' vol. i. 1836, p. 307; on the _Cyprinus carpio_, p. 331; on the _Tinca vulgaris_, p. 331; on the _Abramis brama_, p. 336. See, for the minnow (_Leuciscus phoxinus_), 'Loudon's Mag. of Nat. Hist.' vol. v. 1832, p. 682.

[391] Leuckart quotes Meinecke (Wagner, 'Handwoerterbuch der Phys.' B. iv. 1853, s. 775) that with Butterflies the males are three or four times as numerous as the females.

[392] 'The Naturalist on the Amazons,' vol. ii. 1863, p. 228, 347.

[393] Four of these cases are given by Mr. Trimen in his 'Rhopalocera Africae Australis.'

[394] Quoted by Trimen, 'Transact. Ent. Soc.' vol. v. part iv. 1866, p. 330.

[395] 'Transact. Linn. Soc.' vol. xxv. p. 37.

[396] 'Proc. Entomolog. Soc.' Feb. 17th, 1868.

[397] Quoted by Dr. Wallace in 'Proc. Ent. Soc.' 3rd series, vol. v. 1867, p. 487.

[398] Blanchard, 'Metamorphoses, Moeurs des Insectes,' 1868, p. 225-226.

[399] 'Lepidopteren-Doubblettren Liste,' Berlin, No. x. 1866.

[400] This naturalist has been so kind as to send me some results from former years, in which the females seemed to preponderate; but so many of the figures were estimates, that I found it impossible to tabulate them.

[401] Guenther's 'Record of Zoological Literature,' 1867, p. 260. On the excess of female Lucanus, ibid. p. 250. On the males of Lucanus in England, Westwood, 'Modern Class. of Insects,' vol. i. p. 187. On the Siagonium, ibid. p. 172.

[402] Walsh, in 'The American Entomologist,' vol. i. 1869, p. 103. F. Smith, 'Record of Zoological Literature,' 1867, p. 328.

[403] 'Farm Insects,' p. 45-46.

[404] 'Observations on N. American Neuroptera,' by H. Hagen and B. D. Walsh, 'Proc. Ent. Soc. Philadelphia,' Oct. 1863, p. 168, 223, 239.

[405] 'Proc. Ent. Soc. London,' Feb. 17, 1868.

[406] Another great authority in this class, Prof. Thorell of Upsala ('On European Spiders,' 1869-70, part i. p. 205) speaks as if female spiders were generally commoner than the males.

[407] See, on this subject, Mr. Pickard-Cambridge, as quoted in 'Quarterly Journal of Science,' 1868, p. 429.

[408] I have often been struck with the fact, that in several species of Primula the seeds in the capsules which contained only a few were very much larger than the numerous seeds in the more productive capsules.

[409] 'Principles of Biology,' vol. ii. 1867, chaps. ii.-xi.

[410] 'De l'Espece et de la Class.' &c., 1869, p. 106.

[411] See, for instance, the account which I have given in my 'Journal of Researches,' 1845, p. 7.

[412] I have given ('Geolog. Observations on Volcanic Islands,' 1844, p. 53) a curious instance of the influence of light on the colours of a frondescent incrustation, deposited by the surf on the coast-rocks of Ascension, and formed by the solution of triturated sea-shells.

[413] 'Facts and Arguments for Darwin,' English translat. 1869, p. 20. See the previous discussion on the olfactory threads. Sars has described a somewhat analogous case (as quoted in 'Nature,' 1870, p. 455) in a Norwegian crustacean, the _Pontoporeia affinis_.

[414] See Sir J. Lubbock in 'Annals. and Mag. of Nat. Hist.' vol. xi. 1853, pl. i. and x.; and vol. xii. (1853) pl. vii. See also Lubbock in 'Transact. Ent. Soc.' vol. iv. new series, 1856-1858, p. 8. With respect to the zigzagged antennae mentioned below, see Fritz Mueller, 'Facts and Arguments for Darwin' 1869, p. 40, foot-note.

[415] See a paper by Mr. C. Spence Bate, with figures, in 'Proc. Zoolog. Soc.' 1868, p. 363; and on the nomenclature of the genus, ibid. p. 585. I am greatly indebted to Mr. Spence Bate for nearly all the above statements with respect to the chelae of the higher crustaceans.

[416] 'Hist. Nat. des Crust.' tom. ii. 1837, p. 50.

[417] Fritz Mueller, 'Facts and Arguments for Darwin,' 1869, p. 25-28.

[418] 'Travels in the Interior of Brazil,' 1846, p. 111. I have given, in my 'Journal of Researches,' p. 463, an account of the habits of the Birgos.

[419] Mr. Ch. Fraser, in 'Proc. Zoolog. Soc.' 1869, p. 3. I am indebted to Mr. Bate for the statement from Dr. Power.

[420] Claus, 'Die freilebenden Copepoden,' 1863, s. 35.

[421] 'Facts and Arguments,' &c., p. 79.

[422] 'A History of the Spiders of Great Britain,' 1861-64. For the following facts, see p. 102, 77, 88.

[423] Aug. Vinson ('Araneides des Iles de la Reunion,' pl. vi. figs. 1 and 2) gives a good instance of the small size of the male in _Epeira nigra_. In this species, as I may add, the male is testaceous and the female black with legs banded with red. Other even more striking cases of inequality in size between the sexes have been recorded ('Quarterly Journal of Science,' 1868, July, p. 429); but I have not seen the original accounts.

[424] Kirby and Spence, 'Introduction to Entomology,' vol. i. 1818, p. 280.

[425] Theridion (Asagena, Sund.) serratipes, 4-punctatum et guttatum; see Westring, in Kroyer, 'Naturhist. Tidskrift,' vol. iv. 1842-1843, p. 349; and vol. ii. 1846-1849, p. 342. See, also, for other species, 'Araneae Svecicae,' p. 184.

[426] Walckenaer et P. Gervais, 'Hist. Nat. des Insectes: Apteres,' tom. iv. 1847, p. 17, 19, 68.

[427] Sir J. Lubbock, 'Transact. Linnean Soc.' vol. xxv. 1866, p. 484. With respect to the Mutillidae see Westwood, 'Modern Class. of Insects,' vol. ii. p. 213.

[428] These organs in the male often differ in closely-allied species, and afford excellent specific characters. But their importance, under a functional point of view, as Mr. E. MacLachlan has remarked to me, has probably been overrated. It has been suggested, that slight differences in these organs would suffice to prevent the intercrossing of well-marked varieties or incipient species, and would thus aid in their development. That this can hardly be the case, we may infer from the many recorded cases (see for instance, Bronn, 'Geschichte der Natur,' B. ii. 1843, s. 164; and Westwood, 'Transact. Ent. Soc.' vol. iii. 1842, p. 195) of distinct species having been observed in union. Mr. MacLachlan informs me (vide 'Stett. Ent. Zeitung,' 1867, s. 155) that when several species of Phryganidae, which present strongly-pronounced differences of this kind, were confined together by Dr. Aug. Meyer, _they coupled_, and one pair produced fertile ova.

[429] 'The Practical Entomologist,' Philadelphia, vol. ii. May, 1867, p. 88.

[430] Mr. Walsh, ibid. p. 107.

[431] 'Modern Classification of Insects,' vol. ii. 1840, p. 206, 205. Mr. Walsh, who called my attention to this double use of the jaws, says that he has repeatedly observed this fact.

[432] We have here a curious and inexplicable case of dimorphism, for some of the females of four European species of Dytiscus, and of certain species of Hydroporus, have their elytra smooth; and no intermediate gradations between sulcated or punctured and quite smooth elytra have been observed. See Dr. H. Schaum, as quoted in the 'Zoologist,' vol. v.-vi. 1847-48, p. 1896. Also Kirby and Spence, 'Introduction to Entomology,' vol. iii. 1826, p. 305.

[433] Westwood, 'Modern Class.' vol. ii. p. 193. The following statement about Penthe, and others in inverted commas, are taken from Mr. Walsh, 'Practical Entomologist,' Philadelphia, vol. ii. p. 88.

[434] Kirby and Spence, 'Introduct.' &c., vol. iii. p. 332-336.

[435] 'Insecta Maderensia,' 1854, p. 20.

[436] E. Doubleday, 'Annals and Mag. of Nat. Hist.' vol. i. 1848, p. 379. I may add that the wings in certain Hymenoptera (see Shuckard, 'Fossorial Hymenop.' 1837, p. 39-43) differ in neuration according to sex.

[437] H. W. Bates, in 'Journal of Proc. Linn. Soc.' vol. vi. 1862, p. 74. Mr. Wonfor's observations are quoted in 'Popular Science Review,' 1868, p. 343.

[438] Kirby and Spence, 'Introduction to Entomology,' vol. iii. p. 299.

[439] Robinet, 'Vers a Soie,' 1848, p. 207.

[440] 'Transact. Ent. Soc.' 3rd series, vol. v. p. 486.

[441] 'Journal of Proc. Ent. Soc.' Feb. 4th, 1867, p. lxxi.

[442] For this and other statements on the size of the sexes, see Kirby and Spence, ibid. vol. iii. p. 300; on the duration of life in insects, see p. 344.

[443] 'Transact. Linnean Soc.' vol. xxvi. 1868, p. 296.

[444] 'The Malay Archipelago,' vol. ii. 1869, p. 313.

[445] 'Modern Classification of Insects,' vol. ii. 1840, p. 526.

[446] See Mr. B. T. Lowne's very interesting work, 'On the Anatomy of the Blow-Fly, Musca vomitoria,' 1870, p. 14.

[447] Westwood, 'Modern Class. of Insects,' vol. ii. p. 473.

[448] These particulars are taken from Westwood's 'Modern Class. of Insects,' vol. ii. 1840, p. 422. See, also, on the Fulgoridae, Kirby and Spence, 'Introduct.' vol. ii. p. 401.

[449] 'Zeitschrift fuer wissenschaft. Zoolog.' B. xvii. 1867, s. 152-158.

[450] I am indebted to Mr. Walsh for having sent me this extract from a 'Journal of the Doings of Cicada septemdecim,' by Dr. Hartman.

[451] L. Guilding, 'Transact. Linn. Soc.' vol. xv. p. 154.

[452] Koeppen, as quoted in the 'Zoological Record,' for 1867, p. 460.

[453] Gilbert White, 'Nat. Hist. of Selborne,' vol. ii. 1825, p. 262.

[454] Harris, 'Insects of New England,' 1842, p. 128.

[455] 'The Naturalist on the Amazons,' vol. i. 1863, p. 252. Mr. Bates gives a very interesting discussion on the gradations in the musical apparatus of the three families. See also Westwood, 'Modern Class.' vol. ii. p. 445 and 453.

[456] 'Proc. Boston Soc. of Nat. Hist.' vol. xi. April, 1868.

[457] 'Nouveau Manuel d'Anat. Comp.' (French translat.), tom. i. 1850 p. 567.

[458] 'Zeitschrift fuer wissenschaft. Zoolog.' B. xvii. 1867, s. 117.

[459] Westwood, 'Modern Class. of Insects,' vol. i. p. 440.

[460] Westwood, 'Modern Class. of Insects,' vol. i. p. 453.

[461] Landois, ibid. s. 121, 122.

[462] Mr. Walsh also informs me that he has noticed that the female of the _Platyphyllum concavum_, "when captured makes a feeble grating noise by shuffling her wing-covers together."

[463] Landois, ibid. s. 113.

[464] 'Insects of New England,' 1842, p. 133.

[465] Westwood, 'Modern Classification,' vol. i. p. 462.

[466] Westwood, ibid. vol. i. p. 453.

[467] Landois, ibid. s. 115, 116, 120, 122.

[468] 'Transact. Ent. Soc.' 3rd series, vol. ii. ('Journal of Proceedings, p. 117.)

[469] Westwood, 'Modern Class. of Insects,' vol. i. p. 427; for crickets, p. 445.

[470] Mr. Ch. Horne, in 'Proc. Ent. Soc.' May 3, 1869, p. xii.

[471] The _Oecanthus nivalis_, Harris, 'Insects of New England,' 1842, p. 124.

[472] Platyblemnus: Westwood, 'Modern. Class.' vol. i. p. 447.

[473] B. D. Walsh, the Pseudo-neuroptera of Illinois, in 'Proc. Ent. Soc. of Philadelphia,' 1862, p. 361.

[474] 'Modern Class.' vol. ii. p. 37.

[475] Walsh, ibid. p. 381. I am indebted to this naturalist for the following facts on Hetaerina, Anax, and Gomphus.

[476] 'Transact. Ent. Soc' vol. i. 1836, p. lxxxi.

[477] See abstract in the 'Zoological Record' for 1867, p. 450.

[478] Kirby and Spence, 'Introduct. to Entomology,' vol. ii. 1818, p. 35.

[479] See an interesting article, "The Writings of Fabre," in 'Nat. Hist. Review,' April, 1862, p. 122.

[480] 'Journal of Proc. of Entomolog. Soc.' Sept. 7th, 1863, p. 169.

[481] P. Huber, 'Recherches sur les Moeurs des Fourmis,' 1810, p. 150, 165.

[482] 'Proc. Entomolog. Soc. of Philadelphia,' 1866, p. 238-239.

[483] Quoted by Westwood, 'Modern Class. of Insects,' vol. ii. p. 214.

[484] Pyrodes pulcherrimus, in which the sexes differ conspicuously, has been described by Mr. Bates in 'Transact. Ent. Soc.' 1869, p. 50. I will specify the few other cases in which I have heard of a difference in colour between the sexes of beetles. Kirby and Spence ('Introduct. to Entomology,' vol. iii. p. 301) mention a Cantharis, Meloe, Rhagium, and the _Leptura testacea_; the male of the latter being testaceous, with a black thorax, and the female of a dull red all over. These two latter beetles belong to the Order of Longicorns. Messrs. R. Trimen and Waterhouse, junr., inform me of two Lamellicorns, viz., a Peritrichia and Trichius, the male of the latter being more obscurely coloured than the female. In _Tillus elongatus_ the male is black, and the female always, as it is believed, of a dark blue colour with a red thorax. The male, also, of _Orsodacna atra_, as I hear from Mr. Walsh, is black, the female (the so-called _O. ruficollis_) having a rufous thorax.

[485] 'Proc. Entomolog. Soc. of Philadelphia,' 1864, p. 228.

[486] Kirby and Spence, 'Introduct. Entomolog.' vol. iii. p. 300.

[487] Kirby and Spence, ibid. vol. iii. p. 329.

[488] 'Modern Classification of Insects,' vol. i. p. 172. On the same page there is an account of Siagonium. In the British Museum I noticed one male specimen of Siagonium in an intermediate condition, so that the dimorphism is not strict.

[489] 'The Malay Archipelago,' vol. ii. 1869, p. 276.

[490] '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.

[491] Quoted from Fischer, in 'Dict. Class. d'Hist. Nat.' tom. x. p. 324.

[492] 'Ann. Soc. Entomolog. France,' 1866, as quoted in 'Journal of Travel,' by A. Murray, 1868, p. 135.

[493] Westwood, 'Modern Class.' vol. i. p. 184.

[494] Wollaston, On certain musical Curculionidae, 'Annals and Mag. of Nat. Hist.' vol. vi. 1860, p. 14.

[495] 'Zeitschrift fuer wiss. Zoolog.' B. xvii. 1867, s. 127.

[496] I am greatly indebted to Mr. G. R. Crotch for having sent me numerous prepared specimens of various beetles belonging to these three families and others, as well as for valuable information of all kinds. He believes that the power of stridulation in the Clythra has not been previously observed. I am also much indebted to Mr. E. W. Janson, for information and specimens. I may add that my son, Mr. F. Darwin, finds that _Dermestes murinus_ stridulates, but he searched in vain for the apparatus. Scolytus has lately been described by Mr. Algen as a stridulator, in the 'Edinburgh Monthly Magazine,' 1869, Nov., p. 130.

[497] Schioedte, translated in 'Annals and Mag. of Nat. Hist.' vol. xx. 1867, p. 37.

[498] Westring has described (Kroyer, 'Naturhist. Tidskrift,' B. ii. 1848-49, p. 334) the stridulating organs in these two, as well as in other families. In the Carabidae I have examined _Elaphrus uliginosus_ and _Blethisa multipunctata_, sent to me by Mr. Crotch. In Blethisa the transverse ridges on the furrowed border of the abdominal segment do not come into play, as far as I could judge, in scraping the rasps on the elytra.

[499] I am indebted to Mr. Walsh, of Illinois, for having sent me extracts from Leconte's 'Introduction to Entomology,' p. 101, 143.

[500] M. P. de la Brulerie, as quoted in 'Journal of Travel,' A. Murray, vol. i. 1868, p. 135.

[501] Mr. Doubleday informs me that "the noise is produced by the insect raising itself on its legs as high as it can, and then sinking its thorax five or six times, in rapid succession, against the substance upon which it is sitting." For references on this subject see Landois, 'Zeitschrift fuer wissen. Zoolog.' B. xvii. s. 131. Olivier says (as quoted by Kirby and Spence, 'Introduct.' vol. ii. p. 395) that the female of _Pimelia striata_ produces a rather loud sound by striking her abdomen against any hard substance, "and that the male, obedient to this call, soon attends her and they pair."

[502] Apatura Iris: 'The Entomologist's Weekly Intelligencer,' 1859, p. 139. For the Bornean Butterflies see C. Collingwood, 'Rambles of a Naturalist,' 1868, p. 183.

[503] See my 'Journal of Researches,' 1845, p. 33. Mr. Doubleday has detected ('Proc. Ent. Soc.' March 3rd, 1845, p. 123) a peculiar membranous sac at the base of the front wings, which is probably connected with the production of the sound.

[504] See also Mr. Bates' paper in 'Proc. Ent. Soc. of Philadelphia,' 1865, p. 206. Also Mr. Wallace on the same subject, in regard to Diadema, in 'Transact. Entomolog. Soc. of London,' 1869, p. 278.

[505] 'The Naturalist on the Amazons,' vol. i. 1863, p. 19.

[506] See the interesting article in the 'Westminster Review,' July, 1867, p. 10. A woodcut of the Kallima is given by Mr. Wallace in Hardwicke's 'Science Gossip,' Sept. 1867, p. 196.

[507] See the interesting observations by Mr. T. W. Wood, 'The Student,' Sept. 1868, p. 81.

[508] Mr. Wallace in 'Hardwicke's Science Gossip,' Sept. 1867, p. 193.

[509] See also, on this subject, Mr. Weir's paper in 'Transact. Ent. Soc.' 1869, p. 23.

[510] 'Westminster Review,' July, 1867, p. 16.

[511] For instance, Lithosia; but Prof. Westwood ('Modern Class. of Insects,' vol. ii. p. 390) seems surprised at this case. On the relative colours of diurnal and nocturnal Lepidoptera, see ibid. p. 333 and 392; also Harris, 'Treatise on the Insects of New England,' 1842, p. 315.

[512] Such differences between the upper and lower surfaces of the wings of several species of Papilio, may be seen in the beautiful plates to Mr. Wallace's Memoir on the Papilionidae of the Malayan Region, in 'Transact. Linn. Soc.' vol. xxv. part i. 1865.

[513] 'Proc. Ent. Soc.' March 2nd, 1868.

[514] See also an account of the S. American genus Erateina (one of the Geometrae) in 'Transact. Ent. Soc.' new series, vol. v. pl. xv. and xvi.

[515] 'Proc. Ent. Soc. of London,' July 6, 1868, p. xxvii.

[516] Harris, 'Treatise,' &c., edited by Flint, 1862, p. 395.

[517] For instance, I observe in my son's cabinet that the males are darker than the females in the _Lasiocampa quercus_, _Odonestis potatoria_, _Hypogymna dispar_, _Dasychira pudibunda_, and _Cycnia mendica_. In this latter species the difference in colour between the two sexes is strongly marked; and Mr. Wallace informs me that we here have, as he believes, an instance of protective mimickry confined to one sex, as will hereafter be more fully explained. The white female of the Cycnia resembles the very common _Spilosoma menthrasti_, both sexes of which are white; and Mr. Stainton observed that this latter moth was rejected with utter disgust by a whole brood of young turkeys, which were fond of eating other moths; so that if the Cycnia was commonly mistaken by British birds for the Spilosoma, it would escape being devoured, and its white deceptive colour would thus be highly beneficial.

[518] 'Rambles of a Naturalist in the Chinese Seas,' 1868, p. 182.

[519] Wallace on the Papilionidae of the Malayan Region, in 'Transact. Linn. Soc.' vol. xxv. 1865, p. 8, 36. A striking case of a rare variety, strictly intermediate between two other well-marked female varieties, is given by Mr. Wallace. See also Mr. Bates, in 'Proc. Entomolog. Soc.' Nov. 19th, 1866, p. xl.

[520] Mr. R. MacLachlan, 'Transact. Ent. Soc.' vol. ii. part 6th, 3rd series, 1866, p. 459.

[521] H. W. Bates, 'The Naturalist on the Amazons,' vol. ii. 1863, p. 228. A. R. Wallace, in 'Transact. Linn. Soc.' vol. xxv. 1865, p. 10.

[522] On this whole subject see 'The Variation of Animals and Plants under Domestication,' vol. ii. 1868, chap. xxiii.

[523] A. R. Wallace, in 'The Journal of Travel,' vol. i. 1868, p. 88. 'Westminster Review,' July, 1857, p. 37. See also Messrs. Wallace and Bates in 'Proc. Ent. Soc.' Nov. 19th, 1866, p. xxxix.

[524] 'The Variation of Animals and Plants under Domestication,' vol. ii. chap. xii. p. 17.

[525] 'Transact. Linn. Soc.' vol. xxiii. 1862, p. 495.

[526] 'Proc. Ent. Soc.' Dec. 3rd, 1866, p. xlv.

[527] 'Transact. Linn. Soc.' vol. xxv. 1865, p. 1; also 'Transact. Ent. Soc.' vol. iv. (3rd series), 1867, p. 301.

[528] See an ingenious article entitled, "Difficulties of the Theory of Natural Selection," in the 'Month,' 1869. The writer strangely supposes that I attribute the variations in colour of the Lepidoptera, by which certain species belonging to distinct families have come to resemble others, to reversion to a common progenitor; but there is no more reason to attribute these variations to reversion than in the case of any ordinary variation.

[529] Wallace, "Notes on Eastern Butterflies," 'Transact. Ent. Soc.' 1869, p. 287.

[530] Wallace, in 'Westminster Review,' July, 1867, p. 37; and in 'Journal of Travel and Nat. Hist.' vol. i. 1868, p. 88.

[531] See remarks by Messrs. Bates and Wallace, in 'Proc. Ent. Soc.' Nov. 19, 1866, p. xxxix.

[532] See Mr. Wallace in 'Westminster Review,' July, 1867, p. 11 and 37. The male of no butterfly, as Mr. Wallace informs me, is known to differ in colour, as a protection, from the female; and he asks me how I can explain this fact on the principle that one sex alone has varied and has transmitted its variations exclusively to the same sex, without the aid of selection to check the variations being inherited by the other sex. No doubt if it could be shewn that the females of very many species had been rendered beautiful through protective mimickry, but that this has never occurred with the males, it would be a serious difficulty. But the number of cases as yet known hardly suffices for a fair judgment. We can see that the males, from having the power of flying more swiftly, and thus escaping danger, would not be so likely as the females to have had their colours modified for the sake of protection; but this would not in the least have interfered with their receiving protective colours through inheritance from the females. In the second place, it is probable that sexual selection would actually tend to prevent a beautiful male from becoming obscure, for the less brilliant individuals would be less attractive to the females. Supposing that the beauty of the male of any species had been mainly acquired through sexual selection, yet if this beauty likewise served as a protection, the acquisition would have been aided by natural selection. But it would be quite beyond our power to distinguish between the two processes of sexual and ordinary selection. Hence it is not likely that we should be able to adduce cases of the males having been rendered brilliant exclusively through protective mimickry, though this is comparatively easy with the females, which have rarely or never been rendered beautiful, as far as we can judge, for the sake of sexual attraction, although they have often received beauty through inheritance from their male parents.

[533] 'Proc. Entomolog. Soc.' Dec. 3rd, 1866, p. xlv., and March 4th, 1867, p. lxxx.

[534] See Mr. J. Jenner Weir's paper on insects and insectivorous birds, in 'Transact. Ent. Soc.' 1869, p. 21; also Mr. Butler's paper, ibid. p. 27.

END OF VOL. I.

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TRANSCRIBER'S NOTE

Footnote 292 spells the city Charleston but the text has it as Charlestown. This is as in the original book.

Inconsistent spelling of St. Petersburg/Petersburgh as in the original text.