Mimicry by Female Insects only.

But there is yet another series of phenomena connected with this subject, which considerably strengthens the view here adopted, while it seems quite incompatible with either of the other hypotheses; namely, the relation of protective colouring and mimicry to the sexual differences of animals. It will be clear to every one that if two animals, which as regards "external conditions" and "hereditary descent," are exactly alike, yet differ remarkably in colouration, one resembling a protected species and the other not, the resemblance that exists in one only can hardly be imputed to the influence of external conditions or as the effect of heredity. And if, further, it can be proved that the one requires protection more than the other, and that in several cases it is that one which mimics the protected species, while the one that least requires protection never does so, it will afford very strong corroborative evidence that there is a real connexion between the necessity for protection and the phenomenon of mimicry. Now the sexes of insects offer us a test of the nature here indicated, and appear to furnish one of the most conclusive arguments in favour of the theory that the phenomena termed "mimicry" are produced by natural selection.

The comparative importance of the sexes varies much in different classes of animals. In the higher vertebrates, where the number of young produced at a birth is small and the same individuals breed many years in succession, the preservation of both sexes is almost equally important. In all the numerous cases in which the male protects the female and her offspring, or helps to supply them with food, his importance in the economy of nature is proportionately increased, though it is never perhaps quite equal to that of the female. In insects the case is very different; they pair but once in their lives, and the prolonged existence of the male is in most cases quite unnecessary for the continuance of the race. The female, however, must continue to exist long enough to deposit her eggs in a place adapted for the development and growth of the progeny. Hence there is a wide difference in the need for protection in the two sexes; and we should, therefore, expect to find that in some cases the special protection given to the female was in the male less in amount or altogether wanting. The facts entirely confirm this expectation. In the spectre insects (Phasmidae) it is often the females alone that so strikingly resemble leaves, while the males show only a rude approximation. The male Diadema misippus is a very handsome and conspicuous butterfly, without a sign of protective or imitative colouring, while the female is entirely unlike her partner, and is one of the most wonderful cases of mimicry on record, resembling most accurately the common Danais chrysippus, in whose company it is often found. So in several species of South American Pieris, the males are white and black, of a similar type of colouring to our own "cabbage" butterflies, while the females are rich yellow and buff, spotted and marked so as exactly to resemble species of Heliconidae with which they associate in the forest. In the Malay archipelago is found a Diadema which had always been considered a male insect on account of its glossy metallic-blue tints, while its companion of sober brown was looked upon as the female. I discovered, however, that the reverse is the case, and that the rich and glossy colours of the female are imitative and protective, since they cause her exactly to resemble the common Euploea midamus of the same regions, a species which has been already mentioned in this essay as mimicked by another butterfly, Papilio paradoxa. I have since named this interesting species Diadema anomala (see the Transactions of the Entomological Society, 1869, p. 285). In this case, and in that of Diadema misippus, there is no difference in the habits of the two sexes, which fly in similar localities; so that the influence of "external conditions" cannot be invoked here as it has been in the case of the South American Pieris pyrrha and allies, where the white males frequent open sunny places, while the Heliconia-like females haunt the shades of the forest.

We may impute to the same general cause (the greater need of protection for the female, owing to her weaker flight, greater exposure to attack, and supreme importance)—the fact of the colours of female insects being so very generally duller and less conspicuous than those of the other sex. And that it is chiefly due to this cause rather than to what Mr. Darwin terms "sexual selection" appears to be shown by the otherwise inexplicable fact, that in the groups which have a protection of any kind independent of concealment, sexual differences of colour are either quite wanting or slightly developed. The Heliconidae and Danaidae, protected by a disagreeable flavour, have the females as bright and conspicuous as the males, and very rarely differing at all from them. The stinging Hymenoptera have the two sexes equally well coloured. The Carabidae, the Coccinellidae, Chrysomelidae, and the Telephori have both sexes equally conspicuous, and seldom differing in colours. The brilliant Curculios, which are protected by their hardness, are brilliant in both sexes. Lastly, the glittering Cetoniadae and Buprestidae, which seem to be protected by their hard and polished coats, their rapid motions, and peculiar habits, present few sexual differences of colour, while sexual selection has often manifested itself by structural differences, such as horns, spines, or other processes.

Cause of the dull Colours of Female Birds.

The same law manifests itself in Birds. The female while sitting on her eggs requires protection by concealment to a much greater extent than the male; and we accordingly find that in a large majority of the cases in which the male birds are distinguished by unusual brilliancy of plumage, the females are much more obscure, and often remarkably plain-coloured. The exceptions are such as eminently to prove the rule, for in most cases we can see a very good reason for them. In particular, there are a few instances among wading and gallinaceous birds in which the female has decidedly more brilliant colours than the male; but it is a most curious and interesting fact that in most if not all these cases the males sit upon the eggs; so that this exception to the usual rule almost demonstrates that it is because the process of incubation is at once very important and very dangerous, that the protection of obscure colouring is developed. The most striking example is that of the gray phalarope (Phalaropus fulicarius). When in winter plumage, the sexes of this bird are alike in colouration, but in summer the female is much the most conspicuous, having a black head, dark wings, and reddish-brown back, while the male is nearly uniform brown, with dusky spots. Mr. Gould in his "Birds of Great Britain" figures the two sexes in both winter and summer plumage, and remarks on the strange peculiarity of the usual colours of the two sexes being reversed, and also on the still more curious fact that the "male alone sits on the eggs," which are deposited on the bare ground. In another British bird, the dotterell, the female is also larger and more brightly-coloured than the male; and it seems to be proved that the males assist in incubation even if they do not perform it entirely, for Mr. Gould tells us, "that they have been shot with the breast bare of feathers, caused by sitting on the eggs." The small quail-like birds forming the genus Turnix have also generally large and bright-coloured females, and we are told by Mr. Jerdon in his "Birds of India" that "the natives report that during the breeding season the females desert their eggs and associate in flocks while the males are employed in hatching the eggs." It is also an ascertained fact, that the females are more bold and pugnacious than the males. A further confirmation of this view is to be found in the fact (not hitherto noticed) that in a large majority of the cases in which bright colours exist in both sexes incubation takes place in a dark hole or in a dome-shaped nest. Female kingfishers are often equally brilliant with the male, and they build in holes in banks. Bee-eaters, trogons, motmots, and toucans, all build in holes, and in none is there any difference in the sexes, although they are, without exception, showy birds. Parrots build in holes in trees, and in the majority of cases they present no marked sexual difference tending to concealment of the female. Woodpeckers are in the same category, since though the sexes often differ in colour, the female is not generally less conspicuous than the male. Wagtails and titmice build concealed nests, and the females are nearly as gay as their mates. The female of the pretty Australian bird Pardalotus punctatus, is very conspicuously spotted on the upper surface, and it builds in a hole in the ground. The gay-coloured hang-nests (Icterinae) and the equally brilliant tanagers may be well contrasted; for the former, concealed in their covered nests, present little or no sexual difference of colour—while the open-nested tanagers have the females dull-coloured and sometimes with almost protective tints. No doubt there are many individual exceptions to the rule here indicated, because many and various causes have combined to determine both the colouration and the habits of birds. These have no doubt acted and re-acted on each other; and when conditions have changed one of these characters may often have become modified, while the other, though useless, may continue by hereditary descent an apparent exception to what otherwise seems a very general rule. The facts presented by the sexual differences of colour in birds and their mode of nesting, are on the whole in perfect harmony with that law of protective adaptation of colour and form, which appears to have checked to some extent the powerful action of sexual selection, and to have materially influenced the colouring of female birds, as it has undoubtedly done that of female insects.

Use of the gaudy Colours of many Caterpillars.

Since this essay was first published a very curious difficulty has been cleared up by the application of the general principle of protective colouring. Great numbers of caterpillars are so brilliantly marked and coloured as to be very conspicuous even at a considerable distance, and it has been noticed that such caterpillars seldom hide themselves. Other species, however, are green or brown, closely resembling the colours of the substances on which they feed, while others again imitate sticks, and stretch themselves out motionless from a twig so as to look like one of its branches. Now, as caterpillars form so large a part of the food of birds, it was not easy to understand why any of them should have such bright colours and markings as to make them specially visible. Mr. Darwin had put the case to me as a difficulty from another point of view, for he had arrived at the conclusion that brilliant colouration in the animal kingdom is mainly due to sexual selection, and this could not have acted in the case of sexless larvae. Applying here the analogy of other insects, I reasoned, that since some caterpillars were evidently protected by their imitative colouring, and others by their spiny or hairy bodies, the bright colours of the rest must also be in some way useful to them. I further thought that as some butterflies and moths were greedily eaten by birds while others were distasteful to them, and these latter were mostly of conspicuous colours, so probably these brilliantly coloured caterpillars were distasteful, and therefore never eaten by birds. Distastefulness alone would however be of little service to caterpillars, because their soft and juicy bodies are so delicate, that if seized and afterwards rejected by a bird they would almost certainly be killed. Some constant and easily perceived signal was therefore necessary to serve as a warning to birds never to touch these uneatable kinds, and a very gaudy and conspicuous colouring with the habit of fully exposing themselves to view becomes such a signal, being in strong contrast with the green or brown tints and retiring habits of the eatable kinds. The subject was brought by me before the Entomological Society (see Proceedings, March 4th, 1867), in order that those members having opportunities for making observations might do so in the following summer; and I also wrote a letter to the Field newspaper, begging that some of its readers would co-operate in making observations on what insects were rejected by birds, at the same time fully explaining the great interest and scientific importance of the problem. It is a curious example of how few of the country readers of that paper are at all interested in questions of simple natural history, that I only obtained one answer from a gentleman in Cumberland, who gave me some interesting observations on the general dislike and abhorrence of all birds to the "Gooseberry Caterpillar," probably that of the Magpie-moth (Abraxas grossulariata). Neither young pheasants, partridges, nor wild-ducks could be induced to eat it, sparrows and finches never touched it, and all birds to whom he offered it rejected it with evident dread and abhorrence. It will be seen that these observations are confirmed by those of two members of the Entomological Society to whom we are indebted for more detailed information.

In March, 1869, Mr. J. Jenner Weir communicated a valuable series of observations made during many years, but more especially in the two preceding summers, in his aviary, containing the following birds of more or less insectivorous habits:—Robin, Yellow-Hammer, Reed-bunting, Bullfinch, Chaffinch, Crossbill, Thrush, Tree-Pipit, Siskin, and Redpoll. He found that hairy caterpillars were uniformly rejected; five distinct species were quite unnoticed by all his birds, and were allowed to crawl about the aviary for days with impunity. The spiny caterpillars of the Tortoiseshell and Peacock butterflies were equally rejected; but in both these cases Mr. Weir thinks it is the taste, not the hairs or spines, that are disagreeable, because some very young caterpillars of a hairy species were rejected although no hairs were developed, and the smooth pupae of the above-named butterflies were refused as persistently as the spined larvae. In these cases, then, both hairs and spines would seem to be mere signs of uneatableness.

His next experiments were with those smooth gaily-coloured caterpillars which never conceal themselves, but on the contrary appear to court observation. Such are those of the Magpie moth (Abraxas grossulariata), whose caterpillar is conspicuously white and black spotted—the Diloba coeruleocephala, whose larvae is pale yellow with a broad blue or green lateral band—the Cucullia verbasci, whose larvae is greenish white with yellow bands and black spots, and Anthrocera filipendulae (the six spot Burnet moth), whose caterpillar is yellow with black spots. These were given to the birds at various times, sometimes mixed with other kinds of larvae which were greedily eaten, but they were in every case rejected apparently unnoticed, and were left to crawl about till they died.

The next set of observations were on the dull-coloured and protected larvae, and the results of numerous experiments are thus summarised by Mr. Weir. "All caterpillars whose habits are nocturnal, which are dull coloured, with fleshy bodies and smooth skins, are eaten with the greatest avidity. Every species of green caterpillar is also much relished. All Geometrae, whose larvae resemble twigs as they stand out from the plant on their anal prolegs, are invariably eaten."

At the same meeting Mr. A. G. Butler, of the British Museum, communicated the results of his observations with lizards, frogs, and spiders, which strikingly corroborate those of Mr. Weir. Three green lizards (Lacerta viridis) which he kept for several years, were very voracious, eating all kinds of food, from a lemon cheesecake to a spider, and devouring flies, caterpillars, and humble bees; yet there were some caterpillars and moths which they would seize only to drop immediately. Among these the principal were the caterpillar of the Magpie moth (Abraxas grossulariata) and the perfect six spot Burnet moth (Anthrocera filipendulae). These would be first seized but invariably dropped in disgust, and afterwards left unmolested. Subsequently frogs were kept and fed with caterpillars from the garden, but two of these—that of the before-mentioned Magpie moth, and that of the V. moth (Halia wavaria), which is green with conspicuous white or yellow stripes and black spots—were constantly rejected. When these species were first offered, the frogs sprang at them eagerly and licked them into their mouths; no sooner, however, had they done so than they seemed to be aware of the mistake that they had made, and sat with gaping mouths, rolling their tongues about until they had got quit of the nauseous morsels.

With spiders the same thing occurred. These two caterpillars were repeatedly put into the webs both of the geometrical and hunting spiders (Epeira diadema and Lycosa sp.), but in the former case they were cut out and allowed to drop; in the latter, after disappearing in the jaws of their captor down his dark silken funnel, they invariably reappeared, either from below or else taking long strides up the funnel again. Mr. Butler has observed lizards fight with and finally devour humble bees, and a frog sitting on a bed of stone-crop leap up and catch the bees which flew over his head, and swallow them, in utter disregard of their stings. It is evident, therefore, that the possession of a disagreeable taste or odour is a more effectual protection to certain conspicuous caterpillars and moths, than would be even the possession of a sting.

The observations of these two gentlemen supply a very remarkable confirmation of the hypothetical solution of the difficulty which I had given two years before. And as it is generally acknowledged that the best test of the truth and completeness of a theory is the power which it gives us of prevision, we may I think fairly claim this as a case in which the power of prevision has been successfully exerted, and therefore as furnishing a very powerful argument in favour of the truth of the theory of Natural Selection.

Summary.

I have now completed a brief, and necessarily very imperfect, survey of the various ways in which the external form and colouring of animals is adapted to be useful to them, either by concealing them from their enemies or from the creatures they prey upon. It has, I hope, been shown that the subject is one of much interest, both as regard a true comprehension of the place each animal fills in the economy of nature, and the means by which it is enabled to maintain that place; and also as teaching us how important a part is played by the minutest details in the structure of animals, and how complicated and delicate is the equilibrium of the organic world.

My exposition of the subject having been necessarily somewhat lengthy and full of details, it will be as well to recapitulate its main points.

There is a general harmony in nature between the colours of an animal and those of its habitation. Arctic animals are white, desert animals are sand-coloured; dwellers among leaves and grass are green; nocturnal animals are dusky. These colours are not universal, but are very general, and are seldom reversed. Going on a little further, we find birds, reptiles, and insects, so tinted and mottled as exactly to match the rock, or bark, or leaf, or flower, they are accustomed to rest upon,—and thereby effectually concealed. Another step in advance, and we have insects which are formed as well as coloured so as exactly to resemble particular leaves, or sticks, or mossy twigs, or flowers; and in these cases very peculiar habits and instincts come into play to aid in the deception and render the concealment more complete. We now enter upon a new phase of the phenomena, and come to creatures whose colours neither conceal them nor make them like vegetable or mineral substances; on the contrary, they are conspicuous enough, but they completely resemble some other creature of a quite different group, while they differ much in outward appearance from those with which all essential parts of their organization show them to be really closely allied. They appear like actors or masqueraders dressed up and painted for amusement, or like swindlers endeavouring to pass themselves off for well-known and respectable members of society. What is the meaning of this strange travestie? Does Nature descend to imposture or masquerade? We answer, she does not. Her principles are too severe. There is a use in every detail of her handiwork. The resemblance of one animal to another is of exactly the same essential nature as the resemblance to a leaf, or to bark, or to desert sand, and answers exactly the same purpose. In the one case the enemy will not attack the leaf or the bark, and so the disguise is a safeguard; in the other case it is found that for various reasons the creature resembled is passed over, and not attacked by the usual enemies of its order, and thus the creature that resembles it has an equally effectual safeguard. We are plainly shown that the disguise is of the same nature in the two cases, by the occurrence in the same group of one species resembling a vegetable substance, while another resembles a living animal of another group; and we know that the creatures resembled, possess an immunity from attack, by their being always very abundant, by their being conspicuous and not concealing themselves, and by their having generally no visible means of escape from their enemies; while, at the same time, the particular quality that makes them disliked is often very clear, such as a nasty taste or an indigestible hardness. Further examination reveals the fact that, in several cases of both kinds of disguise, it is the female only that is thus disguised; and as it can be shown that the female needs protection much more than the male, and that her preservation for a much longer period is absolutely necessary for the continuance of the race, we have an additional indication that the resemblance is in all cases subservient to a great purpose—the preservation of the species.

In endeavouring to explain these phenomena as having been brought about by variation and natural selection, we start with the fact that white varieties frequently occur, and when protected from enemies show no incapacity for continued existence and increase. We know, further, that varieties of many other tints occasionally occur; and as "the survival of the fittest" must inevitably weed out those whose colours are prejudicial and preserve those whose colours are a safeguard, we require no other mode of accounting for the protective tints of arctic and desert animals. But this being granted, there is such a perfectly continuous and graduated series of examples of every kind of protective imitation, up to the most wonderful cases of what is termed "mimicry," that we can find no place at which to draw the line, and say,—so far variation and natural selection will account for the phenomena, but for all the rest we require a more potent cause. The counter theories that have been proposed, that of the "special creation" of each imitative form, that of the action of "similar conditions of existence" for some of the cases, and of the laws of "hereditary descent and the reversion to ancestral forms" for others,—have all been shown to be beset with difficulties, and the two latter to be directly contradicted by some of the most constant and most remarkable of the facts to be accounted for.

General deductions as to Colour in Nature.

The important part that "protective resemblance" has played in determining the colours and markings of many groups of animals, will enable us to understand the meaning of one of the most striking facts in nature, the uniformity in the colours of the vegetable as compared with the wonderful diversity of the animal world. There appears no good reason why trees and shrubs should not have been adorned with as many varied hues and as strikingly designed patterns as birds and butterflies, since the gay colours of flowers show that there is no incapacity in vegetable tissues to exhibit them. But even flowers themselves present us with none of those wonderful designs, those complicated arrangements of stripes and dots and patches of colour, that harmonious blending of hues in lines and bands and shaded spots, which are so general a feature in insects. It is the opinion of Mr. Darwin that we owe much of the beauty of flowers to the necessity of attracting insects to aid in their fertilisation, and that much of the development of colour in the animal world is due to "sexual selection," colour being universally attractive, and thus leading to its propagation and increase; but while fully admitting this, it will be evident from the facts and arguments here brought forward, that very much of the variety both of colour and markings among animals is due to the supreme importance of concealment, and thus the various tints of minerals and vegetables have been directly reproduced in the animal kingdom, and again and again modified as more special protection became necessary. We shall thus have two causes for the development of colour in the animal world, and shall be better enabled to understand how, by their combined and separate action, the immense variety we now behold has been produced. Both causes, however, will come under the general law of "Utility," the advocacy of which, in its broadest sense, we owe almost entirely to Mr. Darwin. A more accurate knowledge of the varied phenomena connected with this subject may not improbably give us some information both as to the senses and the mental faculties of the lower animals. For it is evident that if colours which please us also attract them, and if the various disguises which have been here enumerated are equally deceptive to them as to ourselves, then both their powers of vision and their faculties of perception and emotion, must be essentially of the same nature as our own—a fact of high philosophical importance in the study of our own nature and our true relations to the lower animals.

Conclusion.

Although such a variety of interesting facts have been already accumulated, the subject we have been discussing is one of which comparatively little is really known. The natural history of the tropics has never yet been studied on the spot with a full appreciation of "what to observe" in this matter. The varied ways in which the colouring and form of animals serve for their protection, their strange disguises as vegetable or mineral substances, their wonderful mimicry of other beings, offer an almost unworked and inexhaustible field of discovery for the zoologist, and will assuredly throw much light on the laws and conditions which have resulted in the wonderful variety of colour, shade, and marking which constitutes one of the most pleasing characteristics of the animal world, but the immediate causes of which it has hitherto been most difficult to explain.

If I have succeeded in showing that in this wide and picturesque domain of nature, results which have hitherto been supposed to depend either upon those incalculable combinations of laws which we term chance or upon the direct volition of the Creator, are really due to the action of comparatively well-known and simple causes, I shall have attained my present purpose, which has been to extend the interest so generally felt in the more striking facts of natural history to a large class of curious but much neglected details; and to further, in however slight a degree, our knowledge of the subjection of the phenomena of life to the "Reign of Law."



IV.

THE MALAYAN PAPILIONIDAE OR SWALLOW-TAILED BUTTERFLIES, AS ILLUSTRATIVE OF THE THEORY OF NATURAL SELECTION.

Special Value of the Diurnal Lepidoptera for enquiries of this nature.

When the naturalist studies the habits, the structure, or the affinities of animals, it matters little to which group he especially devotes himself; all alike offer him endless materials for observation and research. But, for the purpose of investigating the phenomena of geographical distribution and of local, sexual, or general variation, the several groups differ greatly in their value and importance. Some have too limited a range, others are not sufficiently varied in specific forms, while, what is of most importance, many groups have not received that amount of attention over the whole region they inhabit, which could furnish materials sufficiently approaching to completeness to enable us to arrive at any accurate conclusions as to the phenomena they present as a whole. It is in those groups which are, and have long been, favourites with collectors, that the student of distribution and variation will find his materials the most satisfactory, from their comparative completeness.

Pre-eminent among such groups are the diurnal Lepidoptera or Butterflies, whose extreme beauty and endless diversity have led to their having been assiduously collected in all parts of the world, and to the numerous species and varieties having been figured in a series of magnificent works, from those of Cramer, the contemporary of Linnaeus, down to the inimitable productions of our own Hewitson.[G] But, besides their abundance, their universal distribution, and the great attention that has been paid to them, these insects have other qualities that especially adapt them to elucidate the branches of inquiry already alluded to. These are, the immense development and peculiar structure of the wings, which not only vary in form more than those of any other insects, but offer on both surfaces an endless variety of pattern, colouring, and texture. The scales, with which they are more or less completely covered, imitate the rich hues and delicate surfaces of satin or of velvet, glitter with metallic lustre, or glow with the changeable tints of the opal. This delicately painted surface acts as a register of the minutest differences of organization—a shade of colour, an additional streak or spot, a slight modification of outline continually recurring with the greatest regularity and fixity, while the body and all its other members exhibit no appreciable change. The wings of Butterflies, as Mr. Bates has well put it, "serve as a tablet on which Nature writes the story of the modifications of species;" they enable us to perceive changes that would otherwise be uncertain and difficult of observation, and exhibit to us on an enlarged scale the effects of the climatal and other physical conditions which influence more or less profoundly the organization of every living thing.

[G] W. C. Hewitson, Esq., of Oatlands, Walton-on-Thames, author of "Exotic Butterflies" and several other works, illustrated by exquisite coloured figures drawn by himself; and owner of the finest collection of Butterflies in the world.

A proof that this greater sensibility to modifying causes is not imaginary may, I think, be drawn from the consideration, that while the Lepidoptera as a whole are of all insects the least essentially varied in form, structure, or habits, yet in the number of their specific forms they are not much inferior to those orders which range over a much wider field of nature, and exhibit more deeply seated structural modifications. The Lepidoptera are all vegetable-feeders in their larva-state, and suckers of juices or other liquids in their perfect form. In their most widely separated groups they differ but little from a common type, and offer comparatively unimportant modifications of structure or of habits. The Coleoptera, the Diptera, or the Hymenoptera, on the other hand, present far greater and more essential variations. In either of these orders we have both vegetable and animal-feeders, aquatic, and terrestrial, and parasitic groups. Whole families are devoted to special departments in the economy of nature. Seeds, fruits, bones, carcases, excrement, bark, have each their special and dependent insect tribes from among them; whereas the Lepidoptera are, with but few exceptions, confined to the one function of devouring the foliage of living vegetation. We might therefore anticipate that their species—population would be only equal to that of sections of the other orders having a similar uniform mode of existence; and the fact that their numbers are at all comparable with those of entire orders, so much more varied in organization and habits, is, I think, a proof that they are in general highly susceptible of specific modification.

Question of the rank of the Papilionidae.

The Papilionidae are a family of diurnal Lepidoptera which have hitherto, by almost universal consent, held the first rank in the order; and though this position has recently been denied them, I cannot altogether acquiesce in the reasoning by which it has been proposed to degrade them to a lower rank. In Mr. Bates's most excellent paper on the Heliconidae, (published in the Transactions of the Linnaean Society, vol. xxiii., p. 495) he claims for that family the highest position, chiefly because of the imperfect structure of the fore legs, which is there carried to an extreme degree of abortion, and thus removes them further than any other family from the Hesperidae and Heterocera, which all have perfect legs. Now it is a question whether any amount of difference which is exhibited merely in the imperfection or abortion of certain organs, can establish in the group exhibiting it a claim to a high grade of organization, still less can this be allowed when another group along with perfection of structure in the same organs, exhibits modifications peculiar to it, together with the possession of an organ which in the remainder of the order is altogether wanting. This is, however, the position of the Papilionidae. The perfect insects possess two characters quite peculiar to them. Mr. Edward Doubleday, in his "Genera of Diurnal Lepidoptera," says, "The Papilionidae may be known by the apparently four-branched median nervule and the spur on the anterior tibiae, characters found in no other family." The four-branched median nervule is a character so constant, so peculiar, and so well marked, as to enable a person to tell, at a glance at the wings only of a butterfly, whether it does or does not belong to this family; and I am not aware that any other group of butterflies, at all comparable to this in extent and modifications of form, possesses a character in its neuration to which the same degree of certainty can be attached. The spur on the anterior tibiae is also found in some of the Hesperidae, and is therefore supposed to show a direct affinity between the two groups: but I do not imagine it can counterbalance the differences in neuration and in every other part of their organization. The most characteristic feature of the Papilionidae, however, and that on which I think insufficient stress has been laid, is undoubtedly the peculiar structure of the larvae. These all possess an extraordinary organ situated on the neck, the well-known Y-shaped tentacle, which is entirely concealed in a state of repose, but which is capable of being suddenly thrown out by the insect when alarmed. When we consider this singular apparatus, which in some species is nearly half an inch long, the arrangement of muscles for its protrusion and retraction, its perfect concealment during repose, its blood-red colour, and the suddenness with which it can be thrown out, we must, I think, be led to the conclusion that it serves as a protection to the larva, by startling and frightening away some enemy when about to seize it, and is thus one of the causes which has led to the wide extension and maintained the permanence of this now dominant group. Those who believe that such peculiar structures can only have arisen by very minute successive variations, each one advantageous to its possessor, must see, in the possession of such an organ by one group, and its complete absence in every other, a proof of a very ancient origin and of very long-continued modification. And such a positive structural addition to the organization of the family, subserving an important function, seems to me alone sufficient to warrant us in considering the Papilionidae as the most highly developed portion of the whole order, and thus in retaining it in the position which the size, strength, beauty, and general structure of the perfect insects have been generally thought to deserve.

In Mr. Trimen's paper on "Mimetic Analogies among African Butterflies," in the Transactions of the Linnaean Society, for 1868, he has argued strongly in favour of Mr. Bates' views as to the higher position of the Danaidae and the lower grade of the Papilionidae, and has adduced, among other facts, the undoubted resemblance of the pupa of Parnassius, a genus of Papilionidae, to that of some Hesperidae and moths. I admit, therefore, that he has proved the Papilionidae to have retained several characters of the nocturnal Lepidoptera which the Danaidae have lost, but I deny that they are therefore to be considered lower in the scale of organization. Other characters may be pointed out which indicate that they are farther removed from the moths even than the Danaidae. The club of the antennae is the most prominent and most constant feature by which butterflies may be distinguished from moths, and of all butterflies the Papilionidae have the most beautiful and most perfectly developed clubbed antennae. Again, butterflies and moths are broadly characterised by their diurnal and nocturnal habits respectively, and the Papilionidae, with their close allies the Pieridae, are the most pre-eminently diurnal of butterflies, most of them lovers of sunshine, and not presenting a single crepuscular species. The great group of the Nymphalidae, on the other hand (in which Mr. Bates includes the Danaidae and Heliconidae as sub-families), contains an entire sub-family (Brassolidae) and a number of genera, such as Thaumantis, Zeuxidia, Pavonia, &c., of crepuscular habits, while a large proportion of the Satyridae and many of the Danaidae are shade-loving butterflies. This question, of what is to be considered the highest type of any group of organisms, is one of such general interest to naturalists that it will be well to consider it a little further, by a comparison of the Lepidoptera with some groups of the higher animals.

Mr. Trimen's argument, that the lepidopterous type, like that of birds, being pre-eminently aerial, "therefore a diminution of the ambulatory organs, instead of being a sign of inferiority, may very possibly indicate a higher, because a more thoroughly aerial form," is certainly unsound, for it would imply that the most aerial of birds (the swift and the frigate-birds, for example) are the highest in the scale of bird-organization, and the more so on account of their feet being very ill adapted for walking. But no ornithologist has ever so classed them, and the claim to the highest rank among birds is only disputed between three groups, all very far removed from these. They are—1st. The Falcons, on account of their general perfection, their rapid flight, their piercing vision, their perfect feet armed with retractile claws, the beauty of their forms, and the ease and rapidity of their motions; 2nd. The Parrots, whose feet, though ill-fitted for walking, are perfect as prehensile organs, and which possess large brains with great intelligence, though but moderate powers of flight; and, 3rd. The Thrushes or Crows, as typical of the perching birds, on account of the well-balanced development of their whole structure, in which no organ or function has attained an undue prominence.

Turning now to the Mammalia, it might be argued that as they are pre-eminently the terrestrial type of vertebrates, to walk and run well is essential to the typical perfection of the group; but this would give the superiority to the horse, the deer, or the hunting leopard, instead of to the Quadrumana. We seem here to have quite a case in point, for one group of Quadrumana, the Lemurs, is undoubtedly nearer to the low Insectivora and Marsupials than the Carnivora or the Ungulata, as shown among other characters by the Opossums possessing a hand with perfect opposable thumb, closely resembling that of some of the Lemurs; and by the curious Galeopithecus, which is sometimes classed as a Lemur, and sometimes with the Insectivora. Again, the implacental mammals, including the Ornithodelphia and the Marsupials, are admitted to be lower than the placental series. But one of the distinguishing characters of the Marsupials is that the young are born blind and exceedingly imperfect, and it might therefore be argued that those orders in which the young are born most perfect are the highest, because farthest from the low Marsupial type. This would make the Ruminants and Ungulata higher than the Quadrumana or the Carnivora. But the Mammalia offer a still more remarkable illustration of the fallacy of this mode of reasoning, for if there is one character more than another which is essential and distinctive of the class, it is that from which it derives its name, the possession of mammary glands and the power of suckling the young. What more reasonable, apparently, than to argue that the group in which this important function is most developed, that in which the young are most dependent upon it, and for the longest period, must be the highest in the Mammalian scale of organization? Yet this group is the Marsupial, in which the young commence suckling in a foetal condition, and continue to do so till they are fully developed, and are therefore for a long time absolutely dependent on this mode of nourishment.

These examples, I think, demonstrate that we cannot settle the rank of a group by a consideration of the degree in which certain characters resemble or differ from those in what is admitted to be a lower group; and they also show that the highest group of a class may be more closely connected to one of the lowest, than some other groups which have developed laterally and diverged farther from the parent type, but which yet, owing to want of balance or too great specialization in their structure, have never reached a high grade of organization. The Quadrumana afford a very valuable illustration, because, owing to their undoubted affinity with man, we feel certain that they are really higher than any other order of Mammalia, while at the same time they are more distinctly allied to the lowest groups than many others. The case of the Papilionidae seems to me so exactly parallel to this, that, while I admit all the proofs of affinity with the undoubtedly lower groups of Hesperidae and moths, I yet maintain that, owing to the complete and even development of every part of their organization, these insects best represent the highest perfection to which the butterfly type has attained, and deserve to be placed at its head in every system of classification.

Distribution of the Papilionidae.

The Papilionidae are pretty widely distributed over the earth, but are especially abundant in the tropics, where they attain their maximum of size and beauty, and the greatest variety of form and colouring. South America, North India, and the Malay Islands are the regions where these fine insects occur in the greatest profusion, and where they actually become a not unimportant feature in the scenery. In the Malay Islands in particular, the giant Ornithopterae may be frequently seen about the borders of the cultivated and forest districts, their large size, stately flight, and gorgeous colouring rendering them even more conspicuous than the generality of birds. In the shady suburbs of the town of Malacca two large and handsome Papilios (Memnon and Nephelus) are not uncommon, flapping with irregular flight along the roadways, or, in the early morning, expanding their wings to the invigorating rays of the sun. In Amboyna and other towns of the Moluccas, the magnificent Deiphobus and Severus, and occasionally even the azure-winged Ulysses, frequent similar situations, fluttering about the orange-trees and flower-beds, or sometimes even straying into the narrow bazaars or covered markets of the city. In Java the golden-dusted Arjuna may often be seen at damp places on the roadside in the mountain districts, in company with Sarpedon, Bathycles, and Agamemnon, and less frequently the beautiful swallow-tailed Antiphates. In the more luxuriant parts of these islands one can hardly take a morning's walk in the neighbourhood of a town or village without seeing three or four species of Papilio, and often twice that number. No less than 130 species of the family are now known to inhabit the Archipelago, and of these ninety-six were collected by myself. Thirty species are found in Borneo, being the largest number in any one island, twenty-three species having been obtained by myself in the vicinity of Sarawak; Java has twenty-eight species; Celebes twenty-four, and the Peninsula of Malacca, twenty-six species. Further east the numbers decrease; Batchian producing seventeen, and New Guinea only fifteen, though this number is certainly too small, owing to our present imperfect knowledge of that great island.

Definition of the word Species.

In estimating these numbers I have had the usual difficulty to encounter, of determining what to consider species and what varieties. The Malayan region, consisting of a large number of islands of generally great antiquity, possesses, compared to its actual area, a great number of distinct forms, often indeed distinguished by very slight characters, but in most cases so constant in large series of specimens, and so easily separable from each other, that I know not on what principle we can refuse to give them the name and rank of species. One of the best and most orthodox definitions is that of Pritchard, the great ethnologist, who says, that "separate origin and distinctness of race, evinced by a constant transmission of some characteristic peculiarity of organization," constitutes a species. Now leaving out the question of "origin," which we cannot determine, and taking only the proof of separate origin, "the constant transmission of some characteristic peculiarity of organization," we have a definition which will compel us to neglect altogether the amount of difference between any two forms, and to consider only whether the differences that present themselves are permanent. The rule, therefore, I have endeavoured to adopt is, that when the difference between two forms inhabiting separate areas seems quite constant, when it can be defined in words, and when it is not confined to a single peculiarity only, I have considered such forms to be species. When, however, the individuals of each locality vary among themselves, so as to cause the distinctions between the two forms to become inconsiderable and indefinite, or where the differences, though constant, are confined to one particular only, such as size, tint, or a single point of difference in marking or in outline, I class one of the forms as a variety of the other.

I find as a general rule that the constancy of species is in an inverse ratio to their range. Those which are confined to one or two islands are generally very constant. When they extend to many islands, considerable variability appears; and when they have an extensive range over a large part of the Archipelago, the amount of unstable variation is very large. These facts are explicable on Mr. Darwin's principles. When a species exists over a wide area, it must have had, and probably still possesses, great powers of dispersion. Under the different conditions of existence in various portions of its area, different variations from the type would be selected, and, were they completely isolated, would soon become distinctly modified forms; but this process is checked by the dispersive powers of the whole species, which leads to the more or less frequent intermixture of the incipient varieties, which thus become irregular and unstable. Where, however, a species has a limited range, it indicates less active powers of dispersion, and the process of modification under changed conditions is less interfered with. The species will therefore exist under one or more permanent forms according as portions of it have been isolated at a more or less remote period.

Laws and Modes of Variation.

What is commonly called variation consists of several distinct phenomena which have been too often confounded. I shall proceed to consider these under the heads of—1st, simple variability; 2nd, polymorphism; 3rd, local forms; 4th, co-existing varieties; 5th, races or subspecies; and 6th, true species.

1. Simple variability.—Under this head I include all those cases in which the specific form is to some extent unstable. Throughout the whole range of the species, and even in the progeny of individuals, there occur continual and uncertain differences of form, analogous to that variability which is so characteristic of domestic breeds. It is impossible usefully to define any of these forms, because there are indefinite gradations to each other form. Species which possess these characteristics have always a wide range, and are more frequently the inhabitants of continents than of islands, though such cases are always exceptional, it being far more common for specific forms to be fixed within very narrow limits of variation. The only good example of this kind of variability which occurs among the Malayan Papilionidae is in Papilio Severus, a species inhabiting all the islands of the Moluccas and New Guinea, and exhibiting in each of them a greater amount of individual difference than often serves to distinguish well-marked species. Almost equally remarkable are the variations exhibited in most of the species of Ornithoptera, which I have found in some cases to extend even to the form of the wing and the arrangement of the nervures. Closely allied, however, to these variable species are others which, though differing slightly from them, are constant and confined to limited areas. After satisfying oneself, by the examination of numerous specimens captured in their native countries, that the one set of individuals are variable and the others are not, it becomes evident that by classing all alike as varieties of one species we shall be obscuring an important fact in nature; and that the only way to exhibit that fact in its true light is to treat the invariable local form as a distinct species, even though it does not offer better distinguishing characters than do the extreme forms of the variable species. Cases of this kind are the Ornithoptera Priamus, which is confined to the islands of Ceram and Amboyna, and is very constant in both sexes, while the allied species inhabiting New Guinea and the Papuan Islands is exceedingly variable; and in the island of Celebes is a species closely allied to the variable P. Severus, but which, being exceedingly constant, I have described as a distinct species under the name of Papilio Pertinax.

2. Polymorphism or dimorphism.—By this term I understand the co-existence in the same locality of two or more distinct forms, not connected by intermediate gradations, and all of which are occasionally produced from common parents. These distinct forms generally occur in the female sex only, and their offspring, instead of being hybrids, or like the two parents, appear to reproduce all the distinct forms in varying proportions. I believe it will be found that a considerable number of what have been classed as varieties are really cases of polymorphism. Albinoism and melanism are of this character, as well as most of those cases in which well-marked varieties occur in company with the parent species, but without any intermediate forms. If these distinct forms breed independently, and are never reproduced from a common parent, they must be considered as separate species, contact without intermixture being a good test of specific difference. On the other hand, intercrossing without producing an intermediate race is a test of dimorphism. I consider, therefore, that under any circumstances the term "variety" is wrongly applied to such cases.

The Malayan Papilionidae exhibit some very curious instances of polymorphism, some of which have been recorded as varieties, others as distinct species; and they all occur in the female sex. Papilio Memnon is one of the most striking, as it exhibits the mixture of simple variability, local and polymorphic forms, all hitherto classed under the common title of varieties. The polymorphism is strikingly exhibited by the females, one set of which resemble the males in form, with a variable paler colouring; the others have a large spatulate tail to the hinder wings and a distinct style of colouring, which causes them closely to resemble P. Coon, a species having the two sexes alike and inhabiting the same countries, but with which they have no direct affinity. The tailless females exhibit simple variability, scarcely two being found exactly alike even in the same locality. The males of the island of Borneo exhibit constant differences of the under surface, and may therefore be distinguished as a local form, while the continental specimens, as a whole, offer such large and constant differences from those of the islands, that I am inclined to separate them as a distinct species, to which the name P. Androgeus (Cramer) may be applied. We have here, therefore, distinct species, local forms, polymorphism, and simple variability, which seem to me to be distinct phenomena, but which have been hitherto all classed together as varieties. I may mention that the fact of these distinct forms being one species is doubly proved. The males, the tailed and tailless females, have all been bred from a single group of the larvae, by Messrs. Payen and Bocarme, in Java, and I myself captured, in Sumatra, a male P. Memnon, and a tailed female P. Achates, under circumstances which led me to class them as the same species.

Papilio Pammon offers a somewhat similar case. The female was described by Linnaeus as P. Polytes, and was considered to be a distinct species till Westermann bred the two from the same larvae (see Boisduval, "Species General des Lepidopteres," p. 272). They were therefore classed as sexes of one species by Mr. Edward Doubleday, in his "Genera of Diurnal Lepidoptera," in 1846. Later, female specimens were received from India closely resembling the male insect, and this was held to overthrow the authority of M. Westermann's observation, and to re-establish P. Polytes as a distinct species; and as such it accordingly appears in the British Museum List of Papilionidae in 1856, and in the Catalogue of the East India Museum in 1857. This discrepancy is explained by the fact of P. Pammon having two females, one closely resembling the male, while the other is totally different from it. A long familiarity with this insect (which replaced by local forms or by closely allied species, occurs in every island of the Archipelago) has convinced me of the correctness of this statement; for in every place where a male allied to P. Pammon is found, a female resembling P. Polytes also occurs, and sometimes, though less frequently than on the continent, another female closely resembling the male: while not only has no male specimen of P. Polytes yet been discovered, but the female (Polytes) has never yet been found in localities to which the male (Pammon) does not extend. In this case, as in the last, distinct species, local forms, and dimorphic specimens, have been confounded under the common appellation of varieties.

But, besides the true P. Polytes, there are several allied forms of females to be considered, namely, P. Theseus (Cramer), P. Molanides (De Haan), P. Elyros (G. R. Gray), and P. Romulus (Linnaeus). The dark female figured by Cramer as P. Theseus seems to be the common and perhaps the only form in Sumatra, whereas in Java, Borneo, and Timor, along with males quite identical with those of Sumatra, occur females of the Polytes form, although a single specimen of the true P. Theseus taken at Lombock would seem to show that the two forms do occur together. In the allied species found in the Philippine Islands (P. Alphenor, Cramer = P. Ledebouria, Eschscholtz, the female of which is P. Elyros, G. R. Gray,) forms corresponding to these extremes occur, along with a number of intermediate varieties, as shown by a fine series in the British Museum. We have here an indication of how dimorphism may be produced; for let the extreme Philippine forms be better suited to their conditions of existence than the intermediate connecting links, and the latter will gradually die out, leaving two distinct forms of the same insect, each adapted to some special conditions. As these conditions are sure to vary in different districts, it will often happen, as in Sumatra and Java, that the one form will predominate in the one island, the other in the adjacent one. In the island of Borneo there seems to be a third form; for P. Melanides (De Haan) evidently belongs to this group, and has all the chief characteristics of P. Theseus, with a modified colouration of the hind wings. I now come to an insect which, if I am correct, offers one of the most interesting cases of variation yet adduced. Papilio Romulus, a butterfly found over a large part of India and Ceylon, and not uncommon in collections, has always been considered a true and independent species, and no suspicions have been expressed regarding it. But a male of this form does not, I believe, exist. I have examined the fine series in the British Museum, in the East India Company's Museum, in the Hope Museum at Oxford, in Mr. Hewitson's and several other private collections, and can find nothing but females; and for this common butterfly no male partner can be found except the equally common P. Pammon, a species already provided with two wives, and yet to whom we shall be forced, I believe, to assign a third. On carefully examining P. Romulus, I find that in all essential characters—the form and texture of the wings, the length of the antennae, the spotting of the head and thorax, and even the peculiar tints and shades with which it is ornamented—it corresponds exactly with the other females of the Pammon group; and though, from the peculiar marking of the fore wings, it has at first sight a very different aspect, yet a closer examination shows that every one of its markings could be produced by slight and almost imperceptible modifications of the various allied forms. I fully believe, therefore, that I shall be correct in placing P. Romulus as a third Indian form of the female P. Pammon, corresponding to P. Melanides, the third form of the Malayan P. Theseus. I may mention here that the females of this group have a superficial resemblance to the Polydorus group of Papilios, as shown by P. Theseus having been considered to be the female of P. Antiphus, and by P. Romulus being arranged next to P. Hector. There is no close affinity between these two groups of Papilio, and I am disposed to believe that we have here a case of mimicry, brought about by the same causes which Mr. Bates has so well explained in his account of the Heliconidae, and which has led to the singular exuberance of polymorphic forms in this and allied groups of the genus Papilio. I shall have to devote a section of my essay to the consideration of this subject.

The third example of polymorphism I have to bring forward is Papilio Ormenus, which is closely allied to the well-known P. Erechtheus, of Australia. The most common form of the female also resembles that of P. Erechtheus; but a totally different-looking insect was found by myself in the Aru Islands, and figured by Mr. Hewitson under the name of P. Onesimus, which subsequent observation has convinced me is a second form of the female of P. Ormenus. Comparison of this with Boisduval's description of P. Amanga, a specimen of which from New Guinea is in the Paris Museum, shows the latter to be a closely similar form; and two other specimens were obtained by myself, one in the island of Goram and the other in Waigiou, all evidently local modifications of the same form. In each of these localities males and ordinary females of P. Ormenus were also found. So far there is no evidence that these light-coloured insects are not females of a distinct species, the males of which have not been discovered. But two facts have convinced me this is not the case. At Dorey, in New Guinea, where males and ordinary females closely allied to P. Ormenus occur (but which seem to me worthy of being separated as a distinct species), I found one of these light-coloured females closely followed in her flight by three males, exactly in the same manner as occurs (and, I believe, occurs only) with the sexes of the same species. After watching them a considerable time, I captured the whole of them, and became satisfied that I had discovered the true relations of this anomalous form. The next year I had corroborative proof of the correctness of this opinion by the discovery in the island of Batchian of a new species allied to P. Ormenus, all the females of which, either seen or captured by me, were of one form, and much more closely resembling the abnormal light-coloured females of P. Ormenus and P. Pandion than the ordinary specimens of that sex. Every naturalist will, I think, agree that this is strongly confirmative of the supposition that both forms of female are of one species; and when we consider, further, that in four separate islands, in each of which I resided for several months, the two forms of female were obtained and only one form of male ever seen, and that about the same time, M. Montrouzier in Woodlark Island, at the other extremity of New Guinea (where he resided several years, and must have obtained all the large Lepidoptera of the island), obtained females closely resembling mine, which, in despair at finding no appropriate partners for them, he mates with a widely different species—it becomes, I think, sufficiently evident this is another case of polymorphism of the same nature as those already pointed out in P. Pammon and P. Memnon. This species, however, is not only dimorphic, but trimorphic; for, in the island of Waigiou, I obtained a third female quite distinct from either of the others, and in some degree intermediate between the ordinary female and the male. The specimen is particularly interesting to those who believe, with Mr. Darwin, that extreme difference of the sexes has been gradually produced by what he terms sexual selection, since it may be supposed to exhibit one of the intermediate steps in that process, which has been accidentally preserved in company with its more favoured rivals, though its extreme rarity (only one specimen having been seen to many hundreds of the other form) would indicate that it may soon become extinct.

The only other case of polymorphism in the genus Papilio, at all equal in interest to those I have now brought forward, occurs in America; and we have, fortunately, accurate information about it. Papilio Turnus is common over almost the whole of temperate North America; and the female resembles the male very closely. A totally different-looking insect both in form and colour, Papilio Glaucus, inhabits the same region; and though, down to the time when Boisduval published his "Species General," no connexion was supposed to exist between the two species, it is now well ascertained that P. Glaucus is a second female form of P. Turnus. In the "Proceedings of the Entomological Society of Philadelphia," Jan., 1863, Mr. Walsh gives a very interesting account of the distribution of this species. He tells us that in the New England States and in New York all the females are yellow, while in Illinois and further south all are black; in the intermediate region both black and yellow females occur in varying proportions. Lat. 37 deg. is approximately the southern limit of the yellow form, and 42 deg. the northern limit of the black form; and, to render the proof complete, both black and yellow insects have been bred from a single batch of eggs. He further states that, out of thousands of specimens, he has never seen or heard of intermediate varieties between these forms. In this interesting example we see the effects of latitude in determining the proportions in which the individuals of each form should exist. The conditions are here favourable to the one form, there to the other; but we are by no means to suppose that these conditions consist in climate alone. It is highly probable that the existence of enemies, and of competing forms of life, may be the main determining influences; and it is much to be wished that such a competent observer as Mr. Walsh would endeavour to ascertain what are the adverse causes which are most efficient in keeping down the numbers of each of these contrasted forms.

Dimorphism of this kind in the animal kingdom does not seem to have any direct relations to the reproductive powers, as Mr. Darwin has shown to be the case in plants, nor does it appear to be very general. One other case only is known to me in another family of my eastern Lepidoptera, the Pieridae; and but few occur in the Lepidoptera of other countries. The spring and autumn broods of some European species differ very remarkably; and this must be considered as a phenomenon of an analogous though not of an identical nature, while the Araschnia prorsa, of Central Europe, is a striking example of this alternate or seasonal dimorphism. Among our nocturnal Lepidoptera, I am informed, many analogous cases occur; and as the whole history of many of these has been investigated by breeding successive generations from the egg, it is to be hoped that some of our British Lepidopterists will give us a connected account of all the abnormal phenomena which they present. Among the Coleoptera Mr. Pascoe has pointed out the existence of two forms of the male sex in seven species of the two genera Xenocerus and Mecocerus belonging to the family Anthribidae, (Proc. Ent. Soc. Lond., 1862); and no less than six European Water-beetles, of the genus Dytiscus, have females of two forms, the most common having the elytra deeply sulcate, the rarer smooth as in the males. The three, and sometimes four or more, forms under which many Hymenopterous insects (especially Ants) occur, must be considered as a related phenomenon, though here each form is specialized to a distinct function in the economy of the species. Among the higher animals, albinoism and melanism may, as I have already stated, be considered as analogous facts; and I met with one case of a bird, a species of Lory (Eos fuscata), clearly existing under two differently coloured forms, since I obtained both sexes of each from a single flock, while no intermediate specimens have yet been found.

The fact of the two sexes of one species differing very considerably is so common, that it attracted but little attention till Mr. Darwin showed how it could in many cases be explained by the principle of sexual selection. For instance, in most polygamous animals the males fight for the possession of the females, and the victors, always becoming the progenitors of the succeeding generation, impress upon their male offspring their own superior size, strength, or unusually developed offensive weapons. It is thus that we can account for the spurs and the superior strength and size of the males in Gallinaceous birds, and also for the large canine tusks in the males of fruit-eating Apes. So the superior beauty of plumage and special adornments of the males of so many birds can be explained by supposing (what there are many facts to prove) that the females prefer the most beautiful and perfect-plumaged males, and that thus, slight accidental variations of form and colour have been accumulated, till they have produced the wonderful train of the Peacock and the gorgeous plumage of the Bird of Paradise. Both these causes have no doubt acted partially in insects, so many species possessing horns and powerful jaws in the male sex only, and still more frequently the males alone rejoicing in rich colours or sparkling lustre. But there is here another cause which has led to sexual differences, viz., a special adaptation of the sexes to diverse habits or modes of life. This is well seen in female Butterflies (which are generally weaker and of slower flight), often having colours better adapted to concealment; and in certain South American species (Papilio torquatus) the females, which inhabit the forests, resemble the AEneas group of Papilios which abound in similar localities, while the males, which frequent the sunny open river-banks, have a totally different colouration. In these cases, therefore, natural selection seems to have acted independently of sexual selection; and all such cases may be considered as examples of the simplest dimorphism, since the offspring never offer intermediate varieties between the parent forms.

The phenomena of dimorphism and polymorphism may be well illustrated by supposing that a blue-eyed, flaxen-haired Saxon man had two wives, one a black-haired, red-skinned Indian squaw, the other a woolly-headed, sooty-skinned negress—and that instead of the children being mulattoes of brown or dusky tints, mingling the separate characteristics of their parents in varying degrees, all the boys should be pure Saxon boys like their father, while the girls should altogether resemble their mothers. This would be thought a sufficiently wonderful fact; yet the phenomena here brought forward as existing in the insect-world are still more extraordinary; for each mother is capable not only of producing male offspring like the father, and female like herself, but also of producing other females exactly like her fellow-wife, and altogether differing from herself. If an island could be stocked with a colony of human beings having similar physiological idiosyncrasies with Papilio Pammon or Papilio Ormenus, we should see white men living with yellow, red, and black women, and their offspring always reproducing the same types; so that at the end of many generations the men would remain pure white, and the women of the same well-marked races as at the commencement.

The distinctive character therefore of dimorphism is this, that the union of these distinct forms does not produce intermediate varieties, but reproduces the distinct forms unchanged. In simple varieties, on the other hand, as well as when distinct local forms or distinct species are crossed, the offspring never resembles either parent exactly, but is more or less intermediate between them. Dimorphism is thus seen to be a specialized result of variation, by which new physiological phenomena have been developed; the two should therefore, whenever possible, be kept separate.

3. Local form, or variety.—This is the first step in the transition from variety to species. It occurs in species of wide range, when groups of individuals have become partially isolated in several points of its area of distribution, in each of which a characteristic form has become more or less completely segregated. Such forms are very common in all parts of the world, and have often been classed by one author as varieties, by another as species. I restrict the term to those cases where the difference of the forms is very slight, or where the segregation is more or less imperfect. The best example in the present group is Papilio Agamemnon, a species which ranges over the greater part of tropical Asia, the whole of the Malay archipelago, and a portion of the Australian and Pacific regions. The modifications are principally of size and form, and, though slight, are tolerably constant in each locality. The steps, however, are so numerous and gradual that it would be impossible to define many of them, though the extreme forms are sufficiently distinct. Papilio Sarpedon presents somewhat similar but less numerous variations.

4. Co-existing Variety.—This is a somewhat doubtful case. It is when a slight but permanent and hereditary modification of form exists in company with the parent or typical form, without presenting those intermediate gradations which would constitute it a case of simple variability. It is evidently only by direct evidence of the two forms breeding separately that this can be distinguished from dimorphism. The difficulty occurs in Papilio Jason, and P. Evemon, which inhabit the same localities, and are almost exactly alike in form, size, and colouration, except that the latter always wants a very conspicuous red spot on the under surface, which is found not only in P. Jason, but in all the allied species. It is only by breeding the two insects that it can be determined whether this is a case of a co-existing variety or of dimorphism. In the former case, however, the difference being constant and so very conspicuous and easily defined, I see not how we could escape considering it as a distinct species. A true case of co-existing forms would, I consider, be produced, if a slight variety had become fixed as a local form, and afterwards been brought into contact with the parent species, with little or no intermixture of the two; and such instances do very probably occur.

5. Race or subspecies.—These are local forms completely fixed and isolated; and there is no possible test but individual opinion to determine which of them shall be considered as species and which varieties. If stability of form and "the constant transmission of some characteristic peculiarity of organization" is the test of a species (and I can find no other test that is more certain than individual opinion) then every one of these fixed races, confined as they almost always are to distinct and limited areas, must be regarded as a species; and as such I have in most cases treated them. The various modifications of Papilio Ulysses, P. Peranthus, P. Codrus, P. Eurypilus, P. Helenus, &c., are excellent examples; for while some present great and well-marked, others offer slight and inconspicuous differences, yet in all cases these differences seem equally fixed and permanent. If, therefore, we call some of these forms species, and others varieties, we introduce a purely arbitrary distinction, and shall never be able to decide where to draw the line. The races of Papilio Ulysses, for example, vary in amount of modification from the scarcely differing New Guinea form to those of Woodlark Island and New Caledonia, but all seem equally constant; and as most of these had already been named and described as species, I have added the New Guinea form under the name of P. Autolycus. We thus get a little group of Ulyssine Papilios, the whole comprised within a very limited area, each one confined to a separate portion of that area, and, though differing in various amounts, each apparently constant. Few naturalists will doubt that all these may and probably have been derived from a common stock, and therefore it seems desirable that there should be a unity in our method of treating them; either call them all varieties or all species. Varieties, however, continually get overlooked; in lists of species they are often altogether unrecorded; and thus we are in danger of neglecting the interesting phenomena of variation and distribution which they present. I think it advisable, therefore, to name all such forms; and those who will not accept them as species may consider them as subspecies or races.

6. Species.—Species are merely those strongly marked races or local forms which when in contact do not intermix, and when inhabiting distinct areas are generally believed to have had a separate origin, and to be incapable of producing a fertile hybrid offspring. But as the test of hybridity cannot be applied in one case in ten thousand, and even if it could be applied would prove nothing, since it is founded on an assumption of the very question to be decided—and as the test of separate origin is in every case inapplicable—and as, further, the test of non-intermixture is useless, except in those rare cases where the most closely allied species are found inhabiting the same area, it will be evident that we have no means whatever of distinguishing so-called "true species" from the several modes of variation here pointed out, and into which they so often pass by an insensible gradation. It is quite true that, in the great majority of cases, what we term "species" are so well marked and definite that there is no difference of opinion about them; but as the test of a true theory is, that it accounts for, or at the very least is not inconsistent with, the whole of the phenomena and apparent anomalies of the problem to be solved, it is reasonable to ask that those who deny the origin of species by variation and selection should grapple with the facts in detail, and show how the doctrine of the distinct origin and permanence of species will explain and harmonize them. It has been recently asserted by Dr. J. E. Gray (in the Proceedings of the Zoological Society for 1863, page 134), that the difficulty of limiting species is in proportion to our ignorance, and that just as groups or countries are more accurately known and studied in greater detail the limits of species become settled. This statement has, like many other general assertions, its portion of both truth and error. There is no doubt that many uncertain species, founded on few or isolated specimens, have had their true nature determined by the study of a good series of examples: they have been thereby established as species or as varieties; and the number of times this has occurred is doubtless very great. But there are other, and equally trustworthy cases, in which, not single species, but whole groups have, by the study of a vast accumulation of materials, been proved to have no definite specific limits. A few of these must be adduced. In Dr. Carpenter's "Introduction to the Study of the Foraminifera," he states that "there is not a single specimen of plant or animal of which the range of variation has been studied by the collocation and comparison of so large a number of specimens as have passed under the review of Messrs. Williamson, Parker, Rupert Jones, and myself, in our studies of the types of this group;" and the result of this extended comparison of specimens is stated to be, "The range of variation is so great among the Foraminifera as to include not merely those differential characters which have been usually accounted SPECIFIC, but also those upon which the greater part of the GENERA of this group have been founded, and even in some instances those of its ORDERS" (Foraminifera, Preface, x). Yet this same group had been divided by D'Orbigny and other authors into a number of clearly defined families, genera, and species, which these careful and conscientious researches have shown to have been almost all founded on incomplete knowledge.

Professor DeCandolle has recently given the results of an extensive review of the species of Cupuliferae. He finds that the best-known species of oaks are those which produce most varieties and subvarieties; that they are often surrounded by provisional species; and, with the fullest materials at his command, two-thirds of the species he considers more or less doubtful. His general conclusion is, that "in botany the lowest series of groups, SUBVARIETIES, VARIETIES, and RACES are very badly limited; these can be grouped into SPECIES a little less vaguely limited, which again can be formed into sufficiently precise GENERA." This general conclusion is entirely objected to by the writer of the article in the "Natural History Review," who, however, does not deny its applicability to the particular order under discussion, while this very difference of opinion is another proof that difficulties in the determination of species do not, any more than in the higher groups, vanish with increasing materials and more accurate research.

Another striking example of the same kind is seen in the genera Rubus and Rosa, adduced by Mr. Darwin himself; for though the amplest materials exist for a knowledge of these groups, and the most careful research has been bestowed upon them, yet the various species have not thereby been accurately limited and defined so as to satisfy the majority of botanists. In Mr. Baker's revision of the British Roses, just published by the Linnaean Society, the author includes under the single species Rosa canina, no less than twenty-eight named varieties, distinguished by more or less constant characters and often confined to special localities; and to these are referred about seventy of the species of Continental and British botanists.

Dr. Hooker seems to have found the same thing in his study of the Arctic flora. For though he has had much of the accumulated materials of his predecessors to work upon, he continually expresses himself as unable to do more than group the numerous and apparently fluctuating forms into more or less imperfectly defined species. In his paper on the "Distribution of Arctic Plants," (Trans. Linn. Soc. xxiii., p. 310) Dr. Hooker says:—"The most able and experienced descriptive botanists vary in their estimate of the value of the 'specific term' to a much greater extent than is generally supposed." ... "I think I may safely affirm that the 'specific term' has three different standard values, all current in descriptive botany, but each more or less confined to one class of observers." ... "This is no question of what is right or wrong as to the real value of the specific term; I believe each is right according to the standard he assumes as the specific."

Lastly, I will adduce Mr. Bates's researches on the Amazons. During eleven years he accumulated vast materials, and carefully studied the variation and distribution of insects. Yet he has shown that many species of Lepidoptera, which before offered no special difficulties, are in reality most intricately combined in a tangled web of affinities, leading by such gradual steps from the slightest and least stable variations to fixed races and well-marked species, that it is very often impossible to draw those sharp dividing-lines which it is supposed that a careful study and full materials will always enable us to do.

These few examples show, I think, that in every department of nature there occur instances of the instability of specific form, which the increase of materials aggravates rather than diminishes. And it must be remembered that the naturalist is rarely likely to err on the side of imputing greater indefiniteness to species than really exists. There is a completeness and satisfaction to the mind in defining and limiting and naming a species, which leads us all to do so whenever we conscientiously can, and which we know has led many collectors to reject vague intermediate forms as destroying the symmetry of their cabinets. We must therefore consider these cases of excessive variation and instability as being thoroughly well established; and to the objection that, after all, these cases are but few compared with those in which species can be limited and defined, and are therefore merely exceptions to a general rule, I reply that a true law embraces all apparent exceptions, and that to the great laws of nature there are no real exceptions—that what appear to be such are equally results of law, and are often (perhaps indeed always) those very results which are most important as revealing the true nature and action of the law. It is for such reasons that naturalists now look upon the study of varieties as more important than that of well-fixed species. It is in the former that we see nature still at work, in the very act of producing those wonderful modifications of form, that endless variety of colour, and that complicated harmony of relations, which gratify every sense and give occupation to every faculty of the true lover of nature.

Variation as specially influenced by Locality.

The phenomena of variation as influenced by locality have not hitherto received much attention. Botanists, it is true, are acquainted with the influences of climate, altitude, and other physical conditions, in modifying the forms and external characteristics of plants; but I am not aware that any peculiar influence has been traced to locality, independent of climate. Almost the only case I can find recorded is mentioned in that repertory of natural-history facts, "The Origin of Species," viz. that herbaceous groups have a tendency to become arboreal in islands. In the animal world, I cannot find that any facts have been pointed out as showing the special influence of locality in giving a peculiar facies to the several disconnected species that inhabit it. What I have to adduce on this matter will therefore, I hope, possess some interest and novelty.