Contributions to the Theory of Natural Selection Part 6

IV.

THE MALAYAN PAPILIONIDae OR SWALLOW-TAILED b.u.t.tERFLIES, AS ILl.u.s.tRATIVE 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, s.e.xual, 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 b.u.t.terflies, whose extreme beauty and endless diversity have led to their having been a.s.siduously 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 l.u.s.tre, 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 b.u.t.terflies, 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 b.u.t.terflies" and several other works, | | ill.u.s.trated by exquisite coloured figures drawn by himself; | | and owner of the finest collection of b.u.t.terflies 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 antic.i.p.ate 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 b.u.t.terfly, whether it does or does not belong to this family; and I am not aware that any other group of b.u.t.terflies, 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 a.n.a.logies among African b.u.t.terflies,"

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 Parna.s.sius, 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 b.u.t.terflies may be distinguished from moths, and of all b.u.t.terflies the Papilionidae have the most beautiful and most perfectly developed clubbed antennae. Again, b.u.t.terflies 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 b.u.t.terflies, 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 (Bra.s.solidae) 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 b.u.t.terflies. 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 cla.s.sed 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 cla.s.sed 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 ill.u.s.tration of the fallacy of this mode of reasoning, for if there is one character more than another which is essential and distinctive of the cla.s.s, 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 cla.s.s 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 ill.u.s.tration, 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 b.u.t.terfly type has attained, and deserve to be placed at its head in every system of cla.s.sification.

_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_," const.i.tutes 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 cla.s.s 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, a.n.a.logous 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 cla.s.sing 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 s.e.xes, 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 s.e.x 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 cla.s.sed 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 circ.u.mstances 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 s.e.x. Papilio Memnon is one of the most striking, as it exhibits the mixture of simple variability, local and polymorphic forms, all hitherto cla.s.sed under the common t.i.tle 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. c.o.o.n, a species having the two s.e.xes 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 cla.s.sed 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 circ.u.mstances which led me to cla.s.s 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 cla.s.sed as s.e.xes 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 b.u.t.terfly 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 b.u.t.terfly 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 a.s.sign 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 s.e.xes 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 s.e.x. 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 s.e.xes has been gradually produced by what he terms s.e.xual 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 is approximately the southern limit of the yellow form, and 42 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 lat.i.tude 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 a.n.a.logous 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 a.n.a.logous 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 s.e.x 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 a.n.a.logous 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 s.e.xes of each from a single flock, while no intermediate specimens have yet been found.

The fact of the two s.e.xes 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 s.e.xual 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 acc.u.mulated, till they have produced the wonderful train of the Peac.o.c.k 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 s.e.x only, and still more frequently the males alone rejoicing in rich colours or sparkling l.u.s.tre. But there is here another cause which has led to s.e.xual differences, viz., a special adaptation of the s.e.xes to diverse habits or modes of life. This is well seen in female b.u.t.terflies (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 s.e.xual 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 ill.u.s.trated 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 cla.s.sed 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 princ.i.p.ally 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 const.i.tute 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.