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Darwin and Modern Science (1909)

Edited by A.C. Seward


XV. THE VALUE OF COLOUR IN THE STRUGGLE FOR LIFE.

By E.B. POULTON.
Hope Professor of Zoology in the University of Oxford.

INTRODUCTION.

T
he following pages have been written almost entirely from the historical stand-point. Their principal object has been to give some account of the impressions produced on the mind of Darwin and his great compeer Wallace by various difficult problems suggested by the colours of living nature. In order to render the brief summary of Darwin's thoughts and opinions on the subject in any way complete, it was found necessary to say again much that has often been said before. No attempt has been made to display as a whole the vast contribution of Wallace; but certain of its features are incidentally revealed in passages quoted from Darwin's letters. It is assumed that the reader is familiar with the well-known theories of Protective Resemblance, Warning Colours, and Mimicry both Batesian and Mullerian. It would have been superfluous to explain these on the present occasion; for a far more detailed account than could have been attempted in these pages has recently appeared. (Poulton, "Essays on Evolution" Oxford, 1908, pages 293-382.) Among the older records I have made a point of bringing together the principal observations scattered through the note- books and collections of W.J. Burchell. These have never hitherto found a place in any memoir dealing with the significance of the colours of animals.

INCIDENTAL COLOURS.

Darwin fully recognised that the colours of living beings are not necessarily of value as colours, but that they may be an incidental result of chemical or physical structure. Thus he wrote to T. Meehan, Oct. 9, 1874: "I am glad that you are attending to the colours of dioecious flowers; but it is well to remember that their colours may be as unimportant to them as those of a gall, or, indeed, as the colour of an amethyst or ruby is to these gems." ("More Letters of Charles Darwin", Vol. I. pages 354, 355. See also the admirable account of incidental colours in "Descent of Man" (2nd edition), 1874, pages 261, 262.)

Incidental colours remain as available assets of the organism ready to be turned to account by natural selection. It is a probable speculation that all pigmentary colours were originally incidental; but now and for immense periods of time the visible tints of animals have been modified and arranged so as to assist in the struggle with other organisms or in courtship. The dominant colouring of plants, on the other hand, is an essential element in the paramount physiological activity of chlorophyll. In exceptional instances, however, the shapes and visible colours of plants may be modified in order to promote concealment.

TELEOLOGY AND ADAPTATION.

In the department of Biology which forms the subject of this essay, the adaptation of means to an end is probably more evident than in any other; and it is therefore of interest to compare, in a brief introductory section, the older with the newer teleological views.

The distinctive feature of Natural Selection as contrasted with other attempts to explain the process of Evolution is the part played by the struggle for existence. All naturalists in all ages must have known something of the operations of "Nature red in tooth and claw"; but it was left for this great theory to suggest that vast extermination is a necessary condition of progress, and even of maintaining the ground already gained.

Realising that fitness is the outcome of this fierce struggle, thus turned to account for the first time, we are sometimes led to associate the recognition of adaptation itself too exclusively with Natural Selection. Adaptation had been studied with the warmest enthusiasm nearly forty years before this great theory was given to the scientific world, and it is difficult now to realise the impetus which the works of Paley gave to the study of Natural History. That they did inspire the naturalists of the early part of the last century is clearly shown in the following passages.

In the year 1824 the Ashmolean Museum at Oxford was intrusted to the care of J.S. Duncan of New College. He was succeeded in this office by his brother, P.B. Duncan, of the same College, author of a History of the Museum, which shows very clearly the influence of Paley upon the study of nature, and the dominant position given to his teachings: "Happily at this time (1824) a taste for the study of natural history had been excited in the University by Dr Paley's very interesting work on Natural Theology, and the very popular lectures of Dr Kidd on Comparative Anatomy, and Dr Buckland on Geology." In the arrangement of the contents of the Museum the illustration of Paley's work was given the foremost place by J.S. Duncan: "The first division proposes to familiarize the eye to those relations of all natural objects which form the basis of argument in Dr Paley's Natural Theology; to induce a mental habit of associating the view of natural phenomena with the conviction that they are the media of Divine manifestation; and by such association to give proper dignity to every branch of natural science." ((From "History and Arrangement of the Ashmolean Museum" by P.B. Duncan: see pages vi, vii of "A Catalogue of the Ashmolean Museum", Oxford, 1836.)

The great naturalist, W.J. Burchell, in his classical work shows the same recognition of adaptation in nature at a still earlier date. Upon the subject of collections he wrote ("Travels in the Interior of Southern Africa", London, Vol. I. 1822, page 505. The references to Burchell's observations in the present essay are adapted from the author's article in "Report of the British and South African Associations", 1905, Vol. III. pages 57-110.): "It must not be supposed that these charms (the pleasures of Nature) are produced by the mere discovery of new objects: it is the harmony with which they have been adapted by the Creator to each other, and to the situations in which they are found, which delights the observer in countries where Art has not yet introduced her discords." The remainder of the passage is so admirable that I venture to quote it: "To him who is satisfied with amassing collections of curious objects, simply for the pleasure of possessing them, such objects can afford, at best, but a childish gratification, faint and fleeting; while he who extends his view beyond the narrow field of nomenclature, beholds a boundless expanse, the exploring of which is worthy of the philosopher, and of the best talents of a reasonable being."

On September 14, 1811, Burchell was at Zand Valley (Vlei), or Sand Pool, a few miles south-west of the site of Prieska, on the Orange River. Here he found a Mesembryanthemum (M. turbiniforme, now M. truncatum) and also a "Gryllus" (Acridian), closely resembling the pebbles with which their locality was strewn. He says of both of these, "The intention of Nature, in these instances, seems to have been the same as when she gave to the Chameleon the power of accommodating its color, in a certain degree, to that of the object nearest to it, in order to compensate for the deficiency of its locomotive powers. By their form and colour, this insect may pass unobserved by those birds, which otherwise would soon extirpate a species so little able to elude its pursuers, and this juicy little Mesembryanthemum may generally escape the notice of cattle and wild animals." (Loc. cit. pages 310, 311. See Sir William Thiselton-Dyer "Morphological Notes", XI.; "Protective Adaptations", I.; "Annals of Botany", Vol. XX. page 124. In plates VII., VIII. and IX. accompanying this article the author represents the species observed by Burchell, together with others in which analogous adaptations exist. He writes: "Burchell was clearly on the track on which Darwin reached the goal. But the time had not come for emancipation from the old teleology. This, however, in no respect detracts from the merit or value of his work. For, as Huxley has pointed out ("Life and Letters of Thomas Henry Huxley", London, 1900, I. page 457), the facts of the old teleology are immediately transferable to Darwinism, which simply supplies them with a natural in place of a supernatural explanation.") Burchell here seems to miss, at least in part, the meaning of the relationship between the quiescence of the Acridian and its cryptic colouring. Quiescence is an essential element in the protective resemblance to a stone--probably even more indispensable than the details of the form and colouring. Although Burchell appears to overlook this point he fully recognised the community between protection by concealment and more aggressive modes of defence; for, in the passage of which a part is quoted above, he specially refers to some earlier remarks on page 226 of his Vol. I. We here find that even when the oxen were resting by the Juk rivier (Yoke river), on July 19, 1811, Burchell observed "Geranium spinosum, with a fleshy stem and large white flowers...; and a succulent species of Pelargonium...so defended by the old panicles, grown to hard woody thorns, that no cattle could browze upon it." He goes on to say, "In this arid country, where every juicy vegetable would soon be eaten up by the wild animals, the Great Creating Power, with all-provident wisdom, has given to such plants either an acrid or poisonous juice, or sharp thorns, to preserve the species from annihilation..." All these modes of defence, especially adapted to a desert environment, have since been generally recognised, and it is very interesting to place beside Burchell's statement the following passage from a letter written by Darwin, Aug. 7, 1868, to G.H. Lewes; "That Natural Selection would tend to produce the most formidable thorns will be admitted by every one who has observed the distribution in South America and Africa (vide Livingstone) of thorn- bearing plants, for they always appear where the bushes grow isolated and are exposed to the attacks of mammals. Even in England it has been noticed that all spine-bearing and sting-bearing plants are palatable to quadrupeds, when the thorns are crushed." ("More Letters", I. page 308.)

ADAPTATION AND NATURAL SELECTION.

I have preferred to show the influence of the older teleology upon Natural History by quotations from a single great and insufficiently appreciated naturalist. It might have been seen equally well in the pages of Kirby and Spence and those of many other writers. If the older naturalists who thought and spoke with Burchell of "the intention of Nature" and the adaptation of beings "to each other, and to the situations in which they are found," could have conceived the possibility of evolution, they must have been led, as Darwin was, by the same considerations to Natural Selection. This was impossible for them, because the philosophy which they followed contemplated the phenomena of adaptation as part of a static immutable system. Darwin, convinced that the system is dynamic and mutable, was prevented by these very phenomena from accepting anything short of the crowning interpretation offered by Natural Selection. ("I had always been much struck by such adaptations (e.g. woodpecker and tree-frog for climbing, seeds for dispersal), and until these could be explained it seemed to me almost useless to endeavour to prove by indirect evidence that species have been modified." "Autobiography" in "Life and Letters of Charles Darwin", Vol. I. page 82. The same thought is repeated again and again in Darwin's letters to his friends. It is forcibly urged in the Introduction to the "Origin" (1859), page 3.) And the birth of Darwin's unalterable conviction that adaptation is of dominant importance in the organic world,--a conviction confirmed and ever again confirmed by his experience as a naturalist--may probably be traced to the influence of the great theologian. Thus Darwin, speaking of his Undergraduate days, tells us in his "Autobiography" that the logic of Paley's "Evidences of Christianity" and "Moral Philosophy" gave him as much delight as did Euclid.

"The careful study of these works, without attempting to learn any part by rote, was the only part of the academical course which, as I then felt and as I still believe, was of the least use to me in the education of my mind. I did not at that time trouble myself about Paley's premises; and taking these on trust, I was charmed and convinced by the long line of argumentation." ("Life and Letters", I. page 47.)

When Darwin came to write the "Origin" he quoted in relation to Natural Selection one of Paley's conclusions. "No organ will be formed, as Paley has remarked, for the purpose of causing pain or for doing an injury to its possessor." ("Origin of Species" (1st edition) 1859, page 201.)

The study of adaptation always had for Darwin, as it has for many, a peculiar charm. His words, written Nov. 28, 1880, to Sir W. Thiselton- Dyer, are by no means inapplicable to-day: "Many of the Germans are very contemptuous about making out use of organs; but they may sneer the souls out of their bodies, and I for one shall think it the most interesting part of natural history." ("More Letters" II. page 428.)

PROTECTIVE AND AGGRESSIVE RESEMBLANCE: PROCRYPTIC AND ANTICRYPTIC COLOURING.

Colouring for the purpose of concealment is sometimes included under the head Mimicry, a classification adopted by H.W. Bates in his classical paper. Such an arrangement is inconvenient, and I have followed Wallace in keeping the two categories distinct.

The visible colours of animals are far more commonly adapted for Protective Resemblance than for any other purpose. The concealment of animals by their colours, shapes and attitudes, must have been well known from the period at which human beings first began to take an intelligent interest in Nature. An interesting early record is that of Samuel Felton, who (Dec. 2, 1763) figured and gave some account of an Acridian (Phyllotettix) from Jamaica. Of this insect he says "THE THORAX is like a leaf that is raised perpendicularly from the body." ("Phil. Trans. Roy. Soc." Vol. LIV. Tab. VI. page 55.)

Both Protective and Aggressive Resemblances were appreciated and clearly explained by Erasmus Darwin in 1794: "The colours of many animals seem adapted to their purposes of concealing themselves either to avoid danger, or to spring upon their prey." ("Zoonomia", Vol. I. page 509, London, 1794.)

Protective Resemblance of a very marked and beautiful kind is found in certain plants, inhabitants of desert areas. Examples observed by Burchell almost exactly a hundred years ago have already been mentioned. In addition to the resemblance to stones Burchell observed, although he did not publish the fact, a South African plant concealed by its likeness to the dung of birds. (Sir William Thiselton-Dyer has suggested the same method of concealment ("Annals of Botany", Vol. XX. page 123). Referring to Anacampseros papyracea, figured on plate IX., the author says of its adaptive resemblance: "At the risk of suggesting one perhaps somewhat far- fetched, I must confess that the aspect of the plant always calls to my mind the dejecta of some bird, and the more so owing to the whitening of the branches towards the tips" (loc. cit. page 126). The student of insects, who is so familiar with this very form of protective resemblance in larvae, and even perfect insects, will not be inclined to consider the suggestion far-fetched.) The observation is recorded in one of the manuscript journals kept by the great explorer during his journey. I owe the opportunity of studying it to the kindness of Mr Francis A. Burchell of the Rhodes University College, Grahamstown. The following account is given under the date July 5, 1812, when Burchell was at the Makkwarin River, about half-way between the Kuruman River and Litakun the old capital of the Bachapins (Bechuanas): "I found a curious little Crassula (not in flower) so snow white, that I should never has (have) distinguished it from the white limestones...It was an inch high and a little branchy,...and was at first mistaken for the dung of birds of the passerine order. I have often had occasion to remark that in stony place(s) there grow many small succulent plants and abound insects (chiefly Grylli) which have exactly the same colour as the ground and must for ever escape observation unless a person sit on the ground and observe very attentively."

The cryptic resemblances of animals impressed Darwin and Wallace in very different degrees, probably in part due to the fact that Wallace's tropical experiences were so largely derived from the insect world, in part to the importance assigned by Darwin to Sexual Selection "a subject which had always greatly interested me," as he says in his "Autobiography", ("Life and Letters", Vol. I. page 94.) There is no reference to Cryptic Resemblance in Darwin's section of the Joint Essay, although he gives an excellent short account of Sexual Selection (see page 295). Wallace's section on the other hand contains the following statement: "Even the peculiar colours of many animals, especially insects, so closely resembling the soil or the leaves or the trunks on which they habitually reside, are explained on the same principle; for though in the course of ages varieties of many tints may have occurred, YET THOSE RACES HAVING COLOURS BEST ADAPTED TO CONCEALMENT FROM THEIR ENEMIES WOULD INEVITABLY SURVIVE THE LONGEST." ("Journ. Proc. Linn. Soc." Vol. III. 1859, page 61. The italics are Wallace's.)

It would occupy too much space to attempt any discussion of the difference between the views of these two naturalists, but it is clear that Darwin, although fully believing in the efficiency of protective resemblance and replying to St George Mivart's contention that Natural Selection was incompetent to produce it ("Origin" (6th edition) London, 1872, pages 181, 182; see also page 66.), never entirely agreed with Wallace's estimate of its importance. Thus the following extract from a letter to Sir Joseph Hooker, May 21, 1868, refers to Wallace: "I find I must (and I always distrust myself when I differ from him) separate rather widely from him all about birds' nests and protection; he is riding that hobby to death." ("More Letters", I. page 304.) It is clear from the account given in "The Descent of Man", (London, 1874, pages 452-458. See also "Life and Letters", III. pages 123-125, and "More Letters", II. pages 59-63, 72-74, 76-78, 84-90, 92, 93.), that the divergence was due to the fact that Darwin ascribed more importance to Sexual Selection than did Wallace, and Wallace more importance to Protective Resemblance than Darwin. Thus Darwin wrote to Wallace, Oct. 12 and 13, 1867: "By the way, I cannot but think that you push protection too far in some cases, as with the stripes on the tiger." ("More Letters", I. page 283.) Here too Darwin was preferring the explanation offered by Sexual Selection ("Descent of Man" (2nd edition) 1874, pages 545, 546.), a preference which, considering the relation of the colouring of the lion and tiger to their respective environments, few naturalists will be found to share. It is also shown that Darwin contemplated the possibility of cryptic colours such as those of Patagonian animals being due to sexual selection influenced by the aspect of surrounding nature.

Nearly a year later Darwin in his letter of May 5, 1868?, expressed his agreement with Wallace's views: "Expect that I should put sexual selection as an equal, or perhaps as even a more important agent in giving colour than Natural Selection for protection." ("More Letters", II. pages 77, 78.) The conclusion expressed in the above quoted passage is opposed by the extraordinary development of Protective Resemblance in the immature stages of animals, especially insects.

It must not be supposed, however, that Darwin ascribed an unimportant role to Cryptic Resemblances, and as observations accumulated he came to recognise their efficiency in fresh groups of the animal kingdom. Thus he wrote to Wallace, May 5, 1867: "Haeckel has recently well shown that the transparency and absence of colour in the lower oceanic animals, belonging to the most different classes, may be well accounted for on the principle of protection." ("More Letters", II. page 62. See also "Descent of Man", page 261.) Darwin also admitted the justice of Professor E.S. Morse's contention that the shells of molluscs are often adaptively coloured. ("More Letters", II. page 95.) But he looked upon cryptic colouring and also mimicry as more especially Wallace's departments, and sent to him and to Professor Meldola observations and notes bearing upon these subjects. Thus the following letter given to me by Dr A.R. Wallace and now, by kind permission, published for the first time, accompanied a photograph of the chrysalis of Papilio sarpedon choredon, Feld., suspended from a leaf of its food-plant:

July 9th,
Down, Beckenham, Kent.

My Dear Wallace,

Dr G. Krefft has sent me the enclosed from Sydney. A nurseryman saw a caterpillar feeding on a plant and covered the whole up, but when he searched for the cocoon (pupa), was long before he could find it, so good was its imitation in colour and form to the leaf to which it was attached. I hope that the world goes well with you. Do not trouble yourself by acknowledging this.

Ever yours

Ch. Darwin.

Another deeply interesting letter of Darwin's bearing upon protective resemblance, has only recently been shown to me by my friend Professor E.B. Wilson, the great American Cytologist. With his kind consent and that of Mr Francis Darwin, this letter, written four months before Darwin's death on April 19, 1882, is reproduced here (The letter is addressed: "Edmund B. Wilson, Esq., Assistant in Biology, John Hopkins University, Baltimore Md, U. States.":

December 21, 1881.

Dear Sir,

I thank you much for having taken so much trouble in describing fully your interesting and curious case of mimickry.

I am in the habit of looking through many scientific Journals, and though my memory is now not nearly so good as it was, I feel pretty sure that no such case as yours has been described (amongst the nudibranch) molluscs. You perhaps know the case of a fish allied to Hippocampus, (described some years ago by Dr Gunther in "Proc. Zoolog. Socy.") which clings by its tail to sea-weeds, and is covered with waving filaments so as itself to look like a piece of the same sea-weed. The parallelism between your and Dr Gunther's case makes both of them the more interesting; considering how far a fish and a mollusc stand apart. It would be difficult for anyone to explain such cases by the direct action of the environment.—I am glad that you intend to make further observations on this mollusc, and I hope that you will give a figure and if possible a coloured figure.

With all good wishes from an old brother naturalist,

I remain, Dear Sir,

Yours faithfully,

Charles Darwin.

Professor E.B. Wilson has kindly given the following account of the circumstances under which he had written to Darwin: "The case to which Darwin's letter refers is that of the nudibranch mollusc Scyllaea, which lives on the floating Sargassum and shows a really astonishing resemblance to the plant, having leaf-shaped processes very closely similar to the fronds of the sea-weed both in shape and in colour. The concealment of the animal may be judged from the fact that we found the animal quite by accident on a piece of Sargassum that had been in a glass jar in the laboratory for some time and had been closely examined in the search for hydroids and the like without disclosing the presence upon it of two large specimens of the Scyllaea (the animal, as I recall it, is about two inches long). It was first detected by its movements alone, by someone (I think a casual visitor to the laboratory) who was looking closely at the Sargassum and exclaimed 'Why, the sea-weed is moving its leaves'! We found the example in the summer of 1880 or 1881 at Beaufort, N.C., where the Johns Hopkins laboratory was located for the time being. It must have been seen by many others, before or since.

"I wrote and sent to Darwin a short description of the case at the suggestion of Brooks, with whom I was at the time a student. I was, of course, entirely unknown to Darwin (or to anyone else) and to me the principal interest of Darwin's letter is the evidence that it gives of his extraordinary kindness and friendliness towards an obscure youngster who had of course absolutely no claim upon his time or attention. The little incident made an indelible impression upon my memory and taught me a lesson that was worth learning."

VARIABLE PROTECTIVE RESEMBLANCE.

The wonderful power of rapid colour adjustment possessed by the cuttle-fish was observed by Darwin in 1832 at St Jago, Cape de Verd Islands, the first place visited during the voyage of the "Beagle". From Rio he wrote to Henslow, giving the following account of his observations, May 18, 1832: "I took several specimens of an Octopus which possessed a most marvellous power of changing its colours, equalling any chameleon, and evidently accommodating the changes to the colour of the ground which it passed over. Yellowish green, dark brown, and red, were the prevailing colours; this fact appears to be new, as far as I can find out." ("Life and Letters", I. pages 235, 236. See also Darwin's "Journal of Researches", 1876, pages 6- 8, where a far more detailed account is given together with a reference to "Encycl. of Anat. and Physiol.")

Darwin was well aware of the power of individual colour adjustment, now known to be possessed by large numbers of lepidopterous pupae and larvae. An excellent example was brought to his notice by C.V. Riley ("More Letters" II, pages 385, 386.), while the most striking of the early results obtained with the pupae of butterflies--those of Mrs M.E. Barber upon Papilio nireus--was communicated by him to the Entomological Society of London. ("Trans. Ent. Soc. Lond." 1874, page 519. See also "More Letters", II. page 403.)

It is also necessary to direct attention to C.W. Beebe's ("Zoologica: N.Y. Zool. Soc." Vol. I. No. 1, Sept. 25, 1907: "Geographic variation in birds with especial reference to the effects of humidity".) recent discovery that the pigmentation of the plumage of certain birds is increased by confinement in a superhumid atmosphere. In Scardafella inca, on which the most complete series of experiments was made, the changes took place only at the moults, whether normal and annual or artificially induced at shorter periods. There was a corresponding increase in the choroidal pigment of the eye. At a certain advanced stage of feather pigmentation a brilliant iridescent bronze or green tint made its appearance on those areas where iridescence most often occurs in allied genera. Thus in birds no less than in insects, characters previously regarded as of taxonomic value, can be evoked or withheld by the forces of the environment.

WARNING OR APOSEMATIC COLOURS.

From Darwin's description of the colours and habits it is evident that he observed, in 1833, an excellent example of warning colouring in a little South American toad (Phryniscus nigricans). He described it in a letter to Henslow, written from Monte Video, Nov. 24, 1832: "As for one little toad, I hope it may be new, that it may be christened 'diabolicus.' Milton must allude to this very individual when he talks of 'squat like a toad'; its colours are by Werner ("Nomenclature of Colours", 1821) ink black, vermilion red and buff orange." ("More Letters", I. page 12.) In the "Journal of Researches" (1876, page 97.) its colours are described as follows: "If we imagine, first, that it had been steeped in the blackest ink, and then, when dry, allowed to crawl over a board, freshly painted with the brightest vermilion, so as to colour the soles of its feet and parts of its stomach, a good idea of its appearance will be gained." "Instead of being nocturnal in its habits, as other toads are, and living in damp obscure recesses, it crawls during the heat of the day about the dry sand-hillocks and arid plains,..." The appearance and habits recall T. Belt's well-known description of the conspicuous little Nicaraguan frog which he found to be distasteful to a duck. ("The Naturalist in Nicaragua" (2nd edition) London, 1888, page 321.)

The recognition of the Warning Colours of caterpillars is due in the first instance to Darwin, who, reflecting on Sexual Selection, was puzzled by the splendid colours of sexually immature organisms. He applied to Wallace "who has an innate genius for solving difficulties." ("Descent of Man", page 325. On this and the following page an excellent account of the discovery will be found, as well as in Wallace's "Natural Selection", London, 1875, pages 117-122.) Darwin's original letter exists ("Life and Letters", III. pages 93, 94.), and in it we are told that he had taken the advice given by Bates: "You had better ask Wallace." After some consideration Wallace replied that he believed the colours of conspicuous caterpillars and perfect insects were a warning of distastefulness and that such forms would be refused by birds. Darwin's reply ("Life and Letters", III. pages 94, 95.) is extremely interesting both for its enthusiasm at the brilliancy of the hypothesis and its caution in acceptance without full confirmation:

"Bates was quite right; you are the man to apply to in a difficulty. I never heard anything more ingenious than your suggestion, and I hope you may be able to prove it true. That is a splendid fact about the white moths (A single white moth which was rejected by young turkeys, while other moths were greedily devoured: "Natural Selection", 1875, page 78.); it warms one's very blood to see a theory thus almost proved to be true."

Two years later the hypothesis was proved to hold for caterpillars of many kinds by J. Jenner Weir and A.G. Butler, whose observations have since been abundantly confirmed by many naturalists. Darwin wrote to Weir, May 13, 1869: "Your verification of Wallace's suggestion seems to me to amount to quite a discovery." ("More Letters", II. page 71 (footnote).)

RECOGNITION OR EPISEMATIC CHARACTERS.

This principle does not appear to have been in any way foreseen by Darwin, although he draws special attention to several elements of pattern which would now be interpreted by many naturalists as epismes. He believed that the markings in question interfered with the cryptic effect, and came to the conclusion that, even when common to both sexes, they "are the result of sexual selection primarily applied to the male." ("Descent of Man", page 544.) The most familiar of all recognition characters was carefully explained by him, although here too explained as an ornamental feature now equally transmitted to both sexes: "The hare on her form is a familiar instance of concealment through colour; yet this principle partly fails in a closely-allied species, the rabbit, for when running to its burrow, it is made conspicuous to the sportsman, and no doubt to all beasts of prey, by its upturned white tail." ("Descent of Man", page 542.)

The analogous episematic use of the bright colours of flowers to attract insects for effecting cross-fertilisation and of fruits to attract vertebrates for effecting dispersal is very clearly explained in the "Origin". (Edition 1872, page 161. For a good example of Darwin's caution in dealing with exceptions see the allusion to brightly coloured fruit in "More Letters", II. page 348.)

It is not, at this point, necessary to treat sematic characters at any greater length. They will form the subject of a large part of the following section, where the models of Batesian (Pseudaposematic) mimicry are considered as well as the Mullerian (Synaposematic) combinations of Warning Colours.

MIMICRY,—BATESIAN OR PSEUDAPOSEMATIC, MULLERIAN OR SYNAPOSEMATIC.

The existence of superficial resemblances between animals of various degrees of affinity must have been observed for hundreds of years. Among the early examples, the best known to me have been found in the manuscript note-books and collections of W.J. Burchell, the great traveller in Africa (1810-15) and Brazil (1825-30). The most interesting of his records on this subject are brought together in the following paragraphs.

Conspicuous among well-defended insects are the dark steely or iridescent greenish blue fossorial wasps or sand-wasps, Sphex and the allied genera. Many Longicorn beetles mimic these in colour, slender shape of body and limbs, rapid movements, and the readiness with which they take to flight. On Dec. 21, 1812, Burchell captured one such beetle (Promeces viridis) at Kosi Fountain on the journey from the source of the Kuruman River to Klaarwater. It is correctly placed among the Longicorns in his catalogue, but opposite to its number is the comment "Sphex! totus purpureus."

In our own country the black-and-yellow colouring of many stinging insects, especially the ordinary wasps, affords perhaps the commonest model for mimicry. It is reproduced with more or less accuracy on moths, flies and beetles. Among the latter it is again a Longicorn which offers one of the best-known, although by no means one of the most perfect, examples. The appearance of the well-known "wasp-beetle" (Clytus arietis) in the living state is sufficiently suggestive to prevent the great majority of people from touching it. In Burchell's Brazilian collection there is a nearly allied species (Neoclytus curvatus) which appears to be somewhat less wasp- like than the British beetle. The specimen bears the number "1188," and the date March 27, 1827, when Burchell was collecting in the neighbourhood of San Paulo. Turning to the corresponding number in the Brazilian note- book we find this record: "It runs rapidly like an ichneumon or wasp, of which it has the appearance."

The formidable, well-defended ants are as freely mimicked by other insects as the sand-wasps, ordinary wasps and bees. Thus on February 17, 1901, Guy A.K. Marshall captured, near Salisbury, Mashonaland, three similar species of ants (Hymenoptera) with a bug (Hemiptera) and a Locustid (Orthoptera), the two latter mimicking the former. All the insects, seven in number, were caught on a single plant, a small bushy vetch. ("Trans. Ent. Soc. Lond." 1902, page 535, plate XIX. figs. 53-59.)

This is an interesting recent example from South Africa, and large numbers of others might be added--the observations of many naturalists in many lands; but nearly all of them known since that general awakening of interest in the subject which was inspired by the great hypotheses of H.W. Bates and Fritz Muller. We find, however, that Burchell had more than once recorded the mimetic resemblance to ants. An extremely ant-like bug (the larva of a species of Alydus) in his Brazilian collection is labelled "1141," with the date December 8, 1826, when Burchell was at the Rio das Pedras, Cubatao, near Santos. In the note-book the record is as follows: "1141 Cimex. I collected this for a Formica."

Some of the chief mimics of ants are the active little hunting spiders belonging to the family Attidae. Examples have been brought forward during many recent years, especially by my friends Dr and Mrs Peckham, of Milwaukee, the great authorities on this group of Araneae. Here too we find an observation of the mimetic resemblance recorded by Burchell, and one which adds in the most interesting manner to our knowledge of the subject. A fragment, all that is now left, of an Attid spider, captured on June 30, 1828, at Goyaz, Brazil, bears the following note, in this case on the specimen and not in the note-book: "Black...runs and seems like an ant with large extended jaws." My friend Mr R.I. Pocock, to whom I have submitted the specimen, tells me that it is not one of the group of species hitherto regarded as ant-like, and he adds, "It is most interesting that Burchell should have noticed the resemblance to an ant in its movements. This suggests that the perfect imitation in shape, as well as in movement, seen in many species was started in forms of an appropriate size and colour by the mimicry of movement alone." Up to the present time Burchell is the only naturalist who has observed an example which still exhibits this ancestral stage in the evolution of mimetic likeness.

Following the teachings of his day, Burchell was driven to believe that it was part of the fixed and inexorable scheme of things that these strange superficial resemblances existed. Thus, when he found other examples of Hemipterous mimics, including one (Luteva macrophthalma) with "exactly the manners of a Mantis," he added the sentence, "In the genus Cimex (Linn.) are to be found the outward resemblances of insects of many other genera and orders" (February 15, 1829). Of another Brazilian bug, which is not to be found in his collection, and cannot therefore be precisely identified, he wrote: "Cimex...Nature seems to have intended it to imitate a Sphex, both in colour and the rapid palpitating and movement of the antennae" (November 15, 1826). At the same time it is impossible not to feel the conviction that Burchell felt the advantage of a likeness to stinging insects and to aggressive ants, just as he recognised the benefits conferred on desert plants by spines and by concealment. Such an interpretation of mimicry was perfectly consistent with the theological doctrines of his day. (See Kirby and Spence, "An Introduction to Entomology" (1st edition), London, Vol. II. 1817, page 223.)

The last note I have selected from Burchell's manuscript refers to one of the chief mimics of the highly protected Lycid beetles. The whole assemblage of African insects with a Lycoid colouring forms a most important combination and one which has an interesting bearing upon the theories of Bates and Fritz Muller. This most wonderful set of mimetic forms, described in 1902 by Guy A.K. Marshall, is composed of flower- haunting beetles belonging to the family Lycidae, and the heterogeneous group of varied insects which mimic their conspicuous and simple scheme of colouring. The Lycid beetles, forming the centre or "models" of the whole company, are orange-brown in front for about two-thirds of the exposed surface, black behind for the remaining third. They are undoubtedly protected by qualities which make them excessively unpalatable to the bulk of insect-eating animals. Some experimental proof of this has been obtained by Mr Guy Marshall. What are the forms which surround them? According to the hypothesis of Bates they would be, at any rate mainly, palatable hard-pressed insects which only hold their own in the struggle for life by a fraudulent imitation of the trade-mark of the successful and powerful Lycidae. According to Fritz Muller's hypothesis we should expect that the mimickers would be highly protected, successful and abundant species, which (metaphorically speaking) have found it to their advantage to possess an advertisement, a danger-signal, in common with each other, and in common with the beetles in the centre of the group.

How far does the constitution of this wonderful combination--the largest and most complicated as yet known in all the world--convey to us the idea of mimicry working along the lines supposed by Bates or those suggested by Muller? Figures 1 to 52 of Mr Marshall's coloured plate ("Trans. Ent. Soc. Lond." 1902, plate XVIII. See also page 517, where the group is analysed.) represent a set of forty-two or forty-three species or forms of insects captured in Mashonaland, and all except two in the neighbourhood of Salisbury. The combination includes six species of Lycidae; nine beetles of five groups all specially protected by nauseous qualities, Telephoridae, Melyridae, Phytophaga, Lagriidae, Cantharidae; six Longicorn beetles; one Coprid beetle; eight stinging Hymenoptera; three or four parasitic Hymenoptera (Braconidae, a group much mimicked and shown by some experiments to be distasteful); five bugs (Hemiptera, a largely unpalatable group); three moths (Arctiidae and Zygaenidae, distasteful families); one fly. In fact the whole combination, except perhaps one Phytophagous, one Coprid and the Longicorn beetles, and the fly, fall under the hypothesis of Muller and not under that of Bates. And it is very doubtful whether these exceptions will be sustained: indeed the suspicion of unpalatability already besets the Longicorns and is always on the heels,--I should say the hind tarsi--of a Phytophagous beetle.

This most remarkable group which illustrates so well the problem of mimicry and the alternative hypotheses proposed for its solution, was, as I have said, first described in 1902. Among the most perfect of the mimetic resemblances in it is that between the Longicorn beetle, Amphidesmus analis, and the Lycidae. It was with the utmost astonishment and pleasure that I found this very resemblance had almost certainly been observed by Burchell. A specimen of the Amphidesmus exists in his collection and it bears "651." Turning to the same number in the African Catalogue we find that the beetle is correctly placed among the Longicorns, that it was captured at Uitenhage on Nov. 18, 1813, and that it was found associated with Lycid beetles in flowers ("consocians cum Lycis 78-87 in floribus"). Looking up Nos. 78-87 in the collection and catalogue, three species of Lycidae are found, all captured on Nov. 18, 1813, at Uitenhage. Burchell recognised the wide difference in affinity, shown by the distance between the respective numbers; for his catalogue is arranged to represent relationships. He observed, what students of mimicry are only just beginning to note and record, the coincidence between model and mimic in time and space and in habits. We are justified in concluding that he observed the close superficial likeness although he does not in this case expressly allude to it.

One of the most interesting among the early observations of superficial resemblance between forms remote in the scale of classification was made by Darwin himself, as described in the following passage from his letter to Henslow, written from Monte Video, Aug. 15, 1832: "Amongst the lower animals nothing has so much interested me as finding two species of elegantly coloured true Planaria inhabiting the dewy forest! The false relation they bear to snails is the most extraordinary thing of the kind I have ever seen." ("More Letters", I. page 9.)

Many years later, in 1867, he wrote to Fritz Muller suggesting that the resemblance of a soberly coloured British Planarian to a slug might be due to mimicry. ("Life and Letters", III. page 71.)

The most interesting copy of Bates's classical memoir on Mimicry ("Contributions to an Insect Fauna of the Amazon Valley". "Trans. Linn. Soc." Vol. XXIII. 1862, page 495.), read before the Linnean Society in 1861, is that given by him to the man who has done most to support and extend the theory. My kind friend has given that copy to me; it bears the inscription:

"Mr A.R. Wallace from his old travelling companion the Author."

Only a year and a half after the publication of the "Origin", we find that Darwin wrote to Bates on the subject which was to provide such striking evidence of the truth of Natural Selection: "I am glad to hear that you have specially attended to 'mimetic' analogies--a most curious subject; I hope you publish on it. I have for a long time wished to know whether what Dr Collingwood asserts is true--that the most striking cases generally occur between insects inhabiting the same country." (The letter is dated April 4, 1861. "More Letters", I. page 183.)

The next letter, written about six months later, reveals the remarkable fact that the illustrious naturalist who had anticipated Edward Forbes in the explanation of arctic forms on alpine heights ("I was forestalled in only one important point, which my vanity has always made me regret, namely, the explanation by means of the Glacial period of the presence of the same species of plants and of some few animals on distant mountain summits and in the arctic regions. This view pleased me so much that I wrote it out in extenso, and I believe that it was read by Hooker some years before E. Forbes published his celebrated memoir on the subject. In the very few points in which we differed, I still think that I was in the right. I have never, of course, alluded in print to my having independently worked out this view." "Autobiography, Life and Letters", I. page 88.), had also anticipated H.W. Bates in the theory of Mimicry: "What a capital paper yours will be on mimetic resemblances! You will make quite a new subject of it. I had thought of such cases as a difficulty; and once, when corresponding with Dr Collingwood, I thought of your explanation; but I drove it from my mind, for I felt that I had not knowledge to judge one way or the other." (The letter is dated Sept. 25, 1861: "More Letters", I. page 197.)

Bates read his paper before the Linnean Society, Nov. 21, 1861, and Darwin's impressions on hearing it were conveyed in a letter to the author dated Dec. 3: "Under a general point of view, I am quite convinced (Hooker and Huxley took the same view some months ago) that a philosophic view of nature can solely be driven into naturalists by treating special subjects as you have done. Under a special point of view, I think you have solved one of the most perplexing problems which could be given to solve." ("Life and Letters", II. page 378.) The memoir appeared in the following year, and after reading it Darwin wrote as follows, Nov. 20, 1862: "...In my opinion it is one of the most remarkable and admirable papers I ever read in my life...I am rejoiced that I passed over the whole subject in the "Origin", for I should have made a precious mess of it. You have most clearly stated and solved a wonderful problem...Your paper is too good to be largely appreciated by the mob of naturalists without souls; but, rely on it, that it will have LASTING value, and I cordially congratulate you on your first great work. You will find, I should think, that Wallace will fully appreciate it." ("Life and Letters", II. pages 391-393.) Four days later, Nov. 24, Darwin wrote to Hooker on the same subject: "I have now finished his paper...' it seems to me admirable. To my mind the act of segregation of varieties into species was never so plainly brought forward, and there are heaps of capital miscellaneous observations." ("More Letters", I. page 214.)

Darwin was here referring to the tendency of similar varieties of the same species to pair together, and on Nov. 25 he wrote to Bates asking for fuller information on this subject. ("More Letters", I. page 215. See also parts of Darwin's letter to Bates in "Life and Letters", II. page 392.) If Bates's opinion were well founded, sexual selection would bear a most important part in the establishment of such species. (See Poulton, "Essays on Evolution", 1908, pages 65, 85-88.) It must be admitted, however, that the evidence is as yet quite insufficient to establish this conclusion. It is interesting to observe how Darwin at once fixed on the part of Bates's memoir which seemed to bear upon sexual selection. A review of Bates's theory of Mimicry was contributed by Darwin to the "Natural History Review" (New Ser. Vol. III. 1863, page 219.) and an account of it is to be found in the "Origin" (Edition 1872, pages 375-378.) and in "The Descent of Man". (Edition 1874, pages 323-325.)

Darwin continually writes of the value of hypothesis as the inspiration of inquiry. We find an example in his letter to Bates, Nov. 22, 1860: "I have an old belief that a good observer really means a good theorist, and I fully expect to find your observations most valuable." ("More Letters", I. page 176.) Darwin's letter refers to many problems upon which Bates had theorised and observed, but as regards Mimicry itself the hypothesis was thought out after the return of the letter from the Amazons, when he no longer had the opportunity of testing it by the observation of living Nature. It is by no means improbable that, had he been able to apply this test, Bates would have recognised that his division of butterfly resemblances into two classes,--one due to the theory of mimicry, the other to the influence of local conditions,--could not be sustained.

Fritz Muller's contributions to the problem of Mimicry were all made in S.E. Brazil, and numbers of them were communicated, with other observations on natural history, to Darwin, and by him sent to Professor R. Meldola who published many of the facts. Darwin's letters to Meldola (Poulton, "Charles Darwin and the theory of Natural Selection", London, 1896, pages 199-218.) contain abundant proofs of his interest in Muller's work upon Mimicry. One deeply interesting letter (Loc. cit. pages 201, 202.) dated Jan. 23, 1872, proves that Fritz Muller before he originated the theory of Common Warning Colours (Synaposematic Resemblance or Mullerian Mimicry), which will ever be associated with his name, had conceived the idea of the production of mimetic likeness by sexual selection.

Darwin's letter to Meldola shows that he was by no means inclined to dismiss the suggestion as worthless, although he considered it daring. "You will also see in this letter a strange speculation, which I should not dare to publish, about the appreciation of certain colours being developed in those species which frequently behold other forms similarly ornamented. I do not feel at all sure that this view is as incredible as it may at first appear. Similar ideas have passed through my mind when considering the dull colours of all the organisms which inhabit dull-coloured regions, such as Patagonia and the Galapagos Is." A little later, on April 5, he wrote to Professor August Weismann on the same subject: "It may be suspected that even the habit of viewing differently coloured surrounding objects would influence their taste, and Fritz Muller even goes so far as to believe that the sight of gaudy butterflies might influence the taste of distinct species." ("Life and Letters", III. page 157.)

This remarkable suggestion affords interesting evidence that F. Muller was not satisfied with the sufficiency of Bates's theory. Nor is this surprising when we think of the numbers of abundant conspicuous butterflies which he saw exhibiting mimetic likenesses. The common instances in his locality, and indeed everywhere in tropical America, were anything but the hard-pressed struggling forms assumed by the theory of Bates. They belonged to the groups which were themselves mimicked by other butterflies. Fritz Muller's suggestion also shows that he did not accept Bates's alternative explanation of a superficial likeness between models themselves, based on some unknown influence of local physico-chemical forces. At the same time Muller's own suggestion was subject to this apparently fatal objection, that the sexual selection he invoked would tend to produce resemblances in the males rather than the females, while it is well known that when the sexes differ the females are almost invariably more perfectly mimetic than the males and in a high proportion of cases are mimetic while the males are non-mimetic.

The difficulty was met several years later by Fritz Muller's well-known theory, published in 1879 ("Kosmos", May 1879, page 100.), and immediately translated by Meldola and brought before the Entomological Society. ("Proc. Ent. Soc. Lond." 1879, page xx.) Darwin's letter to Meldola dated June 6, 1879, shows "that the first introduction of this new and most suggestive hypothesis into this country was due to the direct influence of Darwin himself, who brought it before the notice of the one man who was likely to appreciate it at its true value and to find the means for its presentation to English naturalists." ("Charles Darwin and the Theory of Natural Selection", page 214.) Of the hypothesis itself Darwin wrote "F. Muller's view of the mutual protection was quite new to me." (Ibid. page 213.) The hypothesis of Mullerian mimicry was at first strongly opposed. Bates himself could never make up his mind to accept it. As the Fellows were walking out of the meeting at which Professor Meldola explained the hypothesis, an eminent entomologist, now deceased, was heard to say to Bates: "It's a case of save me from my friends!" The new ideas encountered and still encounter to a great extent the difficulty that the theory of Bates had so completely penetrated the literature of natural history. The present writer has observed that naturalists who have not thoroughly absorbed the older hypothesis are usually far more impressed by the newer one than are those whose allegiance has already been rendered. The acceptance of Natural Selection itself was at first hindered by similar causes, as Darwin clearly recognised: "If you argue about the non- acceptance of Natural Selection, it seems to me a very striking fact that the Newtonian theory of gravitation, which seems to every one now so certain and plain, was rejected by a man so extraordinarily able as Leibnitz. The truth will not penetrate a preoccupied mind." (To Sir J. Hooker, July 28, 1868, "More Letters", I. page 305. See also the letter to A.R. Wallace, April 30, 1868, in "More Letters" II. page 77, lines 6-8 from top.)

There are many naturalists, especially students of insects, who appear to entertain an inveterate hostility to any theory of mimicry. Some of them are eager investigators in the fascinating field of geographical distribution, so essential for the study of Mimicry itself. The changes of pattern undergone by a species of Erebia as we follow it over different parts of the mountain ranges of Europe is indeed a most interesting inquiry, but not more so than the differences between e.g. the Acraea johnstoni of S.E. Rhodesia and of Kilimanjaro. A naturalist who is interested by the Erebia should be equally interested by the Acraea; and so he would be if the student of mimicry did not also record that the characteristics which distinguish the northern from the southern individuals of the African species correspond with the presence, in the north but not in the south, of certain entirely different butterflies. That this additional information should so greatly weaken, in certain minds, the appeal of a favourite study, is a psychological problem of no little interest. This curious antagonism is I believe confined to a few students of insects. Those naturalists who, standing rather farther off, are able to see the bearings of the subject more clearly, will usually admit the general support yielded by an ever-growing mass of observations to the theories of Mimicry propounded by H.W. Bates and Fritz Muller. In like manner natural selection itself was in the early days often best understood and most readily accepted by those who were not naturalists. Thus Darwin wrote to D.T. Ansted, Oct. 27, 1860: "I am often in despair in making the generality of NATURALISTS even comprehend me. Intelligent men who are not naturalists and have not a bigoted idea of the term species, show more clearness of mind." ("More Letters", I. page 175.)

Even before the "Origin" appeared Darwin anticipated the first results upon the mind of naturalists. He wrote to Asa Gray, Dec. 21, 1859: "I have made up my mind to be well abused; but I think it of importance that my notions should be read by intelligent men, accustomed to scientific argument, though NOT naturalists. It may seem absurd, but I think such men will drag after them those naturalists who have too firmly fixed in their heads that a species is an entity." ("Life and Letters" II. page 245.)

Mimicry was not only one of the first great departments of zoological knowledge to be studied under the inspiration of natural Selection, it is still and will always remain one of the most interesting and important of subjects in relation to this theory as well as to evolution. In mimicry we investigate the effect of environment in its simplest form: we trace the effects of the pattern of a single species upon that of another far removed from it in the scale of classification. When there is reason to believe that the model is an invader from another region and has only recently become an element in the environment of the species native to its second home, the problem gains a special interest and fascination. Although we are chiefly dealing with the fleeting and changeable element of colour we expect to find and we do find evidence of a comparatively rapid evolution. The invasion of a fresh model is for certain species an unusually sudden change in the forces of the environment and in some instances we have grounds for the belief that the mimetic response has not been long delayed.

MIMICRY AND SEX.

Ever since Wallace's classical memoir on mimicry in the Malayan Swallowtail butterflies, those naturalists who have written on the subject have followed his interpretation of the marked prevalence of mimetic resemblance in the female sex as compared with the male. They have believed with Wallace that the greater dangers of the female, with slower flight and often alighting for oviposition, have been in part met by the high development of this special mode of protection. The fact cannot be doubted. It is extremely common for a non-mimetic male to be accompanied by a beautifully mimetic female and often by two or three different forms of female, each mimicking a different model. The male of a polymorphic mimetic female is, in fact, usually non-mimetic (e.g. Papilio dardanus = merope), or if a mimic (e.g. the Nymphaline genus Euripus), resembles a very different model. On the other hand a non-mimetic female accompanied by a mimetic male is excessively rare. An example is afforded by the Oriental Nymphaline, Cethosia, in which the males of some species are rough mimics of the brown Danaines. In some of the orb-weaving spiders the males mimic ants, while the much larger females are non-mimetic. When both sexes mimic, it is very common in butterflies and is also known in moths, for the females to be better and often far better mimics than the males.

Although still believing that Wallace's hypothesis in large part accounts for the facts briefly summarised above, the present writer has recently been led to doubt whether it offers a complete explanation. Mimicry in the male, even though less beneficial to the species than mimicry in the female, would still surely be advantageous. Why then is it so often entirely restricted to the female? While the attempt to find an answer to this question was haunting me, I re-read a letter written by Darwin to Wallace, April 15, 1868, containing the following sentences: "When female butterflies are more brilliant than their males you believe that they have in most cases, or in all cases, been rendered brilliant so as to mimic some other species, and thus escape danger. But can you account for the males not having been rendered equally brilliant and equally protected? Although it may be most for the welfare of the species that the female should be protected, yet it would be some advantage, certainly no disadvantage, for the unfortunate male to enjoy an equal immunity from danger. For my part, I should say that the female alone had happened to vary in the right manner, and that the beneficial variations had been transmitted to the same sex alone. Believing in this, I can see no improbability (but from analogy of domestic animals a strong probability) that variations leading to beauty must often have occurred in the males alone, and been transmitted to that sex alone. Thus I should account in many cases for the greater beauty of the male over the female, without the need of the protective principle." ("More Letters", II. pages 73, 74. On the same subject--"the gay-coloured females of Pieris" (Perrhybris (Mylothris) pyrrha of Brazil), Darwin wrote to Wallace, May 5, 1868, as follows: "I believe I quite follow you in believing that the colours are wholly due to mimicry; and I further believe that the male is not brilliant from not having received through inheritance colour from the female, and from not himself having varied; in short, that he has not been influenced by selection." It should be noted that the male of this species does exhibit a mimetic pattern on the under surface. "More Letters" II. page 78.)

The consideration of the facts of mimicry thus led Darwin to the conclusion that the female happens to vary in the right manner more commonly than the male, while the secondary sexual characters of males supported the conviction "that from some unknown cause such characters (viz. new characters arising in one sex and transmitted to it alone) apparently appear oftener in the male than in the female." (Letter from Darwin to Wallace, May 5, 1867, "More Letters", II. Page 61.)

Comparing these conflicting arguments we are led to believe that the first is the stronger. Mimicry in the male would be no disadvantage but an advantage, and when it appears would be and is taken advantage of by selection. The secondary sexual characters of males would be no advantage but a disadvantage to females, and, as Wallace thinks, are withheld from this sex by selection. It is indeed possible that mimicry has been hindered and often prevented from passing to the males by sexual selection. We know that Darwin was much impressed ("Descent of Man", page 325.) by Thomas Belt's daring and brilliant suggestion that the white patches which exist, although ordinarily concealed, on the wings of mimetic males of certain Pierinae (Dismorphia), have been preserved by preferential mating. He supposed this result to have been brought about by the females exhibiting a deep-seated preference for males that displayed the chief ancestral colour, inherited from periods before any mimetic pattern had been evolved in the species. But it has always appeared to me that Belt's deeply interesting suggestion requires much solid evidence and repeated confirmation before it can be accepted as a valid interpretation of the facts. In the present state of our knowledge, at any rate of insects and especially of Lepidoptera, it is probable that the female is more apt to vary than the male and that an important element in the interpretation of prevalent female mimicry is provided by this fact.

In order adequately to discuss the question of mimicry and sex it would be necessary to analyse the whole of the facts, so far as they are known in butterflies. On the present occasion it is only possible to state the inferences which have been drawn from general impressions,--inferences which it is believed will be sustained by future inquiry.

(1) Mimicry may occasionally arise in one sex because the differences which distinguish it from the other sex happen to be such as to afford a starting-point for the resemblance. Here the male is at no disadvantage as compared with the female, and the rarity of mimicry in the male alone (e.g. Cethosia) is evidence that the great predominance of female mimicry is not to be thus explained.

(2) The tendency of the female to dimorphism and polymorphism has been of great importance in determining this predominance. Thus if the female appear in two different forms and the male in only one it will be twice as probable that she will happen to possess a sufficient foundation for the evolution of mimicry.

(3) The appearance of melanic or partially melanic forms in the female has been of very great service, providing as it does a change of ground-colour. Thus the mimicry of the black generally red-marked American "Aristolochia swallowtails" (Pharmacophagus) by the females of Papilio swallowtails was probably begun in this way.

(4) It is probably incorrect to assume with Haase that mimicry always arose in the female and was later acquired by the male. Both sexes of the third section of swallowtails (Cosmodesmus) mimic Pharmacophagus in America, far more perfectly than do the females of Papilio. But this is not due to Cosmodesmus presenting us with a later stage of history begun in Papilio; for in Africa Cosmodesmus is still mimetic (of Danainae) in both sexes although the resemblances attained are imperfect, while many African species of Papilio have non-mimetic males with beautifully mimetic females. The explanation is probably to be sought in the fact that the females of Papilio are more variable and more often tend to become dimorphic than those of Cosmodesmus, while the latter group has more often happened to possess a sufficient foundation for the origin of the resemblance in patterns which, from the start, were common to male and female.

(5) In very variable species with sexes alike, mimicry can be rapidly evolved in both sexes out of very small beginnings. Thus the reddish marks which are common in many individuals of Limenitis arthemis were almost certainly the starting-point for the evolution of the beautifully mimetic L. archippus. Nevertheless in such cases, although there is no reason to suspect any greater variability, the female is commonly a somewhat better mimic than the male and often a very much better mimic. Wallace's principle seems here to supply the obvious interpretation.

(6) When the difference between the patterns of the model and presumed ancestor of the mimic is very great, the female is often alone mimetic; when the difference is comparatively small, both sexes are commonly mimetic. The Nymphaline genus Hypolimnas is a good example. In Hypolimnas itself the females mimic Danainae with patterns very different from those preserved by the non-mimetic males: in the sub-genus Euralia, both sexes resemble the black and white Ethiopian Danaines with patterns not very dissimilar from that which we infer to have existed in the non-mimetic ancestor.

(7) Although a melanic form or other large variation may be of the utmost importance in facilitating the start of a mimetic likeness, it is impossible to explain the evolution of any detailed resemblance in this manner. And even the large colour variation itself may well be the expression of a minute and "continuous" change in the chemical and physical constitution of pigments.

SEXUAL SELECTION (EPIGAMIC CHARACTERS).

We do not know the date at which the idea of Sexual Selection arose in Darwin's mind, but it was probably not many years after the sudden flash of insight which, in October 1838, gave to him the theory of Natural Selection. An excellent account of Sexual Selection occupies the concluding paragraph of Part I. of Darwin's Section of the Joint Essay on Natural Selection, read July 1st, 1858, before the Linnean Society. ("Journ. Proc. Linn. Soc." Vol. III. 1859, page 50.) The principles are so clearly and sufficiently stated in these brief sentences that it is appropriate to quote the whole: "Besides this natural means of selection, by which those individuals are preserved, whether in their egg, or larval, or mature state, which are best adapted to the place they fill in nature, there is a second agency at work in most unisexual animals, tending to produce the same effect, namely, the struggle of the males for the females. These struggles are generally decided by the law of battle, but in the case of birds, apparently, by the charms of their song, by their beauty or their power of courtship, as in the dancing rock-thrush of Guiana. The most vigorous and healthy males, implying perfect adaptation, must generally gain the victory in their contests. This kind of selection, however, is less rigorous than the other; it does not require the death of the less successful, but gives to them fewer descendants. The struggle falls, moreover, at a time of year when food is generally abundant, and perhaps the effect chiefly produced would be the modification of the secondary sexual characters, which are not related to the power of obtaining food, or to defence from enemies, but to fighting with or rivalling other males. The result of this struggle amongst the males may be compared in some respects to that produced by those agriculturists who pay less attention to the careful selection of all their young animals, and more to the occasional use of a choice mate."

A full exposition of Sexual Selection appeared in the "The Descent of Man" in 1871, and in the greatly augmented second edition, in 1874. It has been remarked that the two subjects, "The Descent of Man and Selection in Relation to Sex", seem to fuse somewhat imperfectly into the single work of which they form the title. The reason for their association is clearly shown in a letter to Wallace, dated May 28, 1864: "...I suspect that a sort of sexual selection has been the most powerful means of changing the races of man." ("More Letters", II. page 33.)

Darwin, as we know from his Autobiography ("Life and Letters", I. page 94.), was always greatly interested in this hypothesis, and it has been shown in the preceding pages that he was inclined to look favourably upon it as an interpretation of many appearances usually explained by Natural Selection. Hence Sexual Selection, incidentally discussed in other sections of the present essay, need not be considered at any length, in the section specially allotted to it.

Although so interested in the subject and notwithstanding his conviction that the hypothesis was sound, Darwin was quite aware that it was probably the most vulnerable part of the "Origin". Thus he wrote to H.W. Bates, April 4, 1861: "If I had to cut up myself in a review I would have (worried?) and quizzed sexual selection; therefore, though I am fully convinced that it is largely true, you may imagine how pleased I am at what you say on your belief." ("More Letters", I. page 183.)

The existence of sound-producing organs in the males of insects was, Darwin considered, the strongest evidence in favour of the operation of sexual selection in this group. ("Life and Letters", III. pages 94, 138.) Such a conclusion has received strong support in recent years by the numerous careful observations of Dr F.A. Dixey ("Proc. Ent. Soc. Lond." 1904, page lvi; 1905, pages xxxvii, liv; 1906, page ii.) and Dr G.B. Longstaff ("Proc. Ent. Soc. Lond." 1905, page xxxv; "Trans. Ent. Soc. Lond." 1905, page 136; 1908, page 607.) on the scents of male butterflies. The experience of these naturalists abundantly confirms and extends the account given by Fritz Muller ("Jen. Zeit." Vol. XI. 1877, page 99; "Trans. Ent. Soc. Lond." 1878, page 211.) of the scents of certain Brazilian butterflies. It is a remarkable fact that the apparently epigamic scents of male butterflies should be pleasing to man while the apparently aposematic scents in both sexes of species with warning colours should be displeasing to him. But the former is far more surprising than the latter. It is not perhaps astonishing that a scent which is ex hypothesi unpleasant to an insect- eating Vertebrate should be displeasing to the human sense; but it is certainly wonderful that an odour which is ex hypothesi agreeable to a female butterfly should also be agreeable to man.

Entirely new light upon the seasonal appearance of epigamic characters is shed by the recent researches of C.W. Beebe ("The American Naturalist", Vol. XLII. No. 493, Jan. 1908, page 34.), who caused the scarlet tanager (Piranga erythromelas) and the bobolink (Dolichonyx oryzivorus) to retain their breeding plumage through the whole year by means of fattening food, dim illumination, and reduced activity. Gradual restoration to the light and the addition of meal-worms to the diet invariably brought back the spring song, even in the middle of winter. A sudden alteration of temperature, either higher or lower, caused the birds nearly to stop feeding, and one tanager lost weight rapidly and in two weeks moulted into the olive-green winter plumage. After a year, and at the beginning of the normal breeding season, "individual tanagers and bobolinks were gradually brought under normal conditions and activities," and in every case moulted from nuptial plumage to nuptial plumage. "The dull colours of the winter season had been skipped." The author justly claims to have established "that the sequence of plumage in these birds is not in any way predestined through inheritance..., but that it may be interrupted by certain factors in the environmental complex."



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