Studies in the Theory of Descent, Volume I

On the other hand, it is readily conceivable that, from the conditions of life of caterpillars living on trees or shrubs with dense foliage, the habit of resting by day and descending from the tree for concealment would not have been acquired. Such larvæ are sufficiently protected by their green colour among the large and numerous leaves; and I shall have occasion to show subsequently that their markings increase this protective resemblance.

The di- or polymorphism of the larvæ of the Sphingidæ does not therefore depend upon a contemporaneous double adaptation, but upon the replacement of an old protective colour by a new and better one, and therefore upon a successive double adaptation. The adult caterpillars of C. Elpenor are not sometimes brown and sometimes green because some individuals have become adapted to leaves and others to the soil, but because the anciently inherited green has not yet been completely replaced by the newly acquired brown coloration, some individuals still retaining the old green colour.

When, in another place,[137 - “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872, p. 21.] I formerly stated “that a species can become adapted in this or that manner to given conditions of life, and that by no means can only one best adapted form be allowed for each species,” this statement is theoretically correct speaking generally, but not in its application to the present class of cases. A comparison with one another of those caterpillars which repose by day, distinctly shows that they all possess a tendency to abandon the green and assume a dull colour, but that this process of replacement has advanced further in some species than in others. It will not be without interest to follow this operation in some detailed cases, since we may thus obtain an insight into the processes by which polymorphism has arisen, as well as into the connection between this phenomenon and simple variability.

In D. Hippophaës the process has either not yet commenced, or is as yet in its first rudiments. If we may trust the statements of authors, together with the ordinary green form there occurs, rarely, a silver-grey variety, which may be regarded as the beginning of a process of colour substitution. Among thirty-five living specimens of this scarce species which I was able to procure, the grey form did not occur, neither have I found it in collections.

In Macroglossa Stellatarum we see the transforming process in full operation. A large number of individuals (about thirty-five per cent.) are still green; the number of dark-coloured individuals reaches forty-six per cent., these, therefore, preponderating; whilst between the two extremes there are about nineteen per cent. of transition forms, showing all possible shades between light green and dark blackish-brown or brownish-violet, and even, in solitary individuals, pure violet (See Figs. 3–12, Pl. III.). The relatively small number of the intermediate forms, taken in connection with the fact that all the 140 specimens employed in my investigation were obtained from one female, leads to the conclusion that these forms owe their existence to cross-breeding. It would be superfluous to attempt to prove this last conclusion with reference to the before-mentioned case, in which a caterpillar was streaked with brown and green (Fig. 9, Pl. III.).

The process of transformation, as already mentioned, advances in such a manner that the intermediate forms diminish relatively to the dark individuals. This is found to be the case with Sphinx Convolvuli, and almost to the same extent with Chærocampa Elpenor, in both of which species the green caterpillars are the rarest.[138 - I am unfortunately not able to give exact numbers showing the relative proportions of the different forms, since I have never bred S. Convolvuli from eggs, nor C. Elpenor in sufficient numbers.] Forms truly intermediate in colour between green and brown no longer occur, but apparently only different shades of light and dark brown, passing into brownish-black.

The process has again made a further advance in Chærocampa Porcellus and Celerio as well as in Pterogon Œnotheræ. In all these species the green form occurs,[139 - [With reference to C. Porcellus, see note 71 (#cn_74), p. 188 (#Page_188). R.M.]] but so rarely that very few collectors have seen it. The brown form has therefore in these cases nearly become the predominant type, and the solitary green specimens which occasionally occur, may be regarded as reversions to an older phyletic stage.

Deilephila Livornica appears to have reached a similar stage, but the caterpillar of this species has been so imperfectly observed, that it is difficult to determine, even approximately, the relative proportion of the brown to the green individuals. I have only seen one of the latter in Dr. Staudinger’s collection (Compare Fig. 62, Pl. VII.).

In Deilephila Vespertilio, Euphorbiæ, Dahlii, Mauritanica, Nicæa, and Galii, the green form has completely disappeared. The blackish olive-green colour shown by many caterpillars of the two last species, can be considered as a faint retention of the light green colour which they formerly possessed, and which they both show at the present time in their young stages.

Beginning with the appearance of single darker individuals, we pass on in the first place to a greater variability of colouring, and from this, by the greater diminution of the intermediate forms, to polymorphism; the complete extermination of these forms ending in dimorphism. The whole process of transformation has been thus effected: – As the new colouring always prevailed over the old, the latter was at length completely displaced, and the caterpillars, which were at first simply variable, became polymorphic and then dimorphic, finally returning to monomorphism.

We thus see the process of transformation still going on, and no doubt can arise as to its inciting causes. When a character can with certainty be ascribed to adaptation, we can explain its origin in no other way than by the action of natural selection. If, as I believe, it can and has been shown, not only that caterpillars in general possess adaptive colours, but that these colours can change during the lifetime of one and the same species, in correspondence with external conditions, we must certainly gain a very high conception of the power which natural selection exerts on this group of living forms.[140 - [In the class of cases treated of in the foregoing portions of this essay, the external conditions remain unaltered during the lifetime of the caterpillar, but change of habit, and in some cases of colour, occurs when the insect has attained a size conceivable à priori, and are realized by observation, in which the environment itself may undergo change during the lifetime of the individual caterpillar. Thus, in the case of hibernating species, the colour which is adaptive to the autumnal colours of the foliage of their food-trees would not assimilate to that of the newly-opened leaves in the spring. I have already quoted (Proc. Zoo. Soc. 1873, p. 155) as instances of what may be called “seasonal adaptation,” the larvæ of Geometra Papilionaria, Acidalia Degenararia, and Gnophos Obscurata, and many more could be named. These species undergo a change of colour before or after hibernation, the change being always adaptive to the environment.It has long been known that caterpillars which feed on flowers or on plants of variously-coloured foliage, in some cases partake of the colour of their food. See, for instance, Dr. L. Möller’s memoir, “Die Abhängigkeit der Inseckten von ihrer Umgebung,” 1867, and B. D. Walsh “On Phytophagic Varieties and Phytophagic Species,” Proc. Ent. Soc. Philadelph., vol. iii., p. 403. In 1865 Mr. R. McLachlan published a paper entitled “Observations on some remarkable varieties of Sterrha Sacraria, Linn., with general notes on variation in Lepidoptera” (Trans. Ent. Soc. 1865, p. 453), in which he gave many illustrations of this phenomenon. The larva of Heliothis Peltiger, according to Mr. Reading’s description (Newman’s “British Moths,” p. 438), is another case in point. In 1874 a number of instances were published by Mr. Thomas G. Gentry in a paper entitled “Remarkable Variations in Coloration, Ornamentation, &c., of certain Crepuscular and Nocturnal Lepidopterous Larvæ” (“Canadian Entomologist,” vol. vi., p. 85. See also W. H. Edwards’ description of the summer and autumnal larvæ of Lycæna Pseudargiolus; Ibid., vol. x., pp. 12, 13).The caterpillars of the Sphingidæ appear also in some cases to vary in a manner very suggestive of phytophagic influences. The observations upon S. Ocellatus recorded in the previous note (#cn_103) (p. 241 (#Page_241)) may perhaps be interpreted in this sense. In order to get experimental evidence upon this subject, I may add that Mr. E. Boscher was good enough at my request to repeat his observations, and conduct some breeding experiments during the present year (1880). In the same locality as that previously mentioned, seven larvæ were found feeding on Salix viminalis, all of which were the bright green spotted variety; and in the same osier-bed six more were found on another species of Salix, two of these being the bluish-green variety, and the other four the bright green form. Unless we have here a local race, these observations, in connection with those of last year, tend to show that the light green form is associated with Salix viminalis. When found in the natural state feeding on apple, the caterpillar of this species is generally, perhaps invariably, the bluish-green form. In order to try the effect of breeding the larvæ ab ovo on distinct food-plants, a large number of eggs laid by a female Ocellatus in July were divided into three batches, one being supplied with Salix triandra, another with S. viminalis, and the third lot with apple. The experiment unfortunately failed in great part, owing to most of the larvæ dying off, three from the third lot only surviving; but these were all of the bluish-green form, which colour was shown by all the caterpillars of this batch from their earliest stage. The observation is thus so far successful, as it goes to support the view that the variety mentioned is associated with apple (and S. triandra?) My friend Mr. W. J. Argent informs me that he had a number of specimens of Sphinx Ligustri in his possession this autumn, some of which had been found on lilac and others on laurestinus, and he states that all those on the latter plant had the ground-colour distinctly darker than in those feeding on lilac. I learn also from Mr. W. Davis, of Dartford, that he found a number of these larvæ this year feeding on ash, and that they were all differently coloured to those found on lilac or privet, being of a more greyish-green. Another case of colour-variation in larvæ is that Emmelesia Unifasciata, specimens of which I have recently had an opportunity of examining, through the courtesy of Mr. W. Davis. This species feeds on the seeds of a species of Bartsia when the capsules are in various stages of growth, and (omitting details of marking) those caterpillars found on the green capsules were green, whilst those on the brown capsules were of a corresponding colour.On the whole I am inclined to believe that sufficient importance has not hitherto been given to phytophagic variability as a factor in determining larval coloration, and a large field for experimental investigation here lies open for future work. The obscure chemico-physiological processes which may perhaps be shown by such researches to lead to phytophagic variation, cannot, I am persuaded, produce any great divergence of character if unaided; but when such causes of variability play into the hands of natural selection variations of direct protective advantage to the species, we can easily see that this all-important agency would seize upon and perpetuate such a power of adaptability to a variable environment. (See Proc. Zoo. Soc. 1873, p. 158, and “Nature,” vol. xiv., pp. 329 and 330.) R.M.]]

V. Biological Value of Special Markings

The following questions now present themselves: Have the markings of caterpillars any biological value, or are they in a measure only sports of nature? Can they be considered as partially or entirely the result of natural selection, or has this agency had no share in their production?

The problem here offers itself more distinctly than in any other group of living forms, because it presents an alternative without a third possibility. In other words, if it is not possible to show that larval markings have a distinct biological significance, there remains only for their explanation the assumption of a phyletic force, since the direct action of the environment is insufficient to account for such regularity of development throughout a series of forms. The explanation by sexual selection is excluded ab initio, since we are here concerned with larvæ, and not with reproductive forms.[141 - [In 1879 Mr. George Francis, of Adelaide, forwarded from the latter place a number of moths (a species of Anapæa) together with their larvæ (in alcohol) and cocoons (Proc. Ent. Soc. 1879, p. xvi), and in an accompanying note he stated that the male larva when living is of “a bright emerald green, with red and pink markings on the back, and yellow, black, and white streaks on the sides.” The male larva is described as being smaller than the female, and as possessing all the brilliant colours, the latter “having no red markings, but only white, yellow, and green, with a little black.” I was at first disposed to think that we might be dealing here with two distinct species having differently marked larvæ; but Mr. Francis this present year (1880) forwarded a large number of the living cocoons of this species, which I separated according to size, and, on the emergence of the moths (August), I found that all those from the small cocoons were males, and those from the larger cocoons females. There can be no doubt, therefore, that we have but one species in this case, the larva of which presents the remarkable phenomenon of sexual difference of coloration. As an analogous fact I may here mention the well-known case of Orgyia Antiqua, the larva of which differs in the colour of the tufts of hair according to sex. R.M.]]

The biological significance of marking – if such significance it possess – will be most easily investigated by examining whether species with similar markings have any conditions of life in common which would permit of any possible inference as to the significance of the markings.

Among the Sphingidæ we find four chief forms of marking; (1) complete absence of all marking; (2) longitudinal stripes; either a simple subdorsal or this together with a spiracular and dorsal line; (3) oblique stripes; (4) eye-spots and ring-spots, single, paired, or in complete rows.

Now if we consider in which species these four kinds of marking are of general occurrence, not only in the small group of the Sphingidæ but in the whole order Lepidoptera, we shall arrive at the following results: —

1. Complete absence of marking, so common in the larvæ of other insects, such as the Coleoptera, is but seldom found among Lepidopterous caterpillars.

To this category belong all the species of Sesiidæ (the genera Sesia, Trochilia, Sciapteron, Bembecia, &c.), the larvæ of which, without exception, are of a whitish or yellowish colour, and live partly in the wood of trees and shrubs and partly in the shoots of herbaceous plants. Subterranean larvæ also, living at the roots of plants, such as Hepialus Humuli at the roots of hop, and H. Lupulinus at those of Triticum Repens, possess neither colour nor marking. These, like the foregoing, are yellowish-white, evidently because they are deprived of the influence of light.[142 - [I have already given reasons for suspecting that the colour of green caterpillars may be due to the presence of chlorophyll (or some derivative thereof) in their tissues (see Proc. Zoo. Soc. 1873, p. 159). This substance appears to be one of great chemical stability, and, according to Chautard, who has detected it in an unaltered state in the tissues of certain leaf-feeding insects by means of its absorption spectrum (“Comp. Rend.” Jan. 13th, 1873), it resists the animal digestive processes (Ann. Ch. Phys. [5], iii., 1–56). If this view should be established by future observations, we must regard the green colour of caterpillars as having been produced, when protective, from phytophagic variability by the action of natural selection; and the absence of colour in internal feeders, above referred to, is only secondarily due to the exclusion of light, and depends primarily on the absence of chlorophyll in their food. In connection with this I may adduce the fact, that some few species of Nepticula (N. Oxyacanthella, N. Viscerella, &c.) are green, although they live in leaf-galleries where this colour can hardly be of use as a protection; but their food (hawthorn and elm) contains chlorophyll. See also note 130 (#cn_133), p. 293 (#Page_293). Further investigations in this direction are much needed. R.M.]] The larvæ of certain small moths, such as Tortrix Arbutana and Pomonana, which live in fruit, and many case-bearing Tineina, are likewise without marking and devoid of bright colour, being generally whitish. Many of the small caterpillars which feed exteriorly are also – so far as my experience extends – without definite markings, these being among the most minute, such as the greenish leaf-mining species of Nepticula. It is among the larger species that we first meet with longitudinal and oblique stripes. Eye-spots do not occur in any of these larvæ, a circumstance of the greatest importance for the biological significance of this character, as will be shown subsequently. The small size of the caterpillars cannot be the sole cause of the absence of such eye-spots, since in young Smerinthus caterpillars one centimeter long, the oblique stripes are beautifully developed, and the larvæ of many of the smaller moths considerably exceed this size. The surface of these caterpillars therefore, i. e., the field on which markings are displayed, is not absolutely too small for the development of such a character.

Besides the larvæ of the Micro-lepidoptera and of those species living in the dark, there is also a complete absence of marking in the young stages of many caterpillars. Thus, all the Sphingidæ of which I have been able to observe the development, show no markings immediately after emergence from the egg; in many they appear very soon, even before the first moult, and, in other species, after this period.

2. The second category of markings, longitudinal stripes, is very widely distributed among the most diverse families. This character is found among the larvæ of butterflies, Sphingidæ, Noctuæ, Micro-lepidoptera, &c., but in all these groups it is absent in many species. This last fact is opposed to the view that this character is purely morphological, and leads to the supposition that it may have a biological value, being of service for the preservation of the individual, and therefore of the species.

I find that such marking is of service, stripes extending longitudinally along the upper surface of the caterpillar generally making the latter less conspicuous. This, of course, does not hold good under all circumstances, since there are many species with very striking colours which possess longitudinal stripes. Let us consider, however, a case of adaptive colouring, such as a green caterpillar, which, on this account only, is difficult to see, since it accords with the colour of the plant on which it lives. If it is a small caterpillar, i. e., if its length and thickness do not considerably exceed that of the parts of its food-plant, it can scarcely be better concealed – stripes would hardly confer any special advantage unless the parts of the plant were also striped. But the case is quite different if the caterpillar is considerably larger than the parts of the plant (leaves, stalks, &c.). The most perfect adaptive colouring would not now prevent it from standing out conspicuously as a larger body, among the surrounding parts of the plants. It must be distinctly advantageous therefore to such a caterpillar to be striped, since these markings to a certain extent divide the large body into several longitudinal portions – they no longer permit it to be seen as a whole, and thus act more effectively than mere assimilative colouring in causing it to escape detection. This protection would be the more efficacious if the stripes resembled the parts of the plant in colour and size, such, for instance, as the lines of light and shadow produced by stalks or by long and sharp-edged leaves.

If this view be correct, we should expect longitudinal stripes to be absent in the smallest caterpillars, and to be present more especially in those species which live on plants with their parts similarly disposed, i. e., on plants with numerous thin, closely-growing stalks and grass-like leaves, or on plants with needle-shaped leaves.

It has already been mentioned that the smallest species are devoid of longitudinal striping. The larvæ of the Micro-lepidoptera show no such marking, even when they do not live in the dark, but feed either on the surface or in superficial galleries of the leaves (Nepticula, &c.), in which they must be exposed to almost as much light as when living on the surface. The fact that the subdorsal line sometimes appears in very young Sphinx-larvæ is explained, as has already been shown, by the gradual backward transference of adaptational characters acquired in the last stage of development.

It can easily be demonstrated that longitudinally striped caterpillars mostly live on plants, of which the general appearance gives the impression of a striped arrangement. We have only to consider in connection with their mode of life, any large group of adaptively coloured species marked in this manner. Thus, among the butterflies, nearly all the Satyrinæ possess larvæ conspicuously striped – a fact which is readily explicable, because all these caterpillars live on grasses. This is the case with the genera Melanargia, Erebia, Satyrus, Pararge, Epinephele, and Cænonympha, no species of which, so far as the larvæ are known, is without longitudinal stripes, and all of which feed on grasses. It is interesting that here also, as in certain Sphingidæ, some species are brown, i. e., adapted to the soil, whilst the majority are green, and are therefore adapted to living grass. Just as in the case of the Sphingidæ also, the brown species conceal themselves by day on the earth, whilst some of the green species have likewise acquired this habit. I have already shown how this habit originates from the increasing size of the growing larva, which would otherwise become too conspicuous, in spite of adaptive colour and marking. A beautiful confirmation of this view is found in the circumstance that only the largest species of Satyrus, such as S. Proserpinus, Hermione, Phædrus, &c., possess brown caterpillars. I should not be surprised if a more exact investigation of these species, which have hitherto been but seldom observed, revealed in some cases a dimorphism similar to that of the Sphingidæ; and I believe that I may venture to predict that the young stages of all these brown larvæ – at present quite unknown – are, as in the last-named group, green.

Besides the Satyrinæ, most of the larvæ of the Pierinæ and Hesperidæ possess longitudinal stripes, which are generally less strongly pronounced than in the former subfamily. Some of the Pierinæ live on Cruciferæ, of which the narrow leaves and thin leaf- and flower-stalks present nothing but a linear arrangement; other species of this group, however, feed on Leguminosæ (Lathyrus, Lotus, Coronilla, Vicia), and some few on broad-leaved bushes (Rhamnus). This last fact may appear to be opposed to the theory; but light lateral stripes, such for example, as those possessed by Gonepteryx Rhamni, can never be disadvantageous, and may be of use, even on large leaves, so that if we consider them as an inherited character, there is no reason for natural selection to eliminate them. In the case of caterpillars living on vetch, clover, and other Leguminosæ, it must not be forgotten that, although their food-plants do not present any longitudinal arrangement of parts, they always grow among grasses, the species feeding on such plants always resting between grass stems, and very frequently on the grass itself, so that they can have no better protective marking than longitudinal stripes. The striping of the Hesperidæ larvæ, which partly feed on grasses but mostly on species of Leguminosæ, can be explained in a similar manner.

It is not here my intention to go through all the groups of Lepidoptera in this manner. The instances adduced are quite sufficient to prove that longitudinal stripes occur wherever we should expect to find them, and that they really possess the biological significance which I have ascribed to them. That these markings are occasionally converted into an adaptive imitation of certain special parts of a plant, is shown by the larvæ of many moths, such for example as Chesias Spartiata, which lives on broom (Spartium Scoparium), its longitudinal stripes deceptively resembling the sharp edges of the stems of this plant.[143 - [The same applies to Pseudoterpna Cytisaria, also feeding on broom at the same time of the year. The most striking cases of adaptive resemblance brought about by longitudinal stripes are to be found among fir and pine feeders, species belonging to the most diverse families (Hyloicus Pinastri, Trachea Piniperda, Fidonia Piniaria, &c., &c.) all being most admirably concealed among the needle-shaped leaves. R.M.]]

3. Oblique striping. Can the lilac and white oblique stripes on the sides of a large green caterpillar, such as those of Sphinx Ligustri; or the red and white, or white, black, and red stripes of Smerinthus Tiliæ and Sphinx Drupiferarum respectively, be of any possible use? Have we not here just one of those cases which clearly prove that such a character is purely morphological, and worthless for the preservation of the individual? Does not Nature occasionally sport with purposeless forms and colours; or, as it has often been poetically expressed, does she not here give play to the wealth of her phantasy?

At first sight this indeed appears to be the case. We might almost doubt the adaptive importance of the green ground-colour on finding coloured stripes added thereto, and thus – as one might suppose – abolishing the beneficial action of this ground-colour, by making the insect strikingly conspicuous. But this view would be decidedly incorrect, since oblique stripes are of just the same importance as longitudinal stripes. The former serve to render the caterpillar difficult of detection, by making it resemble, as far as possible, a leaf; they are imitations of the leaf-veins.

Nobody who is in the habit of searching for caterpillars will doubt that, in cases where the oblique stripes are simply white or greenish-white, it is extremely difficult to see the insect on its food-plant, e. g.S. Ocellatus on Salix; not only because it possesses the colour of the leaves, but no less because its large body does not present an unbroken green surface, which would bring it into strong contrast with the leaves, and thus arrest the attention. In the case of the species named, the coloured area of the body is divided by oblique parallel stripes, just in the same manner as a willow leaf. In such instances of course we have not presented to us any special imitation of a leaf with all its details – there is not a perfect resemblance of the insect to a leaf, but only an arrangement of lines and interspaces which does not greatly differ from the division of a leaf by its ribs.

That this view is correct is shown by the occurrence of this form of marking. It is on the whole rare, being found, besides in many Sphingidæ, in isolated cases in various families, but is always confined to those larvæ which live on ribbed leaves, and never occurring in species which feed on grasses or on trees with needle-shaped leaves. This has already been shown with respect to the Sphingidæ, in which the oblique stripes are only completely developed in the subfamilies Smerinthinæ and Sphinginæ. The species of Smerinthus all live on trees such as willows, poplars, lime, oak, &c., and all possess oblique stripes. The genus Anceryx also belongs to the Sphinginæ, and these caterpillars, as far as known, live on trees with needle-shaped leaves. The moths of this last genus are very closely allied to the species of Sphinx, not only in form and colour, but also in many details of marking. The larvæ are however different, this distinction arising entirely from their adaptation to needle-shaped leaves, the Sphinx caterpillars being adapted to ordinary foliage. The species of Anceryx, as has been already shown, are brown mixed with green, and never possess even a trace of the oblique stripes, but have a latticed marking, consisting of many interrupted lines, which very effectively serves to conceal them among the needles and brown bark of the Coniferæ.

Of the Sphinginæ living on plants with ordinary foliage, not a single species is without oblique stripes. I am acquainted with ten species of caterpillars and their respective food-plants, viz. Sphinx Carolina, Convolvuli, Quinquemaculata, Prini, Drupiferarum, Ligustri; Macrosila Rustica and Cingulata; Dolba Hylæus and Acherontia Atropos.

Besides among the Sphinginæ, oblique stripes occur in the larvæ of certain butterflies, viz. Apatura Iris, Ilia, and Clytie, all of which live on forest trees (aspen and willows), and are excellently adapted to the leaves by their green colour. In addition to these, I am acquainted with the larvæ of some few moths, viz. of Aglia Tau and Endromis Versicolora, both of which also live on forest trees.

Oblique stripes also occasionally occur in the smaller caterpillars of Noctuæ, Geometræ, and even in those of certain Pyrales, in all of which they are shorter and differently arranged. In these cases also, my theory of adaptation holds good, but it would take us too far if I attempted to go more closely into them. I will here only mention the extraordinary adaptation shown by the caterpillar of Eriopus Pteridis. This little Noctuid lives on Pteris Aquilina; it possesses the same green colour as this fern, and has double oblique white stripes crossing at a sharp angle on each segment, these resembling the lines of sori of the fern-frond so closely, that the insect is very difficult to perceive.

After all these illustrations it can no longer remain doubtful that the oblique stripes of the Sphingidæ are adaptive. But how are the coloured edges bordering these stripes in so many species to be explained?

I must confess that I long doubted the possibility of being able to ascribe any biological value to this character, which appeared to me only conspicuous, and not protective. Cases may actually occur in which the brightly coloured edges of the oblique stripes make the caterpillar conspicuous – just in the same manner as any marking may bring about a conspicuous appearance by presenting a striking contrast of colour. I am acquainted with no such instance, however. As a rule, in all well-adapted caterpillars, considering their colour in its totality, this is certainly not the case. The coloured edges, on the contrary, enhance the deceptive appearance by representing the oblique shadows cast by the ribs on the under-side of the leaf; all these caterpillars rest underneath the leaves, and never on the upper surface.

This explanation may, perhaps, at first sight appear far-fetched, but if the experiment be made of observing a caterpillar of Sphinx Ligustri on its food-plant, not immediately before one’s eyes in a room, but at a distance as under natural conditions, it will be found that the violet edges do not stand out brightly, but show a colour very similar to that of the shadows playing about the leaves. The coloured edges, in fact, produce a more effective breaking up of the large green surface of the caterpillar’s body, than whitish stripes alone. Of course if the insect was placed on a bare twig in the sun, it would be easily visible at a distance; the larva never rests in such a position, however, but always in the deep shadow of the leaves, in which situation the coloured edges produce their peculiar effect. It may be objected that the oblique white stripes, standing simply on a dark green ground-colour, would produce the same effect, and that my explanation therefore leaves the bright colouring of these edges still unaccounted for. I certainly cannot say why in Sphinx Ligustri these edges are lilac, and in S. Drupiferarum, S. Prini, and Dolba Hylæus red, nor why they are black and green in Macrosila Rustica, and blue in Acherontia Atropos. If we knew exactly on what plants these caterpillars fed originally, we might perhaps indulge in comparing with an artistic eye the shadows playing about their leaves, seeing in one case more red, and in another more blue or violet. The coloured stripes of the Sphingidæ must be regarded as the single strokes of a great master on the countenance of a human portrait. Looked into closely, we see red, blue, or even green spots and strokes; but all these colours, conspicuous when close, disappear on retreating, a general effect of colour being then produced, which cannot be precisely described by words.

Quite in accordance with this explanation, we see caterpillars with the brightest coloured stripes concealing themselves in the earth by day, and betaking themselves to their food-plants only in the dusk of the evening or dawn of morning and even during the night; i. e. in a light so faint that feeble colours would produce scarcely any effect. The bright blue of Acherontia Atropos, for example, would give the impression of oblique shadows without any distinctive colour.

It is precisely the case of this last caterpillar, which formerly appeared to me to present insurmountable difficulties to the explanation of the coloured stripes by adaptation, and I believed that this insect would have to be classed with those species which are brightly coloured because they are distasteful, and are avoided by birds. But although we have no experiments on this point, I must reject this view. Unfortunately, we know scarcely anything of the ontogeny of this caterpillar; but we know at least that the young larvæ (stage four) are greener than the more purely yellow ones of the fifth stage (which, however, are also frequently green), and we know further that some adults are of a dark brownish-grey, without any striking colours. From analogy with the dimorphism of the species of Chærocampa and Sphinx, fully considered previously, it must therefore be concluded that in this case also, a new process of adaptation has commenced – that the caterpillar is becoming adapted to the soil in and on which it conceals itself by day.[144 - The geographical distribution of the dark form indicates that in the case of this species also, the form referred to is replacing the yellow (green) variety. Whilst in the middle of Europe (Germany, France, Hungary) the dark form is extremely rare, in the south of Spain this variety, as I learn from Dr. Noll, is almost as common as the yellow one. I hear also from Dr. Staudinger that in South Africa (Port Natal) the dark form is somewhat the commoner, although the golden-yellow and, more rarely, the green varieties, occur there. I have seen a caterpillar and several moths from Port Natal, and these all agree exactly with ours. The displacement of the green (yellow) form by the dark soil-adapted variety, appears therefore to proceed more rapidly in a warm than in a temperate climate. [Eng. ed. Dr. Noll writes to me from Frankfort that the caterpillar of Acherontia Atropos in the south of Spain does not, as with us, conceal itself by day in the earth, but on the stems underneath the leaves. “At Cadiz, on the hot, sandy shore, Solanum violaceum grows to a height of three feet, and on a single plant I often found more than a dozen Atropos larvæ resting with the head retracted. It can easily be understood why the lateral stripes are blue when one has seen the south European Solaneæ, on which this larva is at home. Solanum violaceum is scarcely green: violet tints alternate with brown, green, and yellow over the whole plant, and between these appear the yellow-anthered flowers, and golden-yellow berries of the size of a greengage. Thus it happens that the numerous thorns, an inch long, between which the caterpillar rests on the stem, pass from violet into shades of blue, red, green, and yellow.”]] An insect which acquires undoubted protective colours cannot, however, be classed with those which possess an immunity from hostile attacks.

That the coloured edges are correctly explained as imitations of the oblique shadows of the leaf-ribs, may also be proved from another point of view. Let us assume, for the sake of argument, that these coloured stripes are not adaptive, and that they have not been produced by natural selection, but by a hypothetical phyletic force. We should then expect to see them appear at some period in the course of the phyletic development – perhaps at first only in solitary individuals, then in several, and finally in all; but we certainly could not expect that at first single, irregular, coloured spots should arise in the neighbourhood of the oblique white stripes – that these spots should then multiply, and fusing together, should adhere to the white stripes, so as to form an irregular spot-like edge, which finally becomes formed into a straight, uniformly broad stripe. The phyletic development of the coloured edges takes place, however, in such a manner, the species of Smerinthus, as has already been established, showing this with particular distinctness. In S. Tiliæ the course of development can be followed till the somewhat irregular red border is formed. In the species of Sphinx this border has become completely linear. It is very possible that the ontogeny of S. Ligustri or Drupiferarum would reveal the whole process, although it may also be possible that owing to the contraction of the development, much of the phylogeny is already lost.

I have now arrived at the consideration of the last kind of marking which occurs in the Sphingidæ, viz.: —

4. Eye-spots and Ring-spots.– These markings, besides among the Sphingidæ, are found only in a very few caterpillars, such as certain tropical Papilionidæ and Noctuæ. I know nothing of the conditions of life and habits of these species, however, and without such knowledge it is impossible to arrive at a complete explanation.

With Darwin, I take an eye-spot to be “a spot within a ring of another colour, like the pupil within the iris,” but to this central spot “concentric zones” maybe added. In the Chærocampa larvæ and in Pterogon Œnotheræ, in which complete ocelli occur, there are always three zones – a central spot, the pupil, or, as I have called it, the “nucleus;” then a light zone, the “mirror;” and, surrounding this again, a dark zone (generally black), the “ground-area.”

As ring-spots I will consider those ocelli which are without the nucleus (pupil), and which are not therefore, strictly speaking, deceptive imitations of an eye, but present a conspicuous light spot surrounded by a dark zone.

Between these two kinds of markings there is, however, no sharp boundary, and morphologically they can scarcely be separated. Species with ring-spots sometimes have nuclei, and ocellated larvæ in some cases possess only a pale spot instead of a dark pupil. I deal here with the two kinds separately, because it happens that they appear in two distinct genera, in each of which they have their special developmental history. Ring-spots originate in a different position, and in another manner than eye-spots; but it must not, on this account, be assumed without further inquiry, that they are called into existence by the same causes; they must rather be investigated separately, from their origin.

Eye-spots are possessed by the genera Chærocampa and Pterogon; ring-spots by the genus Deilephila. In accordance with the data furnished by the above-given developmental histories, the origination of these markings in the two genera may be thus represented: —

In the genera named, eye-spots and ring-spots are formed by the transformation of single portions of the subdorsal line.

In Chærocampa the primary ocelli originate on the fourth and fifth segments by the detachment of a curved portion of the subdorsal, this fragment becoming the “mirror,” and acquiring a dark encircling zone (“ground-area”). The nucleus (pupil) is added subsequently.

In Deilephila we learn from the development of D. Hippophaës, that the primary annulus arises on the segment bearing the caudal horn (the eleventh) by the deposition of a red spot on the white subdorsal line, which is somewhat enlarged in this region. The formation of a dark “ground-area” subsequently occurs, and with this, at first the partial, and then the complete, detachment of the mirror-spot from the subdorsal line takes place.

In both genera the spots arise at first locally on one or two segments, from which they are transferred to the others as a secondary character. In Chærocampa this transference is chiefly backwards, in Deilephila invariably forwards.