The general prevalence of protective colouring among caterpillars has already been so frequently treated of that it is not here my intention to recall particular instances. In order to judge of the effect of marking, however, it will be well to bear in mind that these insects, being generally defenceless and thus requiring protection, have acquired the most diverse means of rendering themselves in some measure secure from their foes.
The sharp spines which occur on the caterpillars of many butterflies (Vanessa,Melitæa,Argynnis), and the hairs on those of many moths, serve for protective purposes. Among other means of protection—although in a different sense—we have in all the species of the great family of thePapilionidæthe strikingly coloured (yellowish red) odour-emitting tentacles concealed near the head, and suddenly protruded for terrifying foes; and likewise the forked horn at the tail of the caterpillars of the genus of mothsHarpyia, thetentacles of which can be suddenly protruded in a similar manner. Adaptive colours and forms combined with certainhabits129are, however, muchmore common than defensive weapons. Thus, the caterpillars of theNoctuæbelonging to the genusCatocalaand its allies, feed only at night on the green leaves of various forest-trees; by day they rest in crevices of the bark on the tree trunk, which they resemble so perfectly in the colour of their peculiar glossy dull grey or brownish skin beset with small humps, that only sharp eyes candetect them, even when we are familiar with theirhabits.130
The striking resemblance of many moths to splinters of wood is well known, and to this is added a habit which helps their disguise, viz., that of remaining stiff and motionless on the approach of danger, just like a splinter projecting from thebranch.131Among the moths coming under this category may be mentionedCucullia Verbasci, and particularly those of the genusXylina, which, when at rest, closely resemble a broken splinter of wood in the colour and marking of their fore wings, and when touched, have a habit of drawing in their legs and falling without opening their wings as though dead.
That simple adaptive colouring prevails widelyamong caterpillars is shown by the large number of greenspecies.132It may be fairly said that all caterpillars which possess no other means of protection or defence are adaptively coloured. These facts are now well known; so also is the explanation of the varied and striking colours of many caterpillars given byWallace.133There is, however,novelty in the proof contained in the foregoing descriptions of larval development, as tothe manner in which the di- and polymorphism of caterpillars can be explained from the external phenomena which they present, these phenomena being well adapted for showing the great importance of protective colouring to the larvæ. We have here presented a double adaptation, although not quite of the nature of that which I formerly admitted on hypotheticalgrounds.134In the first place, from the developmental history there results the conclusion that all Sphinx-larvæ which, in the adult state, are di- or polymorphic, are unicolorous when young. Thus, the caterpillars ofChærocampa Elpenorall remain green till the fourth stage, when they mostly become light or dark brown, and only very seldom retain their green colour.Chærocampa Porcellusbehaves in a precisely similar manner; as also doesPterogon Œnotheræ, which inhabits the same localities, and is found on the same food-plant, but is not very closely related to theChærocampa. In this species also (P. Œnotheræ) the brown is more common than the green form in the adult state, both varieties showing a complicated marking. The young larvæ possess only a light green colour, and a pure white subdorsal line as the only marking; they are so well adapted to the leaves of their food-plants,Epilobium Hirsutum, andE. Rosmarinifolium, that they can only be detectedwith great difficulty. After the third moult they become brown, and can be easily seen when at rest on their food-plant.
Now in all known caterpillars brown colours are adaptive, sometimes causing a resemblance to the soil, and at others to dead leaves or branches. As soon, therefore, as the caterpillars have attained a considerable size, they remain concealed byday.135The truth of this observation not only appears from various entomological notes, but I have frequently convinced myself of its accuracy. I well remember from the earliest times thatC. Elpenor, especially when the larva is adult, always rests by day among the dead branches and leaves of its shrub-like food-plant,Epilobium Hirsutum; and even when this species lives on the low-growingEpilobium Parviflorum, it conceals itself by day on the ground, among the tangled leaves and branches. I have observed thatSphinx Convolvulihas a precisely similar habit, for which reason it is difficult to obtain, even in localities where it occurs very commonly.
In the neighbourhood of Basle I once found at mid-day a brown caterpillar ofPterogon Œnotheræon an isolated dead branch ofEpilobium Rosmarinifolium, and I was informed by Herr Riggenbach-Stähelin—a collector of great experience who accompanied me—that these caterpillars always rest (by day) on withered plants as soon as theybecome brown, but before this change they are only to be found on green plants.
Thus, it cannot well be doubted that the change of colour is associated with a change in the habits of life, and the question arises as to which has been the primary change.
If the view here entertained, that the later brown coloration is adaptive, be correct, the species must have first acquired the habit of concealing itself by day on the ground and among dead herbage, before the original green colour could have been changed into brown by natural selection. This must represent the actual facts of the case.
Nearly allied species which at an advanced age are not dimorphic, but are darkly coloured in all individuals, are especially calculated to throw some light on this point. For instance, the caterpillar ofDeilephila Vespertilio, which comes under this denomination, is light green when young, and rests both by day and night on the leaves of the plant on which it feeds. As soon as it acquires its dark colour—after the third moult—it changes its habits, concealing itself by day on the ground and feeding only by night. For this reason collectors prefer seeking for it in the evening, or with a lantern by night.
The most instructive case, however, is that ofDeilephila Hippophaës, in which no change of colour is associated with age, the caterpillar,throughout its whole life, remaining of a greyish green, which exactly matches the colour of the leaves of its food-plant,Hippophae Rhamnoides. Nevertheless this species also possesses the habit of feeding only at night as soon as it has attained to a considerable size, hiding itself by day at the root of its food-plant. Collectors expressly state that this larva can scarcely be found by day, and recommend that it should be sought for at night with a lantern.
From the foregoing facts and considerations it may fairly be concluded, that the habit of hiding by day, possessed by these and other allied caterpillars, was acquired when they resembled the leaves in colour, and that the adaptation to the colour of the soil, or dead foliage and withered branches, ensued as a secondary consequence.
But why have these caterpillars acquired such a habit, since they appear to be perfectly protected by their resemblance in colour to the green leaves? The answer to this question is easily given when we consider in which species this habit generally occurs.
Does the habit prevail only among the species of the one genusDeilephila, and in all the species of this genus? This is by no means the case, since, on the one hand, many species ofDeilephila, such asD. Euphorbiæ,Galii,Nicæa, andDahlii, do not possess the habit, and, on the other hand, it occurs in species of other genera, such asMacroglossaStellatarum,Sphinx Convolvuli, andAcherontia Atropos.
The habit in question must therefore be the result of certain external conditions of life common to all those species which rest by day. The mode of life common to them all is that they do not live on trees with large leaves or with thick foliage, but on low plants or small-leaved shrubs, such as the SeaBuckthorn.136I believe I do not err when I attribute the habit possessed by the adult larvæ, of concealing themselves by day, to the fact that the green colour is protective only so long as they are small—or, more precisely speaking, as long as their size does not considerably exceed that of a leaf or twig of their food-plant. When they become considerably larger, they must become conspicuous in spite of their adaptive colour, so that it would then be advantageous for them to conceal themselves by day, and to feed only by night. This habit they have acquired, and still observe, even when the secondary adaptation to the colour of the soil, &c., has not been brought about. We learn this fromD. Hippophaës, whichremains green throughout its whole larval existence; and no less from the green forms of the adult larvæ ofSphinx Convolvuli,Chærocampa Elpenor, andPorcellus, all of which conceal themselves by day in the same manner as their brown allies.
It may be objected that there are Sphinx-larvæ—instances of which I have myself adduced—which live on low small-leaved plants, and which nevertheless do not hide themselves by day. This is the case with the spurge-feedingD. Euphorbiæ, so common in many parts of Germany. This caterpillar must, however, be classed with those which, on account of their distastefulness, or for other reasons to be subsequently considered, are rejected by birds and other larger foes, and which, as Wallace has shown, derive advantage from being coloured as vividly as possible. I shall return to this subject later, when treating of the biological value of special markings.
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 theSphingidædoes not therefore depend upon acontemporaneousdouble adaptation, but upon the replacement of an old protective colour by a new and better one, and therefore upon asuccessivedouble adaptation. The adult caterpillars ofC. Elpenorare 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 anotherplace,137I 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 connectionbetween this phenomenon and simple variability.
InD. Hippophaësthe 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.
InMacroglossa Stellatarumwe 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-mentionedcase, 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 withSphinx Convolvuli, and almost to the same extent withChærocampa Elpenor, in both of which species the green caterpillars are therarest.138Forms 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 inChærocampa PorcellusandCelerioas well as inPterogon Œnotheræ. In all these species the green formoccurs,139but 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 Livornicaappears to have reached a similar stage, but the caterpillar of this species has been so imperfectly observed, that it isdifficult 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.).
InDeilephila Vespertilio,Euphorbiæ,Dahlii,Mauritanica,Nicæa, andGalii, 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 ofnatural 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 livingforms.140