Part 15

A day or two previously to each change of its skin, the larva ceases eating altogether; it becomes languid and feeble, its beautiful colours fade, and it seeks for a retreat in which it can undergo this important and sometimes dangerous and even fatal operation in security. Here, either fixing itself by its legs to the surface on which it rests, or, as is the case with many caterpillars, by its prolegs, to a slight web spun for this purpose, it turns and twists its body in various directions, and alternately swells and contracts its different segments. The object of these motions and contortions seems to be, to separate the exterior skin, now become dry and rigid, from the new one just below it. After continuing these operations for some hours, resting at intervals without motion, as if exhausted by their violence, the critical moment arrives: the skin splits in the back, in consequence of the still more violent swelling of the second or third segment: the opening thus made is speedily increased by a succession of swellings and contractions of the remaining segments: even the head itself often divides into three triangular pieces, and the inclosed larva by degrees withdraws itself wholly from its old skin. All larvæ, however, do not force their way through this skin in precisely the same place. Thus, those of the hawthorn butterfly (_Pieris Cratægi_), according to Bonnet[477] make their way out by forcing off what may be called their skull, or the horny part of their head, without splitting the skin, which remains entire; others have been observed to make their way out at the side and the belly. Reaumur noticed the larva of _Zygæna Filipendulæ_, previously to its last moult, actually biting off and detaching several portions of its old skin; and before this, drops of a fluid resembling water were seen to exude from it[478].

The skin when cast is often so entire, that it might be mistaken for the larva itself; comprising not only the covering of the main trunk with the hairs which clothed it, but of the very skull, eyes, antennæ, palpi, jaws, and legs; which, if examined from within, are now found to be hollow, and to have incased, like so many sheaths, similar parts in the new skin. That the feet of the newly-coated larva were actually sheathed, as fingers in a glove, in the same parts of the exuviæ, may be proved by a very simple experiment: if a leg of one just ready to cast its skin be cut off, the same limb will be found mutilated when that change has ensued. The anal horns, also, of the larvæ of the hawk-moth (_Sphinx_ L.) and other similar protuberances, are incased in each other in like manner; but hairs are laid flat between the two skins, and contribute considerably towards their more easy separation. Thus, if you saved the skins cast by the larva of _Callimorpha Caja_, for instance, you would appear to have ten different specimens of caterpillars, furnished with every external necessary part, and differing only in size, and the colour perhaps of the hairs, and all representing the same individual.

But further changes than this take place. Swammerdam says, speaking of the moult of the grub of _Oryctes nasicornis_, a beetle common in Holland, but not satisfactorily ascertained to inhabit Britain, "Nothing in all nature is in my opinion a more wonderful sight than the change of skin in these and other the like worms. This matter, therefore, deserves the greatest consideration, and is worthy to be called a specimen of nature's miracles; for it is not the external skin only that these worms cast, like serpents, but the throat and a part of the stomach, and even the inward surface of the great gut, change their skin at the same time. But this is not the whole of these wonders; for at the same time some hundreds of pulmonary pipes within the body of the worm cast also each its delicate and tender skin. These several skins are afterwards collected into eighteen thicker, and, as it were, compounded ropes, nine on each side of the body, which, when the skin is cast, slip gently and by degrees from within the body through the eighteen apertures or orifices of the pulmonary tubes before described, having their tops or ends directed upwards towards the head. Two other branches of the pulmonary pipes that are smaller, and have no points of respiration, cast a skin likewise." ... "If any one separates the cast little ropes or congeries of the pulmonary pipes with a fine needle, he will very distinctly see the branches and ramifications of these several pipes, and also their annular composition[479]."--Bonnet makes a similar observation with regard to _caterpillars_; but he appears to have observed it more particularly, at least the change of the intestines, previously to the metamorphosis of the insect, when he says with the excrements it casts the inner skin of the stomach and viscera[480]. Both these great men appear to have recorded the result of their own actual observations with regard to the proceedings of two very different kinds of insects; the one the grub of a beetle, and the other the caterpillars of _Lepidoptera_. The account of the former is given quite in detail, as that of a person who is describing what he has actually _seen_: yet by a later and very able physiologist, Dr. Herold, it is affirmed that the inner skin of the intestinal canal is never cast, that canal constantly retaining its two skins. He further affirms, that they are only the _large_ trunks of the Tracheæ that cast their skins, none being detached from their _smaller_ ramifications[481]. When men so eminent for their anatomical skill and nicety, and for their depth and acumen, disagree, the question must be regarded as undecided till further observations throw sufficient weight into one scale or the other.

The larva which has undergone this painful process is at first extremely weak: all its parts are soft and tender; even the corneous ones, as the head and the legs, are then scarcely more than membranous, and are all bathed with a fluid, which, before the moult, intervenes between the two skins, and facilitates their separation[482]: and it is only after some _hours_, or in some cases even _days_, during which it lies without motion, that this humidity evaporates, all its parts become consolidated, and it recovers its strength sufficiently to betake itself to its wonted food. Its colour, too, is usually at first much paler than before, and its markings indistinct, until their tints have been enlivened by exposure to the air, when they become more fresh, vivid, and beautiful to appearance than ever. When a few meals have invigorated its languid powers, the renovated animal makes up for its long abstinence by eating with double voracity. . A similar preparatory fast, and succeeding state of debility, accompany every change of the larva's skin. Each time except the last, the old skin is succeeded by a new one, with few exceptions, similar to the one it has discarded. Previously to the final change, which discloses the pupa, it quits the plant or tree on which it had lived, and appears to be quite unsettled, wandering about and crossing the paths and roads, as if in quest of some new dwelling. It now abstains from food for a longer time than before a common moult, empties itself copiously, and as I have just said, if Swammerdam and Bonnet are to be depended upon, casts the skin that lines the stomach and intestines, as well as that of the Tracheæ.

I have observed above, that all larvæ, with few exceptions, change their skins in the manner that I have described. These exceptions are principally found in the order _Diptera_, of which those of the Linnean genera _Musca_, _Œstrus_, and probably all that, like the maggot of the common flesh-fly, have membranous contractile heads, never change their skin at all, not even preparatory to their becoming pupæ. The skin of the pupa, though often differing greatly in shape from that of the larva, is the same which has covered this last from its birth, only modified in figure by the internal changes that have taken place, and to which its membranous texture readily accommodates itself. The larvæ of the Dipterous genera _Tipula_, _Culex_, and those which have corneous heads, like other larvæ change their skins several times previously to becoming pupæ[483]. The grubs, also, of bees, wasps, ants? and probably many other _Hymenoptera_, do not change their skin till they assume the pupa, nor the larva of the female _Coccus_[484].

If you feel disposed to investigate the reasons of that law of the Creator which has ordained that the skins of the higher animals shall be daily, and imperceptibly, and as it were piece by piece renewed, while those of insects are cast periodically and simultaneously,--the proximate cause must be sought for in the nature of the two kinds of skin, the one being more pliable and admitting a greater degree of tension than the other, and being so constructed as to scale off more readily. If, ascending higher, you wish to know why the skins of insects are so differently circumstanced from our own, the most apparent reason is, to accommodate the skin to the very rapid growth of these animals, which a gradual and slower change would have impeded too much, or the skin have suffered constant dilapidation and injury; therefore their Beneficent Creator has furnished them with one which will stretch to a certain point, and during a certain period, and then yield to the efforts of the inclosed animal, and be thrown aside as a garment that no longer fits the wearer.

viii. And this leads me to a subject to which I am desirous now to bespeak your attention,--the _Growth_, I mean, and size of Insects in this state. As to _size_, larvæ differ as much as insects in their perfect state: these last, however, never grow after their exclusion from the pupa, while larvæ increase in bulk in a proportion, and often with a rapidity, almost without a parallel in the other tribes of animals. Thus Lyonnet found, that the caterpillar of the great goat-moth (_Cossus ligniperda_ F.) after having attained its full growth is at least 72,000 times heavier than when it was first excluded from the egg[485]; and of course had increased in size in the same proportion. Connected with the size of larvæ, is the mode in which their accretion takes place. This, with respect to the more solid parts, as the head, legs, &c., is not, as in other animals, by gradual and imperceptible degrees, but suddenly and at stated intervals. Between the assumption of a new skin and the deposition of an old one, no increase of size takes place in these parts, while the rest of the body grows and extends itself, till, becoming too big for these solid parts, nature restores the equilibrium between them by a fresh moult[486], in which the augmentation of bulk, especially in these parts, is so great, that we can scarcely credit the possibility of its being cased in so small an envelope. Malpighi declares, that the head of a silk-worm that has recently cast its skin is four times larger than before the change[487]. It is very probable, also, that when the outer skin becomes rigid, it confines the body of the larva within a smaller compass than it would expand to if unconfined, so that, when this compression is removed, the soft and elastic new integument immediately swells out, and the animal appears all at once much larger than it was before the moult. In fact, the proximate cause of the rupture and rejection of the old skin is the expansion of the included body, which at length becomes so distended as to split its envelope, aided, indeed, as before described, by the contortions of the creature itself.

The larvæ most notorious for the _rapidity_ of their growth are those of _Musca carnaria_ and other flesh-flies: some of which Redi found to become from 140 to more than 200 times heavier in twenty-four hours[488]: an increase of weight and size in so short a time truly prodigious, but essential for the end of their creation--the rapid removal of dead and putrescent animal matter. As the skins of these larvæ are never changed, we may conclude, if the cause of the change of skin in other larvæ above surmised be accurate, that their skins are more contractile and capable of a greater degree of tension than those of larvæ that are subject to moulting. And two peculiarities observable in them confirm this idea: in the first place, their head is not hard and corneous, as that of the others, but capable of being shortened or lengthened; and in the next, their breathing-pores are not in the sides, but at the extremities of the body, while in the _moulting_ larvæ there are two in almost every segment, which must form so many callous points that impede the stretching of the skin to the utmost. The hairs, spines, and tubercles, that are so often found on caterpillars, must also form so many points of resistance that prevent that full extension of the integument which it might otherwise admit.

There is not always that proportion between the size of larvæ and of the insects that proceed from them that might have been supposed, some small larvæ often producing perfect insects larger than some of those proceeding from such as are of greater size.

ix. As insects often live longest in the state we are treating of, I shall say something next upon the _age_ of larvæ, or the period intervening between their exclusion from the egg and their becoming pupæ. This is exceedingly various, but in every case nicely adapted to their several functions and modes of life. The grubs of the flesh-fly have attained their full growth, and are ready to become pupæ, in _six_ or _seven_ days; the caterpillar of _Argynnis Paphia_, a butterfly, in _fourteen_ days; the larvæ of bees in _twenty_ days; while those of the great goat-moth (_Cossus ligniperda_) and of the cockchafer (_Melolontha vulgaris_) live _three_ years, or at least survive three winters, before the same change. That of another lamellicorn beetle (_Oryctes nasicornis_ F.) is said to be extended to _four_ or _five_; that of the wire-worm (_Elater segetum_) to _five_. That of the stag-beetle (_Lucanus Cervus_) is affirmed by Rösel to be extended to _six_ years; but the most remarkable instance of insect longevity is recorded by Mr. Marsham in the _Linnean Transactions_[489]. A specimen of _Buprestis splendida_, a beautiful beetle never before found in this country, made its way out of a deal desk in an office in London in the beginning of the year 1810, which had been fixed there in the year 1788 or 1789; so that according to every appearance it had existed in this desk more than twenty years. Ample allowance being made for its life as a pupa, we may conclude that it had existed as a larva at least half the above period. The grubs of the species of the genus _Cynips_ L. attain their full size in a short time; but they afterwards remain five or six months in the gall before they become pupæ[490].

With few exceptions it may be laid down, that those larvæ which live on dead animals, in fungi, in dung, and in similar substances, are of the shortest duration in this state; and that those which live under the earth, on the roots of grass, &c. and in wood, the longest: the former becoming pupæ in a few days or weeks, the latter requiring several months, or even years, to bring them to maturity. The larvæ which live on the leaves of plants seem to attain a middle term between the one and the other,--seldom shorter than a few weeks, and rarely longer than seven or eight months. Aquatic larvæ appear to be subject to no general rule: some, as the larvæ of _Gnats_, becoming pupæ in two or three weeks; and others, as those of the _Ephemeræ_, which are thus compensated for their short life as flies, in as many years[491]. The cause of all these differences is obviously dependent on the nature of the food, and the purposes in the economy of creation to which the larvæ are destined.

x. The last part of the history of larvæ relates to their _Preparations for assuming the pupa state_. When they have acquired their full size, after having ceased to take food, by a copious evacuation they empty the intestinal canal, even rejecting the membrane that lines it and the stomach[492]; their colours either change totally, or fade; and they make themselves ready for entering upon a new stage of their existence. Some merely rest in a state of inactivity in the midst of the substances in which they feed, as if conscious of their inability to select any safer abode. Of this description are most Coleopterous, Hymenopterous, and Dipterous larvæ, that feed under ground, or in the interior of trees, fruits, and seeds.

But a still larger tribe, those which feed on leaves, animals, &c. act as if more sensible of the insecurity of this to them important epoch. They are about to exchange their state of vigour and activity for a long period of deathlike sleep. The vigilant caution which was wont to guard them from the attack of their enemies will be henceforward of no avail. Destitute of all the means of active defence, their only chance of safety during their often protracted night of torpor must arise from the privacy of their place of repose. About this, therefore, they exhibit the greatest anxiety. Many, after wandering about as if bewildered, retire to any small hole on the surface of the earth, covering themselves with dead leaves, moss, or the like, or to the chinks of trees, or niches in walls and other buildings, or similar hiding-places. Many penetrate to the depth of several inches under ground, and there form an appropriate cavern by pushing away the surrounding earth; to which they often give consistence by wetting it with a viscid fluid poured from the mouth. The larvæ of other insects undertake long and arduous journeys in search of appropriate places of shelter. Those of flesh-flies, now satiated with the mass of putridity in which they have wallowed, leave it, and conceal themselves in any adjoining heap of dust. The grubs of the gad-fly (_Œstrus_) creep some of them out of the backs of cattle, in tumours of which they have resided, and suffer themselves to fall to the earth; while others, which have fed in the stomach of horses, quit their hold, and by a still more extraordinary and perilous route are carried through the intestines the whole length of their numerous circumvolutions, and are discharged at the anus. And without enumerating other instances, various aquatic larvæ, as that of a common fly (_Elophilus pendulus_), &c. leave the water, now no longer their proper element, and betake themselves to the shore, there to undergo their metamorphosis.

Most of these, having reached their selected retreat, require no other precaution; but another large tribe of larvæ have recourse to further manœuvres for their defence before they assume the pupa. Those of the aphidivorous flies (_Syrphus_ F. &c.), of the various lady-birds (_Coccinella_ L.), and tortoise-beetles (_Cassida_ L.), &c. fix themselves by the anus with a gummy substance to the leaves or twigs under which they propose to conceal themselves during their existence in that state. Others previously suspend themselves by a silken thread fixed to the tail, or passing round the body; by which also, when become pupæ, they are afterwards pendent in a similar position; and lastly, a very great number of larvæ wholly inclose themselves in cases or cocoons, composed of silk and various other materials, by which during their state of repose they are protected both from their enemies and the action of the atmosphere. As these two last-mentioned processes are extremely curious and interesting, I shall not fear tiring you by entering into some further detail respecting them: explaining _first_ the mode by which larvæ _suspend_ themselves, both before and after they are become pupæ, by silken threads; and _next_, the various _cases_ or _cocoons_ in which others inclose themselves, and their manner of operating in the formation of them.

1. The larvæ which suspend themselves and their pupæ, with the exception of the tribe of _Alucitæ_, and some _Geometræ_ of the family of _G. pendularia_, _punctaria_, &c. are almost all _butterflies_[493]. No others follow this mode. They may be divided into two great classes--those which suspend themselves _perpendicularly_ by the _tail_, and those which suspend themselves _horizontally_ by means of a thread girthed round their _middle_. In both cases it should be observed, that the suspension of the pupa is the object in view; but as the process is the work of the larva, this seems the proper place for explaining it. To begin with the _first_ case.

You are aware that all _lepidopterous_ larvæ have the faculty of spinning silk threads from their mouths, and it will readily occur to you that it is by means of these threads that they suspend themselves. But how? How is a caterpillar to hang itself by the tail to threads spun from the mouth? Even suppose this difficulty overcome, others still greater remain. Suppose the caterpillar to be suspended by its tail,--this is but a preparatory operation,--what is required is, that the pupa shall hang in the same position: now when you take into consideration that it is incased _within_ the skin of the larva, and without feet or other external organs; that it has to extricate itself from this skin; to hang itself in its place, and to detach the skin from the threads which hold it--this will appear no trifling task. Indeed at first view it seems impossible. Country-fellows for a prize sometimes amuse the assembled inhabitants of a village by running races in sacks: take one of the most active and adroit of these, bind him hand and foot, suspend him by the bottom of his sack with his head downwards, to the branch of a lofty tree; make an opening in one side of the sack, and set him to extricate himself from it, to detach it from its hold, and suspend himself by his feet in its place. Though endowed with the suppleness of an Indian juggler, and promised his sack full of gold for a reward, you would set him an absolute impossibility: yet this is what our caterpillars, instructed by a beneficent Creator, easily perform. Their manœuvres I shall now endeavour to explain.