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Chapter VIII: Letter XXX: States of Insects (3)

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Of the pilose larvæ, some, like most of those of the smaller moths (_Geometra_, _Tortrix_, _Pyralis_, &c.), have merely a few scattered short hairs, scarcely perceptible except through a lens: others (_Odenesis potatoria_, _Lasiocampa Rubi_) are covered with down more or less thick: in others (_Eriogaster lanestris_, _Lasiocampa Neustria_) the hair is slenderer, and more like wool; the body of two species which I purchased from the collection of Mr. Francillon is covered with woolly hairs, so long as to give them the appearance of a shock-dog; and Madam Merian has figured a similar one, which she could not bring to the perfect state[417]. The hairs of many _Bombycidæ_, known commonly by the name of _hairy caterpillars_, as _Arctia erminea_, &c. are stiffer, resembling bristles; sometimes, as in _Arctia chrysorhea_, mixed with shorter ones. The hairs either spring immediately from the skin (_Noctua Aceris_, _leporina_), or, as is more general, proceed only from certain tubercular elevations, usually subhemispherical, but sometimes conical; of which a number varying from four to twelve are found on each segment of different species. They seem to issue from these tubercles, as little diverging streams from the rose of a watering-pot. In both cases, they form a coating usually so dense as to conceal the body, but sometimes more thinly set, and admitting the skin to be seen more or less between them. In a caterpillar of the beautiful _Arctia ocularia_, the hairs are set upon tubercles alternately nearer the anterior and posterior margin of each segment, so as to form a dense band, the rest being naked; and in the lovely green and black one of _Saturnia Pavonia_, each tubercle bears but six hairs, diverging like a star, the central one being the longest and capitate, so that the chief part of the body appears naked. This diverging position of the hairs is most common in the thick-clothed larvæ also, but many have them placed differently: thus, in those of _Callimorpha Caja_ and _Arctia villica_[418] they are all directed towards the tail, like the quills of a porcupine: in some others the anterior ones point towards the head: in that of _Eriogaster Quercus_ half of the tuft of hairs of each tubercle is directed downwards, the other half upwards: in that of _Arctia Salicis_ all the hairs point downwards, so that the belly is thickly covered, while the back is bare. Another variation is, that the hairs of half the tubercle are sometimes very long, while those of the other half are very short, and even of a different colour[419]. In the larva of Tussuck moths (_Laria pudibunda_, _fascelina_, &c.) the hairs are collected into tufts of a singular appearance, those on the intermediate segments of the back being quite level at the top, so as to resemble so many brushes; while those on the first and last segments are longer, and composed of feathered hairs converging to a point at their extremity, like a common camel-hair-pencil[420]. This last mode of arrangement prevails also in the larva of _Noctua Aceris_; but in this the pencils are shorter, exactly wedge-shaped, and distinguished by another particularity, that of springing directly from the skin, and not from a tubercle. This is also the case with the large caterpillars of _Odenesis potatoria_, which has a double row of short bundles of black hairs on the back, intermixed with larger ones: at each end of the body is a pencil of converging hairs, and the sides are spotted with bundles of white ones, which with longer tawny ones are bent downwards, so as to cover the sides of the creature[421]. Some have the anterior aigrettes disposed like the arms of a cross, of which the body of the caterpillar is the stem[422]. But not only is there considerable variety in the general arrangement of the hairs that clothe our little larvæ, the hairs themselves differ much in their kind and structure, of which I will now, before I proceed to consider spines, give you some account. Several of them are feathered like the plumes of a bird: this is the case with those of _Morpho Idomeneus_, on each segment of the body of which are three blue tubercles, like so many little turquois beads, from each of which proceeds a long black plume[423]. Other hairs terminate in a club; those of the larva of _Noctua Alni_, a specimen of which I possess taken in England, are flat and incrassated at the apex, something like the antennæ of some _Sphingidæ_. Mad. Merian has figured the caterpillar of another moth which feeds upon the Papaw-tree (_Carica Papaya_) with similar hairs[424]. But the most remarkable larva for the shape of its hairs is that of _Anthrenus Musæorum_, the little pest of our cabinets, which I noticed in a former letter[425]. All the hairs of its body are rough with minute points; but those of six diverging long tufts or aigrettes, laid obliquely on the anal extremity of the body, which the animal when alarmed erects as a porcupine does its quills, are of a most singular structure: every hair is composed of a series of little conical pieces, placed end to end, the point of which is directed towards the origin of each hair, which is terminated at the other extremity by a long and large conical mass, resembling somewhat the head of a pike[426].

Besides the one lately mentioned, other caterpillars are rendered striking by the brilliant colour of the tubercles from which their hairs emerge. A remarkable instance of this is the thick large caterpillar of a Bombyx, which feeds upon the _Psidium pyriferum_, or white Guava, figured by Madame Merian. This caterpillar, which is white, with transverse black stripes, and which has two singular long converging curved bunches of hairs near the tail, is splendidly adorned on each side with fifty red tubercles, shining like coral, from which proceed six or seven long diverging hairs. Leeuwenhoeck took these tubercles for eyes[427]. Another figured by the same lady, who mistakes it, with her usual inaccuracy, for the larva of a _Lygæus_ F., and which seems by her description to be between the _onisciform_ and _limaciform_ types, has the apparently fleshy mamillæ that project from its sides and back crowned with little hairy red globes, which give the animal a most singular and unique appearance[428]. Having thus described some of the principal modes in which the All-wise Creator has decked and defended these creatures with _hairs_, I shall next give you a short account of the _spines_ with which he has armed others. The spinous larvæ are principally _lepidopterous_, and more particularly conspicuous in some tribes of the genus _Papilio_ L., though some saw-flies and _Diptera_ are also distinguished by them. _Vanessa Io_[429], _Atalanta_ and _Urtica_, _Argynnis Paphia_, _Urania Leilus_, and many other Butterflies, &c. are clothed with long sharp points, which claim the denomination of spines, rather than that of hairs or bristles; being horny and hard, and so stiff at the point as readily to pierce the skin. Those of the last-mentioned species, Madame Merian says, are as stiff as iron-wire[430]. They are sometimes entirely _simple_, and look like spikes rather than spines, as in the caterpillar of _Nymphalis Amphinome_ and _Morpho Menelaus_[431]; but ordinarily they are beset with _hairs_, or more commonly with shorter spines, which often give them the appearance of plumes, as in _Urania Leilus_ just mentioned: sometimes these lateral spines are so long as to have the appearance of a branch of a tree; this is strikingly the case with a small caterpillar which Captain Hancock brought from Brazil; its body is so thickly planted with spines of this description, that it absolutely wears the appearance of a forest or thicket in miniature. A singular circumstance attends the spines of this species: in many cases a smaller and very slender hair-like spine issues from them, resembling a sting; and this accounts for an observation of Abbott's, that many American caterpillars _sting_ like a nettle, raising little white blisters on the skin when accidentally or slightly touched[432]. Lewin has described the caterpillar of a moth found in New Holland, which he names _Bombyx vulnerans_, that, like these Americans, has also the power of wounding, but in a different way. It darts out, he says, when alarmed by the approach of any thing, from as many knobs or protuberances in its back eight bunches of little stings, with which it inflicts a very painful and venomous wound[433]. The caterpillar of _Papilio Protesilaus_ F., if Madame Merian's account and figure of it are correct, has its body armed with hairy spines, the extreme point of which is surmounted by a star-shaped appendage[434]. Those of a few saw-flies (_Tenthredo Pruni_ L.), and another figured by Reaumur[435], are covered with a little forest of spines without lateral branches, but divided into a fork at the apex. Some spines are merely rough, with very short points, as those round the head, which give so terrific an appearance to the caterpillar of the _Bombyx regalis_, of some proceedings of which I gave you an account in one of my former letters[436].

I must now say something upon the _arrangement_ of these spines. Though in a few instances so thickly set as entirely to conceal the body of the animal, as in the case of the Brazil one lately mentioned, yet generally speaking, even when they are most numerous, they permit the skin to be distinctly seen. Their arrangement is various, though always orderly: in the majority they are planted singly, but in some caterpillars in bundles. In that of _Saturnia Io_, on each segment there are six bundles of longish, quill-shaped, sharp, slender, diverging spines, which also appear to sheath aculei. Madame Merian has figured this larva, or one very near it, as the grub of a _Euglossa_[437], with which, though she affirms she traced it to the fly, it can have no connection. With regard to _number_, some larvæ have only four spines on each segment; others five, others again six, and others seven, or even eight: they are planted on the sides and back only, never on the belly. They are often more numerous on the intermediate than on the anterior and posterior segments; but sometimes the reverse of this takes place; in that of _Attacus Erythrinæ_ only the head and tail are armed with spines, the rest of the body being without any[438]; and in that of _Morpho Teucer_ there is only a single spine on the four intermediate segments[439]. They are usually all nearly of equal length; but in some cases those of the head and tail are much longer than the rest, and remarkably so in the caterpillar of _Urania Leilus_, also beautifully plumose, and gracefully waved[440]. Those in the second and third segments are much longer than any of the rest in that of _Bombyx regalis_; which circumstance gives it the terrific appearance lately alluded to. In the family to which _Argynnis Paphia_ belongs, the larva is adorned with two on the back of the first segment twice as long as the rest, and resembling at first sight two antennæ.

The spines, as well as the hairs of the new skin, are concealed under the old one, and not incased in its spines; but Bonnet ascertained, that if cut off very closely, the larva sometimes died in consequence, whilst no such result followed a similar operation on hairy larvæ. We learn from Reaumur[441], that some spinous larvæ of saw-flies (_Tenthredo_ L.) lose their spines at the last change of their skin; and from Madame Merian, that that of _Attacus Erythrinæ_ before mentioned loses also at the same period the six tremendous black spikes that arm its black and yellow larvæ. The grubs of ants that are destined to pass the winter in the larva state are hairy, but are not so in summer[442]. The spines found in the grubs of some gad-flies (_Œstrus_ L.) are of a different kind from those above described, being very minute triangular flat plates, arranged in different and contrary directions[443], and serving the insect merely to change its place and fix itself[444].

Two other kinds of clothing, if so they may be called, neither coming under the description of _hairs_ nor _spines_, are found in some other larvæ, not only amongst the _Lepidoptera_, but also in some of the other orders. _Nymphalis Populi_ and others of the same family have larvæ furnished on the back of each segment with cylindricoconical processes of a fleshy substance, obtuse at the apex and surrounded with capitate hairs. In that of _N. Sybilla_, which has on each segment two fleshy protuberances, they are bifurcate or trifurcate, and also encircled at the base with a hairy tuft[445]. Others, as those of _Melitæa Artemis_, _Cynthia_, &c. have each segment beset on the back with from seven to nine fleshy, pubescent, wedge-shaped protuberances; two larger ones projecting over the head. Under this head, too, may be noticed, the glutinous secretion which clothes the grub of _Cionus Scrophulariæ_, a little weevil; and of _Tenthredo Cerasi_ L. a saw-fly, and that waxy or powdery substance which transpires through the skin of the larvæ of several _Aphides_, _Chermes_, _Cocci_, _Hylotoma ovata_ F., &c. The _Aphis_, whose extensive ravages of our apple-trees (_A. lanata_) were before described to you[446], is covered and quite concealed by this kind of substance, so that the crevices in the bark which they inhabit look as if they were filled, not with animals, but with cotton. The insect, also, that forms those curious galls produced upon the spruce fir, and which imitate its cones (_Chermes Abietis_ L., _Aphis_ De Geer) secretes a similar substance. In these and other cases of the same kind, this matter seems to be, if I may so speak, wire-drawn through numerous pores in certain oval plates in the skin, more depressed than the rest of the back, arranged regularly upon the segments, and exhibiting minute tuberosities. When young, these animals have more of this secretion than when more advanced: it then hangs from their anal extremity in locks[447].

But the insects most remarkable for a covering of this nature are those _Coccidæ_ of which Bosc has made a genus under the name of _Dorthesia_. De Geer is the first author that notices them, and has given a description and figure of one species under the name of _Coccus floccosus_[448]. It was discovered by Modeer upon some sere fir-leaves in a thick bed of moss. Panzer has figured a second found upon _Geranium sanguineum_, which from the figure appears distinct from De Geer's, under the name of _Coccus dubius_[449]. Fabricius regards this as synonymous with the _Dorthesia characias_ of Bosc, inhabiting _Euphorbia characias_ in South Europe[450]. Olivier found a species upon the bramble[451]. I once took one, which appears to differ in some respects from the preceding species, upon _Melampyrum cristatum_, and our indefatigable friend Mr. Sheppard has sent me another, on what plant found I do not remember, which does not agree with any that I have mentioned. The body of the animals of this genus is covered by a number of cottony or waxy laminæ which partly cover each other, and are arranged usually in a triple series: in De Geer's figure the series appears quadruple, the lateral ones being placed obliquely. The anterior one in my specimen covered the head, and they are all canaliculate. Above the anus are four diverging ones: the whole are of the most dazzling whiteness. When these laminæ are removed, the body appears divided into segments.

With respect to those larvæ which imitate slugs by the viscid covering that besmears them and issues from their pores, we learn from Professor Peck that this exudation takes place as soon as they are hatched; that the animal retains its humidity although exposed to the fiercest heat of the sun, and that at the last moult the skin becomes quite clean, and free from all viscidity[452]. It is probable that the other limaciform larvæ are similarly circumstanced. Madame Merian has figured an _onisciform_ one, the legs of which, she says, are covered with a viscid skin: this produced a _Noctua_. Those of _Papilio Anchises_ also are slimy, and adhere to each other[453].

v. Amongst other qualities which attach to larvæ, we must not omit to say something concerning their _Colour_. For though those which live in darkness, in the earth, in wood, in fruits, &c. are, with few exceptions[454], of an uniform whitish colour, yet such as are exposed to the influence of the light are usually adorned with a vast variety of tints, sometimes the most vivid that can be imagined. That the white colour of the former may be attributed to the absence of light is proved by an experiment of M. Dorthes, who having forced some to live under glasses, exposed to the light, found that they gradually became brown[455]. To attempt any classification of coloured larvæ would be in vain, since they are tinged with almost every possible shade that can be conceived, of many of which it would be difficult to find examples elsewhere; and infinitely diversified as to the arrangement and figure of their multiform markings and spots. A few general remarks, therefore, are all that you will expect on this head. Many are of one uniform colour; while a variety of tints, very different, and very vivid and distinct, ornament others. Sometimes they are distributed in longitudinal rays or bands, at others in transverse ones. Sometimes they are waved or spotted, regularly or irregularly; at others they are sprinkled in dots, or minute streaks, in every possible way. Various larvæ are of the colour of the plant on which they feed, whence they are with difficulty discovered by their enemies. Thus, a large proportion of _Lepidoptera_ are green of different shades, sometimes beautifully contrasted with black bands; a circumstance which renders the caterpillars of two of our finest insects of this order as lovely as the fly: I mean that of _Papilio Machaon_ and _Saturnia Pavonia_. Very frequently the larvæ of quite different species resemble each other so exactly, in colour as well as shape, as scarcely to be distinguishable: this sometimes takes place even where they belong to different genera, as in those of _Bombyx versicolor_ a moth, and _Smerinthus Populi_ a hawk-moth. And it sometimes happens, very fortunately for distinguishing allied species, that where the perfect insects very nearly resemble each other, the larvæ are altogether dissimilar. Thus, the female of _Pieris Rapæ_ is so much like the same sex of _Pieris Brassicæ_, that it might be taken for a variety of it, did not the green caterpillar of the one, and the spotted one of the other, evince the complete distinction of these butterflies. _Noctua Lactuca_, _N. umbratica_, and several other species of the same tribe, which includes _N. Absinthii_, _Verbasci_, _Chamomillæ_, _Abrotani_, are so extremely alike, that the most practised eye can scarcely discover a shade of difference between them, though their larvæ in colour and markings are constantly distinct[456]. The markings of species belonging to the same family are usually different; but in some cases the latter may be prejudged from the former. The larvæ of many of the genus _Sphinx_ L., for example, have their sides marked by oblique streaks running from the back in a direction towards the head; and by this last circumstance they are distinguished from those of _Bombyx versicolor_, _Attacus Tau_, and others of the same tribe, which have also lateral oblique striæ, but running from the back towards the tail[457]. The colours of individual larvæ of the same species are usually alike, but in _Sphinx Elpenor_ and some others they vary exceedingly. Many, like those of _Lasiocampa Rubi_, _Saturnia minor_, &c., are of one colour when first disclosed, and assume others quite different in riper age. Just previously to _changing their skin_, the tints of most larvæ become as dull and obscure, as they are fresh and vivid when the change has fully taken place; and in some instances the new skin is quite differently marked from the old one. This is remarkably the case with the last skin of some of the larvæ of the genus _Tenthredo_ L., which is entirely different from all the preceding ones. As people when they advance far in years usually become more simple in their dress than when they were young, so the larvæ in question change an agreeably variegated skin for one of a uniform and less brilliant colour[458]. Madame Merian has observed with respect to _Attacus Erythrinæ_, that its caterpillar is at first yellowish, with nine black striæ on each side: when arrived at one third of its size, they become orange; the striæ are obliterated, and in their place a round black spot appears on each of the eight intermediate segments[459]. Mr. Sheppard has remarked to me, that the skin of that of _Sphinx Ligustri_, after being under ground four days, was changed from a vivid green to a dull red. Very rarely, however, it becomes of a more brilliant hue just before entering the pupa state: thus, that of another hawk-moth (_Smerinthus Tiliæ_) changes to a bright violet; and the yellow hairs of that of _Laria pudibunda_ then become of a lovely rose colour. And here I may observe, that the hairs and spines also, of larvæ, vary greatly in colour. They are to be met with brown, black, red, yellow, violet, white, &c. De Geer found, that in the larva of _Cimbex nitens_ the two sides of the body were of a different colour, the left being of a deep green, whilst the right side and the rest of the body were paler[460]; but as he saw only a single individual, this was probably an accidental circumstance. Though the caterpillars, as I lately said, of one of the most beautiful butterflies and moths that inhabit Britain contend with the perfect insect in loveliness, yet in general no judgement can be formed of the beauty of the future fly from the colour of the larva; and the young Aurelian must not flatter himself always with the hope, because the caterpillar excites admiration by its colours and their arrangement, that the butterfly or moth it is to produce will do the same; nor ought he to despise and overlook a sombre or plain-coloured individual of the former, under the idea that it will produce one equally plain of the latter, for it often happens that the splendid caterpillar gives a plain butterfly or moth, and _vice versâ_. De Geer, however, gives us two instances of conformity between the colours of the caterpillar and those of the future moth; the one is that of the common currant-moth (_Phalæna G. grossulariata_ L.), the caterpillar of which is white, ornamented with several black spots varying in size. At the two extremities it is yellowish, with a longitudinal ray of the same colour on each side, the head and legs being black. These colours are all to be found in the fly, the ground of its wings being white ornamented with many black spots of different sizes. Its upper wings are traversed by a yellowish band; and towards their base is a spot of the same colour. Its body is yellowish, with black spots; but the head and legs are black[461]. The other is that of a green caterpillar, which gives a green moth, figured by Reaumur (_Pyralis prasinaria_ Fab.)[462]. Sometimes, also, the sex of the future perfect insect may be predicted from the colour it exhibits in its first state: thus, the brown caterpillars of _Noctua Pronuba_ produce males, and the green ones females[463]. The sexes, also, of _N. exoleta_ and _Persicariæ_ differ in that state.

vi. To the full account of the _Food_ of insects given in a former letter[464], which had reference chiefly to their larva state, it is only necessary in this place to add a few particulars not there noticed. Many larvæ when first excluded, as those of _Pieris Cratægi_, &c. devour the shells of the eggs from which they have proceeded[465]; and others (_Cerura Vinula_, _Sphinx Euphorbiæ_, _Noctua Verbasci_), though their usual food is of a vegetable nature, eat with great apparent satisfaction the skins which they cast from time to time, not leaving even the horny legs. This strange repast seems even a stimulating dainty, which speedily restores them to vigour, after the painful operation by which they are supplied with it. Under this head it will not be out of place to mention, that some larvæ of insects, which feed only on the juices of animals, or the nectar and ambrosia of flowers, have no anal passage, and of course no feces. This is said to be the case with the grubs of bees, wasps, the larvæ of _Myrmeleon_, &c.[466]

vii. You will require no stimulus to induce you to attend to the subject I am next going to enter upon,--the _Moulting_, namely, of Larvæ; or their changes of skin. This, indeed, is a subject so replete with interest, and which so fully displays the power, wisdom, and goodness of the Creator, affording at the same time such large occasion for nice investigation, that a pious and inquisitive mind like yours cannot but be taken with it. In the higher orders of animals, though the _hairs_ of quadrupeds and the _feathers_ of birds are in many cases annually renewed, the change, or scaling and increment of the _skin_, is gradual and imperceptible; no simultaneous rejection of it, in which it is stripped off by the animal itself like a worn shirt, being observable, till you descend in the scale to the Serpent tribe[467], which at certain periods disengage themselves from their old integument, and start forth with that new and deadly beauty so finely described by the Mantuan bard:--

"So from his den, the winter slept away,
Shoots forth the burnished snake in open day;
Who, fed with every poison of the plain,
Sheds his old spoils and shines in youth again:
Proud of his golden scales rolls tow'ring on,
And darts his forky _tongue_[468], and glitters in the sun."
PITT.

In these the new skin, I imagine, is formed under the old from the _rete mucosum_; but in _insects_, as I formerly stated[469], since the time of Swammerdam it has generally been believed by entomologists, that the larva includes a series of cases or envelopes, one within the other, containing in the centre the germe of the future perfect insect, whose development and final exclusion take place only when these cases have been successively cast off. This hypothesis, as was explained to you on a former occasion[470], has been controverted by a late writer, Dr. Herold; who affirms that the skins of caterpillars are also successively produced out of the _rete mucosum_. I have however, I hope, satisfied you that the old system is most consonant to nature and probability: but as we are now to enter at large upon the _Moults_ of insects, it will not be without use if I add a few additional reasons which seem to me still further to prove the correctness of Swammerdam's system, as far as it relates to that subject. With regard to the mere formation of the skin from the _rete mucosum_, were this the whole question few would hesitate to adopt the sentiments of M. Herold; but when we come to consider further--that the number of moults of individuals of the _same_ species is always the same, and that it varies in _different_ species, and takes place at certain periods,--we begin to suspect that something more than the mere formation of a new skin upon an old one being cast is to be accounted for; and that the law which prescribes its own definite number of skins to each species, must begin to act in the primordial formation of the larva. Again, the _hairs_ observable in the higher animals do not take their origin from the epidermis solely, but are planted below it in the _rete mucosum_, or deeper[471]; so that the change of skin does not affect them; but in the larvæ of insects they are a continuation of that integument, since, when the moult takes place, they always remain on the rejected skin[472]: this is the case, also, even with spines. If you shave a caterpillar ready to change its skin, either partially or generally, you will find that the parts in the new skin that correspond with those that are denuded, are equally hairy with those that were not[473]; and if you pay attention to the new-clad animal, you will find further, that the hairs never grow after a moult. From hence it follows, that the _hairs_ have their place and take their whole growth between the new skin and the old[474]. Whether the _spines_, simple or compound, lately described to you, that arm some larvæ are similarly circumstanced, seems not as yet to have been ascertained; but as the spinous ones of certain _Tenthredines_ L. and _Lepidoptera_ at their last moult have no spines, the presumption is, that, whether incased or not, they are mere appendages of the skin on which they appear. A new set of hairs, therefore, and probably of spines in spinous larvæ, accompanying each skin, and these varying very much in size, composition, &c. though a new membrane may be admitted to be formed from an action in the _rete mucosum_ without a pre-existent germe of it, it seems not easy to conceive how these hairs and spines can spring up and grow there, each according to a certain law, without existing previously as a kind of _corculum_ or _punctum saliens_; and that the germes of the tubercles, in which the hairs are so generally planted, according to a certain arrangement and in a given number, should also pre-exist, seems most consonant to reason. These and the several skins may all co-exist in their primordial germes, and remain beyond the discovery of our highest powers of assisted vision, till a certain period when they may first enter the range of the microscope-aided eye. It does not therefore follow, because these _primordia semina rerum_ are not discoverable, that therefore they may not exist. Our faculties and organs are too limited and of too little power to enable us to see the essences of being.

Upon the supposition that the hypothesis of Swammerdam is the true one, we may imagine that the envelope that lies within all the rest is that which covers the insect in its pupa state. Above this are placed several others, which successively become external integuments. These changes or casting of the skin in larvæ, analogous, as before observed, to that of serpents, are familiar to every breeder of silk-worms, in which _four_ such changes occur: the first at the end of about _twelve_ days from its birth, and the three next each at the end of _half_ that time from the moulting which preceded it. With some exceptions[475], similar changes of the skin take place in all larvæ, not however in the same number and at the same periods. Most indeed undergo this operation only three or four times; but there are some that moult oftener, from five up to eight (_Arctia villica_), nine (_Callimorpha Dominula_), or even ten times; for so often, M. Cuvier informs us, the caterpillar of the tiger-moth (_Callimorpha Caja_) casts its exuviæ. It has been observed that the caterpillars of the day-flying _Lepidoptera_ (_Papilio_ L.) usually change only _three_ times, while those of the night-flying ones (_Phalæna_ L.) change _four_[476]. The periods that intervene between each change depend upon the length of the insect's existence in the larva state. In those which live only a few weeks or months, they are from eight to twenty days; while in those that live more than a year, as the cockchafer, &c. they are probably proportionably longer: though we know very little with regard to the moult of any insects besides the _Lepidoptera_.

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.

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An Introduction to Entomology: Vol. 3Chapter VIII: Letter XXX: States of Insects (3)

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