All these cocoons, however, must yield in point of singularity of construction, materials, and ingenuity, to one formed by a small caterpillar, described by the illustrious naturalist lately quoted, which feeds upon the oak. This cocoon is wholly composed of small rectangularstrap-shaped pieces of the fine upper skin, or epidermis of the twig upon which it rests, regularly fastened to each other in a longitudinal direction with very slender silken cords. But the mode of its construction is even more remarkable than the substance of which it is fabricated. The caterpillar's first process is to form its slips of bark into two flat triangular wing-like pieces, projecting opposite to each other from each side of the twig, somewhat like the feathers of an arrow. It does not, perhaps, require any great degree of intelligence in a larva to give its cocoon the usual oval form, when it begins to arrange its materials in that shape from the very first, and round so good a mould as its own bent body; but we surely must admit that it is a task to which no stupid artist would be competent, to form first a multitude of strap-shaped laminæ into two triangular plates, and then to bend these plates into a case resembling the longitudinal section of a cone, with an elliptical and protuberant base,—the figure which the cocoon of this insect assumes. All the minutiæ of the manœuvres which it employs in this nice operation could not be comprehended without a more diffuse explanation than I have here room to give: suffice it to say, that the caterpillar fastens silken lines to each exterior opposite and longer side of the laminæ, and by applying all the weight of its body forces them to bend and approach each other, in which position it secures them by other shorter lines. It next repeats the same process with the upper and shorter sides of the plates; which when joined form the base of the cocoon. Both these tasks are accomplished in less than an hour, and the seams are so nicely joined as to be imperceptible. A fine inner tapestry of silk, covering all the asperities of the exteriorwalls, concludes its labours[544]. It is to be lamented that Reaumur was unacquainted with the moth that proceeds from the pupæ inclosed in these ingenious cocoons; which being small, and precisely of the same colour as the bark of the twig that supports them, are not to be discovered but by a very narrow inspection. It would seem, however, to beNoctua Strigulaof Berkhausen,Pyralis strigulalisof Hubner[545]. The larva, he informs us, is found in May: its body is flatter than common, of a yellowish flesh-colour, clothed with tufts of red hair on each segment, and furnished with fourteen feet. Should this description enable you to detect it upon your oaks, a view of its ingenious procedures would amply repay you for the trouble of seeking for it. The larvæ ofCerura vinula,Stauropus Fagi, and several other moths, form their cocoons of grains of wood gnawed from the trees on which they feed. These grains they masticate, mixed with a glutinous fluid secreted from the mouth, into a paste, which forms a covering of an uniform smooth texture, and so hard as not readily to yield to a knife. Of a substance apparently nearly similar is composed the cocoon of a weevil related toLiparus Pini; which with its inhabitant was given me by the ingenious Mr. Bullock. A little moth, whose ravages have been before noticed[546], lines the interior of the grain of barley, of which it has devoured the contents, with silk; divides it into two apartments, into one of which it pushes the excrement it had voided, and in the other assumes the pupa[547].
These, and the other larvæ mentioned above, commonlyform their cocoons of the substances I have indicated; but when by any cause they are prevented from access to them, they often substitute such other materials as are at hand. Reaumur fed a larva that formed its cocoon of minute fragments of paper, which with its mandibles it had cut from the piece that covered the glass vessel that contained it[548]: and the same circumstance happened to Bonnet.
Upon a former occasion I described to you the cases of various kinds formed and inhabited by the insects of theTrichopteraOrder (PhryganeaL.) commonly called case-worms[549]. As these serve for the pupa as well as the larva, they may be regarded as a kind of cocoon. I shall not repeat here what I then said; but having purchased from the collection of the late Mr. Francillon some that seem to belong to this or some cognate tribe, that are of a curious construction, I shall give you some account of two or three of them in this place. The first is not quite three inches long, of a sublanceolate shape, but rather widest towards one end. It consists of an internal tough and thick bag or cocoon, of a silk resembling fine wool of a dirty white colour, which is closely covered transversely by pieces of the stalk of a plant, about three-fourths of an inch in length, and crossing each other at an obtuse angle. The next is thicker and shorter: the internal bag is just covered with small fragments of wood like sawdust; over these are fastened irregularly, short stout pieces of a pithy stick or stalk, and the whole is clothed with a very close-woven ash-coloured web. It seems difficult to conceive how the inclosed animal could contrive to cover her habitation withthis web without going wholly out of it. The third is the most curious and remarkable of all. It is nearly six inches long, and about four-fifths of an inch in diameter. It consists of a bag of thick cinereous silk web, to which are fastened, in a sextuple series, pieces of stick about an inch long, the end of one mostly resting upon the base of another: between each series a space of about three-tenths of an inch intervenes, but at the apex they all converge. This probably imitates the branch or stem of some tree or plant, in which the leaves are linear, and diverge but little from the stem. A label upon it states its country to be New Holland. I suspect the inhabitants of the two last cocoons to be terrestrial animals: the first is probably a true aquatic case-worm.
The same purpose for which the cocoons above described serve, is answered in the case of numerousDipterousinsects, by a humble and less artificial contrivance—the skin, namely, of the larva; which, as was before observed[550], is never cast, but, when the insect is about to enter into the pupa state, assumes a different form and colour; becomes of a thicker and more rigid texture; and defends the included pupa, which is separate from it, till its exclusion. In this case the mouth of the larva is constantly different from that of the perfect insect, or at least has not with it those relations as to number and kind of organs, which have been observed in the mouth of other larvæ compared with the insects that they produce. The animal, immediately after it is clothed with this skin, if it is opened, exhibits only a soft gelatinouspulp, in the surface of which the exterior organs of the adult insect cannot yet be detected. Nature requires more time for their elaboration, or at least for the appearance of their outline, and to consolidate them. This pulp first takes an oblong form (Boule allongéeReaum.), and afterwards that of the insect it is destined to give birth to[551]. The skin of the larva also serves for a cocoon to the pupæ of maleCocci[552]. The grub of the genusAnthrenus, so destructive to our cabinets of natural objects[553], when it assumes the pupa does not quit its skin, but only splits it open longitudinally on the back, and when it becomes an imago makes its exit through the orifice[554]. Some Lepidopterous larvæ even (Alucita pentadactyla,Callimorpha rosea, &c.) assume the pupa state within their last skin[555].
When a larva has finished its cocoon,—which withsome species, that proceed so earnestly as though they had not a moment to lose, is the work of a few hours, of others about two or three days,—after a certain interval it casts its last skin, which is usually suffered to remain in the cocoon (but which one moth,Geometra lacertinaria, ejects through an opening purposely left in its bottom), and the pupa makes its appearance[556]. This interval is exceedingly various. Most larvæ assume the pupa state within a few days after they have formed their cocoons; but some not for several weeks, or even months. The caterpillar ofBombyx cæruleocephala, according to Rösel, lies three weeks in the cocoon before this change is effected; those of manyPupivoraandDiplolepariæLatr., according to Reaumur, six months[557]; that ofPhalæna urticatanine months[558]; and that ofCimbex lutea, according to De Geer, sometimes eighteen months[559]. Brahm observes, that such larvæ of the double-brooded moth,Hepialus Testudo, as form their cocoons in autumn, do not become pupæ until the following spring; while those which form them in summer undergo this change in a few days[560]. From this fact it might be conjectured, that the degree of heat prevailing at the time the insect incloses itself determines the period of the pupa's appearance; but this supposition seems contradicted by what Reaumur observed of a brood of the larvæ ofPhalæna urticata, just mentioned, which, though they formed themselves cocoons in September, did not become pupæ till the June following[561]. I am unable, therefore, to assign any plausible cause for these extraordinary variations.The difficulty of comprehending how animals before so voracious can live so long without food may be partly surmounted, by adverting to the circumstance of its having attained its full growth, and laid up a store of nutriment for the development of the perfect insect. It is consequently no more wonderful that it should not have need of any further supply without casting off its upper integument, than that it should not eat after having done so and become a pupa.
We have now traced our little animals through their egg and larva states, and have arrived at the third stage of their existence,the Pupa State. This, to include all, can only be defined,—that state intervening between the larva and imago, in which the parts and organs of the perfect insect, particularly those of sex, though in few cases fully developed, are prepared and fitted for their final and complete development in the last-mentioned state; and in which the majority of these animals are incapable of locomotion, or of taking food.
Pupæ, like larvæ, may be separated into two great divisions:—
I. Those which, in general form, more or less resemble the larvæ from which they have proceeded.II. Those which are wholly unlike the larvæ from which they have proceeded.
I. Those which, in general form, more or less resemble the larvæ from which they have proceeded.
II. Those which are wholly unlike the larvæ from which they have proceeded.
I. To the first division belong, with some exceptions[562],theDermaptera,Orthoptera,Hemiptera, and mostAptera, with the neuropterous tribes ofLibellulina,Ephemerina, and the genusTermes, in the classInsecta; and the majority of theArachnida. This, like the first division of larvæ, may be subdivided into two corresponding smaller sections; the first including those pupæ which resemble the larvæ, except in the relative proportion and number of some of their parts; and the second those that resemble them, except in having the rudiments of wings, or of wings and elytra.
i. The first subdivision will include the pupæ, if they may be so called[563], of insects of theApteraorder, and of the classArachnida: as, lice,Poduræ,Lepismidæ, centipedes, millipedes, mites, harvest-men, spiders, scorpions, &c. These mostly differ from their larvæ only in that the relative length or number of their legs, the number of the segments of the body in some, or the development of their palpi, more nearly approach the characters of the perfect insect; and in that while in their larva state they have two or more skins to cast, previously to their assumption of the imago, in their pupa state they have but one. In fact, this last circumstance is the only one which, strictly speaking, characterizes the pupæ of this subdivision; as the changes which take place in the number and proportion of the organs are partly produced with each change of the larva's skin. And hence, as it is not easy to ascertain what number of skins a spider, for example, has yet to cast, and as both the larva and pupa differ so little from the perfect insect, it is very difficult to determine in what state insects of this division are. Fromthis difficulty has probably arisen the too great multiplication of species in some of these tribes, particularly theArachnida, the larva and pupa having been mistaken for perfect insects. The pupæ of this subdivision were named by Linnécomplete, from the near resemblance which they bear to the imago.
ii. The second subdivision will include the pupæ of theDermaptera,OrthopteraandHemipteraorders, with few exceptions; as likewise theLibellulina,Ephemerina, andTermitina? amongst theNeuroptera: including the well-known tribes of earwigs, cockroaches, crickets, grasshoppers, locusts, lanthorn-flies, froghoppers (CicadaL.), bugs, plant-lice, dragon-flies, day-flies, white ants, &c. Of these, as in the former subdivision, the pupæ are equally capable of eating and moving with the larvæ, which they resemble, except in having the rudiments of wings, or of wings and elytra. The pupæ of the three orders first enumerated differ from those of theNeuropterain resembling the perfect insect in most instances, both as to shape and the organs for taking their food; and in all other respects, except in not having their wings and elytra fully developed[564]. The resemblance of the pupæ of theLibellulinaandEphemerinato the perfect insects is more distant, and the above organs in the two states are very dissimilar; for the pupæ of the former are furnished with a prehensory mask similar to that of the larvæ before described[565], which the perfect insect hasnot; and those of the latter with the usual oral organs of masticating insects, of which the imago has scarcely the rudiments.
I have applied the termrudimentsto the wings and elytra in this state, not in a strict sense, but merely to denote their appearance; for in fact the wings, &c. are complete, but only folded up longitudinally and transversely, and inclosed in membranous cases, which when the last change takes place remain attached to the puparium or pupa-case. The tegmina or hemelytra in this state usually cover the wings, and the upper wings the under; but in theLibellulinaboth are usually visible. Though commonly very small compared with the instruments of flight in the perfect insect, some of these rudiments, contrasted with the majority, are of considerable magnitude. This is the case with those of some species ofChermes, as we learn from De Geer[566].
II. The second grand division comprises by far the largest number of pupæ: those of allcoleopterous,strepsipterous,lepidopterous,hymenopterous,dipterous, andaphanipterous, and by far the majority ofneuropterousinsects, as well as thehemipterousgenusAleyrodes, and one sex ofCoccusof the same order. These pupæ, however, though agreeing in the circumstance of being unlike the larvæ from which they proceed, differ from each other in several respects, and require to be divided into three great sections, as under:—
i. Those pupæ in which thepartsof the future insect, being folded up under a membranous skin closely applying to each, are distinctlyvisible. To this head belonggenerally, the pupæ ofcoleopterous[567]andhymenopterousinsects; those of theneuropterousgeneraMyrmeleonandHemerobius, &c.; theTrichoptera; amongst theDiptera,Culex,TipulaL.,Tabanus,Bombylius, &c.; and that of the flea (Pulex). These were theincompletepupæ of Linné.
ii. Those pupæ in which thepartsof the future insect, being folded up under a harder skin, arelessdistinctlydiscoverable. To this subdivision belong the pupæ of allLepidoptera, and of them alone. These are what Linné termedobtectedpupæ.
iii. Those pupæ which are inclosed in the thick and opaque skin of the larva, through which notraceof the perfect insect can bediscovered. These, which Linné termedcoarctatepupæ, include a large proportion of thedipterousgenera; asŒstrusL.,MuscaL.,EmpisL.,ConopsL., &c. &c.[568]
I shall next advert, chiefly to the pupæ of the grand division last described, under the distinct heads ofsubstance,figure, andparts;colour,age,sex,motions, andextrication of the perfect insect.
i. As to theirsubstance—at first interiorly all pupæ consist of a milky fluid, in which the unformed members of the future perfect insect may be said to float, and inwhich they may be discerned, and separated with the point of a pin[569]. In proportion as these acquire consistency, and are more and more developed by the absorption of the surrounding fluid, they occupy its place, and fill up the cavity of the puparium. The rest of this fluid passes off by transpiration[570]. Reaumur is of opinion that it is from the epiploon, orcorps graisseux, that this matter is prepared, which he regards as analogous to the white of an egg[571]. Incoarctatepupæ the included animal, or the pulp that contains its germes (in which the limbs and body at first are not discernible), fills at this period the whole skin-cocoon; but in proportion as the above evaporation takes place, and the consolidation of the body and parts proceeds, it shrinks at each end, so that when near assuming the imago, a considerable cavity appears both at the head and tail of the cocoon[572]. At this period of its existence, from the quantity of fluid included in the puparium, the animal weighs usually considerably more than it does when become a perfect insect[573].
Theexteriorintegument orskinof pupæ, which is usually lined with a very thin white pellicle, is of different consistence in different orders. In theColeopteraandHymenopterait is, with a few exceptions, of a soft and membranous texture; in theLepidoptera(especially those that are not defended by cocoons), andDiptera, it is more rigid and harder, being either coriaceous or corneous. Lepidopterous pupæ, however, are not excluded fromthe last skin of the larvæ with this hard covering. At the moment of this change the envelope is nearly as soft and membranous as in the order first mentioned. But they are besides covered with a viscous fluid, which appears to ooze out, chiefly from under the wings, and which very soon drying, forms the exterior hard shell[574]. At first the antennæ, wings, and legs, like those ofColeopteraandHymenoptera, can be each separated from the body; and it is only after these parts have been glued together by the fluid just mentioned, which takes place in less than twenty-four hours[575], that they are immoveably attached to the body of the pupa, as we usually see them. In fact, the essential difference between incomplete and obtected pupæ seems to be, that in the former the limbs and body are only covered each with a single membranous integument, whereas in the latter they are besides glued together by a substance which forms an additional and harder envelope. It is not easy to explain the alteration that takes place in the texture of the skin of suchdipterouspupæ as retain the skin of the larva. In the latter this is generally a transparent and very fine membrane: yet the very same integument becomes to the pupa an opaque and rigid case.
The surface of the skin of the greater number of pupæ is smooth, but in those of manyPapilionidæit is rugose and warty: this you may see, particularly in that ofPapilio Machaon. In many of the hawkmoths (SphinxL.) it is covered with impressed puncta. InAttacus Iothe upper side of the channels that separate the intermediate segments of the abdomen are curiously striated with transversestriæ, formed of very minute granula, the lower side being transversely sulcated. In some few instances, as inArctia Salicis,Laria pudibundaandfascelina, the skin of the pupa is clothed with hair[576]: as is also that ofHesperia Bixæ, according to Madame Merian[577]. De Geer has described a little beetle under the name ofTenebrio lardarius (LatridiusLatr.,CorticariaMarsh.), the pupa of which is beset with very fine hairs, terminating in a spherical or oval button[578].
ii. I shall include under the same head both thefigureor shape, andpartsof pupæ, as the latter in most kinds are either the same or nearly the same as those of the larva, or merely incasing those of the imago, so as not to require that detailed notice that I judged necessary when treating of the parts of larvæ.
With regard toincompletepupæ, nothing further can be said of their extremely variousfigure, than that it has a general resemblance to that of the perfect insect. The head, trunk, abdomen, and their respective external organs, are alike visible in both; but in the pupæ, the latter, instead of occupying their natural situation, are all closely folded under the breast and abdomen: or, as in the case of the long ovipositors of some Ichneumons, laid along the back. In a specimen of some coleopterous insect now before me, the following is the order of the arrangement of the parts:—The head is inflexed; the mandibulæ are open; between them are seen the labium and labial palpi; these appear to cover and conceal the maxillæ, and the maxillary palpi extend on each side beyond them; theantennæ pass above the thighs of the two anterior pair of legs, and then turning down over the breast between them and the posterior legs, repose upon the base of the wings; which also are turned down between the intermediate and posterior pair of legs, and rest upon the latter; the tibiæ are bent in and folded upon the thigh, and the tarsi turn outwards[579]. In another coleopterous species, the wings and elytra are placed under the hind-legs. InHymenopterouspupæ the antennæ appear usually to lie between the legs[580]. In manyTipulæthe long legs are bent into three folds in the pupæ; but the tarsi are extended, and lie close to each other, the anterior pair being the shortest[581]. In a specimen belonging to this tribe in my cabinet, which I think containedCtenocera pectinicornis, the six leg-cases are of the same length, exactly parallel and adjacent, and being annulated wear the appearance of tracheæ[582]. These parts have each their separate case, so that a pin may be introduced between them and the body: which cases, as well as the general envelope, are usually formed of a fine soft transparent membrane; but sometimes, as in the lady-bird (Coccinella), the tortoise-beetle (Cassida), the crane-fly (Tipula), &c. it is harder and more opaque, so that though it is usually easy for a practised Entomologist from an examination of the pupa, particularly in theHymenoptera, to predict to what genus the insect to be disclosedfrom them will belong, yet in these cases the organs being not so conspicuous, a less experienced examiner might be perplexed, and unable to come to a conclusion.
Althoughhymenopterouspupæ have usually no parts but what are afterwards seen in the perfect insect, this is not the case with severalcoleopterousanddipterousones, which are furnished with various temporary appendages, indispensable to them to bring about their final change, or for other purposes. Thus, the pupa of the male ofLucanus Cervushas two short, jointed anal processes[583]. That ofHydrophilus caraboideshas a pedunculated lunulate one; and moreover, the sides of the abdominal segments, and the top of the thorax, are beset with hairs, which are not seen in the perfect insect[584]. The abdomen of many, also, is armed with spines. That, the arrangement of whose organs I lately described, has a quadruple series in the back of this part; viz. on each of the first five segments, 3, 2, 2, 3. The five first ventral segments also have on each side three spines; the inner are incurved, the intermediate nearly upright, and the outer one recurved. These spines, except those of the innermost ventral series, terminate in a bristle. In another coleopterous species the back part of the head is armed with a pair of lateral spines, and that of the thorax with three processes, the external ones armed with a single spine, and the intermediate one with a pair. De Geer has figured the pupa of anAsilus, the head of which is armed with eight spines—two robust ones in front, and three smaller ones, connected at the base on each side. The abdominal segments, also, are fringed with spines[585]. The abdomen of the pupa ofCtenocera pectinicornisis armed with several strong conical spines, pointing mostly towards the tail, which is likewise the case with that ofTipula lunata[586]. As the above pupæ are usually subterranean or subcortical, the spines assist in pushing them out of the ground, &c. The respiratory horns that proceed from the thorax of the pupæ of many of the aquatic gnats will be noticed in another place. Those ofCorethra culiciformisand of some other aquatic gnat-likeDiptera, have their anus furnished with a pair of oars, or natatory laminæ, by which they rise to the surface[587].
The figure ofobtectedpupæ, or chrysalises, is more uniform. They are commonly obtuse at the anterior extremity, and gradually contracted to a point at the posterior, or tail. The outline usually inclines to a long oval or an ellipse; but in some, asAttacus IoandLuna, the pupa is shorter and more spherical. InGeometra sambucariait represents an elongated cone, and inHepialusit is nearly cylindrical. In the butterfly tribe (PapilioL.) the outline is frequently rendered angular by various protuberances.
In all these pupæ may be distinguished the following parts:—first, theHead-case(Cephalo-theca), or anterior extremity;secondly, theTrunk-case(Cyto-theca), or intermediate part; andthirdly, theAbdomen-case(Gastro-theca).
1. TheHead-casecovers and protects theheadof the inclosed imago. From its sides behind proceed theantennæ-cases (Cera-theca); and before from the middle, the tongue-case (Glosso-theca). Just below the base of the antennæ-case you may discern the eye-cases (Ophthalmo-theca), surrounded on their inner side by a crescent-shaped lævigated piece, which may perhaps transmit some light to the inclosed prisoner.
2. TheTrunk-case, divided into thethorax, oruppersurface, extending from the head to the dorsal segments of the abdomen, and consisting of three pieces, answering to theprothorax,mesothorax, andmetathoraxof the perfect insect: the first answering to the prothorax small, the second covering the mesothorax very large, and the two next representing the metathorax, at first appearing to belong to the abdomen, but having no spiracle; and the breast (pectus) or under-surface reaching from the head to the ventral abdominal segments, from which proceed the wing-cases (Ptero-theca) and leg-cases (Podo-theca), which organs, with the antenna-cases and tongue-case, entirely cover, or rather form, the breast. The arrangement of the whole is as follows:—The wing-cases, which are more or less triangular, and exhibit the larger nervures of the wings, are a lateral continuation of the mesothorax, which turn downwards from the sides of the breast, and cover, or replace, the three first ventral segments of the abdomen. Theantenna-cases, united to the anterior portion of the head just behind the eye-cases, repose immediately next to those of the wings running parallel with their inner margin. Then follow the legs, the tibiæ forming an angle with the thigh, and the case of the anterior pair being innermost, and representing the breast-bone in the pupa. The tongue lies over the forelegs,except in the case of some sphinxes, which I shall notice afterwards: so that the glosso-theca covers both them and it.
3. The abdomen-case consists ofninesegments when viewed on the back, and of onlysixwhen viewed below; so that it might be said to have ten dorsal and six ventral segments: but the fact is, that the place of the three anterior ventral segments, or rather ventral portions of the segments, (for they form complete rings without any lateral suture,) are replaced by the wings and other organs: in consequence of this, the fourth segment, which is less covered than the three first, at its posterior margin forms an annulus or ring. In counting the abdominal segments of a pupa, you must be careful not to include the piece that represents themetathorax, which looks as if it belonged to the abdomen[588]. In the pupæ ofbutterfliesyou will discover evident traces oftendorsal segments; but in manymoths, and somehawk-moths, you will perceive at first onlyeight, or evenseven, but a closer examination will enable you to discover the line that marks out the others; and if you divide the puparium longitudinally, and inspect its internal surface, you will see very visible sutures between them. The intermediate segments are sometimes separated from each other and the preceding and subsequent ones by deep channels. In the pupa ofPapilio Machaonthere is one such channel between the third and fourth segments. InBombyxregalisthe channel is between the sixth and seventh, and inB. imperatoriathere are three, namely, a channel between the third and fourth, and fourth and fifth, and fifth and sixth segments. The way in which insects with an exserted sting fold it in the pupa seems not to have been noticed; but from an observation of De Geer upon one species ofIchneumon, it appears to be turned up over the back of the abdomen[589].
These little animals, thus swathed and banded, exhibit no unapt representation of an Egyptian mummy; though Lamarck applies the termMumiatoincompletepupæ[590], to which it seems less happily applicable.
Chrysalises, as to the modifications of their general figure, may be conveniently divided into two great classes:first, those that have no angular projections, the anal mucro of some excepted, on different parts of their body; andsecondly, those which have such projections. Each of these classes affords variations in its peculiar characters which require to be noticed.
1. The first of these are calledangularpupæ[591], and are confined to theButterflyor diurnal tribes. In some the head projects into one short conical protuberance: this you may see in the chrysalis of the common cabbage butterfly (Pieris Brassicæ), and others of the same genus[592]; in the brimstone-butterfly (Colias Rhamni[593]), and in the beautiful purple emperor or high-flier (Apatura IrisF.[594]): though in this last it is not conspicuous. Butthe most remarkable instance of a single eminence from the head is exhibited by the pupa of a tropical butterfly (Morpho IdomeneusLatr.), figured by Madame Merian. In this the head projects into a long incurved obtuse horn[595]. In others the head is armed with two mucros, or conical eminences. This is the case with the common butterfly of the nettle (Vanessa UrticæF.[596]), and with that of the beautifulPapilio Machaon[597]. In these the prominences are trigonal. These processes, which in some, as in the peacock-butterfly (Vanessa Io), stand upright[598], and in others diverge (Papilio Machaon), form the eye-cases of the included imago; and in their outer base is planted the crescent-shaped piece I lately mentioned, which seems intended to convey light into it. In many the prothorax, besides a lateral angular projection, has in the middle another triangular or trigonal one, somewhat resembling a Roman nose; on each side of which is a smaller elevated black point: so that it requires no great stretch of imagination to find out in it a sort of resemblance to the human face, which, though not quite so striking as honest Goedart figures it[599], is however very considerable. In the pupa ofMorpho Menelaus, figured by Madame Merian[600], this nasiform prominence of the prothorax is extended into a long arched horn, reaching to the middle of the abdomen. The pupa of the silver-washed fritillary (Argynnis PaphiaF.), and others of the same genus, exhibit beneath this nasiform prominencea very deep depression, itself beset with one or more series of smaller angular elevations. The back of the abdomen is often furnished with two rows of protuberances, in some species larger, in others smaller[601]; sometimes sharp and conical, and sometimes flat, and in some instances resembling the fins of fishes[602]. These bosses usually decrease in size towards the tail.
2. The second kind of chrysalises are denominatedconical[603]. These, which include thecrepuscularandnocturnal Lepidoptera, and the butterflies withonisciformlarvæ, have no protuberances, and are less variable in their form—their anterior extremity being almost constantly oval and rounded, and their posterior conical and acute. An exception to this form is met with in the pupa of a moth long celebrated (Lasiocampa Pithyocampa)[604], which has the head acute and the tail obtuse, and armed with two points[605]. Another occurs in that of the Cossus, which has two points on the head, by which it makes an opening in its cocoon: when it assumes the imago, one of these is placed below the other[606]. And some few have the anterior end nearly flat instead of rounded. The pupa of the orange-tip butterfly (Pieris Cardamines) seems intermediate between the angular and conical kinds: it is somewhat boat-shaped, and distinguished by a fusiform process from the head and tail[607]. Other modifications of the usual figure are met with, but are for the most part so slight as not to require notice. Oneor two, however, should not be passed over. The pupæ of many hawk-moths (SphinxL.) have the anterior piece of the head-case elongated into a sort of cylindrical proboscis, which is incurved beneath the breast: you will find this formation inS. ConvolvuliandLigustri[608]. In some, as in a species figured by Madame Merian, that feeds upon theAnnona squamosa, it is rolled up like a serpent in many folds[609]. InNoctua Linariæthe tongue-case turns upwards, and is prominent laterally beyond the body[610]. This singular appendage is one of those beautiful instances of compensating contrivances, as Dr. Paley calls them, which perpetually occur in the insect tribes. The tongue of these hawk-moths is of very great length, often three inches, while the pupa itself is scarcely two; it could not possibly, therefore, have been extended at length, as it is in common cases, but is coiled up within the above protuberance. When the tongue is but a little longer than the breast, the ordinary plan is adhered to, but the apex of the breast projects a little over the abdomen into a sort of nose, in which the end of the tongue is contained. This conformation may be seen in the pupa ofNoctua Gamma,Verbasci, and many other species. Sometimes, as inN. LinariæF., this projection is recurved into a short horn.
I have before adverted to theadminiculaor short spines looking towards the anus, with which the dorsal segments of the abdomen of some pupæ are armed; and by which, when the time for their exclusion is arrived, they are enabled to push themselves upwards or outwards fromtheir several places of confinement[611]: you will find these in the pupa of the great goat-moth (Cossus ligniperda); and in the cylindrical pupa of the moth called the ghost (Hepialus HumuliF.) there are two rows of sharp triangular spines on the back of each segment. These are not laid flat, but, as they do also in theCossus, form an acute angle with the body; which gives them greater power of resistance. Those that constitute the row nearest the base of the segment are longer than the anterior row, the middle spines than the lateral ones. The first and last segment are without them, and the last segment but one has a sharp ventral transverse ridge, armed with many sharp teeth[612]. The abdominal spines lately mentioned, of semicomplete pupæ, are alsoadminicula.
The tail of this description of pupæ is in many instances armed with a mucro, or sharp point, emerging from its upper side. You will see this in most hawk-moths. In the pupa ofHesperia Proteusthe mucro is truncate at the apex; in that ofBombyx imperatoriait is long, and terminates in two diverging points. In the majority of chrysalises of both descriptions the tail is acute, and usually furnished with hooks of different kinds. These are so various in shape and number, &c. that they would probably afford good characters for discriminating many allied species. In some there are but two or three, in others five or six, in others they are more numerous[613]. Sometimes they are quite straight[614], but most commonly recurved, so as to form a hook. The hawk-moths, and afew others, asBombyx Pini,Cerura Vinula, &c., have no anal hooks whatever. Under this head I shall observe, that in many conical pupæ below the anal angle or mucro, is the appearance of a vertical foramen or passage: this is particularly conspicuous inHepialus, in which it is surmounted by a bifid ridge, and has under it a pair of minute black tubercles.
A pretty accurate judgement of the division to which the perfect insect when disclosed will belong, may usually be formed from the figure of itschrysalis. All theangularones, with scarcely any exception, inclosebutterflies. The converse, however, does not hold; for some that are not angular, as those ofParnassius ApolloandMnemosyne, and most of the LinneanPlebeii urbicolæ, also inclose flies of that description. With these exceptions, allconicalchrysalises give birth tomothsorhawkmoths. An idea even of the family or genus under which the perfect insect will arrange, may be generally formed from the figure of the chrysalis; less distinctly, however, in the conical or rounded, than in the angular kinds, in which the prominences of the head and trunk, as before explained, usually vary in different families. Even the sex of some moths may be judged from the pupæ: those of females being thicker; and those also of the females that have no wings, or only the rudiments of them, will of course vary somewhat from the ordinary form: but there is a still more striking difference in that ofCallimorpha? vestitaF., and others of the singular tribe before noticed[615], called by the GermansSacktrager(sack-bearers), from the sack-like cases in which the larva resides. Thefemales of these having not only no wings, but no antennæ, and legs not longer than those of the larva, their pupa more resembles that of adipterousthan of alepidopterousinsect, it being not easy to determine which is the head and which the tail[616].
In these too we can often learn from the outline of the wing-cases, whether the inhabitant of the chrysalis has these organs indented or intire. If the former, the margins of these cases are sinuate, as in that ofVanessa C. album; if the latter, they are intire, as inPieris Brassicæ. Even in conical pupæ,—the size, the shape of the antennæ, which may be distinguished through the skin that covers them, and slight modifications of the ordinary form,—give indications of the genus of the included insect sufficiently conclusive to a practised eye.
The true figure ofcoarctatepupæ when they are mature, the parts of the future fly being very visible, and each being included in a separate case[617], is that of those that belong to theincompletedivision; but as this is a character not cognizable without dissection, it is customary, in speaking of pupæ of this description, to refer solely to the shape of the exterior covering, which is in fact a cocoon formed of the dried skin of the larva moulded into a different form. In this sense the figure of coarctate pupæ is extremely various. The majority of them are more or less oval or elliptical, without any distinct parts, were it not that they usually retain traces of the segments which composed the larva's body[618]. Of this figure are the pupæ of the common cheese-maggot[619], and many other flies. Others(SepedonLatr.) have the pupa shaped like a boat. That ofScæva PyrastriF. assumes the figure of a flask; or, according to Reaumur's more accurate comparison, of atear[620]. The tail of many of these pupæ, particularly of aquatic species, is elongated into a sort of beak, either simple or forked, or is beset with spines variously arranged. The pupa ofStratyomis Chamæleon, and other allied species, differs from all the rest of this subdivision in retaining the exact form of the larva[621]; and hence constitutes an exception to the general character of our second great Division.
iii. There is much less variety in thecolourof pupæ than in that of larvæ. The majority of coleopterous and hymenopterous pupæ are white, or whitish; of lepidopterous and dipterous, brown of various shades, often verging on black in the former and on red in the latter. The angular lepidopterous ones, however, are more gaily decorated. Some,Pieris Brassicæ, are of a greenish yellow, marked with spots of black; others are of a uniform green,Apatura Iris,Pieris Cardamines; others, reddish,Vanessa C. album; others again red with black spots,Urania Leilus[622]. A still greater number shine as though gilded with burnished gold—either applied in partial streaks,Vanessa Cardui; or covering the entire surface,Vanessa Urticæ. It was from this gilded appearance in someobtectedpupæ that the termsChrysalisandAureliawere applied to the whole. The alchemists mistook this for real gold; and referred to the case as an argument in favour of the transmutation of metals. But Reaumur has satisfactorily shown, that in this instance the old proverb is strictly applicable—"All is not gold that glitters." He found that this appearance is owing to the shining white membrane immediately below the outer skin, which being of a transparent yellow gives a golden tinge to the former; in the same way that tinfoil, when covered with a yellow varnish, assumes the metallic appearance which we see in gilt leather. He mentions, too, that for the production of this effect—it is essential that the inner membrane be moist: whence may be explained the disappearance of the gilding as soon as the butterfly is ready to escape from the pupa. The shade of colour in these gilded chrysalises is various: some are of a rich yellow, like pure gold; others much paler; and some nearly as white as silver. That ofHipparchia CassiæF. is red with silver spots[623].
Though by far the greater number of the chrysalises of moths are of an uniform chestnut, brown, or black,—a few are of other colours; as that ofGeometra alniaria, which is of a glaucous blue; ofNoctua sponsa, lilac; and ofNoctua pacta, of a lovely blue, caused by a kind of bloom, like that of a plum, spread upon a brown ground. A similar bloom is found on that ofParnassius Apollo, and on the anterior part of that ofPlatypterix cultariaandsicula; in which last, Kliemann observed it to thebe renewed when rubbed off[624]Many pupæ have the sheaths of the wings of a different colour from that of the rest of the body; a few are variegated with paler streaks or bands, asClostera Anastomosis, which has two red longitudinal stripes down its dark-brown back; and that of the common gooseberry and currant moth, which may be found in every garden, has alternate rings of black and yellow[625].
A few pupæ vary in their colour, as the painted lady-butterfly (Vanessa Cardui), some of which are light-brown with gray streaks and golden dots, others wholly of a golden yellow or brown, others of a light green[626].
Almost all at their first assumption of the pupa state have a different colour from that which they take a few days afterwards. This last they retain until the disclosure of the perfect insect; except some that have transparent skins, which a few days previously to this period exhibit the colours of the included animal.
iv. There is as great variety in the length of theageof Insects in their pupa as in their larva state. Some species continue in it onlytwoorthree days(Aleyrodes ChelidoniiLatr.,Tinea proletellaL.); others, as manyweeks, ormonths, or evenyears. Each, however, has in general a stated period, which in ordinary circumstances it neither much exceeds nor falls short of. The only general rule that can be laid down is—thatsmallpupæ continue in that state a shorter time than those oflargerbulk. Thus, amongst coleopterous genera, the more minute species ofCurculioL.; amongst theHymenoptera,theIchneumones minutiL.; amongst theLepidoptera, the subcutaneous tribes; and the majority of theDiptera,—remain as pupæ only a few days or weeks: while the larger species in all these orders commonly exist in the same state several months—many even upwards oftwoyears. There are, however, numerous exceptions to this rule; for some large pupæ are disclosed in a much shorter time than some others not a twentieth part of their bulk.
The reasons both of the rule and of the exceptions to it are sufficiently obvious. And first, as to the rule:—If you open a pupa soon after its assumption of that state, you will find its interior filled with a milky fluid, in the midst of which the rudiments of its future limbs and organs, themselves almost as fluid, swim. Now the end to be accomplished during the pupa's existence is, the gradual evaporation of the watery parts of this fluid, and the development of the organs of the inclosed animal by the absorption and assimilation of the residuum. Reaumur, by inclosing a pupa in a stopped glass tube, collected a quantity of clear and apparently of pure water, equal to eight or ten large drops, which had evaporated from it, and was condensed against the sides of the tube, and it was found to have lost an eighteenth part of its weight[627]. It is plain, therefore, that this necessary transpiration, other circumstances being alike, must take place sooner in asmallthan in alargepupa. Next, as to the exceptions:—Since the more speedy or more tardy evaporation of fluids depends upon their exposure to a greater or less degree of heat, we mightà prioriconclude, that pupæexposed to a high temperature would sooner attain maturity, even though larger in bulk, than others exposed to a low one:—and this is the fact. The pupa of a large moth, which has assumed that state in the early part of summer, will often disclose the perfect insect in twelve or fourteen days; while that of an Ichneumon, not one hundredth part of its size, that did not enter this state till late in autumn, will not appear as a fly for seven or eight months. But this is not the whole. The very same insect, according as it has become a pupa at an earlier or later period of the year, will at one time live but a few weeks, at another several months, in that state. Thus, if the caterpillar ofPapilio Machaon, one of those which has annually a double brood, becomes a pupa in July, the butterfly will appear inthirteen days: if not until September, it will not make its appearance until the June following; that is, not in less thannineorten months: and the case is the same with the pupæ ofNoctua Psi, and of a vast number of other insects. To put beyond all doubt the dependence of these remarkable variations on temperature merely, it was only necessary that they should be effected, as Lister long ago advised[628], by artificial means. This Reaumur accomplished. In the month of January he placed the chrysalises of several moths and butterflies, which would not naturally have been disclosed until the following May, in a hothouse: the result was, that the perfect insects made their appearance in less than a fortnight, in the very depth of winter; and by other numerous and varied experiments he ascertained, that in this heated atmosphere five or sixdayshastened their maturity more than as manyweekswould have done in the open air. The disclosed insects were in every respect perfect, and the females, after pairing, laid their eggs, and then died, just as if they had not been thus prematurely forced into existence. The converse of this experiment equally succeeded:—by keeping pupæ the whole summer in an icehouse, Reaumur caused them to produce the fly one full year later than their ordinary period[629].
This extraordinary fact leads us to a very singular and unexpected conclusion—that we have the power of lengthening or shortening the life of many insects at pleasure; that we can cause one individual to live more than twice as long as another of the same species, andvice versâ. Had Paracelsus made this discovery, it would have led him to pursue his researches after the elixir of immortality with redoubled confidence, and would have supplied him with an argument for the possibility of prolonging the life of man beyond its usual term, which his sceptical opponents would have found some difficulty in rebutting. Even the logical Reaumur seems inclined to infer from it, that this object of the alchemists was not so chimerical as we are wont to conclude[630]. He confesses, however, if it were to be attained only by the same process as effects the extension of an insect's life—by prolonging its state of torpor and insensibility,—that few would choose to enjoy it on such conditions. The man of pleasure, blunted by excess of use to all modern stimuli, might perhaps not object to a sleep of a hundred years, in the hope of finding something new under thesun when he waked; and an ardent astronomer would probably commit himself with scientific joy to a repose as long and as sound as that of the seven sleepers, for the chance of viewing his predicted return of a comet, on stepping out of his cave: but ordinary mortals would consign themselves to the perils of so long a night with reluctance, apprehending a fate no better than what befel the magician, who ordered himself to be cut in small pieces and put in pickle, with the expectation of becoming young again[631].
The duration, then, of an insect's existence in the pupa state, depends upon its bulk, upon the temperature to which it is exposed, and upon a combination of these two circumstances. This experiment appears very simple. We seem to ourselves to have accomplished what is so often undertaken in vain—to have found an entrance into the cabinet of Nature, and to have made ourselves masters of the contents of one of the pages of her sealed and secret book. We deceive, ourselves, however: this book, when it seems most legible, is often interlined withsympathetic inks, if I may so speak, which require tests unknown to us for their detection. If you lay up a considerable number of the pupæ of a moth now calledEriogaster lanestris, the larva of which is not uncommon in June on the black-thorn, selected precisely of the same size, and exposed to exactly the same temperature, the greater number of them will disclose the perfect insect in the February following; some not till the February of the year ensuing, and the remainder not before the samemonth in the third year[632]. Mr. Jones of Chelsea, a most acute lepidopterist, in one of his excursions captured a female ofArctia mendica, another moth, which laid a number of eggs, thirty-six of which produced caterpillars: all these fed, spun their cocoons, and went into the pupa state in the usual manner, but at the proper season only twelve produced the fly. As this was no uncommon circumstance, he concluded that the rest were dead: to his great astonishment, however, in the next season twelve more made their appearance; and the following year the remainder burst into life, equally perfect with the foregoing[633]. In this extraordinary result, which also occasionally has been observed to take place in the emperor-moth (Saturnia pavonia), the privet-hawkmoth (Sphinx Ligustri), and that of the spurge (S. Euphorbiæ)[634], and other species,—it is clear that something besides mere size and temperature is concerned: for, these circumstances being precisely alike, one pupa arrives at maturity in six months, and another of the same brood requires between two and three years. We can guess, that the end which the All-wise Creator has in view, in causing this remarkable difference, is the prevention of all possibility of the destruction of the species.Eriogaster lanestrisandArctia mendica, &c., for instance, are doomed, for some reasonunknown to us[635], to be disclosed from the pupa in the cold and stormy months of February and March, almost every day of which in certain years is so ungenial that few insects could then survive exposure, much less deposit their eggs and ensure the succession of a progeny. Now, were all these to make their appearance in the perfect state in thesameyear, it might happen that the whole race in a particular district would be destroyed. But this possibility is effectually guarded against by the beautiful provision under consideration, it being very improbable that three successive seasons should be throughout unfavourable; and without such occurrence, it is clear that some of the race of this moth will be preserved. In the case of other moths, whose pupæ though disclosed in the summer are governed by the same rule, the prevention of the extinction of the species, by any extraordinary increase in a particular year of their natural enemies, seems the object in view[636]. But though the intention be thus obvious, the means by which it is effected are impenetrably concealed. What physiologist would not be puzzled with the eggs of a bird, of which one-third should require for their hatching to be sat upon only a fortnight, another third a month, and the remainder six weeks? Yet this would be an anomaly exactly analogous to that observed by Mr. Jones with respect to the pupæ ofA. mendica. Reaumur found that when the skin of pupæ was varnished, so as to prevent absorption, the appearance ofthe fly happened nearly two months later than in ordinary circumstances. Are we to conjecture that those of the moth just mentioned, or ofE. lanestris, that are latest matured, from a greater degree of viscidity in the fluid that forms them[637], have thicker and more impervious skins than those disclosed at an earlier period? Or are we to refer the difference to some unknown peculiarity of organization? On any supposition, the fact remains equally wonderful; and I know of none the illustration of which is more worthy of the patient investigation of the physiologist.
As the period of maturity of the perfect insect is thus in some cases not fixed even to years, and as in many it seems dependent upon such variable causes; nothing appears more improbable than that it should ever be so strictly determined, that even the week in which the fly will leave its pupa-case can be pretty accurately predicted. Such, however, is the fact with regard to theEphemeraso interestingly described by Reaumur, the myriads of which that issue from the banks of the Seine all appear in two or three days, somewhere between the 10th and 18th of the month of August[638]in every year; at which time the fishermen regularly expect them. A like regularity attends the appearance of those described by Swammerdam, which every year, for three days about the feast of St. John, issue in clouds from the Rhine[639]—Not only is the week fixed, but in several instances eventhe hour. The Ephemeræ observed by Reaumur appear at no other time than betweeneightandteno'clock in theevening; and so unalterably is their exclusion fixed, that neither cold nor rain can retard it. Between these hours, in the evenings on which they appear, you may see them fill the air, but an hour before or after, you will in vain look for one[640]. So also the silkworm-moth and the hawkmoth of the evening primrose (Sphinx Œnotheræ) constantly break forth from the pupa atsunrise: and the hawkmoth of the lime (Smerinthus Tiliæ) as certainly atnoon[641]. Schroeter states, that of sixteen specimens of the death's-head-hawkmoth (S. Atropos) which he bred, every one was disclosed betweenfourandseveno'clock in theafternoon[642].
Before I conclude this head, I must observe, that after a caterpillar or gnat has spun its cocoon, it sometimes remains for a considerable period before it incloses itself in the pupa-case, and casts off the form of a larva. Thus the little parasite (Ichneumon glomeratusL.) that destroys the caterpillar of the common cabbage-butterfly, remains a larva in its cocoon for many months, but it becomes a perfect insect a few days after it has put on its puparium[643]; and the caterpillars of the great goat-moth (Cossus ligniperda), if they spin their cocoon in the autumn, remain in it through the winter in the larva state; whereas, if they inclose themselves in the month of June, they assume the pupa, so as to appear as flies in three or four weeks[644]. It is not therefore easy to state preciselythe age of those pupæ which are produced from larvæ that spin cocoons.