Volume 1 — LETTER XXXI.
Reading edition
STATES OF INSECTS.
PUPA STATE.
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. To the first division belong, with some exceptions[562], the Dermaptera, Orthoptera, Hemiptera, and most Aptera, with the neuropterous tribes of Libellulina, Ephemerina, and the genus Termes, in the class Insecta; and the majority of the Arachnida. 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 the Aptera order, and of the class Arachnida: 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. From this difficulty has probably arisen the too great multiplication of species in some of these tribes, particularly the Arachnida, the larva and pupa having been mistaken for perfect insects. The pupæ of this subdivision were named by Linné complete, from the near resemblance which they bear to the imago.
ii. The second subdivision will include the pupæ of the Dermaptera, Orthoptera and Hemiptera orders, with few exceptions; as likewise the Libellulina, Ephemerina, and Termitina? amongst the Neuroptera: including the well-known tribes of earwigs, cockroaches, crickets, grasshoppers, locusts, lanthorn-flies, froghoppers (Cicada L.), 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 the Neuroptera in 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 the Libellulina and Ephemerina to 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 has not; 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 term rudiments to 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 the Libellulina both 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 of Chermes, as we learn from De Geer[566].
II. The second grand division comprises by far the largest number of pupæ: those of all coleopterous, strepsipterous, lepidopterous, hymenopterous, dipterous, and aphanipterous, and by far the majority of neuropterous insects, as well as the hemipterous genus Aleyrodes, and one sex of Coccus of 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 the parts of the future insect, being folded up under a membranous skin closely applying to each, are distinctly visible. To this head belong generally, the pupæ of coleopterous[567] and hymenopterous insects; those of the neuropterous genera Myrmeleon and Hemerobius, &c.; the Trichoptera; amongst the Diptera, Culex, Tipula L., Tabanus, Bombylius, &c.; and that of the flea (Pulex). These were the incomplete pupæ of Linné.
ii. Those pupæ in which the parts of the future insect, being folded up under a harder skin, are less distinctly discoverable. To this subdivision belong the pupæ of all Lepidoptera, and of them alone. These are what Linné termed obtected pupæ.
iii. Those pupæ which are inclosed in the thick and opaque skin of the larva, through which no trace of the perfect insect can be discovered. These, which Linné termed coarctate pupæ, include a large proportion of the dipterous genera; as Œstrus L., Musca L., Empis L., Conops L., &c. &c.[568]
I shall next advert, chiefly to the pupæ of the grand division last described, under the distinct heads of substance, figure, and parts; colour, age, sex, motions, and extrication of the perfect insect.
i. As to their substance—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 in which 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, or corps graisseux, that this matter is prepared, which he regards as analogous to the white of an egg[571]. In coarctate pupæ 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].
The exterior integument or skin of pupæ, which is usually lined with a very thin white pellicle, is of different consistence in different orders. In the Coleoptera and Hymenoptera it is, with a few exceptions, of a soft and membranous texture; in the Lepidoptera (especially those that are not defended by cocoons), and Diptera, it is more rigid and harder, being either coriaceous or corneous. Lepidopterous pupæ, however, are not excluded from the 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 of Coleoptera and Hymenoptera, 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 such dipterous pupæ 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 many Papilionidæ it is rugose and warty: this you may see, particularly in that of Papilio Machaon. In many of the hawkmoths (Sphinx L.) it is covered with impressed puncta. In Attacus Io the upper side of the channels that separate the intermediate segments of the abdomen are curiously striated with transverse striæ, formed of very minute granula, the lower side being transversely sulcated. In some few instances, as in Arctia Salicis, Laria pudibunda and fascelina, the skin of the pupa is clothed with hair[576]: as is also that of Hesperia Bixæ, according to Madame Merian[577]. De Geer has described a little beetle under the name of Tenebrio lardarius (Latridius Latr., Corticaria Marsh.), 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 the figure or shape, and parts of 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 to incomplete pupæ, nothing further can be said of their extremely various figure, 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; the antennæ 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. In Hymenopterous pupæ the antennæ appear usually to lie between the legs[580]. In many Tipulæ the 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 contained Ctenocera 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 the Hymenoptera, to predict to what genus the insect to be disclosed from 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.
Although hymenopterous pupæ have usually no parts but what are afterwards seen in the perfect insect, this is not the case with several coleopterous and dipterous ones, 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 of Lucanus Cervus has two short, jointed anal processes[583]. That of Hydrophilus caraboides has 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 an Asilus, 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 of Ctenocera pectinicornis is armed with several strong conical spines, pointing mostly towards the tail, which is likewise the case with that of Tipula 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 of Corethra culiciformis and of some other aquatic gnat-like Diptera, have their anus furnished with a pair of oars, or natatory laminæ, by which they rise to the surface[587].
The figure of obtected pupæ, 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, as Attacus Io and Luna, the pupa is shorter and more spherical. In Geometra sambucaria it represents an elongated cone, and in Hepialus it is nearly cylindrical. In the butterfly tribe (Papilio L.) the outline is frequently rendered angular by various protuberances.
In all these pupæ may be distinguished the following parts:—first, the Head-case (Cephalo-theca), or anterior extremity; secondly, the Trunk-case (Cyto-theca), or intermediate part; and thirdly, the Abdomen-case (Gastro-theca).
1. The Head-case covers and protects the head of the inclosed imago. From its sides behind proceed the antennæ-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. The Trunk-case, divided into the thorax, or upper surface, extending from the head to the dorsal segments of the abdomen, and consisting of three pieces, answering to the prothorax, mesothorax, and metathorax of 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. The antenna-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 of nine segments when viewed on the back, and of only six when 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 the metathorax, which looks as if it belonged to the abdomen[588]. In the pupæ of butterflies you will discover evident traces of ten dorsal segments; but in many moths, and some hawk-moths, you will perceive at first only eight, or even seven, 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 of Papilio Machaon there is one such channel between the third and fourth segments. In Bombyx regalis the channel is between the sixth and seventh, and in B. imperatoria there 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 of Ichneumon, 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 term Mumia to incomplete pupæ[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; and secondly, 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 called angular pupæ[591], and are confined to the Butterfly or 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 Iris F.[594]): though in this last it is not conspicuous. But the most remarkable instance of a single eminence from the head is exhibited by the pupa of a tropical butterfly (Morpho Idomeneus Latr.), 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 Urticæ F.[596]), and with that of the beautiful Papilio 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 of Morpho 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 Paphia F.), and others of the same genus, exhibit beneath this nasiform prominence a 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 denominated conical[603]. These, which include the crepuscular and nocturnal Lepidoptera, and the butterflies with onisciform larvæ, 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. One or two, however, should not be passed over. The pupæ of many hawk-moths (Sphinx L.) 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 in S. Convolvuli and Ligustri[608]. In some, as in a species figured by Madame Merian, that feeds upon the Annona squamosa, it is rolled up like a serpent in many folds[609]. In Noctua Linariæ the 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 of Noctua Gamma, Verbasci, and many other species. Sometimes, as in N. Linariæ F., this projection is recurved into a short horn.
I have before adverted to the adminicula or 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 from their 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 Humuli F.) 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 the Cossus, 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 also adminicula.
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 of Hesperia Proteus the mucro is truncate at the apex; in that of Bombyx imperatoria it 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 a few others, as Bombyx 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 in Hepialus, 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 its chrysalis. All the angular ones, with scarcely any exception, inclose butterflies. The converse, however, does not hold; for some that are not angular, as those of Parnassius Apollo and Mnemosyne, and most of the Linnean Plebeii urbicolæ, also inclose flies of that description. With these exceptions, all conical chrysalises give birth to moths or hawkmoths. 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 of Callimorpha? vestita F., and others of the singular tribe before noticed[615], called by the Germans Sacktrager (sack-bearers), from the sack-like cases in which the larva resides. The females 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 a dipterous than of a lepidopterous insect, 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 of Vanessa C. album; if the latter, they are intire, as in Pieris 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 of coarctate pupæ 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 the incomplete division; 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 (Sepedon Latr.) have the pupa shaped like a boat. That of Scæva Pyrastri F. assumes the figure of a flask; or, according to Reaumur's more accurate comparison, of a tear[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 of Stratyomis 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 the colour of 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 some obtected pupæ that the terms Chrysalis and Aurelia were 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 of Hipparchia Cassiæ F. 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 of Geometra alniaria, which is of a glaucous blue; of Noctua sponsa, lilac; and of Noctua 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 of Parnassius Apollo, and on the anterior part of that of Platypterix cultaria and sicula; in which last, Kliemann observed it to the be 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, as Clostera 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 the age of Insects in their pupa as in their larva state. Some species continue in it only two or three days (Aleyrodes Chelidonii Latr., Tinea proletella L.); others, as many weeks, or months, or even years. 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—that small pupæ continue in that state a shorter time than those of larger bulk. Thus, amongst coleopterous genera, the more minute species of Curculio L.; amongst the Hymenoptera, the Ichneumones minuti L.; amongst the Lepidoptera, the subcutaneous tribes; and the majority of the Diptera,—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 of two years. 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 a small than in a large pupa. 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 might à priori conclude, that pupæ exposed 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 of Papilio Machaon, one of those which has annually a double brood, becomes a pupa in July, the butterfly will appear in thirteen days: if not until September, it will not make its appearance until the June following; that is, not in less than nine or ten months: and the case is the same with the pupæ of Noctua 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 six days hastened their maturity more than as many weeks would 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, and vice 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 the sun 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 with sympathetic 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 called Eriogaster 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 same month in the third year[632]. Mr. Jones of Chelsea, a most acute lepidopterist, in one of his excursions captured a female of Arctia 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 lanestris and Arctia mendica, &c., for instance, are doomed, for some reason unknown 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 the same year, 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 of A. mendica. Reaumur found that when the skin of pupæ was varnished, so as to prevent absorption, the appearance of the fly happened nearly two months later than in ordinary circumstances. Are we to conjecture that those of the moth just mentioned, or of E. 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 the Ephemera so 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 even the hour. The Ephemeræ observed by Reaumur appear at no other time than between eight and ten o'clock in the evening; 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 at sunrise: and the hawkmoth of the lime (Smerinthus Tiliæ) as certainly at noon[641]. Schroeter states, that of sixteen specimens of the death's-head-hawkmoth (S. Atropos) which he bred, every one was disclosed between four and seven o'clock in the afternoon[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 glomeratus L.) 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 precisely the age of those pupæ which are produced from larvæ that spin cocoons.
v. I have not much to say with regard to the sex of pupæ. The male is probably to be distinguished from the female by being smaller; but in the first great division of pupæ, those which resemble the larvæ, and are locomotive, the female in numerous cases may be known by the Ovipositor, or instrument for depositing her eggs in their proper station: and the male also has his anal instruments. Sometimes in this state the animal is so matured, as to be capable of continuing its kind. I have found the pupæ both of a Gryllus L. and of a Cimex L. in coitu.
vi. Though the pupæ of the second great division are usually not locomotive, yet I must not omit some notice of their motions. As the legs of insects in this state are folded within a common or partial integument, of course none of the pupæ now under consideration, with the exception of those of the Trichoptera order, can walk: coarctate ones are even incapable of the slightest motion, and exhibit no symptom whatever of animation. Some of those that are termed incomplete, however, and most chrysalises, have the power of communicating to their bodies a slight movement, extending more or less in different species, which is effected by the abdominal segments solely. The latter, during the first twelve hours of being pupæ, when their skin is soft, frequently turn themselves, that the side on which they lie may not be flattened; afterwards by far the majority merely wriggle or twist their abdomen when touched, or in any way incommoded or disturbed. We learn from De Geer, that the pupa of the ghost-moth (Hepialus Humuli), the cocoon of which is more than twice the length of the chrysalis, moves in it from one end to the other[645]. Bonnet observed one of a moth (perhaps Lasiocampa Quercus), which alternately fixed itself at the top and bottom of its spacious and obliquely-fixed cocoon; descending slowly, but ascending as quickly, and almost in the same manner, as a chimney-sweeper in a chimney[646]. The pupa of the weevil of the water-hemlock (Lixus paraplecticus) will move from one end of the interior of a branch to another by means of its adminicula, aided by the motion of its abdominal segments[647]. But the most locomotive of pupæ of the second division are those of gnats, and many Tipulidans, which pass this state in the water. These will move from the bottom to the surface, and back again, with great facility and velocity. I have before mentioned several other motions of pupæ[648], which I shall not repeat here, by which they extricate themselves from their several places of intermediate repose, before they leave the puparium: if the imago were to be disclosed in the interior of a tree, or in the earth, its wings would be materially injured in forcing its way out. The object of several of the above motions may be to alarm insects that might attack these defenceless beings. The twirling motion in particular, formerly noticed[649], in some species, by causing a rustling against the sides of the cocoon, makes a considerable noise—so singular in that of a red underwing-moth (Noctua pacta), that Rösel tells us, (who by the by was more timid than becomes a philosopher,) that the first time he heard it, he had nearly thrown away the box that contained it, in his fright[650].
vii. We are next to consider The extrication of the perfect insect from the puparium, or pupa-case, and from the cocoon. The period when the pupa has attained maturity, and the inclosed insect is ready to burst the walls of its prison, may be often ascertained. Just at this time the colour frequently undergoes an alteration, the golden or silver tint of the gilded chrysalises vanishes; and those which are transparent, usually permit the form and colours of the insect within and the motions of their limbs to be distinctly seen through them. In the Libellulina the eyes become more brilliant[651]. The mature pupæ of the moth lately mentioned (Eriogaster lanestris) have a particular swell of the abdominal segments, not apparent in those that are to continue till another season, or longer[652]. Those of the case-worms (Trichoptera) push off the grates from the cases which they have hitherto inhabited, and swim about[653]. Other signs and motions doubtless predict the approach of this great change in other species, which have not been recorded.
The mode in which insects make their way out of the puparium differs in different orders. In obtected pupæ, the struggles of the included butterfly or moth first effect a longitudinal slit down the middle of the thorax, where there is usually a suture for the purpose. The slit rapidly extends along the head, and down the parts which compose the breast, and the insect gradually withdraws itself from its case. It is not, however, from the outer skin merely that it has to disengage itself, but also from a series of inner membranous cases, which separately inclose the antennæ, proboscis, feet, &c., as a glove does the fingers; and similar cases inclose the parts of the perfect insect in pupæ of all the other orders. This is sometimes a work of difficulty, but ordinarily it is effected with ease.
Incomplete and semicomplete pupæ undergo nearly the same process, save that in them the body is not swathed up in a common case; and therefore they have only to liberate themselves from the partial cases that envelop the several parts of their body.
In coarctate pupæ, as those of Muscidæ, Syrphidæ, Œstridæ, &c., the process is different. Their outer-case is ordinarily more rigid and destitute of the sutures, which in the former tribes so easily yield to a slight effort. Yet in these, at the anterior end under which the head of the fly lies, and from which it always issues, there is commonly a sort of lid, joined by a very indistinct suture to the rest, which can be pushed off, leaving a sufficient opening for the egress of the insect. In the pupæ of many of this tribe this lid is composed of two semicircular pieces, which can be separately removed. Many species seem to be able to force off the lid of their puparium, by merely pushing against it with their heads: but the common flesh-fly and many other Muscidæ, which are perhaps too feeble to effect this, or whose puparia are stronger than ordinary, are furnished with a very remarkable apparatus for this express and apparently sole purpose. They are gifted with the power of introducing air under the middle part of the head, to which the antennæ are fixed, and of inflating that part into a sort of membranous vesicle as big as the head itself; by the action of which against the end of the pupa-case, the lid is soon forced off. So powerful is this singular lever, that it is even sufficient to rupture the fibrous galls in which the pupæ of the gay-winged Tephritis Cardui[654] are inclosed. That it is designed by Creative Wisdom to answer this sole purpose seems proved, from its disappearing soon after the disclosure of the fly, whose head shortly becomes all alike hard. Reaumur suspects that it may also be intended to promote the circulation of the insect's fluids; but to me his reasons appear not conclusive[655]. In one instance a mode still more unexpected obtains. The illustrious naturalist just named found that the fly which proceeded from one of the rat-tailed grubs (Elophilus Latr.) had actually the power of completely reversing its situation in its narrow case; and that it then employed its tail in pushing off the lid, which other species remove by means of their heads[656].
The extrication of insects whose pupæ are above ground, like those of butterflies, many beetles, flies, &c., is comparatively a simple operation. But what, you will ask, becomes of those species whose pupæ are concealed deep in the earth, or in the heart of the trees on which their larvæ have fed? Of this you shall be informed.—Coleopterous insects disclosed from pupæ thus circumstanced, wait until their organs have acquired strength, and their elytra are sufficiently hardened to protect their filmy wings from damage in forcing their way through the earth or wood which covers them. Thus Oryctes nasicornis, a rhinoceros beetle common on the Continent, is a full month before it reaches the surface of the earth, after quitting its puparium. But it is evident that no delay would enable lepidopterous or dipterous insects, which are without elytra, to make their way out of such situations, without irreparable injury to their delicate wings. Many of these, therefore, while still within the hard case of the pupa, have the precaution, a few days previously to their exclusion, to force themselves up to the surface of the earth, or, when they reside in the interior of trees, to the entrance of their hole. This is effected by a successive wriggling of the abdominal segments, which in several species, of the Coleoptera, Lepidoptera, and Diptera orders, for this purpose, as has been more than once observed[657], are furnished with sharp points (adminicula), admitting a progressive, but not a retrograde motion. The puparia of the great goat-moth (Cossus ligniperda) may be often seen projecting from orifices in willow-trees; and those of the common crane-fly (Tipula oleracea) from the surface of the earth, to which they have thus made their way from a depth of several inches.
In all the preceding instances the exclusion of the perfect insect is complete, as soon as it has withdrawn itself from the puparium. But to a very large number, even after this is effected, the arduous task still remains of piercing the cocoons of leaves, of thick silk, of tough gum, or even of wood, in which the pupæ are incased. We can readily conceive how the strong jaws of coleopterous and hymenopterous species may be employed to release them from their confinement. But what instruments can be used for this purpose by moths in a state of great debility, whose mouth has nothing like jaws—merely a soft membranous proboscis? How shall the silkworm-moth (B. Mori) force its way through the close texture of a silken ball, through which the finger could not be easily pushed? Or the puss-moth (Cerura Vinula) pierce the walls of its house of glue and wood, which scarcely yield to the knife? You will not doubt that these difficulties have been foreseen by Infinite Wisdom, and provided against by Infinite Power. The egress of moths from their cocoons is secured in two ways;—either by some peculiarity in the first construction of the cocoon by the caterpillar, or by some process which the pupa or perfect insect is instructed to perform. As examples of each, several curious instances may be cited.
The larva of the moth which about 1760 made such havoc in the province of Angoumois in France, becomes a pupa in the interior of the grain of wheat which it has excavated; but the opening by which it first entered is not bigger than a pin's point, and is quite insufficient for the egress of the moth. How, then, is the latter to force its way through the tough skin which surrounds it? The larva, previously to assuming the pupa state, gnaws out a little circular piece at that end of the grain where the head of the future moth would lie, taking care not to detach it entirely. At this little door, which is sufficient to protect it from intruders, the moth has but to push, when it falls down, and leaves a free passage for its exit. A contrivance almost similar is adopted by a caterpillar which feeds in the interior of the heads of a species of teazel (Dipsacus L.), for a minute and interesting history of which we are indebted to Bonnet. This caterpillar previously to its metamorphosis actually cuts a circular opening in the head, sufficiently large for the egress of the future moth; but to secure this sally-port during its long sleep, it artfully closes it with fibres of the teazel, closely but not strongly glued together[658]. Another small caterpillar described by the same author, resides in the leaf of an ash curiously rolled up into a cone, and then assumes the pupa, which is inclosed in a silken cocoon, ingeniously suspended by two threads like a hammock in the middle of its habitation, and of so slight a texture that it presents no obstacle to the extrication of the moth. It is the closely-joined sides of its leafy dwelling that form a barrier, which, were it not for the precaution of the larva, would be impenetrable to so small and weak an animal. The little provident creature, before its change to a pupa, gnaws in the leaf a round opening, taking care not to cut through the exterior epidermis. This door is to serve the moth for its exit, like that formed by the wheat-caterpillar. But in proportion to its bulk its verdant apartment is of considerable size. How then shall the moth know the exact place where its outlet has been traced? How, without a clue, shall it discover in its dark abode the precise circle which requires only a push to throw it down? Even this is foreseen and provided against. Out of twenty positions in which its hammock might have been slung, the caterpillar has been directed so to place it, that the silken cord that suspends the head is fastened close to the side of the door which it has previously constructed; and the moth, guided by this filum ariadneum, at once makes its way out of an apartment which, but for this contrivance, might have been to it a labyrinth as inextricable as that of Minos[659].
The mode in which other caterpillars provide for their extrication, when become moths, from their silken cocoons, is not less ingenious. Those of Eriogaster lanestris (of which I have lately said so much,) and others, form oblong cocoons, which, viewed externally, you would at the first glance assert were of one solid piece: but on examining them more narrowly, you perceive one end of them to be a distinct lid, of a size large enough to permit the moth to issue out; and that it is kept in its place by a few slight threads, easily broken by pressure from within[660]. A few pages back[661] I mentioned a cocoon formed by the larva of Tortrix prasinana, of the shape of a boat reversed, composed of two inclined walls fastened together at the top and ends. In constructing this cocoon, it firmly glues to each other the top and one end, so as to form an impermeable suture; but the other end, at which the moth is to issue, though externally it seems as strong as the rest, is merely drawn close by a slender thread or two fastened on the inside, and easily broken from within. And, what is particularly singular in the construction of this ingenious habitation, the sides forming the end last mentioned, though originally requiring force to draw them into their required position, become so elastic as to close again when the moth has passed between them and made her escape; the cocoon preserving its usual shape, even when deprived of its inhabitant[662]. A similar cocoon is constructed by another leaf-rolling caterpillar, that of Tortrix chlorana[663]. Many similar proofs of contrivance in the construction of silken cocoons might be adduced, but I shall confine myself to one more only—I mean that furnished by the flask-shaped brown one of Saturnia Pavonia, and some other moths. If you examine one of these cocoons, which are common enough in some places on the pear-tree or the willow, you will perceive that it is generally of a solid tissue of layers of silk almost of the texture of parchment; but at the narrow end, or that which may be compared to the neck of the flask, that it is composed of a series of loosely-attached longitudinal threads, converging, like so many bristles, to a blunt point, in the middle of which is a circular opening[664]. It is through this opening that the moth escapes. The silk of its cocoon is of so strong a texture and so closely gummed, that had both ends been similarly closed, its egress would have been impracticable; it finds, however, no difficulty in forcing its way through the aperture of a sort of reversed funnel, formed of converging threads that readily yield to pressure from within. But an objection will here probably strike you. You will ask, Is not this facility of egress purchased at too dear a rate? Must not a chrysalis in an open cocoon be exposed to the attacks of those ichneumons of which you have said so much, and of numerous other enemies, which will find admittance through this vaunted door? Our caterpillar would seem to have foreseen your dilemma; at least, under heavenly guidance, she has guarded against the danger as effectually as if she had. If you cut open the cocoon longitudinally, you will see that within the exterior funnel-shaped end, at some distance she has framed a second funnel, composed of a similar circular series of stiff threads, which, proceeding from the sides of the cocoon, converge also to a point, and form a sort of cone exactly like the closed peristome of a moss; or, to use a more humble though not less apt illustration, like the wires of certain mousetraps[665]. In this dome not the slightest opening is left, and from its arched structure it is impenetrable to the most violent efforts of any marauders from without; whilst it yields to the slightest pressure from within, and allows the egress of the moth with the utmost facility. When she has passed through it, the elastic threads resume their former position, and the empty cocoon presents just the same appearance as one still inhabited. Rösel relates with amusing naïvété how this circumstance puzzled him the first time he witnessed it: he could scarcely help thinking that there was something supernatural in the appearance of one of these fine moths in a box in which he had put a cocoon of this kind, but in which he could not discover the slightest appearance of any insect having escaped from it, until he slit it longitudinally[666]. But from an observation of Meinecken, it appears that these converging threads serve a double purpose; being necessary to compress the abdomen of the moth as it emerges from the cocoon, which forces the fluids to enter the nervures of the wings, and give them their proper expansion. For he found, that when the pupa is taken out of the cocoon, the moth is disclosed at the proper time, but remains always crippled in its wings; which never expand properly, unless the abdomen be compressed with the finger and thumb, so as to imitate the natural operation[667].
I am next to give you some account of the second mode in which the release of the perfect insect from its cocoon is effected—that, namely, wherein its own exertions chiefly accomplish the work. I shall from a large number select only a few instances. The texture of the cocoon of the silkworm-moth is uniform in every part, and the layers of silk are equally thick at both ends. The moth makes its way out by cutting or breaking these threads at the end opposite to its head: an operation which, as it destroys the continuity of the silk, those who breed these insects are particularly careful to guard against, by exposing the cocoon to heat sufficient to destroy the included pupa. The question is—What instruments does the moth employ to effect this? And this we are not able to answer satisfactorily. Malpighi asserts that the animal first wets the silk with a liquid calculated to dissolve the gum that connects the threads, and then employs its lengthened head to push them aside and make an opening[668]. But, as Reaumur has observed, besides that so obtuse a part as the head of a moth is but ill fitted to act as a wedge, we find the threads not merely pushed to each side, but actually cut asunder. He therefore infers that the eyes, which are the only hard organs of the head, are the instruments by which the threads are divided—their numerous minute facets serving the purpose of a fine file[669]. It should be observed, however, that Mr. Swayne confirms Malpighi's assertion, that the silkworm does not cut, but merely pushes aside, the threads of its cocoon; and he informs us that he has proved the fact, by unwinding a pierced cocoon, the thread of which was entire[670]. Yet Reaumur's correctness cannot be suspected: and he affirms, that from observation there can scarcely be a doubt that most of the threads are broken[671]; which is further confirmed in an account of the breeding of silk-worms published in the American Philosophical Transactions: in which it is expressly stated, that cocoons out of which the fly has escaped, cannot be wound[672]. Analogy, it must be confessed, is against Reaumur's opinion; since other kinds of silkworms make their escape by means of a fluid. Thus we are informed by Dr. Roxburgh, that Attacus Paphia, when prepared to assume the imago, discharges from its mouth a large quantity of liquid, with which the upper end of the case is so perfectly softened, as to enable the moth to work its way out in a very short space of time,—an operation which, he says, is always performed in the night[673]. Perhaps the two opinions may be reconciled, by supposing the silkworm first to moisten and then break the threads of its cocoon. In those that are of a slighter texture, a mere push against the moistened end is probably sufficient: and hence we find in so many newly disclosed moths the hair in that part wet, and closely pressed down[674]. If it be apparently difficult for the silkworm-moth to effect an opening in its cocoon, how much harder must seem the task of the puss-moth (Cerura Vinula) to pierce the solid walls of its wood-thickened case? Here the eyes are clearly incompetent; nor could any ordinary fluid assist their operation, for the gum which unites the ligneous particles is indissoluble in aqueous menstrua. You begin to tremble for the fate of the moth incarcerated in such an impervious dungeon—but without cause: what an aqueous solvent cannot effect, an acid is competent to: and with a bag of such acid our moth is furnished. The contents of this she pours out as soon as she has forced her head through the skin of the chrysalis, and upon the opposite end of the cocoon. The acid instantly acts upon the gum, loosens the cohesion of the grains of wood, and a very gentle effort suffices to push down what was a minute ago so strong a barrier. How admirable and effectual a provision! But there is yet another marvel connected with it. Ask a chemist, of what materials a vessel ought to be to contain so potent an acid: he will reply,—of glass. Yet our moth has no glass recipient: her bottle is a membranous bag; but of so wonderful a fabric as not to be acted upon by a menstruum which a gum, apparently of a resinous nature, is unable to resist! This fact can only be explained by the analogous insensibility of the stomach to the gastric juice, which in some animals can dissolve bone,—and it is equally worthy of admiration. In both cases, the vitality of the membranous or fleshy receptacle secures it from the action of the included fluid; but how—who shall explain?
Ordinarily it is the moth that breaks the cocoon; but in the goat-moth and many Tortrices it is the pupa itself that performs the work, either wholly or partially. The pupa of the former is for this purpose furnished with sharp points upon the head, capable of effecting this object[675]. The locust-moth, another species of Cossus (C. Robiniæ Peck), whose history has been admirably detailed by Professor Peck, has a different process. "In the silk-moth," says he, "and all others which I have had opportunity to observe, the chrysalis bursts in the cocoon, and the fluid which surrounded the new insect in it escaping at the same time, so weakens or dissolves the fibre and texture of the silk, that the moth is able to extricate itself, leaving the chrysalis behind it; but this is not the manner in the locust-moth. After remaining till all its parts are fully grown and it is ready to quit its prison, a certain quantity of exercise is necessary, to break the ligaments which attach the moth to the shell of the chrysalis, and to loosen the folds of the abdomen. In taking this exercise, it can only move the abdomen in various directions: as one side of the rings is moved forward, the hooks in the serrated lines above mentioned (the adminicula) take hold of the silk, and prevent their sliding back; the next flexure brings forward the opposite side of the rings, which are prevented by the points on that side from slipping back in the same manner, and the chrysalis is forced out of the slightly woven extremity of the cocoon, and through the silk-lined cavity, till it is protruded for about one-third of its length out of the opening in the bark, and into the air[676]."
An exception to the general rule—that the rupturing of the cocoon is the business of the inclosed insect itself—is met with amongst ants; the workers of which not only feed the young, but actually make an aperture in their cocoons, cutting the threads with their mandibles with admirable dexterity and patience, one by one, at the time they are ready to emerge, the precise period for which these indefatigable nurses are well aware of, that they may meet with no obstacle. Without this aid, the young ant would be unable to force its way through the strong and dense coating of silk that infolds it[677]. And a proceeding somewhat akin to this was observed by the Hon. Captain Percy, R.N., who himself related it to me. Being fond of the study of insects, he was in the habit of attending to their motions; and in the beginning of September 1821 noticed those of a number of female Tipulæ, probably T. oleracea L., busily engaged in depositing their eggs amongst the roots of grass. While observing these proceedings, he at the same time saw one quitting its pupa-case, which had already by its own efforts got its head, thorax, and anterior legs out of it. It was then joined by two male flies; which, with their anal forceps and posterior legs taking hold of the pupa-case, appeared with their mouths and anterior legs to push the little prisoner upwards, moving her backwards and forwards; and as they kept raising her, shifting their hold of the skin till she was entirely extricated, when they left her to recover her strength by herself. Probably the extreme length of the two pair of hind-legs of these animals may render such assistance necessary for their extrication.
There remains yet to be explained under this head the manner in which the perfect insect is excluded from certain aquatic pupæ; such as those of Phryganeæ, gnats, and one of those Tipulidæ that resemble gnats. These pupæ (perhaps that they may be safe from the attack of birds) are destined to remain during the greater part of their existence in this state at the bottom of the water. But it is obvious that if the perfect insects were there to be disclosed, their wings would be wetted, and they would be drowned. It is the provision by which this result is obviated that now calls for your attention.
You have already been told that the larvæ of Phryganeæ inclose themselves in cases of different materials, open at each end[678]. You have also learned, that in becoming pupæ, they secure each end of their cases with a grating of silk[679]. When that change has occurred, they remain motionless at the bottom of the water. Now how are these pupæ, encased in tubes of a greater specific gravity than the surrounding fluid, to make their way to the surface when the time has arrived for their becoming denizens of the air? This they accomplish in the following manner:—The pupa is furnished with two strong exterior moveable mandibuliform processes, and has the power of moving its four anterior legs and antennæ while in the pupa-case. With these temporary jaws it makes an opening in one of the silken doors of its case, forces its way out at that end, and then by moving its legs, the cases of which in some species are ciliated for this very purpose, swims to the surface, where its skin splits, and discloses the included insect. That these jaws are given for the express and exclusive purpose of being thus applied, seems undeniable. The pupa eats nothing—they are therefore in every other point of view superfluous. They are given to it alone of all other similar pupæ, because unnecessary to all others; and they are cast off along with the rest of the puparium, the perfect insect having no vestige of jaws[680].
The gnat has to undergo its change on the surface of the water—How is it to accomplish this without being wetted? In the pupa state they usually remain suspended with the posterior end of the body turned downwards: but when the period for its change is arrived, it stretches it out upon the surface, above which its thorax is elevated. Scarcely has it been a moment in this position, than, swelling out the interior and anterior parts of the thorax, it causes it to split between the two respiratory horns. Through this opening the anterior part of the gnat then emerges. As soon as the head and trunk are disengaged, it proceeds with its labour, and gets out more and more; elevating itself so as to appear in the puparium like a mast in a boat. As it proceeds, the mast is more and more elevated and lengthened, till it becomes nearly perpendicular—just as the mast of a boat is gradually raised from a nearly horizontal to a vertical position: at this period a very small portion of the abdomen remains in the puparium. Neither its legs nor wings are of any use in maintaining it in this position. The latter are too soft, and, as it were, folded; and the former are stretched out along the abdomen—the segments of this last part are the only agents. The observer who sees how the little boat gradually sinks, and how its margin approaches the water, forgets the mischievous insect it contains, which at another time he would crush without remorse, and becomes interested for its fate; especially should wind agitate the water. A very little is sufficient to drive about rapidly the little voyager, since it catches the wind in some degree as a sail. If it should be upset, it would be all over with it;—and numbers do thus perish. The gnat, after having fixed itself thus perpendicularly, draws first its two anterior legs out of their case, and moves them forward, and next the two intermediate ones; then inclining itself towards the water, it rests its legs upon it, for water is to them a soil sufficiently firm and solid to support them, although surcharged with the weight of the insect's body. As soon as it is thus upon the water, it is in safety; its wings unfold themselves and are dried, and it flies away. All this is the work of an instant[681].
The pupæ of Chironomus plumosus proceed from those red worm-like larvæ so common throughout the summer in tubs of rain-water, &c., described by Reaumur[682]. They are not inclosed in cases, but are of a greater specific gravity than the water at the bottom of which they reside, until within a few hours of the exclusion of the fly. They have the power of swimming, however; and by moving the tail alternately backwards and forwards, can slowly raise themselves to the top of the water. But here occurs a difficulty. For the extrication of the imago it is necessary that they should remain quietly suspended at the surface; and moreover that the thorax, in which the opening for its exit is to be made, should be at least level with it: and this is precisely what takes place. If you watch one of these pupæ when it ascends from the bottom, you will see that as soon as it has reached the top it remains suspended there motionless; and that its thorax is the highest part of the body, and level with the surface. Now the question is, in what way this is accomplished? How can a pupa of greater specific gravity than water, remain suspended without motion at its surface? and how can its thorax, which is at its heaviest end, be kept uppermost?—By a most singular and beautiful contrivance, which I shall explain; the more particularly because it has escaped Reaumur, and, as far as I know, all other entomological observers. The middle of the back of the thorax has the property of repelling water—apparently from being covered with some oily secretion. Hence, as soon as the pupa has once forced this part of its body above the surface, the water is seen to retreat from it on all sides, leaving an oval space in the disk, which is quite dry. Now though the specific gravity of the pupa is greater than that of water, it is but so very slightly greater, that the mere attraction of the air to the dry part of the thorax, when once exposed to it, is sufficient to retain it at the surface; just as a small dry needle swims under similar circumstances. That this is a true solution of the phænomenon, I am convinced by the result of several experiments. If, when the pupa is suspended at the surface, a drop of water be let fall upon the dry portion of the thorax, it instantly sinks to the bottom,—the thorax, which belongs to the heaviest half, being the lowest; and if the pupa be again brought to the surface, so that the fluid is repelled from its disk, it remains suspended there without effort, as before. Just previously to the exclusion of the fly, the dry part of the thorax is seen to split in the middle. The air enters, and forms a brilliant stratum resembling quicksilver, between the body of the insect and its puparium; and the former pushing forth its head and forelegs, like the gnat, rests the latter upon the water, and in a few seconds extricates itself wholly from its envelope.
Before I close this letter, I must state a fact connected with the subject of it that deserves to be recorded. It is a general rule, that one pupa-case incloses only one insect; but Kleesius, a German entomologist, asserts that he had once two specimens of Gastropacha quercifolia produced from one pupa; which was large, being full two inches long, and one thick.